ill

CONTENTS

No. 1. Published January 1980.

A revision of the systematics of Australian sipun-

culans. (Sipuncula). (S. J. Edmonds) ......... 1

Nos. 2-6. Published May 1980,

No, 2. The distribution in Australia of the grass bugs

of the tribe Stenodemini (Heteroptera-Miridae-

Mirinae). (Jose C. M. Carvalho and Gordon F.

a ay a MOR Egy ae PO Ne Ares MOP er 75

No. 3. Two new genera of the subfamily Pentato-

minae (Heteroptera: Pentatomidae) from the

Australian region. (Imtiaz Ahmad and Naseer

Ahmad Khan). . eis | Se Ss

No. 4. Some reanaries 0 on » the breeding distribution

and taxonomy of the prions (Procellariidae:

Pachypoiek 10. Bo Geox. on cca ede sat 9]

No. 5. Evolutionary systematics of Yemylla: XI.

Species from the Australian region (Insecta:

Collembola). (Maria Manuela da Gama).. 123

No. 6. Macropodid skeletons, including Simasthe-

nurus Tedford, from an unusual “drowned cave”

deposit in the South East of South Australia.

(Neville :S. Pledge}... .c.ce cee ae ess GH

Nos. 7-9, Published January 1981.

No. 7. The genera Laccobius and Nothydrus

(Coleoptera, Hydrophilidae) in Australia and New

Zealand, (Elio Gentili),........+..,.+-.. 149

No. 8 The Fabrician types of the Australian and

New Zealand Coleoptera in the Banks collection

at the British Museum (Natural History).

{Winifred P, K. Radford) ............... 155

No. 9. Sarcoptiformes (Acari) of South Australian

soils. 1. Notation; 2. Bifemorata and Ptyctima

(Cryptostigmata). (David C. Lee)

Nos. 10-13. Published September 1981,

No. 10. Evolutionary systematics of Xenylla. XII.

Redescription of X. occidentalis Womersley and

a comparison of the chaetotaxy of . arenosa

Uchida and Tamura and ¥. fittoralis Womersley

(Insecta: Collembola). (Maria Manuela da Gama)

Ne eee OR Ae ee i a at Se peg) ct fs 223

No. Il. A new species of ce i genus Cafacanthus

Spinola (Heteroptera: Pentatomidac: Pentato-

minae) from the New Hebrides with morpho-

logical notes on two other Australasian species and

their relationships. (Imtiaz Ahmad and Syed

Bearivalchany he os cG a nd eed ¢ ples te ce kee 227

No, 12. The fossil pelicans of Australasia. (Pat

Vickers Rich and G. F, Van Tets)........ 235

No, 13. On some oviparous filarial nematodes

mainly from Australian birds. (Odile Bain and

Patricia M. Mawson). ......-.-..000ee0 265

Nos. 14-16, Published June 1982.

No, 14. Observations on Chyzeria Canestrini and

some related genera (Acarina: Trombidioidea) with

remarks on the classification of the superfamily

and description of a pygmephorid mite phoretic

on Chyzeria. (R. V. Southcott) .......... 285

No. 15. Sarcoptiformes (Acari) of South Australian

soils. 3. Arthronotina (Cryptostigmata). (David C.

Jie? Tig eye ays mega, a ey 1 ee a 327

No, 16. Polyclad turbellarians from the southern

coasts of Australia. (Stephen Prudhoe)... 361

Nos. 17-18. Published April 1983.

No. 17. Helminth type specimens in the South

Australian Museum. 1. Nematoda. (Lesley R.

TU TREE po Mg en ee ed et eRe Mk end 385

No, 18. Reproductive biology of the frogs of the

Magela Creek system, Northern Territory.

(Michael J. Tyler, Graeme A. Crook and Margaret

RSH OR Up en antlers ets Baal oh cetyten MA SVG PEN :

Nos. 19-23. Published May 1984.

No, 19. The taxonomy of the Atrax adelaidensis

species-group (Macrothelinae: Mygalomorphae)

with notes on burrowing behaviour. (M. R. Gray)

Be Pe ey ih ta ae, He ae eg gt 44]

No. 20. The Psocoptera (Insecta) of South Australia.

(€> DE Siithersh ae ee rte ee

No, 21. Helminth type-specimens in the South

Australian Museum. II. Acanthocephala and

Cestoda. (Lesley R. Smales)............. 492

No, 22. Helminth type-specimens in the South

Australian Museum. Ill. Trematoda. (Lesley R.

Smales) 503

No, 23. A revision of the genus Aplerotus Dallas

(Heteroptera: Pentatomidae: Pentatominae) with

description of a new species from South Australia,

(Imtiaz Ahmad, Naseer Ahmad Khan and Sved

Kaimaltiddin) 5... 0 ces 06.05snueaeneae 513

CONTRIBUTORS

AHMAD, IMTIAZ and KAMALUDDIN, SYED

A new species of the genus Cafacanthus Spinola

(Heteroptera: Pentatomidae: Pentatominae) from the

New Hebrides with morphological notes on two

other Australasian species and their relationships

“TEE A A a eg aes Cott Sener Coie 227

AHMAD IMTIAZ and KHAN, NASEER AHMAD

Two new genera of the subfamily Pentatominae

(Heteroptera: Pentatomidae) from the Australian

PUNT ceases Pee aes cee en 83

AHMAD, IMTIAZ; KHAN, NASSER AHMAD and

KAMALUDDIN, SYED

A revision of the genus Aplerorus Dallas (Hetero-

ptera: Pentatomidae: Pentatominae) with description

of a new species from South Australia...... 313

BAIN, ODILE and MAWSON, PATRICIA M.

On some oviparous filarial nematodes mainly from

Puistraam GINS seat Peed ew cess 265

CARVALHO, JOSE C. M. and GROSS, GORDON F.

The distribution in Australia of the grass bugs of

the tribe Stenodemini (Heteroptera-Miridae-

UTI AG WaT Neh de Weg bt 8 4 ales of sescdln oseg tee 74

COX, J. B.

Some remarks on the breeding distribution and

taxonomy of the prions (Procellariidae: Pachyprila)

CROOK, GRAEME A.

Bee rMichsel THe cic ek ce gene a bees

DA GAMA, MARIA MANUELA

Evolutionary systematics of Nenvila: XI. Species

from the Australian region (Insecta: Collembola)

Bee SP gh ct rt eke jc ec is pee 123

Evolutionary systematics of Xenylla. II. Redeserip-

tion of X. occidentalis Womersley and a comparison

of the chaetotaxy of X. arenosa Uchida and Tamura

and X. fittoralis Womersley (Insecta; Collembola)

If te a or etal oi Se aaa ear ea 223

DAVIES, MARGARET

Seeiichael Jo Tyler. .6 cs. vsea~ eee Soe

EDMONDS, S. J.

A revision of the systematics of Australian sipuncu-

fans TSP MTCUIST . oo. ands ee ee ee dee Fe wey l

GENTILI, ELIO

The genera Laccobius and Nothydrus (Coleoptera,

Hydrophilidae) in Australia and New Zealand 143

GRAY, M. R.

The taxonomy of the Atray adelaidensis species-

group (Macrothelinae: Mygalomorphae) with notes

on burrowing behaviour ................-. 44]

415

GROSS, GORDON F.

See Jose C. M. Carvalho ... 022 04 ve cues 75

KAMALUDDIN, SYED

See Imtiaz ARMA... os ee esas eres Bly OS

KHAN, NASEER AHMAD

See Imtiaz Almad... 1.00.50 caeeesaes 83, 313

LEE, DAVID C.

Sarcoptiformes (Acari) of South Australian soils. 1.

Notation; 2. Bifemorata and Ptyctima (Cryptostig-

STUN Atala se Pegg Se MS Sate sence 199

Sarcoptiformes (Acari) of South Australian soils. 3.

Arthronotina (Cryptostigmata) ............ 327

MAWSON, PATRICIA M.

See Odile Bain . . 265

PLEDGE, NEV ILLE 5.

Macropodid skeletons, including Simosrhenurus,

from an unusual “drowned cave” deposit in the

South East of South Australia............. 131

PRUDHOE, STEPHEN

Polyclad turbellarians from the southern coasts of

GET AU AM aie cipotter ats hits A, sepsis nest. oan te Dalene 361

RADFORD, WINIFRED P. K.

The Fabrician types of the Australian and New

Zealand Coleoptera in the Banks collection at the

British Museum (Natural History) ......... 155

RICH, PAT VICKERS and G. F. VAN TETS

The fossil pelicans of Australasia .......... 235

SMALES, LESLEY R.

Helminth type specimens in the South Australian

Museum. 1. Nematoda ...........-00.0005 385

Helminth type-specimens in the South Australian

Museum. II. Acanthocephala and Cestoda.. 492

Helminth type-specimens in the South Australian

Museum. III. Trematoda 503

SMITHERS, C. N.

The Psocoptera (Insecta) of South Australia 453

SOUTHCOTT, R. Y.

Observations on Chyzeria Canestrini and some

related genera (Acarina: Trombidioidea) with

remarks on the classification of the superfamily and

description of a pygmephorid mite phoretic on

Chyzeria.. A Mas Hee Be

TYLER, MICHAEL I, ‘GRAEME A. CROOK and

MARGARET DAVIES

Reproductive biology of the frogs of the Magela

Creek system, Northern Territory .......... 415

VAN TETS, G. F.

See PAL VICBETS. PUEB lr» siere t «8 SoA & oe ete 235

ACAI CSC esc cote cama v Sleha aiea ww ncele sn eres

PEA Wetec ing cet cataaete laud. mira. c'esi, wii tal 199,

POT aL Gee ee ee en 2 rsa a

adelaidensis, Atrax (species-group) ...........

PRReIN Ere ETN les ee co cee aye yp Spare ay tee hes

MST ETi WT ahs ae gee ee etn ae

PURI DEWCUEAIN No tre tcode-drm Seid sce eee a ws

Atrax adelaidensis ......505.-0c. 00004 eeeeeas

Bre Te AU eye cere g Aniedsa SG xe le eM ic eS

Birds (nematodes from) .........0..0000000%

UNECE Torres eo eats Sarr pra ee oar Cee Ce

Burrowing behaviour (of spiders) ............

RUC NTICC ate 2... vote cis ele oe teks. pce Sune

RCHISMEOA: «ss oa cn daied ev aass Re eee ee 123,

Cy PMOSMEMISIA. os. ke ace ew eels eee 199,

Fabrician types (of Coleoptera) ..............

Frogs (reproductive biology)..............+..

MEATS GO TUES tg Pe ets ew cn wae w Ve age pene de

Helminths .......0..080000005 265, 385, 493,

RSDETOPIEDA G14, ra s.s 9 2 cies ee Poe 75, 83, 227,

Biireniine sess... ciehae eae) scacee, rea or

Insecta... 75, 83, 123, 143, 135, 223, 227, 453,

Macropodidde. .. 0. scares abcececnnerereeeee

PRU ERE RESINS Se cctac, cloner es leaneica Walon mde 48

Vil

INDEX

493 Magela Creek System (frogs of) ............. 415

Bap, MUMMED IEE ale kG ss in. or an, 9 oo ere pamper eel ba 75

aE TNE rena ta Sart: vss “ean “eto ns a cen nar mcned TM neem eR eee 75

ed OTe ce Mein Socal Arac 9a four Glee Woe oe eee 285

223 Mypalomorphiae 2.2... sks eee eee 44]

- PUCREIAL OMI 28 crbat ons ana pecte iaa treed she Letom 385

44] Nematodes (of birds)... 0... .c6ecsunieee wes 265

Put Ceres re ee ee, SN nna < lain 4 cer Pre ie 143

199 occidentalis, Xenylla 223

265 f Peete os RES ea roe

BTS! Sei Tel nye] Sk Bate toot, ee ee -a oaSA , |

441 “Pelicans: (f0ssil) 5. cs. oe ae soe ee ee ee 235

285 Pentatomidae ......2s00ce cc eaeeceee 83, 227, 513

Penigtominge 6. G8 otk cae cise Wap mel, eS

227 :

Ps CEA oo Gis ids et hanes = eer ahaha ane et 361

493 :

293 PLIOMS.-. +--+ eee eee e eee e ees |

285 Procellartidae .......0ss005 settee eee eees |

155 Psocoptera (of South Australia) ............. 453

533 ELE AS it Tie RO a OR ror PO oe le eee 199

327 Reproductive biology (of frogs).............. 415

155. Sareopiformes soy ees eas eae ede ces 199, 327

405. SimOgeHenrys’, co es csc eg ee aes + ee ee a 131

75 SULT CUDA se 2 en Pian y Hae he cat oars en and abies 1

Sipunculans (see Sipuncula)

503° Spiders (burrowing behaviour of)............ 441

AAD S, PSTN EMMULIN yin ass aid ori vipat aw aad Byes Sar 75

= dha t: biw'r b: hae ee ee ae er rere) as 403

S13. Tromibigigndlea, 25 50 fica we bee cs ne men ale eee ee

143 sTUie ee Sav TeAMNE! whe. drapes a, edge ees dom y otis ws kensel ton ae 361

293 Type-specimens (helminth) ......... 385, 493, 503

131 ROTIWILE LS. © re Teer caste Soe ee ee ae 123, 223

RECORDS oF THE

SOUTH AUSTRALIAN

MUSEUM

VOLUME 18 NUMBER 1 January, 1980

A REVISION OF THE

SYSTEMATICS OF AUSTRALIAN

SIPUNCULANS (SIPUNCULA)

By S. J. EDMONDS

N MUSEUM

Adelaide

South Australia 5000

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS

(SIPUNCULA)

BY S. J. EDMONDS

Summary

The author lists 57 species and subspecies of sipunculans reported from Australia and has examined

Australian specimens of 43 of them. Three new species are described — Phasolion cronullae from

the shell of Gazameda gunnii (Reeves), Themiste variospinosa and Paraspidosiphon johnstoni. The

other species examined are redescribed. The Australian sipunculans belong to 4 families and 12

genera. The genus Themiste is considered to comprise 3 new subgenera: Themiste s.s., Lagenopsis

and Stephensonum. The genus Centrosiphon Shipley, 1903, because the types is thought to be a

Golfingia, is considered to be invalid. Specimens identified by Edmonds (1956) as Aspidosiphon

klunzingeri Selenka & de Man are now considered to be a new species, Paraspidosiphon johnstoni.

Specimens identified by Edmonds (1956) as Aspidosiphon steenstrupii Diesing are now thought to

be Paraspidosiphon formosanus (Sato, 1939), Phascolosoma heronis Edmonds, 1956 is now

considered to be a junior synonym of Phascolosoma stephensoni (Stephen, 1942) and Phascolosoma

dunwichi Edmonds, 1956 a junior synonym of Phascolosoma scolops (Selenka & de Man, 1883).

Aspidosiphon elegans elegans (Chamisso & Eysenhardt, 1821) is considered to include

Aspidosiphon exilis Sluiter, 1902. Suggestions about narcotising and dissecting specimens are

given and the ecology and habits of the group are discussed. Maps showing the distribution of

sipunculans in Australia are also included,

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (STPUNCULA)

8S. J. EDMONDS

Honorary Associate, South Australian Museum,

ABSTRACT

A revision of the systematics of

Aust, Mus, 18(1):

EDMONDS, §. J. 197%.

ee Sipunculuns (Siputcula). Ree. S-

The author jists 357 species and subspecies of

sipuneulans reported from Australia and has

examined Australian specimens of 43 of them. Three

new species are deseribed—Phascolion cronullae

fram the shell of Gazameda gunnii (Reeves),

Themiste variospinasa and Paraspidosiphon john-

ston. The other species examined are redescribed.

The Australian sipunculans belong to 4 families and

12 venera. The genus Themiste is considered to

comprise 3 new subgenera, Themiste $.5., Lagenopsis

and Stephensonum. The genus Centrosiphon Shipley,

1903, because the type is thought to be a Golfingia,

is Considered to be invalid. Specimens identified by

Edmonds (1956) as Axspidosiphon klunzingeri

Selenka & de Man are now considered to be a new

species, Paraspidesiphon johnstoni. Specimens iden-

tified hy Fdmonds (1954) as Aspidasiphan steen-

strupii Diesing are now thought to be Peraspidosiph-

on formosanus (Sato, 1939), Phascalasoma heronts

Edmonds. 1956 is now considered to be a junior

synonym Of Phascolosoma stephensoni (Stephen,

1942) and Phascolosama dunwichi Edmonds, 1956 a

junior synonym of Phascolosoma scolops (Selenka &

de Man, 1883), Aspidosiphon elegans elegans

(Chamisso & Fysenhardt, 1821) is considered to

include Aspidosiphon exilis Sluiter, 1902. Sugges-

tlons about parcotising and dissecting specimens are

given and the ecology and habits of the group are

discussed, Maps showing, the distribution of

sipunculans in Australia are also included,

CONTENTS

!, INTRODUCTION Page

Aim of the study... 06... 00y 0. 1d 1

Characters of the phylum Sipuncula Las 2

Sipunculans, echiurans, ete. ,..... 0.5. 2

Where sipunculans are found |... ..- 2

The ecological rale of sipunculans 2

Narcotisation and dissection of sipuncu-

FetRS sHatelata se! the atalely eat elele a vuslniele 4

Previous systematic studies of Austra-

lian sipunculans ... 2. ..;.0y eyes 5

Australian collections of sipunculans . - 7

Acknowledgments ..........0.00.04- 7

Januar, [9s

Adelaide, South Australia, S000

1. SYSTEMATICS

Phylum Sipuncula and list of species

reported from Australia .... _.. 7

Key to families of Sipuncula ....,.... 8

Family Sipunculidae and key to genera 8

Sipunculus and key to species .._... 8

Xenosiphon and key to species... , 2

Siphonosoma and key to species .. .. 14

Family Golfingiidae and key ta genera 18

Golfingia and key taspecies ._,_... 18

Phascolion and key ta species ...... 29

Onchnesoma .....00.-..-, wpe 32

Themiste and key ta species ..,...., 32

Family Aspidosiphonidae and key to

PENESH i ke tom epee rete 43

Aspidosiphon and key to species - 43

Paraspidosiphon and key to species. . 49

Cloeosiphon ..,..,.. ce atta bls, 54

Lithacrosiphon .......: 54

Family Phascolosomatidae and ey to

BPENGra’s; yb seth b es eb eeh tes a]

Phascolosonia and ‘identification of

SPCCIOS pg ores ets psdeeete 55

Ill. REFERENCES ............0...0-. 71

I. INTRODUCTION

Aim of the study

It is twenty years since the last general work on

the systematics of the Australian sipunculans was

published (Edmonds 1955, 1956). In the meantime

more specimens have been collected and changes

have been made in the systematics of the phylum.

Some of the more recently collected specimens are

new species and others are species not previously

reported from Australia. Further, the additional

specimens enable information about some of the

previously described species to be extended and

imperfections in their descriptions, and even errors

in their identification, to be corrected. The aim of

the study is to bring up to date, as far as it is possible.

the systematics of the Australian sipunculans. Seeing

that many parts of the coast, especially the northern

ones, have still not been visited by collectors it seems

almost certain that more species will be found in the

future.

In identifying the animals I have relied mostly on

the description of their external and internal

anatomy, Little or no attempt has been made to use

some of the more modern methods that are

sometimes employed to separate closely related

species and to determine new ones, for instance the

2 REC. §& AUST, MUS, 18 (1): 174

use of electrophoretic techniques (Manwell 1977 p.

331) or the study of chromosones. The difficulties of

using some of these methods when most of one’s

material is already fixed are great. As far as 15

possible | have worked with the concept in mind that

a species consists of a population of animals and that

it is not just a single animal or type specimen,

Most af the specimens have been collected

intertidally or from the upper sublittoral zone and

only a few have been dredged. J have included

specimens from Lord Howe [sland but not from

Norfolk Island nor the eastern regions of the

Tasman Sea, although it is possible that the range of

the species may extend to the hame waters of

Australia. | have included the reeards of Australian

species not seen by me but which have been

identified by other workers. ! have made little or no

attempt to examine these records eritically but |

have included the species in the keys that are given

in the paper,

Characters of the phylum Sipunculat

Sipunculans are a group of unsegmented, caelo-

mate, bilaterally symmetrical, marine invertebrates.

Their body consists of two chief parts, a cylindrical,

globular, flasks or sac-like, highly muscular trunk

and a highly extensible and comparatively slender

introvert, that is capable of complete cetraction

within the trunk. The mouth opens at the anterior

extremity of the introvert and is usually wholly or

partly surrounded by « group of tentacles or a

tentacular fold. The introvert may be armed with

hooks ar spines (Sometimes both); it may, however,

be unarmed. Small glandular openings and hemis-

pherical, conical or rather flat papillae are usually

present on the trunk and introvert. Setae are absent.

The alimentary canal is Jong and usually wound into

a spiral for most of its length, The anus is. usually

situated dorsally on the anterior surface of the trunk,

In most genera two nephridia open to the exteriar on

each side of the nerve cord near the anus but only

one nephridiuin is found in the genera Onchnesoma

and Phascolion. A contractile yessel is attached to

the anterior region of the oesophagus. Antertarly it

branches into the tentacles but posteriorly it ends

blindly, Tt is usually single but sometimes double; it

may be simple or give off few to many tubules which

may be long or short. The blood cells contain the

Tespitatary pigment haemerythrin. The ventral

nerve cord is unsegmented and the brain lies dorsally

near the anterior extremity of the introvert. The

gonads develop at the base of the retractors and the

sexes, although Separate, are indistinguishable

externally, Fertilisation is external, cleavage of the

zygote is spiral and the larva is a trochophore. No

Pootnotet, Based on the characters given in Stephen & Edmonds

(1972); pp. 18-19

Taniary. LOST

segmentation appears during development. One

species is known to be a protandrous hermaphrodite

and two species are known to reproduce asexually,

Sipunculans, echiurans, holethurians ete,

Holothurians, sand-burrowing anemones, nemer-

tines and especially echiurans are often mistaken lor

sipunculans. ft is not difficult, however, to

distinguish between them. The introvert of a

sipunculan can be completely retracted into the body

cavily but the proboscis of an echiuran cannot. The

mouth of a sipunculan is placed at the anterior tip of

the introvert but in an echiuran it lies on the trunk at

the hase of the proboscis, The aius of asipunculan is

usually situated anteriorly on the dorsal surface of

the trunk but in an echiuran it is at the posterior

extremity of the trunk. Tentacles are usually

associated with the mouth of a sipunculan but are

lacking in an echiuran. Setae are absent in

sipunculans but usually a pair of them protrudes

from the surface of an echiuran just posterior to the

mouth.

Although they possess tentacles, anemones lack

an introvert, a tubular alimentary canal and

nephridia. Nemertines possess a posteriorly placed

anus, they Jack a body cavity and are usually without

tentacles, Holothurians lack an introvert and the

anus is posterior.

Where sipunculans are found

Sipunculans live in tropical, ternperate and polar

seas and their bathymetric range is wide. Many have

been found intertidally and others have been

dredged at depths of 6 860 m (Murina, 1964a p. 250,

Stephen & Edmonds, 1972 p. 4 and 79; Cutler, 1977

p. 155). Fisher (1952 p, 371) thinks that, being soft

bodied and defenceless creatures, they will prabably

live in any protected place that provides access to

reasonably clear water and food.

In Australia most sipunculans have been collected

intertidally or from shallow waters, They have been

found in limestone reefs, in reef-forming and solitary

corals, in mangrove flats, under stones, in fissures of

non-caleareous rocks, in clumps of mussels, in

masses of serpulid worms, in cracks in wooden jetty

piles, in the holdfasts of algae, amongst the routs of

niariné angiosperms, in firm sand, in fine mud, in

the empty shells of 4 number of molluses and in the

tubes and tests of larger foraminiferans.

Records of some species dredged off the

Australian coast are given in Murina (1972) and

Cutler (1977).

The ecological importance of sipunculans

Sipunculans perform a number of roles in the

environment. Most are detritus feeders,

some assist in the breaking down of limestone and

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPLINCULA) »

coral reefs, some Serve as food for other animals,

same live commensally with solitary catrals,

anemones and molluscs. and others act as either the

intermediate or final hosts of parasites.

The gut of sipunculans usually contains sand,

mud. small particles of coral or limestone rock,

Pieces of algae or marine angiosperms, fragments of

mollusean shells and echinoderm exoskeleton, the

frustules of diatoms and the skeletal parts of

foraminiferans, The information suggests that

sipynculans are detritus feeders and that they extract

any food contained in whatever they ingest. Chin

and Wu (1950) claimed that diatoms were an

important constituent of the food of a number of

sipunculans collected al Amoay. China. In her

experiments on the development of sipunculans Rice

(1970; p. 142) fed the larvae on algal cultures of

Phaeodactylum tricoraurum and Isechrysis galbana

and the larvae of Phascelosama agassizii on “mixed

diatoms and dinoflagellates collected from tidal

pools” (Rice 1973 p. 3).

Very little experimental work, however, has been

done on the feeding habits of the group. Peebles and

Fox (1933) observed (1) that Themiste (= Dendros-

tamu) zosleticola burrows in perfeetly clean sand

which contains no food but that in these conditions

the anima! does. not ingest the sand, (2) that the

tentacles are sensitive to small particles of food and

to traces of chemicals and (3) that the anterior

region of the introvert, the tentacles and the collar

are the most senstitive to contact with foreign

substances. It is possible, therefore, that sipunculans

do not indiscriminately swallow their environment

buc that they are able to exercise some selection as ta

what they eat, Gardiner (1903; p. 333) claims that

some sipunculans break down coral fragments into

smaller particles. They do noi, however. possess 4

erop and gizzard like earthworms or a gastric mill

like some crustaceans,

Rice (1976 p, 126) reports that there are at least

two patterns of feeding behaviour in rock-dwelling

sipunculans. Species with long, extensible introverts

and shor digiuform tentacles feed by extending

their introvert fram the mouth of the burrow and

grazing the surtace of the rock near them. She says,

"Phascolasoma perlucens,. maintained in the laborat-

ary in intact burrows has been observed to feed trom

the surface of the rock on sediment and detritus.

Some particles adhere direetly to the tentacles while

athers seem to be seraped off the rack by the small

hooks of the introvert.” On the other hand rack-

dwelling species with long filiform tentacles and

relatively short Introverts such as Phascolosama

antillarum and Themiste (agentformis make use of a

ciliary-mucus. mechanism, “The tentacular crown 1s

extended above the mouth of the burrow and

particles adhering to the sticky tentacles are directed

by ciliary currents into the digestive tract.”

Although sipunculans may mbhabit the tubes and

cavities made by other animals, it is known that

some species, especially those belonging to the

genera Phascolosoma, Aspidosiphon, Paras-

pidasiphon, Cloeosiphon, Lithacrosiphon and

Themuste are able to construct \heir own homes in

calcareous rock. Such species play an important part

in the breaking down of coral and limestone reefs,

The importance of sipunculans as members of a

coral reef community is discussed in Rice (1976: p.

122), How soft bodied creatures are able to bore into

hard rock is a baffling problem, The fesearches of

Rice (1969), Rice and Metntyre (1972) and Wilhams

and) Margolis (1974) suggest that mechanical

abrasion and chemical action are involved.

Sipunculans are 4 source of food for 4 number of

different animals. Kohn (1975 pp. 313-331) has

reyiewed the topic in an article an predatian on

sipunculans, Pallas (1774) reported that

Siphonosoma edule was used as food by the

Batavians and Sato (1935) that Sipunculus indicus

Was eaten by the natives of Palau Is. Chin (1947)

reported that Sipunculus nudusy was eaten at Amoy,

China and went as far as to supply a recipe for its

preparatian as a dish, Many fish eat sipunculans,

Kohn (1975) lists two orders of elasmobranchs and

six orders (17 families) of teleosts. He says that “the

fishes are all generalized predators on benthic

invertebrates but sipunculans, mainly Aspidosiphon,

are the major prey of the stingray Dasyatix

americanus and the margate Haeumulon album in

the Caribbean. In the trunktish Lactophyrs triqueler

and the economically important haddock, Melanog-

rammus aeglefinnus, in the Barents Sea,

Aspidasiphon spinoscutalus and Golfingia tar-

garifacea are only second to polychaetes in the diets

in nature. In other fishes sipunculans represent

subsidiary or incidental components of the diet." In

South Austraha small specimens of Siprunculus

robusms are sometimes found in the stomach of the

local King George (spotted) whiting Sillaginodes

punctaris, Some sipunculans collected jn such a way

by W. Zeidler (3/1/76) are in fhe callection of the

South Australian Museum (reg. no. E 1072).

Kohn says that only a few invertebrates are known

to prey on sipunculans in nature. Kohn (1970, 1975)

reports that the gastropod Mitra literata preys on

sipunculans that burrow in the intertidal limestone

reef at Oahu, Hawail. Sixteen of the 24 specimens of

Mitra that he examined were each found ta contain

the remains of one sipunculan. The species eaten

were Pheascolosoma scolops, P. stephensani (=P

heronis) and Aspidosiphon elegans, [is believed thar

the animals were eaten mostly during the night.

The sipunculan Aspidosiphon jukesii Baird lives

commensally in the base of the solitary corals,

Heteropsammia and Heterocyathus; the association is

well documented and described (Bouvier 1895;

Sluiter 1902; Stephen and Robertson 1952; Goreau

and Yonge 1968; Yonge 1975; Rice 1976), Some of

Rice’s observations are referred to in the present

paper on p, 49, Specimens of Golfingia hespera

(Chamberlin) were found as commensals in the

tubes of Cerianthus (Fisher 1952) and syllids are

sometimes found in association with specimens of

Phascolin (see p, 30), A list of same molluscs

associated with sipunculans is given in Stephen &

Edmonds, 1972 p, 342.

Sipunculans may contain parasites in their gut,

body fluids or tissues. The most commonly found

parasitic protozoans are sporozoans, especially

gregarines. Some of the records are listed in Stephen

& Edmonds (1972 p. 341) and further information is

given in Jones (1975b p. 349). Rhabdococles have

been reported from the gut of at least four different

sipunculans and encysted metacercaria of uniden-

tified trematodes from the intestine, brain, tenta-

cles, gonads and contractile vessel of several species

(Stephen & Edmonds 1972 p. 341). Nematodes have

been reported from the body cavity of at least three

species (Augener, 1903 p. 361; Edmonds, 1976 p.

222; Jones, 1975a p. 343).

Narcolisation and dissection of sipunculans

Sipunculans are usually much more easily

indentified if they have been relaxed before they are

preserved. The nature and arrangement of the

tentacles and the presence of hooks and spines are

readily observed if the specimen has been

narcotised. One way of doing this is to place them in

a dish containing sea water to which is carefully

added (drop by drop) some 80% alcohol in which

some menthol has been previously dissalved. It may

take from a half to six hours before the animals do

not respond to touch, The specimens should then be

left overnight in 5% neutral formalin and stored in

70% alcohal.

REC. S. AUST, MUS, 18 (1)-

1-74 January, L980

Not many sipunculans can be identified from their

external appearance; most require to be dissected,

One method is [o pin them out under water in a dish

containing a layer of solidified paraftin wax about

20 mm thick. Place the dorsal (anal) side uppermost.

With the aid of a sharp scalpel or fine scissors and

forceps cut the body wall longitudinally along a line

just to one side of the anus. Lift up the body wall

ahead of the cut so that the jncjsion does as little

damage as possible to the structures underneath.

Cut the whole length of the trunk and pin back the

flaps. Wash away coagulated blood with the aid of a

stream of water from a wash bottle,

In order to make a mount of the introvert hooks

or body papillae, snip off a small section of the skin

and place it on a slide with a drop of glycerine. Tease

up the tissue containing the hooks with the aid of

two fine needles. When the preparation is now

examined under low power single hooks or groups of

hooks can often be seen. Remove unwanted tissue

and cover with a caver slip, Only hooks that lie flat

should be drawn. There is no need to tease the piece

of lissue containing the papillae,

Figures 1-3 show some structures of a dissected

sipunculan.

A glossary of terms used in sipunculan taxonomy

is given in Stephen and Edmonds (1972 pp. 8-11).

Previous systematic studies of Australian sipunculans

Records of sipunculans from Australia are

contained in the following: Stimpson (1855),

Keferstein (1865), Baird (1868), Selenka & de Man

(1883), Augener (1903), Kesteven (1903), Fischer

(1914, 1919a, 1921, 1927), Monro (1931), Wheeler

(1938), Edmonds (1955, 1956), Murina (1964a,

1972), Cutler (1977) and Gibbs (1978),

More general information about the systematics of

the phylum is given in Hyman (1959), Tétry (1959),

Fisher (1952), Stephen & Edmonds (1972), Cutler

(1973) and Cutler & Murina (1977),

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA)

FIGS, 1-3, Diagrams showing some of the external and internal characters of sipunculans. The specimen in Fig. | has been dissected

from the dorsal side, a, anus; as, anal shield; c, caecum; cm, circular musc!e; cs, caudal shield; ev, contractile vessel: dr, dorsal

retractor; f, fastening muscles; g, gonad; h, hooks; i, introvert; in, intestine; Im, longitudinal muscle: m, mesentery; md, mid-

dorsal line; mo, mouth; n, nephridia; ne, nerve cord; np, nephridiopore; o, oesophagus; p, papillae; r, rectum: sm, spindle muscle;

t, tentacles; tf, tentacular fold; tp, triangular papillae; tr, trunk; vr, ventral retractor; wm, wing muscle.

6 REC. S. AUST. MUS. 18 (1); 1-74 January, 1980

FIGS. 4-17. Diagrams of Australian genera of sipunculans; 4, Sipunculus; 5, Xenosiphon; 6, Siphonosoma; 7, Golfingia, 8,

Onchnesoma; 9 & 10, Themiste; 11, Phascolion (in shell); 12, Phascolion (removed from shell); 13, Aspidosiphon; 14,

Paraspidosiphon; 15, Cloeosiphon; 16, Lithacrosiphon; 17, Phascolosoma (The numbers along the scale give its length in mm)

A REVISION OF THE SYSTEMATICS OF ALISTRALIAWN SIPUNCULANS (SIPUNCL/LA) 7

Australian collections of sipunculans

Collections of sipunculans are held in the

Museums of all the States of Australia, The largest

are those in the Austrahan Museum, Sydney and the

South Australian Museum, Adelaide, A list of all

the specimens in the latter Museum and their

registered numbers is available on application to the

Museum.

In the present paper the following abbreviations

are used in referring to the collections: AMS—Aus-

tralian Museum. Sydney: SAM—South Australian

Museum, Adelaide; NMV—National Museum Vie-

toria, Melbourne, WAM—Western Australian

Museum, Perth; TM--Tasmanian Museum and Art

Gallery, Hobart; QM—Queensland Museum, Bris-

bane, The number in brackets in the reference gives

the number of specimens in the collection,

Acknowledgements

1 am indebted to the following for the help that

they have given me: Dr. P. Hutchings and Miss E.

Pope (Aust. Mus,, Sydney), Dr, B, Smith (Nat-

Mus,, Melbourne), Dr. A. Green (Tas. Mus...

Hobart), Mr. W. Zeidler (Sth. Aust. Mus,), Dr, R-

George and Mrs. L. Marsh (West, Aust. Mus.), Mr.

A. Dartnall (Mus., Darwin), Mr. W, Green, Dr, D.

Fielder, Prof. W, Stephenson and Mr, 8, Cook

(Dept. of Zoology, Univ. of Queensland), Miss 1.

Bennett (Univ, of Sydney), Prof. E. Cutler (Utica,

U.S.A.), Dr. M. Rice (Smithsonian Instit.. U.S.A, ).

Dr. V. G, Murina (Sevastopol, U.S.S.R.), Mr, R,

Sims, and Mr. E. Easton (Brit. Mus. Nat. Hist..

London), Dr. van der Spoel (Zool, Mus.,

Amsterdam), Dr, G. Flarwich (Humbolt Mus..

Berlin, D,D.R,) and particularly Dr. P, E. Cribbs

(Marine L.ab., Plymouth, England) who generously

sent me duplicates of specimens collected by him at

the Great Barrier Reef. Mr. P. Kempster (Univ. of

Adelaide) took the photographs and Miss Lesley

Howard drew some of the figures,

Wf. SYSTEMATICS

Phylum Sipuncula and list of species reported from

Australia

The characters of the phylum are stated on p, 2,

(Specific names are in alphabetical order and

Australian specimens of the species marked * have

not been seen by me).

Family Sipunculidae

Sipunculus indicus Peters

Sipunculus robustus Keferstein

Sipunculus titubans filubans Selenka & de Man

Xenosiphon mundanus (Selenka & de Man)

*Siphonosoma australe (Keferstein)

Siphonosoma boholense (Selenka & de Man)

Siphonosoma cumanense cunidnense (Keferstein}

Siphonosoma novaepommeraniae Fischer

Siphonosoma rotumantium (Shipley)

Siphonosoma vastum (Selenka & de Man)

Family Golfingiidae

*Golfingia

Golfingia

*Golfingia

Golfingia

*Golfingia

Galfingia

*Golfingia

“Golfingia

Golfingia

*“Golfingia

coriacea (Keferstein)

herdmani (Shipley)

impravisa (Théel)

margaritacea adelaidensis Edmonds

minuta (Keferstein)

misakiana (Ikeda)

murinae mturinge Cutler

ohlini (Théel)

pellucida (Keferstein)

schuerei (Augencr)

semperi (Selenka & de Man)

Golfingia trichocephala (Sluiter)

Golfingia vulgaris queenslandensis Edmonds

Phascolion collare Selenka & de Man

Phascolion cronullae n.sp.

*Phascolion dentalicolum Sato

*“Phascolion pacificum Murina

Themiste cymodoceae (Edmonds)

Themiste dehamata (Kesteven)

Themiste fusca (Edmonds)

Themiste huttoni (Benham)

Themiste lageniformis Baird

Themiste variospinosa n.sp.

*Onchnesoma steenstrupit Koren & Danielssen

Family Aspidosiphonidae

Aspidosiphon (Chamisso &

Eysenhard)

“Aspidosiphon exhaustus (Sluiter)

Aspidosiphon gracilis Baird

Aspidosiphon hartmeyeri Fischer

Aspidosiphon inquilinus Sluiter

Aspidosiphon jukesii Baird

*Paraspidosiphon cumingli (Baird)

Paraspidosiphon formosanus (Sato)

Paraspidosiphan johnstonii n.sp.

*Paraspidosiphon steenstrupii (Diesing)

Cloeosiphon aspergillus (Quatrefages)

Lithacrosiphon cristatus (Sluiter)

elegans elegans

Family Phascolosomatidae

Phascolosoma

Phascalosoma

*Phascolosoma

Phascolosoma

Phascalosoma

Phascolosoma

albolineatum Baird

annulatum Hutton

arcuatum (Gray)

nigrescens Keferstein

nigritorquatum (Sluiter)

noduliferum Stimpson

pacificum Keferstein

perlucens Baird

rottnesti Edmonds

scolops (Selenka and de Man)

stephensoni (Stephen)

KEY TO FAMILIES OF SIPUNCULA

(based on Fisher, 1952 and Stephen and Edmonds, 1972)

1, Horny, ¢hunous or calcareous shield. cone or cap present at

anterior extremitw of trunk. Longitudinal musculature oF

beady wall may form bands or be continuaus

Aspidosiphonidae™ (p. 43)

No horny orealcarequs shield, cone or cap present at anterior

region of trunk. 6.4.45, 2

Tentacles basically surround moulh ‘and f may be simple or

branched, lie in groups, be reduced to a few lobes of every

be absent but not lying in a horseshoe-shaped ring dorsal tq

mouth. Nuchul organ, if present, Is dorsal to tentacles und

not enclosed by them vount oe ee

Tentacles arranged ina horseshoe- Séagedtri ring which does not

surround mouth bur lies dorsal to it and which encloses the

michal organ, if present, Longitudinal museles grouped

into bundles, except in one genus of four species. Skin

usually bearing comeal to hemispherical papilhform

glands, usually largest and mast promiment at anterior and

posterior extremities oF trunk. Phascolosomatidae (p. 35)

Longitudinal musculature of body wall thickened to form well

déetined bands. Integumentary canals ar caclomic sacs

present in body wall (except in that of one spevies)

Sipunculidae (p. 8)

Longitudinal musculature of body wall continuous and not

thickened into bands. - Golfingidae (p, 18)

ine}

tot

Family Sipunculidae and key to genera

Sipunculidae Baird, 1868 (in part); Sedgwick, L898

(in part); Stephen and Edmonds, 1972,

Description: Adult specimens large and cylindri-

eal, Longitudinal musculature always in bundles.

Retractor muscles always four (except in

Siphonomecus). Mouth surrounded by a ring of

tentacles or a tentacular fold. Body wall of adults

with inteégumental canals or coelomic sacs (except in

Phascolopsis), Nephridia two. Type genus: Sipun-

culus Linnaeus.

Key to genera of the Sipunculidae

1, Introvert with sub-triangular, scale-like papillae but without

hooks, Skin of (runk marked into rectangles by intersecting

bands of longitudinal and circular muscles ..,.,... aA

Introver! without sub-triangular, seale-like pupillae. Hooks

or spines may he present, Skin not strongly marked oft into

rectangles (except in Siphonomecus) ...0 000 ee A

2. Four retractor muscles and an addifional pair of protractor

muscles, the latter arising from body wall near anus and

connecting with introvert near brain, Spindle muscle arises

anteriorly from wall of rectum, Gonad in torm of a fine

loop attached to body wall and rectum

XMenosiphon Fisher (p. 12)

Four retractor muscles but no protractors. Spindle muscle

arising from body wall anterior to anus. Gonads at base of

ventral retractors and not in form of a filamentous loop

Sipunculus Linnaeus (p. 4)

*Dr. P_E, Gibbs of the Marine Laboratory at Plymouth, England

has correctly pointed out to mé (personal communication) that

the tentacles of Aspielosiphon and Paraspidosiphon lie ins

horseshoe-shaped ring dorsal to the mouth, as in Phas-

colosamu. The statements of Stephen and Edmonds (1972: p,

19, 215-216, 238) about the condition of the tentacles in

Aspidosiphonidae, Aspidasiphon and Paraspidasiphon are

therefore Wrong und must be corrected. Lam indebted ws Dr.

Gibbs for his information. Dr. Gibbs (1977: p, 30) hus recently

referred 10 the condition of the tentacles in. Aspidosiphan

mueller’ Diesing,

REC, S. AUST, MUS.

wy 74 Vareey, PORE

3. Four retructor muscles - 215 cite on ef

Two retractor miuiscles | - . Siphanamecus Fisher™

4, Spindle muscle not fixed to body wall posteriorly. No

coelomic extensions i bods wall, Imtrayert withoul hooks

Or spines Phascolopsis Fisher*

Spindle nnrusele fixed posteriorly. Coelomic sacs or canals

present itt body wall, (qtiewert spines ar hooks present of

absent - - Siphonusoina Spengel (p- l4)

Genus Sipunculus |.innacus

Sipuneulis Linnaeus, 1776 p. LTS: Fisher. 1952 p

375; Stephen & Edmonds, 1972 p. 21.

Description: Usually large animals, often stout,

Trunk cylindrical and usually divided into squures or

rectangles by intersection of longitudinal and

circular muscles, Introvert short, clearly differenti-

ated from truok and carrying numerous, flat

triangular papillae. Mouth surrounded by a

tentacular fold, the margins of which in living

specimens form more or less distinct tentacles,

Introvert lacks hooks and spines. Spindle muscle

attached anteriorly in front of anus. Coelomic

extensions present in body wall, Contractile vessel

double. Post-oesophageal or “sipunewlus” loop

present in alimentary canal anterior (o intestinal

spiral. Paraneural muscle well developed. Type

species: Sipunciuluy midus Linnieus 1776.

Key to species of Sipuaculus known from Australia

\. Longitudinal musele in 37-43 bands, trunk very long and

rather slender se . S frylicey (p. 9)

Longiludinal ayuscles in less than 37 bands; trunk usually

SLOUY yy ee:

2. Longitudinal neces in 2: 30 (usually 29) hands, Uigitate

processes af brain rather threadlike a lulera!l in

position. - S. robustus (p.9)

Longitudinal muscles in 23. 26 bands; digitate processes short,

finwer- to leaf-like rather (han threadlike

S. nimagns fitbans (p. LO)

Remarks: Triangular, scale-like papillae are

always present on the tatrovert of Sipanculus and

Xenosiphion but not Siphonosoma. In Nenasiphon

two protractor muscles are present and the ganads

jie on a tilamentous Joop but notin Sipuncalis nor

Siphonesoma. The spindle muscle af Siphonasoma

arises dnteriorly from three roots and is fixed

posteriorly to the trunk wall, In Sipuneulus and

Xenosiphon tt arises anteriorly froma single root and

is not fixed posteriorly.

Fischer (1914 p. 1) and Murina (1972 p. 307)

identified single specimens from St. Vincent Gulf,

South Australia as Sipuaculus nudus Linnaeus.

Because §$. robustus is commonly collected in this

Gulf (see p. 10) there is some doubt about their

records,

* Species of this genus mit reported from Australia.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 9

Sipunculus indicus Peters

Sipunculus indicus Peters, 1850 pp, 382-383;

Keferstein, 1865 p, 421, pl. 31, fig. 1; Selenka

and de Man, 1883 pp, 111-112; Edmonds, 1971

p. 137; Stephen and Edmonds, 1972 pp. 27-28.

Location af type: not known to author; specimen

from Mozambique.

Description: Specimen long and cylindrical,

Length of trunk 410 mm and width, almost uniform,

12mm. Posterior extremity slightly rounded and

swollen: invaginated terminal organ present. Surface

of trunk divided into small rectangular areas by

intersection of circular and longitudinal muscles.

Circular annulations of trunk very noticeable.

Surface of introvert bears sub-triangular papillae.

Longitudinal musculature in 37-42 anastomosing

bands. Four retractor muscles arise at about the

same level, the ventral pair from muscles 2-4 and the

dorsal pair from 9-13 or 10-14. Alimentary canal

with well developed post-o¢sophageal loop. Anus

opens anteriorly to nephridiopores. Rectum long

and fixed to body wall for most of its length. Small

caecum attached to rectum, Two long nephridia,

Opening between muscles 4-5, extend back past

point of fixation of reetractors and ate attached to

body wall for most of their length. Brain simple and

lacking a tufted organ. Contractile vessel double.

Systematic position: Johnston (1969: p, 43)

described Nenosiphon (Xenopsis) indicus from the

Laceadive Is, The species has two rectractor and two

protractor muscles arising from the body wall at

different levels (Johnston 1969, fig. 3). Since he does

not refer to Sipunculus indicus Peters and does not

raise the question of synonymy it looks as if

Johnson's “indicus and Peter's “indicus” are

different.

The Western Australian specimen, although it

shows similarities with Xenosiphon indicus Johnson,

is being described as S. indicus Peters because it

lacks protractor muscles. Its four retractors arise

from the body wall at about the same level, In

addition the brain is simple and lacks the “tufted

organs’ described for XN. indicus.

S. indicus is long and cylindrical, It possesses

about 40 longitudinal muscles and the anus lies in

front of the nephridiopores. Brain without proces-

ses, thus differing from S. robusrus and S, titubans.

Good illustrations of the species are given in

Keferstein 1865, pl. 31, fig. 1 and in Sato 1939, pl.

19, lig. 4. No previous Australian record,

Distribution: (1) in Australia: Western Australia

at Exmouth Gull,

(2) elsewhere; Zanzibar, Madagascar, South

Africa. Billiton, South China Sea, West Caroline Is..

Coral Sea (AMS W5734),

Specimens examined and locality: Western

Australia—Yardia Creek near Exmouth Gulf (1)

AMS W9250.

Sipunculus robustus Keferstein

(Figs. 18, 20-22)

Sipunculus robustus Keferstein, 1865 p. 421; Stephen

and Edmonds, 1972 pp. 36-37.

Sipunculus angasii Baird, 1868 p. 80; Edmonds 1955

pp. 83-86, figs. 1-4, pl. J.

Location of type: Hamburg Museum; specimen

from Uvea (Wallis Is.), Pacific Ocean.

Description; Specimens long, cylindrical and often

stout; frequently washed up on beaches near

Adelaide after early winter storms. Colour pale pink

and slightly iridescent, Trunk 60-260 mm long, 8-

20mm wide and usually divided into small,

rectangular areas by intersection of circular and

longitudinal muscles. Introvert short, clearly dif-

ferentiated from trunk, 15-30 mm long and 5-10 mm

wide, with numerous, posteriorly directly, scale-like

papillae but no hooks. Mouth surrounded by a

wrinkled fold, which in living specimens more

closely resembles finger-like tentacles. Posterior

extremity of trunk usually rounded and swollen but

sometimes pointed. Terminal organ present.

(specimen

from Tasmunia), (scale measurements

are in mm)

FIG. 18. Sipuneulus robustys,

10 REC. & AUST. MUS. 18 (1);

Longitudinal muscles in 27-30 (usually less than

30), sometimes anastomosing, bundles, Four stout,

equal retractors arise at same level in anterior third

of trunk, ventral pair from muscles 3-5 (2-5, 2-6, 3-4)

and dorsal from 9-11 (9-12, 10-12, 8-13), Anterior

region of oesophagus fastened to dorsal retractors by

thin mesenteries and rest of gut to body wall by thin

threads. Post-ocsophageal or “‘sipunculus’’ loop

present in foregut, Racemase glands and caecum

present. Spindle muscle arises anterior ta anus but

not fixed posteriorly. Contractile vessel double.

Nephridia attached between muscles.4 and 5 (5-6, 6-

7) anterior to anus, Bilobed brain gives off dorso-

laterally a number of delicate, thread-like digitate

processes,

Systematic position and remarks: Edmonds (1955)

identified a common South Australian sipunculan as

Sipunciilus angasti Baird 1868, a species described

from, Port Lincoln, South Australia. After an

examination of the holotype of S. robustus, Stephen

and Edmonds (1972: p. 37) concluded that S.

robustus and §, angasti are conspecific, the name

rabustus having priority. S, robustus ts well known in

the Indo-Pacific region.

A comparison made between about 50 specimens

af 8. mudus collected by me at Morgat, Brittany

(France) during 1961 and about 50 specimens of 8.

robustus: collected from South Australian beaches

confirms that the two species, although closely

related, are different. In S nudus there are 28-32

(usually over 30) longitudinal muscles while in 5,

robustus there are 27-30 (usually 28-29). In S.

robustus the digitate processes are rather threadlike

and somewhat lateral in position, In S, nudus they

are fewer, short, rounded to fingerlike and situated

more dorsally, The retractors in S. nudus usually

span six or seven longitudinal muscles while in S,

rabustus it is usually three or four,

Fischer (1914 p. 1) identified S. mudus from St.

Vincent Gulf, South Australia and Murina (1972 p.

307) S. nudus from Adelaide, Both records are of

single specimens. It seems likely that the specimens

are what IT would call S. rabustus-

S, robustus in South Australia lives below the level

of low tide on sandy beaches, The gut contents are

usually sand. Two smail specimens were collected

from the put contents of a fish, Sillaginodes

punctatus, caught at Marian Bay, South Australia

(SAM E1072), The body fluid of S, rabustus like that

of S. audus contains, in addition to blood cells and

amoebocytes, free swimming, ciliated urn cells

which are thought to help in the removal of some

waste materials from the animal,

1-74 January, {980

Previous Australian records: Monro (1931).

Augener (1903). Edmonds (1955), Gibbs (1978),

Distribution; (1) in Australia; Oucensland,

Victoria, Tasmania, South Australia and West

Australia,

(2) elsewhere; Palau, Wallis Is,, West Caroline

Is., Marshall Is., Philippines, Indonesia, Maldive

and Laceadive Is,, Madras, Madagascar, Zanzibar,

off South Africa (at 2.720 m) and Malacea Sc. (at

1 140 m).

Specimens examined and localities: Queensland

—Low Is. (1) SAM E1060 and (1) (Dr. P. Gibbs)

SAM E1057; Turtle Is. (1) (Dr. P. Gibbs) SAM

E1057; Townsville (1) EL052; Line Is. AMS G11388:

Hayman Is, (1) AMS W3131. Victoria—Port

Arlington (1) NM G1125;, Black Rock (1) NM

G1126; Hobsons Bay (1); Queenscliff (1) NM

G1124; Rosebud (1) NM G1128; Brighton (1) NM

G1123; Portland (1) SAM E1049. Tasmania—Seven

Mile Beach (1) SAM E1048 and (2) TM K103/154530.

South Australia—St, Vincent Gulf (washed up after

storms), especially at Aldinga and Sellicks Beaches

(80) SAM E1050, E1051, E1053 and E1055; Spencer

Gulf at Minlacowie (2) SAM E1054; Aldinga Beach

(2) AMS W3600, Western Australia—Rottnest Is.

(1) WAM 125/76, Woodmans Point (1) WAM

172/76.

Sipunculus titubans titubans Selenka & Bulow

(Figs. 23-25)

Sipuncilus titubans Selenka & Bulow, 1883 pp. [00-

LOL, Stephen & Edmonds, 1972 pp. 37-38.

Location of type: Zoological Museum, Humbolt-

University of Berlin, DDR.; speamen from

Puntarenas (coll, Griibe), cat, no, 1036,

Description: This. account is based on 12

specimens, two larger ones from the Gulf of

Carpentaria and 10 smaller ones from Moreton Bay,

Queensland. Al are cylindrical with body wail

divided into rectangles by the crossing of Jongi-

tudinal and circular museles, Body wall of larger

specimen thick but of smaller pink and semi-

transparent. Trunk of larger 80-120 mm long and

8-11 mm wide, of smaller 18-55 mm long and 2-5 mm

wide. Posterior extremity pointed, rounded or

formed into a glans. [nvaginated terminal organ

present in two, Introvert short (maximum length 10

mm), much narrower than trunk and bearing

numerous, fleshy, sub-triangular papillae. Tentacles

not exposed in any specimen but dissection shows

that a tentacular fold is present.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 11

Longitudinal muscles in 23-26 bands (usually 24 at

hase of retractors), anastomosatan slight. Four

introvert retractors arising at same level in anterior

fourth of trunk. Base of retractors usually extended

laterally by strands of muscular tissue (making it

difficult sometimes to specify the span of the

muscles). Ventral retraetors arising from muscles

2-5, 1-4, 1-5 or 1-6, dorsal retactars trom 6-10. 6-11,

7-11 or 8-11, Anterior oesophagus attached to dorsal

retractors by thin mesenteries and post-oesaphageal

loop present im foregut, Rectum short and rectal

caecum presente in lwo larger specimens and twa

smaller ones. Two racemose glands attached to

strands of tissue connecting rectum and base of

dorsal retractors. Contractile vessel double with

roughened or vesiculated surface but no tubular villi.

Spindle muscle arises anterior to anus but not fixed

posteriorly. Nephridia about one fourth or filth as

long as trunk, arising between muscles 4-5 well in

front of anus and attached for about.a fourth of fifth

of their length. Brain bilobed with a number of short

digilate processes along its anterio-dorsal margin;

processes may be simple, stubby or leal-ltke and

those laterally placed may branch slightly.

Systematics: At first 1 thought that these

specimens might be Sipunculus aequabilis S)uiter,

1902 desctribed from Indonesia, A re-examination of

the holoiype (Zoological Museum, Amsterda-

m)—an already dissected specimen—confirms that it

has 21-22 longitudinal muscles, that its ventral

retractors arise [rom musele number 3 and the dorsal

pair from 8-9. [thas a small caecum but no racemose

glands. The specimens from Queensland however,

possess a greater number of longitudinal muscles

and their retractars span more muscles. One point

not previously reported is that at least three stout

swellings or protuberances arise from the fore-brain

of the holotype of S. aequabilis. L am inclined to

think that they correspond to digitate processes.

Although resembling Sipynculus norvegicus

(Danielssen, 1869) in many respects the Queensland

specimens differ because they possess digitate

processes, Recently ] examined two specimens of S,

norvegicus from the collection of the British Museum

(Nat. Hist.)—ane from Norway (1922-5-22-1) and

the other from the Bay of Biscay (97-4-21-4). The

brain of both lacks digitate processes. Confirmatory

evidence on this point is given by Akesson (1958 p.

138); “At about the place where the other species

(S_ nudus and S. robustus) have digitate processes S,

norvegicus has a muscular strand, a rudiment of the

larval protractor muscle, There is certainly no

histological similarity between the two structures

and they are not connected in any way.’

Sipunculus titubans Selenka & Biilaw, 1883 was

described from a Single specimen collected at

Puntarenas. Recently [ examined the holotype, a

dissected specimen. The trunk is about 48 mm long

and the introvert 6 mm, A tentacular fold surrounds

the mouth and triangular papillae are present on the

introvert. Longitudinal muscles 26 at base of

retractors and 26 posteriorly, Ventral retractors

arise from muscles 1-5, 1-6 (type description says 3-5

or 1-5) and dorsals from 8-11, 7-11. Racemose

glands and caecum present. Contractile vessel

double. Nephridia opening anteriorly to anus

between muscles 4-5 and fixed far half their length

As stated in the type description. no spindle muscle

iS present but the intestine has been damaged and

part of it is missing. Brain bilobed and giving off

from its anterio-dorsa] margin a number of short

tufted (possibly leaf-like) digitate processes, some-

thing not mentioned by Selenka (unless included in

the statement “Nervensystem demjenigen S, nudies

sehr abnlich’),

Consequently 1 consider the specimens from

Queensland ta be 8. titubany. The chief differences

are that in the holotype (1) the digitate processes

seem to form a tufted or brush-like structure while in

the others the processes gre stouter and (2) the

nephridia are fixed for half and nota fifth of their

length. The absence of a spindle muscle in the

holotvpe is so unusual in Sipunculus that it may

reasonably be regarded as accidental and not

general. Unfortunately no reference ta the absence

or presence of a spindle nmuscle in their specimens of

S. titubans has been made, as far as T can see, hy

other authors.

The Queensland specimens differ from Sipunculus

robustus Keferstein which has 27-29 longitudinal

muscles and longer and more thread-like digitate

processes and from Sipunculus mudus Linnaeus

which has 30-33 muscles. and finger-like processes,

Sipunculus zenkevitcht Murina, 1969, dredged from

the Pacific Ocean. has 25 longitudinal muscles, an

oval brain with two eyespats but apparently no

processes, Sipunculus longipapillosus Murina, 1968

has 24-26 longitudinal muscles but trunk papillae

with long extensions. Sipunculus delphinus Mutina,

1967 has 25 longitudinal muscles, retractors arising

from muscles 2-5 (2-4) and 7-8 (8-11), a caecum and

tacemose glands and seems very closely related to 8.

titubans,

What evolutionary or adaptive value the different

kinds of digitate processes have in the Sipunculidae

is not known, Metalnikoff (1900) considered that the

processes Were sensory structures. Akesson (1958 p.

136) maintains that they are not sensory but “forma

part of a Secretory organ, In the organ js

accumulated the neurosecretory substance from the

bipolar neuroseeretory cells in the anterior dorsal

margin of the brain. From there the secretion ts

given off to the coelomic fluid”.

12 REC. S. AUST. MUS. 18 (1):

Digitate processes have been found, at least, in

the following Sipunculus species: S. nudus Linnaeus

(finger-like), S. robustus Keferstein (thread-like), S,

timbans Selenka & Bulow (brush-to finger-like), S.

galapagensis Fisher (irregularly folded sheets of

tissue), S. marcusi Ditadi (leaf-like), S. natans

Fisher (numerous slender lobes forming a conspicu-

ous tuft) and S. polymyotus Fisher (very slender,

flattened, thin and digitate). Three or four swellings

or protuberances on the forebrain of the holotype of

§. aqeuabilis Sluiter are probably processes. The

processes are lacking in §. norvegicus Damelssen.

Fischer, 1895 described a subspecies, S, nmubans

diptychus, from West Africa which has 30-33 muscle

bands, retractor muscles attached to bands 2-5 and

11-14 and a spindle muscle.

No previous record {rom Australia.

Distribution: (1) in Australia: Queensland at

Weipa and Moreton Bay.

(2) elsewhere: North-west Africa, Canary Is.,

Gold Coast Gulf of Guinea, Senegal, Zanzibar,

Madagascar, Gulf of Siam, Thailand, West Indies,

southern Chile,

Specimens examined and localities: Queensland

—Weipa (Albatross Bay, Gulf of Carpentaria) (1)

AMS W9241; mouth of Embley River (Albatross

Bay) (1) WAM 231/76; Middlebanks (Moreton Bay)

dredged in sandy mud at LO m (10) SAM E1069,

Genus Xenosiphon Fisher

Nenostphon Fisher, 1947 p. 360; 1954 p. 312;

Stephen & Edmonds, 1972 p. 38; Johnston,

1969 p. 43,

Type species; Xenosiphon branchiatus: Fisher,

1947,

Description: Large, cylindrical and closely

resembling Sipunculus. Unlike Sipunculus in posses-

sing an extra pair of protractor or retractor muscles,

atising from posterior border of introvert near brain.

Introvert short and with flat, triangular papillae,

Longitudinal and circular muscles banded as in

Sipunculus. Rectum long and anus lies anterior to

nephridiopores, Gonads form a filamentous loop.

Integumental canals or sa¢s present

Type species: Sipunculus mundanus branchiatus

Fischer, 1895.

Remarks; Four species have been described for

the genus. They fall into three subgenera; (1)

Xenosiphon (sensu stricta) Fisher, 1954 p, 312,

Subcutaneous coelomic system consists of inde-

pendent, irregular sacs, some of which carry

papilliform gills. No accessory intestinal loop,

1-74 January. 198()

Nephridia long and attached to body wall by

mesentery, X, caribaes. Fisher, 1954 also belongs to

this subgenus.

(2) Austrosiphon Fisher, 1954

Type species: Xenasiphon mundanus (Selenka &

Bulow, 1883).

Subcutaneous system in form of longitudinal

canals lying between longitudinal ridges of body

wall, Accessory or ‘‘sipunculus’’ loop present in

foregut, Nephridia small and free,

(3) Xenopsis Johnson, 1969

Type species: Xenosiphon indicus Johnson, 1969.

Subcutaneous system as in Austrosiphon., Two

protractors and only one pair of retractors.

“Sipunculus’’ loop present in foregut, No cephalic

tube.

In Australia only one

mundanus, is known.

species, Xenosiphon

Xenosiphon (Austrosiphon) mundanus (Selenka and

de Man)

(Figs. 26-27)

Sipunculus mundanus Selenka and de Man, 1883 pp,

198-109, pl. 12, fig. 174.

Xenosiphon mundanus Fisher, 1954 p. 314;

Edmonds, 1955 pp, 87-89; Edmonds, 1960 p,

160: Cutler, 1977 p. 138.

Location of type: British Mus. (Nat. Hist.),

London; specimen from Sow and Pig Shoal, Port

Jackson, New South Wales.

Description: Specimens long, cylindrical and

superficially resembling a Sipunculus. Trunk 130-

270 mm long and 7-12 mm wide. Body wall divided

into réctangular areas by intersecting bands. of

longitudinal and circular muscles. Introvert short,

10-20 mm long and 4-7 mm wide, beating many,

posteriorly directed, scale-like papillae and a

tentacular pad oar fold. Annulations caused by

circular muscles usually less marked on dorsal

surface of postenar third of worm. Longitudinal

muscles in 28-31, slightly anastamosing bands.

Terminal organ present al posterior extremity of

trunk

Four retractor muscles arise at about same level, a

ventral pair from muscles 2-4, 1-3, 1-4 and dorsal

pair from 7-10, 7-11, 8-11. Two ribbon-like

protractor muscles arise fron body wall of anterior

part of trunk and are attached to introvert near

brain, Oesophagus long and straight; short post-

oesophageal or “sipunculus” loop present, Spirals of

intestine fastened by many short mesenteric threads,

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 13

FIGS. 19-31, Fig, 19. tentacular fold of a Sipunculus (after Gibbs), Figs. 20-22, Sipunculus rebustus; 20, anterior region dissected; 21,

brain and digitate processes; 22, subtriangular papillae from introvert, Figs. 23-25. Sipunculus titubans titbans; brain and

processes of three specimens fram Queensland: Figs. 26-27, Xenosiphon mundanus; 26, anterior region dissected; 27, brain and

tufted pracesses (after Fisher). Fig. 28, Siphonosoma cumanense: anterior region dissected. Fig. 29, Siphonosoma australe; hooks.

Fig. 30, Siphonosama rotumanum: hooks. Fig. 31, Siphonosoma vastum; anterior region dissected,

O-1

i4 REC. 8. AUST. MUS. 18 (1):

Anal aperture lies well anterior to nephridiopores,

Rectum long and attached to body wall, Smull

caecum present, Contractile vessel double.

Gonads in form of a fine, delicate Joop of tissue

arising on each side of rectum and running across

body wall to base of retractors and nerve cord.

Nephridia short, stout and free, opening between

muscles 4-5. Integumental canals lie longitudinally

between ridges of muscles. Brain bilobed and

capped anteriorly with a pad of tufted tissue.

Systematic position; This species is well known in

south-eastern Australia and New Zealand, Its

distinctive features are its two protractor muscles, its

long rectum, its short nephridia (in X. branchiatus

they are long) and the position of the anus well

anterior to the nephridiopores. X. indicus Johnson,

1969 from the Laccadive Is. possesses 40-45

longitudinal muscles.

Previous Australian records: Selenka and de Man

(1883); Edmonds (1955); Cutler (1977).

Distribution: (1) in Australia: New South Wales,

Lord Howe Is., Victoria and off coast of South

Australia at 1320 m (Cutler 1977),

(2) elsewhere: New Britain, New Zealand; Chile

(Tarifino and Tomicic, 1973).

Specimens examined and localities: New South

Wales—Gunnamatta Bay (1) SAM E1083; Pig and

Sow Shoal. Port Jackson (2) SAM EJ080; Lord

1-74 January, L980

Howe Is. (2) AMS G3950 and W1873; Bermagui (2)

AMS W2534; Botany Bay (1) AMS W2535;

Balmoral Beach (1) W1665, Victoria—Queenscliff

(1) SAM E1085; Portland (1) NMM G1217, Port

Fairy (4) AMS W9249. Tasmania—Binalong Bay (1)

TM KS25,

Genus Siphonosoma Spenge!

Siphonosoma Spengel, 1912 p. 264; Fisher 1950a p.

805; 1952 pp. 380-381; Stephen and Edmonds

1972 p. 43,

Type species: Siphonosoma australe (Keferstein,

1865).

Description: Adults usually large and eylindrical,

Introvert offen not well differentiated from trunk

and lacking seale-like, triangular papillae but

sometimes possessing hooks and spines. Tentacles

thread-or finger-like; no tentacular fold, Longitudin-

al muscles always divided into bands (sometimes not

readily noticed from outside). Coelomic sacs in body

wall. Four retractors. Spindle muscle arises

anteriorly from three roots and attached posteriorly

to body wall, Contractile vessel single and dorsal:

small contractile tubules usually present. Two

nephridia (usually with large crescentic nephros-

tomes). No post-oesophageal loop,

Subgenera: Fisher (1950a p. 805) divided the

genus into three subgenera.

TABLE 1.

SUBGENERA OF SIPHONOSOMA

Numerous rectal

Type species

S, australe (Keterstein)

S, vusnom (Selenka & de Man)

S runtinense (Kelerstem)

ee eee mn oo ovmmvvvwvoO oo

Subgenus Transverse

dissepiments caevat

Siphonosentd ss absent absent

Hesperosiphon absent presen

Dainostiphon present whsent

Remarks: Species of Siphonosoma can be

distinguished from those of Sipunculus because (1)

the introvert lacks scale-like papillae (2) the

tentacles do not lorm a fold (3) the spindle muscle

arises anteriorly from three roots and is fixed

posteriorly and (4) no post-oesaphageal loop is

present in the foregut.

KEY FO AUSTRALIAN SPECIES OF SIPHONOSOMA

i. Numerous transverse dissepiments attached to coelomic wall

oftrunk., 8. (Siphonosoma) cumanense cumanense (p, 14)

No transverse dissepiments attached to coelomic wall

. Rectum with very numerous, finger-like caeca or villi

S. (Hesperosiphon) vasium (p15)

Rectum without finger-like caeca or villi .... ;

3. Introvert with hooks, spines or spine-like papillne .....,

Introvert without hooks, spines or spine-like papillae

= Ww.

4. Brown-black, slightly bent hooks present

S. (Siphonosoma) australe (p.

Light-brown, blunt, papilla-like spines present

S. (Siphonosoma) rotumanum (p, 18)

5. Longitudinal muscles 30-32

§, (Siphonosoma) bololense (p.

Longitudinal muscles about 20

S. (Siphanosoma) navaepammeraniae (p, 17)

Siphonosoma (Damosiphon) cumanense cumanense

(Keferstein)

(Fig. 28)

Phascolosoma cumansense Keferstein, 1867 pp. 53-

SS, pl. 6, figs 19-21.

Sipunculus deformis Baird, 1868 pp. 80-81, pl, 9, fig,

2: Edmonds, 1955 p. 90.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPLINCULA) \

Siphonasoma cumanense: Edmonds 1955 pp. 90-92;

Stephen and Edmonds, 1972 pp. 46-49.

Location of type: Not known by author; specinien

from Venezuela,

Description: Specimens of various size und shape:

usually Jong and cylindrical. Length of trank and

maximum width of two largest specimens 410 x (10-

20) mm, 450 x (12-21) mm, and of smallest 50x (4-6)

mm, Anterior region of trunk usually swollen

Introvert not strongly differentiated from trunk and

for such a large species comparatively short:

maximum length 60 mm, minimum [0 mm and

lacking hooks and spines. Mouth surrounded by

finger-like tentacles which, although nurnerous,

seem smal] lor the size of the animal. Anterior

surface of trunk may be divided by furrows into

small areas enclosing flat, glandular openings.

Hemispherical papillae, with diameter up to 0-6 mn,

on anterior and posterior surfaces of trumk; lying

more or less in longitudinal rows and composed of

long narrow plates with their long axes placed along

the radii of the papillae. Body wall thick except

Where if is not traversed by cireular and longitudinal

musculature, such areas containing prolongations or

extensions of the coelam.

Longitudinal muscles in 18-25 (usually 20-22)

anastomosing bands, Four retractors arise in

anterior fourth of trunk at about same level

(sometimes dorsal pair u little more anteriorly), a

ventral pair from muscles 2-3, 3-4, 1-3, 1-4 and a

dorsal from 6-8, 7-8, 8-9. Sometimes retractors may

lic obliquely along one or two muscles rather than

transversally across a wider band,

Anterior region of ocsophagus fixed to ventral

retractors by thin mesenteries, Stout spindle muscle

arises anteriorly from 3 roots, one anterior to anus

and two from muscles 8, 9 or 10 near point of

fixation of dorsal retractors. Two tasteners, arising

from muscle 1 on each side of nerve cord, run to last

spiral of intestine. Rectal caecum usually situated

near point where 3 roots of spindle muscle meet.

Two strands of muscle arise from muscle 1 at about

level of nephrostomes, stretch across body wall and

attach to wing muscle. Contractile vessel with

numerous small villi. A series of thin crescentic

shaped dissepiments lie transversally on each side of

nerve card between base of retractors and postertor

extremity of trunk, spanning muscles 1-8, 3-7, 4-10

or 3-9, Two nephridia, attached ta body wall for

about a fifth of their length and extending to the

base of the retractors, open between muscles 2-3 or

3-4 just anterior to anus. Tufted organs on coelomic

wall near nephrostames, Oval bodies, 0-8-1-5 mm in

diameter, attached to inner body wall. Gonads at

base of ventral retractors, Terminal organ present,

Brain without processes.

Systematic position; These specimens possessing

body dissepiments but lacking introvert spines and

numerous rectal caeca fall into the subgenus

Damosiphon Fisher, 1950a. They are being iden-

tified as S. cumanense cumanense, a species

originally described from Venezuela and since

reported from other tropical and warm waters.

Siphonosoma edule Pallas, 1774, however, is a very

closely related species, so close that it is nat easy to

decide on a character that distinguishes the two. The

introvert af S, edyle is said to be shorter than that of

§. cumanense. The differences are discussed but not

tesalved in Stephen & Edmonds, 1972 P. 46 and 51.

As S. edule was described from Batavia it is possible

that some of these Australian specimens.are S, edule

(especially the specimen SAM E1090), A careful

study of specimens collected from Batavia and

Venezucla needs to be carried out, I think, before a

decision about the two specjes can be made.

Previous Australian records: Baird (1868). Fischer

(1921), Monro (1931), Edmonds (1955) and

Gibbs (1978),

Distribution; (1) in Australia: East coast from

Cape York to the mouth of the Clarence River,

including the Great Barrier Reef; North West coast

near Broome; West coast near Fremantle,

(2) elsewhere: East coast of America, Venezuela,

Florida; Indian Ocean, Madagascar. Zanzibar, Red

Sea, Arabian Sea. Laceadive and Maldive Is.,

Amboina; Pacific Ocean, New Guinea, Loyalty Is.,

New Britain, Philippines, Japan, Tahiti and Indo-

China,

Specimens examined and localities: Queensland

—Dunwich (in mud flats) (1) SAM E1090; same

locality (1) SAM E1091 (2) SAM E1092.

(1) SAM E1093, (1) SAM E1094: Low Is..

(4) SAM E1096; Sir Charles Hardy Is., (11° 55° 8S,

143° 28 E) British Museum, New South Wales

—Shelly Beach (mouth of Clarence River)

(1) AMS W3194; Port Denman (3) AMS. W1006.

Western Austraha—Kwinana (1) WAM _ 196/76;

Fremantle Harbor (1) WAM 199/76; North West

coast (1) Dept, of Zoology, Uniy, of W.A,

Siphonosoma (Hesperasiphon) vastum (Selenka & de

Man)

(Fig. 31)

Sipunculus vastus Selenka & de Man. 1883 pp. 103-

104, fig. 171, fig. 179.

Siphonosema crassum Spengel in Fischer, 1919a p-

279,

Siphonosoma vastum: Fischer, 1927 p, 199;

Edmonds, 1955 pp, 92-95, figs §-9; Stephen &

Edmonds, 1972 p. 55-56.

16 REC, 8. AUST. MUS. 18 (1),

Location of type: Zoological Museum of Humboldt

University of Berlin; specimen from Jaluit

(Marshall Is., Pacific Ocean),

Description: Trunk of Queensland specimen

about 260 mm long and 10-13 mm wide and of

Western Australia 57 mm long and 7 mm wide,

Introvert, retracted in both specimens, 15-50 mm

long; armed with numerous rows of curved, yellow-

brown, rather blunt hooks or spinelets, the largest of

which is about 0:1 mm long. Skin between rows of

spines divided by fine furrows into small rectangular

areas, each containing a small glandular opening.

Surface of introvert and trunk bears hemispherical

papillae, the largest (at anterior of trunk) being up

to 0-5 mm in diameter,

Longitudinal muscles in 22-26 anastomosing

bands, Two stout ventral retractors arise from

muscles 2-6 or 1-6, and two dorsal retractors more

anteriorly from muscles 9 or 9-10. Oesophagus

attached to ventral retractors by fine mesenteries

and gut filled with calcareous fragments, Contractile

vessel with numerous small tubules or villi,

Numerous larger fingerlike villi or caeca J-1-5 mm

long, attached to rectum), their function 1s unknown,

Anal aperture between muscles 11-12 at about level

of nephridiopores, Wing muscle strong. Stout

spindle muscle arises anteriorly from 3 roots, one

from muscle 12 anterior to anus and others from

muscles 11 and 8. Another fastener from last whorl

of intestine bifurcates, each root being attached to

muscle | on each side of nerve cord, Spindle muscle

attached posteriorly to body wall. A sac-like,

globular, rectal or intestinal caecum, feadily

distinguishable from the numerous fingerlike villi,

also present, Two nephridia, possessing prominent

semi-lunar nephrostomes open between muscles 3

and 4, are attached for about a third of their length.

Brain small and lacking processes.

Systematic position: The internal anatomy of

these specimens resembles more closely that of S.

vastum as shown in fig. 171 of Selenka & de Man,

1883 tather than that of S. parvum Fischer, 1928,

Previous Australian record: Shark Bay, Western

Australia (Fischer, 1919a, 1927); Queensland

(Edmonds, 1955).

Queensland at

Reef; Western

(1) Australian:

Barrier

Distribution:

Heron Is, and Great

Australia at Pt. Cloates,

(2) elsewhere; Pacific Ocean; Marshall Is,,

Funafuti, Rotuma, New Britain, Guam and

Solomon Is, Indian Ocean; Mauritius, Laccadive fs.,

Maldive Is, {ndonesia-

1-74 Tunuary, 1980

Specimens examined and localities; Queensland

—Outer Barrier Reef (1) AMS W1606; Heron Is.

(1) NMY coll.; Gillet Cay (4) AMS W9245. Western

Australia—Point Cloates (1) WAM coll,, Ningalloo

Exped.

Siphonosoma (Siphonosoma) australe (Keferstein)

(Fig. 29)

Phascolosoma australe Keferstein, 1865 pp. 422-3,

figs. 12 and 13.

Sipunculus australis Selenka and de Man, 1883 p. 90,

figs, 180-183,

Siphonosoma ansirale Fisher, SOT:

Edmonds, 196] pp. 217-220.

1950a p.

Sipunculus aeneus Baird, 1868 p, 76; Edmonds. 1961

pp. 217-220,

Location of type: Not known by author; specimen

from Sydney, Australia.

Description: Although specimens of S. ausirale

from New Zealand, Salomon Is. and south India

have been examined by the authar none are

available from Australia. The following information

is based on that of Keferstein (1865), Selenka and de

Man (1883), Fischer (1922), Fisher (1950a) and

Edmonds(1961).

Specimens long and cylindrical; trunk 130 mm

(Keferstein)—220 mm (Fischer). Ratio of length of

introvert to that of trunk half (Keferstein). third ta

half (Fischer) and almost three-quarters (Fischer),

Prominent, hemispherical papillae on surlace of

trunk. Introvert armed with numerous rows of dark

brown hooks or spines, (45 mm long (Keferstein),

0):22-0-34 (Edmonds), 0:3 (Fisher) and 0:2 (Fischer).

Longitudinal muscles in 15 bands (Keferstein) but

13-18 according to other authors. Four retractor

muscles arising at different levels in anterior third af

trunk, a ventral pair from muscles 1-3 of 3-5 and a

dorsal pair from 5-6 or 4-6. Nephridia long and free.

Spindle muscle arises anteriorly from 3 roots and

fastened posteriorly to body wall, Rectal caecum

present.

Systematics! This species lacks both body

dissepiments and numerous rectal cueca. It possesses

rather long, dark spines. The differences between S.

australe and the closely allied S. eniwefoki ure

discussed and illustrated in Fisher, 1950a, The hooks

of S. rotumanum, another allied species, are stubby

and rounded while in S, australe they are longer and

sharper. Previous Australian record: Sydney. New

South Wales (Keferstein, 1865).

Distribution: (1) in Australia; Sydney, New South

Wales.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCLILANS (SIPUNCULA) 17

(2) elsewhere: Pacific Ocean; New Zealand,

Samoa, Fiji. New Britain, Solomon Is., Philippines

and Tasman Sea (at 610m). Indian Ocean;

Zanzibar, Madagascar, [India and Indonesia.

Siphonosoma (Siphonosoma) boholense (Selenka and

de Man)

Sipuneulus boholensis (Semper!) Selenka and de

Man 1883 pp. 109-111, pl, 12, figs. 175-177.

Siphonosoma hoholense Stephen and Edmonds 1972

p. 63,

Location of type, Not known; specimen from Bohol

(Philippines).

Description; Specimens very large and very stout.

Trunk 270-4140 mm long and 15-25 mm_ wide;

maximum width anteriorly. Three specimens dark

purple and fourth yellow brown. Body wall very

thick, Introvert completely or partly contracted in all

specimens; its maximum length about 80 mm. and

width 10-13 mm. A dissected introvert shows that

numerous rows of brown papillae ring the organ but

that hooks are absent, Base of introvert usually

more darkly pigmented than rest of body, Skin of

trunk anterior to anus rough and warty.

Longitudinal muscles in 29-33 anastomosing

bands, Prominent rather flat papillae or skin bodies

present on surface of trunk, Posterior extremity of 2

specimens is shghtly bulbous and a terminal organ

present. Four strong retractors arise in anterior third

of trunk, a ventral pair fram bands 4-5, 6-7, 4-6 anda

dorsal pair more anteriorly from 9-12, 8-11, 8-13.

Contractile vessel, with numerous very smail villi,

appears bushy. Spindle muscle stout: arises

anteriorly from three roots, one being fixed in front

of anus and others near base of dorsal retractors.

Wing muscle strong. Rectal caecum in one specimen

but not in second. Two nephridia, fixed to body wall

for about two thirds of their length, open just in

front of or at about same level as anus.

Nephrostomes prominent and surrounded by

numerous, small, tufted projections of the body

wall. Prolongations of coelom extend into body wall,

No fastener to last whorl of intestine found.

Systematic position: This is one of the largest

Species of sipunculans. It differs from other species

of the same genus most nouceably in the number of

longitudinal bands. In addition it lacks transverse

dissepiments and hooks. This is the first Australian

record.

Distribution: (1) in Australia; Queensland,

(2) elsewhere: Bohol, North Borneo,

Specimens examined and localities: Queens-

land—Yule Point, Townsville (3) SAM E1123,

Dunwich (1) SAM E1124.

Siphonosoma (Siphonosoma) novaepommeraniae

Fischer

Siphonasoma novaepontmeraniae Fischer, 1926 pp,

104-106, pl, 3, figs 2-4; Wesenberg-Lund, 1959b

p. 55; Edmonds, 1971] p. 140; Cutler, 1977 p,

139.

Location of type: Not known; specimen from New

Britain.

Description: Specimen stout, cylindrical, narrow-

ing slightly towards. posterior and dark purple in

colour in preserved condition, Body wall thick,

Trunk 190 mm long and about 12-15 mm wide.

Specimen contracted and longitudinal muscle bands

not apparent externally, Prominent hemispherical

papillae scattered on anterior eighth and posterior

third of trunk, their diameter being 0-27-0-55 mm.

Introvert, in retracted condition, about 45 mm long.

Dissection of introvert shows presence of numerous

rows of flat, circular papillae but no hooks,

Longitudinal muscles 17-18, with only slight

anastomosation, Four short retractor muscles arise

from two levels, a ventral pair from muscles 2-3 or 3-

4 and a dorsal pair more anteriorly fram bands 4-5.

Oesophagus shart but coiled intestine very long,

Spindle muscle arising anteriorly {rom three roots,

one being attached in front of anus, one to muscle 7

on right side and another to muscle 5 on the left near

base of dorsal retractors. Spindle muscle fixed

posteriorly, A fixing muscle connects posterior

region of intestine to body wall. Wing muscle very

strong. Gut filled with tightly packed coral

fragments. Caecum not observed but rectal region

damaged during dissection. Two nephridia, free for

most of their length, arise at about same level as

anus or just anterior to if. Contractile vessel with

very numerous shart villi, giving it a tufted

appearance, No transverse dissepiments in body

cavity. Brain not noticeable.

Systematics: This species can be distinguished

from other Australian species of the same genus

because (1) it lacks transverse body dissepiments,

(2) it lacks hooks, and (3) it has less (17-18)

longitudinal muscles than S, boholense which

possesses about 30.

Previous Australian record; Coral Sea (30° 05'S,

154° 33°F) at 1 560 m (3) (Cutler, 1977 p, 139),

Distribution: (1) in Australia: Queensland

(Heron Is.); Coral Sea (at 1 560 m).

(2) elsewhere: New Britain; Guam; Mauritius.

Specimens examined and locality: Queensland—

Heron Is., (1) SAM E1125.

18 REC. S. AUST. MUS. 18 (1):

Siphonosoma (Siphonosoma) rotumanum (Shipicy)

(Fig, 30)

Sipunculus roturmanus Shipley, 1898 pp. 469-470, pl.

37, figs 1-3.

Siphonosoma hawaiense Edmonds, 1966 pp. 386-

388, figs 1-4,

Siphanesoma rotumanum: Edmonds, 1971 p. 143, fig

4; Stephen & Edmonds, 1972 p, 71.

Location of type; British Museum (Nat, Hist.),

London; specimen fram Rotuma.

Description; Specimen cylindrical and slightly

curved ventrally. Trunk about 65-7 mm long and 8

mm wide, Introvert about 30 mm long, armed with

transverse rings of pale-coloured, blunt hook-like

structures, Although they seem larger than thase

described by Edmonds (1966, 1971), hooks closely

resemble those of S. retumanum. Papillae of two

kinds; one, found mostly on introvert, is small,

circular, about 0-1 mm in diameter, with a clear pore

at its centre and the other, found mostly on trunk

and few in numbers, is larger, hemispherical and

about 0:4 mm in diameter. None of papillae black,

as reported by Edmonds (1971).

Longitudinal muscles 15-16 and slightly anas-

tomosing. Four retractors arise at different levels, a

long ventral pair in mid-region of trunk from

muscles 2-3 and a shorter dorsal pair more anteriorly

from muscles 4-5; all fuse anteriorly. Alimentary

canal held in position by fine fastening strands and a

spindle muscle. Spindle muscle attached anteriorly

well in front of anus and posteriorly by several

radial-like extensions. Spindle muscle with two very

prominent lateral roots connecting with muscle 7 on

each side of nerve cord. Additional fastener runs

from muscle 1 (at a point mid-way between base of

dorsal and yentral retractors) to penultimate spiral

of intestine, Wing muscle strong, Rectal caecum

present. Nephridia slender, fixed to muscle 2 for

most of their length and opening at about same level

as anus; nephrostomes prominent and crescentic.

Tufted bodies present on bady wall near nephros-

tomes.

Systematic position: The specimen corresponds

closely with specimens of S. rotumanum in my

possession from Rotuma, Hawaii and Guam. The

“hooks” or adhesive structures on the introvert are a

little larger than those previously described and only

one fastener was found arising between the dorsal

and ventral retractors,

Much of the internal anatomy of S. ratumanum is

like that of 5, qusirale (Keferstein). The hooks of the

latter, however, are sharp. S$. rotumanum differs

from S, novaepommeraniae (Fischer, 1926) in the

possession of hooks.

1-14 January, 1980

Previous Australian record: Low Is., Queensland

(Gibbs, 1978).

Distribution: (1) In Australia; Queensland.

(2) elsewhere: Pacific Ocean at Rotuma, Hawaii,

Soloman Is. and Guam.

Specimen examined: Queensland—Low Is, (1)

SAM E1128.

Family Golfingiidae and key to genera

Golfingiidae Stephen & Edmonds, 1972 p. 77.

Description: Tentacles basically surround mouth

and may be finger-like, thread-like, branched or

dendritic; they may be reduced to a few lobes or

even be absent. No coelomic extensions in body

wall. Longitudinal and circular musculature continu-

ous. Tentacles usually interrupted mid-dorsally by a

nuchal organ, Type genus; Golfingia Lankester,

1885,

KEY TO GENERA OF GOLFINGIIDAE

1. One nephridium.._..

Two nephridia --.- -... --.., 2... | occe ap

2. Anus on anterior region of introvert near mouth. Single

retractor muscle. Introvert very long . Qnchnesoma (p. 32)

Anus on trunk and not on introvert, One or two retractors.

Animals usually inhabiting discarded shells of gastropods

and scaphopods, sometimes the tubes of annelids and more

rarely solitary corals ........ veeees Phascalion (p 29)

3. Tentacles arising from 4-8 stems and branching or dendritic.

Retractors usually two, very rarely four. Contractile vessel

with well-developed tubules or villi ..... Themiste (p. 32)

Tentacles usually finger-or thread-like and not branching;

sometimes reduced to a few lobes or more rarely absent.

Retractor muscles two or four... ..... 6: Golfingia (p, 18)

Genus Golfingia Lankester

Golfingia Lankester, 1885 p, 469; Fisher, 1950b p,

548; 1952 pp. 388-9; Stephen and Edmonds,

1972 pp, 77-80; Cutler and Murina, 1977 pp,

173-187,

Description: Often bottle- or flask-shaped,

sometimes cylindrical or sub-cylindrical. Longitudi-

nal and circular musculature continuous. One or

more rows of finger- or thread-like tentacles usually

surround mouth; in few species tentacles much

reduced or even absent, Nuchal organ usually

present mid-dorsally in ring of tentacles. Hooks

present or absent. One or two pairs of retractor

muscles. Spindle muscle may or may not be attached

posteriorly ta body wail.

Type species: Sipunculus vulgaris de Blainyille,

1827,

Remarks: The genus is large, containing more

than 100 species. Only about six of these are present

in collectians in Australia; mast of the species listed

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPLINCULA) WwW

in this paper are fram the records of other workers.

The genus has been divided into subgenera by Fisher

(1950b), Wesenberg-Lund (1959a), Stephen (1964),

Cutler and Murina (1977) and Murina (1967; 1975a;

1977). In dealing with the taxonomy of the genus [

have largely followed Cutler and Murina (1977) who

reviewed the group and synonymised a number ot

species. T received Murina’s latest subdivision of the

genus (Murina, 1977) only after the present paper

had been typed.

KEY TO SUBGENERA OF GOLFINGIA

i. One pair of retractor musciey.........., 2

Two pairs of retractor muscles _._.. . 4

2. Spindle muscle not attached posteriorly, ,-.. .-. ----. .3

Spindle muscle attached posteriorly .......... Siphonoides*

_ Phascoleides (p, 24)

.. Phy sanacardia (p27)

3. Contractile vessel without villi

Contractile vessel with villi.

4. Nephridiabilobed......... ., 2... Mitosiphon (p, 22)

Nephridia single-lohed ............... Pereere, rer)

5. Spindle. muscle attached posteriorly .. - Galfingiella (p, 27)

Spindle muscle not attached posteriorly ..Galfingia s.s. (p. 12)

Subgenus Golfingia s.s.

Golfingia sensu stricto Fisher, 1950b p, 549; 1952 p,

390; Stephen and Edmonds, 1972 p, 81; Cutler

and Murina, 1977 p. 179.

Description; Two pairs of retractor muscles.

Hooks may or may not be present, Contractile vessel

simple and without villi, Spindle muscle not fixed

posteriorly.

Type: Sipunculus vulgaris de Blainville-

KEY TO SPECIES OF SUBGENUS GOLFINGIA

REPORTED FROM AUSTRALIA

1. No hooks of spines on introvert

G. margaritacea adelaidensis (p. 21)

Introvert with hooks or spines .-.-..-... 0.24.4, em aly

2. Anterior and posterior extremities of irunk. of unrelaxed

specimens forming 4 cap or shield-like structure; globose

papillae presenton shield... .... _. G. herdmani (p. 19)

No cap or shield-like structure al extremities of trunk ,,.... 3

3. Spines tend to be straight; smal! swelling at hase of spine

G. vulgaris queenslandensis (p. 21)

Spines claw-like; no swelling at base of spines G, ohlini(p, 22)

Golfingia (Golfingia) herdmani (Shipley)

(Figs. 32, 36-38)

Centrosiphon herdmani Shipley, 1903 pp. 171-174,

pl. 1, figs, 4-10.

Location of type: not known by author; type

locality, Sri Lanka (= Ceylon).

Description: Specimens light to dark brown in

colour and cylindrical, When freshly collected or

fixed without relaxation they usually assume the

+ No species of this subgenus has been reported from Australia,

characteristic shape. anterior and posterior regions

of trunk looking rather like aspidosiphonid caps or

shields which are surrounded at their junction with

trunk by a fold of body wall, Caps usually dark

brown or rust coloured in contrast with light brown

trunk, [n relaxed specimens caps and folds. less

noticeable or may be difficult to discern, then

resembling more closely a typical polfingiid. Caps

covered with small, rather globose, glandular, dark

rust-brown, slightly stalked papillae tending to lie on

elevated, radial ridges of body wall which suggests

that longitudinal musculature in these areas could lie

in bundles, An examination of transverse sections of

the area, however, shaws that this is notsa, and that

the ridging is in layers external to the dermis, Nor do

sections show hardened or cornified structures found

in cap of some aspidosiphonids (Hyman 1959, fig.

219H),

Length of trunk of 20 unrelaxed specimens 25-65

mim, width 3-8 mm_ Introvert about a quarter to half

length of trunk, 2-3-5 mm wide and arising centrally

from cap and not ventrally as in Aspidosiphon;

anterior region slightly bulbous, Mouth surrounded

by numerous finger-like tentacles. Anteniorly

introvert armed with randomly arranged, hight to

dark brown hooks or spines 0:15-0:20 mm long.

Associated with spines at or near their base is a small

bulbous swelling of body wall which sometimes

appears to le between the wings of the spines.

Papillae on introvert base and on anterior cap

bulbose (with rendency to be stalked), dark brown

and 0:12-0:19 mm in diameter. Papillae on trunk

wall flatter, less pigmented, smaller and less

noticeable. Trunk may feel and appear smooth

although covered with small, white, glandular

openings. On posterior cap papillae dark brown-rust

coloured and about same size as those on anterior

cap.

Four retractor muscles separate for most of their

length, a strong ventral pair arising from about

middle of trunk and a weaker dorsal pair more

anteriorly. Oesophagus fastened for some of its

length to ventral retractars by thin mesentries.

Intestine Jong (about 20 double spirals). Rectal

caecum present. Three fixing muscles, either two ta

oesophagus and one to fectlum or one to

oesophagus, one to posterior spiral and one to

rectum. Wing muscle strong, sometimes extending

almost fo nerve cord on each side. Spindle muscle

strong, arising under rectum either from a triangular

flap of tissue, the apex of which connects with

spindle muscle or (more frequently) from two strong

roots (perhaps the lateral parts of the triangular

flap). Spindle muscle not fixed posteriorly. Nep-

hridia tubular but may be swollen near nepheidio-

pore, arising just in front of anus, about a third as

long as trunk and free for most of their length,

20 REC. 8. AUST. MUS. 18 (1):

1-74 January, 1980

FIG, 32. Golfingia herdmani, (specimens from Proper

Bay, South Australia).

Gonads at base of ventral retractors. Contractile

vessel tubular and without villi. Nuchal organ

present but not always readily noticeable. Two

eyespots.

Systematic position: The specimens fit well the

description of Centrosiphon herdmani Shipley, 1903

described from two specimens from Sri Lanka

(formerly Ceylon). No other record of the species

exists. Shipley placed them in a new genus because

they possessed, anteriorly and posteriorly, thick-

ened, ‘‘chitinoid” shields and because the introvert

arose from the centre of the shield and not ventrally

to it as in Aspidosiphon. Transverse sections of the

“shields’’, however, show that structurally they are

not differentiated from the rest of the bady wall and

that no chitinous bodies are present. When the

specimens are relaxed the “‘shields” usually become

less evident and sometimes disappear. Because the

presence of the shields or caps is an uncertain

character it seems unwise to use it to separate off a

new genus. The tentacles are arranged like those in

Golfingia, the papillae, the introvert hooks and the

internal anatomy of the specimens are golfingiid and

not aspidosiphonid-like. At present, it seems better

to me to call the species Golfingia (Golfingia)

herdmani rather than Centrosiphon herdmani. The

transference of the type of the genus Centrosiphon to

Golfingia makes the genus Centrosiphon invalid. I

FIG. 33. Golfingia margaritacea

adelatdensts, (specimen from

Aldinga Bay, South Australia).

have not been able to find Shipley’s type; it is not

deposited in either the Nat. Hist. Mus., London or

the Museum of the Zool. Dept. of the University of

Cambridge (U.K.), the places where most of his

types were lodged.

For some time I was uncertain as to whether or

not G. herdmani and G. margaritacea adelaidensis

were conspecific. I have decided against this

possibility because G. margaritacea adelaidensis

lacks hooks (they are present in all my specimens of

G. herdmani), its anterior and posterior trunk is not

modified to look like shields and globose papillae

are not present on its body wall.

The taxonomic position of the specimens,

however, is still unsatisfactory. Species of Galfingia

with shield-like structures at one or both extremities

of the trunk have been previously described, G.

(Phascoloides) rutilofusca (Fischer), redescribed by

Murina (1967) and by Cutler (1977), being one.

Golfingia sp. of Cutler (1977 p. 140) as shown in his

fig. 2 is closely related to G. herdmani and matches

some of my Australian specimens. Murina (1975Sa p.

1085) created Dushana, as a subgenus of Golfingia

to contain G. scutigera and G. adriatica both of

which she said, possessed shield-like modifications

at the extremities of the trunk. G. herdmani might

possibly fit in this subgenus. Both Cutler and Murina

now appear to have doubts about the subgenus

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 21

because neither of them in their most recent reviews

of the genus (Cutler & Murina 1977; Murina, 1977)

mention Dushana. Murina (1977 p, 212) now places

G. scutigera (Roule), the type of Dushana, in the

subgenus Golfingia s,s. No previous Australian

record,

Distribution: (1) in Australia: Proper Bay, near

Port Lincoln, South Australia.

(2) elsewhere: Sri Lanka (Ceylon).

Specimens examined and localities: South Aus-

tralia—Proper Bay (part of Port Lincoln at base of

Eyre Peninsula); amongst roots of Posidonia about

25 m from base of low cliffs at point where railway

line to Coffin Bay crosses road to Sleaford Bay. To

find the animals it is necessary ta dig up a square or

sod of Posidonia and pul] it apart. The animals are

well camouflaged. (15) SAM E1149;

(10) SAM E1158; (3) SAM E1143.

Golfingia (Golfingia) margaritacea adetaidensis

Edmonds

(Figs. 33, 39)

Golfingta margaritacea adelaidensis Edmonds, 1956

pp. 302-3, pl. 2, fig. 2.

Location of type: Australian Museum, Sydney,

specimen from Aldinga reef, St. Vincent Gulf,

South Australia.

Description; Specimens long, slender, cylindrical

and rather active when alive; yellow-brown and

tending to glisten. Trunk 60-100 mm Jong, maximum

width 4-7 mm. Posterior extremity usually tapering

to a sharp point, Anterior and posterior extremities

of trunk show no sign of ridging and no tendency to

form caps as in Golfingia herdmani. Body surface

bearing numerous small, rather flat papillae best

developed and most thickly packed on anterior and

posterior regions of trunk. They appear as white,

circular to elliptical structures about 0:1-0-2 mm in

diameter, in a yellow matrix of body wall. Introvert

slender, comparatively short, 20-25 mm long, 2-

3 mm wide and lacking hooks. Anterior extremity of

introvert of living and dead specimens usually

swollen into a bulb-like structure 3-6 mm in

diameter, Tentacles short, digitiform, white and up

to 80 in number. Body wall generally thin.

Two veniral retractors arise close to nerve cord in

anterior third of trunk and two dorsal retractors

more anteriorly near level of anus, Retractors

separate for most of their length. Oesophagus

fastened to ventral retractors by two mesenteries;

intestine long, consisting in one specimen of 34

double spirals. Spindle muscle arising from a

triangular flap of tissue beneath a strong wing

muscle and not fastened posteriorly. Fastening

muscles three or four, One (Fi) arises from body

wall just below left dorsal retractor and connects

with oesophagus. A second (F2), arising near origin

of Fl, connects with last spiral of intestine (and in a

few specimens gives off a short branch to posterior

oesophagus), In two specimens Fl and F2 arise from

same point, F3 (when present) and F4 arise hetween

right dorsal and ventral retractors and connect with

the last and penultimate intestinal whorls.

These specimens differ from Golfingia herdmani

(Shipley) from Proper Bay because (1) they do not

form a cap or ridge al the extremities of the trunk (2)

they lack introvert spines and (3) they lack rusty

coloured, globose papillae on the anterior and

posterior surfaces of the trunk and the ridges on

which they lie, both of which are present in G.

herdmani.

Edmonds (1956 p, 303) says that “a caecum is

lacking or possibly rudimentary’, Recent study of

more specimens shows that a small caecum is usually

present. The specimens are close anatomically to G.

margatitacea (Sars), described from Norway. and

reported widely from the northern hemisphere and

from the Antarctic and sub-Antarctic. Recently

Cutler (1977 p. 139) has reported G. margaritacea

from off the South Australian coast, Whether his

specimen is the same as mine I do not know, Murina

(1977 p. 230) has placed G, mtargaritacea adelaiden-

sis in the synonymy of G. margaritacea mar

garitacea. | find it difficult, however, to lump the

South Australian material with that described by

Theel and Wesenberg-Lund from Norway and

Greenland and that from Alaska described by Fisher

(1952). The spindle musele of my specimens is strong

and arises from a triangular flap of tissue under and

separate from the wing muscle and not from the

rectum as shown in pl. 23 fig. 3 of Fisher (1952).

Théel’s fig. 174 shows only two fixing muscles (there

are three or four in mine) and no staut fixing muscle

runs tO the rectum near the anus, as shown in Théel

(1905) fig. 174. In addition the specimens are always

yellowish-brown in colour,

Previous Australian records: Edmonds (1956),

Distribution: Known only trom Aldinga, Sauth

Australia.

Specimens examined and locality: In sand and

debris amongst the roots of the marine angiosperm

Amphibolis antarctica on the reef at Aldinga Bay,

St. Vincent Gulf, South Australia. (1) SAM E1159,

(2) SAM E1148, (1) SAM E1176, (1) SAM E1177,

(1) AMS W3603.

Golfingia (Golfingia) vulgaris

Edmonds

Golfingia vulgaris queenslandensis Edmonds, 1956

pp. 303-4, pl, 17.

queenslandensis

22 REC, §. AUST, MUS. 18 (1):

Location of type: Australian Museum, Sydney;

specimens from Heron Is,, Queensland,

Remarks: No additional specimens have been

collected since Edmonds’ 1956 record. There is still

doubt about the identity of these specimens, They

possess two pairs of retractors, hooks, three

fastening muscles and a spindle muscle not attached

posteriorly. The introvert is swollen anteriorly and

there is a weakly developed rim between the base of

the introvert and the trunk, much like that found in

Golfingia herdmani. The hooks, however, are

different from those of the latter species,

Distributians; Known only from Queensland,

Specimens examined: (2) Heron Is., Queensland,

AMS W362.

*Galfingia (Golfingia) ohlini (Théc!)

Phascolosama ohlini Théel, 1911 p. 29, pl, 2. figs 21-

23, pl, 3, figs 24-27. pl. 5, figs 69-70,

Golfingia ohlini: Fisher, 1950b p. 550.

Location af type: Not known; type locality South

Georgia,

Description: T have not seen Austrahan specimens

of the species. Théel’s account of the type specimen

reads, “Total length of body 16 or 17 mm, Body

elongate, subcylindrical, with posterior extremity

pointed, Tentacles slender, 16 in number, arranged

in groups on each side of the median line. Distinct

ciliated sense pads present, separating the dorsal

tentacles, Skin whitish, shining with small cylindrical

papillae, crowded at the posterior extremity of the

body and scarce at its middle, Behind the tentacles

mamillary wart-ltke papillae and scattered hooks,

both directed backwardly. Two free segmental

organs. Muscular layers of the body-wall contiau-

ous, not separated into bands, Two ventral and two

dorsal retractors. Intestinal spiral composed of

about 14 double turns and not attached posteriorly,”

Australian record: New South Wales, off Broken

Bay (33° 34°5 8, 152° 06'S E) (1) Murina, 1972 pp.

301-302,

Distribution: (1) in Australia: off coast of New

South Wales.

(2) elsewhere: Antartica and sub-Antartica;

Mauritius and South Africa: Chile.

Subgenus Mitosiphon Fisher

Mitosiphon Fisher 1950b p. 550; 1952 p. 393:

Stephen & Edmonds 1972 p. 113; Cutler &

Murina 1977 p. 180,

Description; Two pairs of retractor muscles,

Introvert hooks (if present) have accessory comb of

1-74 January, TOS)

spinelets at base. Nephridia bilobed. Spindle muscle

nat attached posteriorly,

Type Phascolosoma hesperum Chamberlin,

KEY TO SPECIES QF SUBGENUS MITOSOPHON

REPORTED FROM AUSTRALIA

1, Introvert hooks with4-5 accessory teeth

G_ misakiqna (p. 22)

Introvert without hooks... G. richocephala (p. 23)

(Murina, 1977 has now assigned G. (richocephala to

a subgenus Apionosoma Sluiter, 1902; G. misakiana

has been left in Mitosiphon).

Golfingia (Mitosiphon) misakiana (Tkeda)

Phascolasoma misakianum Ikeda, 1904 pp. 7-9,

pl. 1, fig. 3, pl, 3, figs 30 -33; Fischer, 1919a p, 281;

1927 p. 204.

Golfingia misakiana: Fisher, 1952 p. 393, Cutler,

1973 p. 144; Murina, 1970 p, 66, 1977 p, 236.

Location of type: Not known to author; specimen

from eoast of Misaki Bay, Japan.

Description; Four specimens, two in very good

condition and with introvert extended, were

examined. Specimens small and slender and

introvert almost thread-like. Trunk of largest 11 mm

x 15-18 mm and introvert about 40 mm x (:4-

0-7 mm. Bady wall thin and transparent enaugh to

show most of internal anatomy without dissection.

Mouth surrounded by 5 (26) swellings or protuber-

ances, probably corresponding to tentacles. About

20 complete rows of clear, almost transparent hooks

surround anterior region of introvert, Rows then

became incomplete and ultimately hooks sparsely

scattered, Hooks small, 0-022-0-027 mm tall, with

sharply bent tips and with usually five sharp

accessory teeth near base. More posteriorly placed

hooks smaller and may lack accessory teeth.

Introvert bearing very numerous, small urn-shaped

glandular papillae. Posterior trunk papillae about

0-20-0-25 mm in diameter and rather flat, consisting

of a few rounded plates arranged like the petals of a

flower, In mid-trunk, papillae are larger (up to

0-1 mm in diameter) and the arrangement of plates

less definite. One feature of specimens is that both

trunk and introvert are covered by a very great

number of very small “points’’ (possibly glandular

pores), placed very closely to each other, lying in

both longitudinal and transverse rows and making

the surface appear striated.

The most transparent specimen shows four

retractor muscles, two long, bilabed nephridia and a

posteriorly fastened spindle muscle.

Systematic positian; The specimens are consi-

dered to be G. miisakiana (Ikeda). The question

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SiPUNCULA) 23

arises whether G. misakiai a and G. trichocephala

(Sluiter), which js also being recorded in this. paper

from the eastern coast of Australia, are the same

species, a species with a genetically determined set

of hooks which for some reason concerned with the

animals’s environment may either be lost or never

appear. Cutler (1973 pp. 139-144) discusses the

question, He states that his specimens of G,

Irichocephala and apparently the type specimen

(since he examined it) lack hooks. Murina (1970;

1977) describes G, trichocephala as being hookless.

Neither can | find hoaks on some specimens from

Moreton Bay, Queensland which | have called G.

trichocephala nor on some from Madagascar, sent to

me by Dr. Thomassin, Murina must consider the two

species valid because she described both of them in

her 1977 paper and | think it best to regard them so,

at present. If the two are the same it is remarkable

that specimens with and without hooks have not

been found together in the same collection.

Nevertheless many characters of my specimens of G,

misakiana correspond with those of G. trichocephala

e.g., size and shape, anterior region of introvert and

papillae (Murina, 1977, fig. 162), bilobed nephridia,

tour retractor muscles and even the fine striations on

the surface of the trunk and introvert, Which are

present in both my specimens of the two species.

The figures of the species were drawn from the

specimen SAM E1161. The specimens lack the large

papillae on the posterior surface of the trunk,

present in G. murinae Cutler, 1969

Previous Australian teeord: Western Australia

(Fischer, 19194).

Distribution: (1) in Australia: Western Australia

at Shark Bay, 6-9 m (Fischer 1919a); N.S.W. at

Broughton Is.

(2) elsewhere; Japan; New Guinea, Tanzania and

Easter Is. (Murina 1977 p.. 238).

Specimens examined and localities: N.S.W. at

Broughton I... amongst roots of marine angiosperm

Posidonia (Collet and Pat Hutchings) (2) AMS

W13157, (1) AMS W1299], (1) SAM E1161.

Golfingia (Mitosiphon) trichocephala (Sluiter)

(Figs. 34, 40)

Apionsoma irichocephalum Sluiter, 1902 pp. 42-44,

pl. 4, figs. 8-11; Murina, 1977 p, 236.

Golfingia trichocephala Murina, 1972 p. 303; Cutler,

1973 p. 139; Cutler and Murina, 1977 p. 180.

Location of type: Zoological Museum, Amsterdam;

specimen from near Surdbaya, Java, Indonesia.

Description: Small, white-brown, slender species

with a long, almost thread-like introvert. Trunk 3:0-

S:lmm long, maximum width usually less than

1 mm; curved ventrally in most specimens. Posterior

extremity pointed. Bedy wall thin and internal

structures often visible under strong light. Introvert

45 mm long in specimen with trunk 4-1 mm long,

lacking hooks and tentacles,

FIG.

Moreton Bay, Queensland), Scale measurements are

in mm

34. Golfingia wmichacephala, (specimen from

Longitudinal musculature continuous. Four slen-

der retractor muscles, a ventral pair arising very

close to nerve cord in middle of trunk and a dorsal

pair more anteriorly. Intestine long, coiled and

slender, No rectal caecum observed in 2 dissected

specimens. Spindle muscle fixed posteriorly to body

wall. Rectum attached to body wall near anus by a

strand of tissue. Suspension at this point accounts for

what Culter (1973) described as a “penuflection in

the rectum’’, Contractile vessel inconspicuous. Two

yellow-brown, free, bilobed nephridia of equal

length opening anteriorly to anal aperture.

Systematic position; Apiosoma trichocephalum

Sluiter, 1902 was dredged. at 56 m from Indonesia

(lat, 7° 25’ §, long. 113° 16'E) and described as

possessing nephridia with single lobes, Cutler (1973

p. 138) re-examined Sluiter’s type and found that the

nephridia were bilobed. After a careful study of the

type specimens he concluded that G. tenuissima

Wesenberg-Lund, 1959a and G, longirostris Wesen-

berg-Lund 1959a were junior synonyms of G

trichacephala, Murina (1977 p. 180) removed it froin

the subgenus Mitosiphon and placed it in Apiosoma

which she now regards as a subgenus.

Fischer (19194) reported Golfingia misakiana

(Ikeda) from Shark Bay, Western Australia, G,

misakiana and G. trichocephala are alike in many

respects. The former, however, is described as

wu REC, 5. AUST. MUS. 18 (1):

possessing introvert hooks with accessory teeth, The

specimens in the present collection from Rottnest

Is., like those from Queensland, lack hooks.

G. trichocephala is identified by its small size, a

very long slender introvert lacking hooks and

tentacles, the presence of four retractors and two

bilobed nephridia.

Previous Australian record; off coast of New

South Wales (33° 34’ S, 152° 16’ EB) by Murina

(1972).

Distribution: (1) in Australia: Queensland, New

South Wales and Western Australia.

(2) elsewhere: South Africa and West Africa

(Wesenberg-Lund, 1959a, 19596, 1963); Western

Atlantic (Cutler, 1973); Indonesia (Sluiter, 1902);

Coral Sea and Tasman Sea (Cutler, 1977); Gulf of

Aden (Murina, 1970),

Specimens examined and localities; Queensland

—Moreton Bay (8), dredged by 8S. Cook (Dept, of

Zoology, Univ. of Queensland), SAM E1152; same

locality (5) coll. W, Green (Univ. of Queensland)

SAM E1150 and SAM E1154, Western Aus-

tralia—William Bay, Rottnest Is. (2) AMS W10565.

Subgenus Phascoloides Fisher

Phascoloides Fisher 1950b p, 550; 1952 p. 395:

Stephen and Edmonds, 1972 p. 131; Cutler and

Murina 1977 p. 182; Murina, 1977 p. 180.

Description; One pair of retractor muscles.

Introvert with or without hooks. Contractile vessel

simple and without villi. Spindle muscle, if present.

not atlached posteriorly,

Type: Sipiinculus eremitus Sars, 1851,

(Murina 1977 p. 180 has replaced Phascoloides

Fisher, 1950b by Nephasoma Pergament 1946 on the

grounds of priority, Her argument is that since

Nephasoma marinki Pergament, 1946 = Golfingia

glacialis Danielssen and Koren, 1881 and because

Fisher (1950b) placed G. glacialis in his subgenus

Phascoloides the term Nephasoma is an older name

in the group).

KEY TO SPECIES OF PHASCOLOJDES RECORDED

FROM AUSTRALIA

1. Tentacles absent or reduced to leaf-like structures or lobular

projections... .-. “GG. minuta (p. 24)

SS ee Geet ir pegh te ss fs ROR PIEE ND By

Tentacles numerous and finger-like ... Cockle rare

. Wing muscle well developed and spindle muscle arising from

!Wwostrong roots... L. _ G. schuettet (p. 25)

Wing muscle aot well developed and spindle muscle not

arising from twa strong roots , ."G. pellucida (p. 26)

1-74 January, 1980

*Golfingia (Phascoloides) improyisa (Théel)

Phascolosoma improvisum Théel, 1905 pp. 82-83, pl.

5, figs. 51-58, pl. 12, figs, 177-182, pl. 14, figs.

202-3.

Golfingia improvisa Murina, 1958 pp. 1625-1628,

Stephen and Edmonds, 1972 p. 145; Cutler,

1973 p, 155; Murina,1977 pp, 184-6, figs, 122a-c.

Location of type: Not known to author; type

locality, West coast of Sweden.

Description; Théel's account reads, ‘“Total length

of largest specimen 15 mm. Proboscis about half of

the total length, or shorter. Trunk cylindrical,

slightly tapering anteriorly and behind. Tentacles

absent and replaced by some irregular, rounded

prominences on the oral disc. Skin hyaline and

provided with distinct papillae especially on the

posterior end of the trunk and on the proboscis. A

girdle of hooks always present. Twa ventral

retractors embracing the nerve cord with their roots,

and varying greatly in length. They are attached ta

the body-wall either at the middle of the trunk, or at

its anterior part or else considerably posteriorly.

Two free segmental organs. Intestinal spiral

composed of about 13 or more double turns and not

attached posteriorly. Body cavity never contains

eggs.”

Remarks: This species has also been reported

from New Zealand in the tube of a foraminiferan

(Edmonds, 1976). G. improvisa is closely related ta

G. minuta (Keferstein) and thought to be

synonymous with it by some workers (see under G,

minuia), Murina (1977 pp, 184-8) discusses the

question.

Australian record: Tasman Sea (39° 35'S, 153! 45’

E) ‘in shell of a foraminiferan” (Murina, 1972 pp.

300) at 3.970 m.

Distribution: (1) in Australia: Tasman Sea.

(2) elsewhere: Sweden, Greenland, cast coast of

New Jersey, U,S.A., France, British Isles, north-

west Pacifie, South Africa,

* Golfingia (Phascoloides) minuta (Keferstein)

Phascalasoma pinutint Keferstein, 1863: p, 40, pl.

3, figs 7-10, Théel, 1911 p. 32, pl, 3, figs 42-45,

pl. 4. figs 46-49.

Golfingia minura: Murina, 1957a pp. 994-995- 1956

pp, 1628-1634; 1977 pp. 186-188, fig. 123:

Cutler, 1973 pp. 155-159,

Location of type: Not known to author: specimen

from St. Vaast la Houge, Normandy. France,

Description: A small species, According to

Keferstein trunk about 6 mm long and introvert &

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) rh

mm. Tentacles reduced to two leaf-like structures or

4-6 “lobular projections” (Cutler, 1973 p. 158).

Museulature thin and continuous, Four retractors

arising in posterior third of trunk (Cutler says *‘a

wide variety of locations’). Spindle muscle not fixed

posteriorly. Nephridia short, sac-like, !ree and

opening a little behind anus. Small caceum usually

present according to Cutler. Akesson (1958) says

that G. minuta is hermaphroditic.

Systematics: G, minute is very closely related to

G. improvisa. Cutler (1973 p. 158) thinks that they

are synonymous but Murina (1958 p, 1624: 1977 p.

186) considers them different.

Australian record: New South Wales. off Broken

Bay (33° 34°5 S, 152° 06°5 FE) (5) Murina, 1972 p.

400.

Distribubon: (1) in Ausiralia; off coast of New

South Wales,

(2) elsewhere: Artic Sea. North Sea, western-

north Atlantic, Mediterranean, off South Africa,

north-west Pacific, Chile, and Falkland Is.

*Golfingia (Phascoloides) pellucida (Keferstcin)

Phascolosoma pellucidum Keferstein, 1865 p. 433,

pl. 32, figs 26-27; Fischer, 1919a p. 281.

Golfingia pellucida; Fischer, 1950b p. 550; Cutler,

1973 pp. 159-162.

Location of type: Not known by author; specimen

from St. Thomas, Antilles, West Indies.

Description: According to Keferstein the body is

8-9 times as long as wide, and the introvert is a half

to three quarters as long as the body. Skin rhin and

transparent, often iridescent. with uniformly distri-

buted small, wart-like papillae which posteriorly are

pointed and yellow. Hooks few and irregularly

distributed between the papillae and folds of skin on

the anterior part of the introvert; they are seen only

with the aid of a microscope and are ()-032 mm high

and (1-044 mm long, Tentacles numerous and rather

long.

Musculature thin, continuous and without band-

ing. Two retractors arising in the middle third of the

trunk. Two large eyespots, Intestine of about 14

spirals, without a posteriorly fastened spindle

muscle, Three fasteners to the first spiral. Rectum

short. Contractile vessel not observed. Nephridia

short and free, Length of bady 32-45 mm, of

introvert 15-23,

Several specimens from St. Thomas, in coral at

depth of 2 feet.

Remarks: Judging from the records. of Keferstein

(1865), Selenka & de Man (1883 p. 32), Augener

(1903 p. 299). Fischer (1914 p. 8), Cutler (1973 p.

159) and Murina (1968 p, 421; 1977 p. 196), G.

pellucida is a variable species. Fischer (1919 ap. 281)

reported it from Cockburn Sound, Western

Australia, Unfortunately he gave no information

about the specimens, Recently I have examined twa

other specimens from Cockburn Sound and have

decided that, although closely related to G,

pellucida, they are different from it. I have referred

them to G, schuettet (Augener, 1903), a species

described from Sydney, New South Wales. Whether

G. schuetiei and G, pellucida are synonymous | am

not able to say.

Australian records: Cockburn Sound, W.A.

(Fischer, 1919a); Torres St., Q. (Selenka & de Man,

1883).

Distribution: Antilles, Costa Rica, Jamaica,

eastern coast of South America, Amboina, western

North Atlantic, Philippines, Singapore, Torres St..

Gulf of Mexico, Gulf of Siam.

Golfingia (Phascoloides) schuettei (Augener)

(Figs. 35, 44-45)

Phascolosoma schutiei Augener, 1903 pp. 335-337,

figs 17-18.

Golfingia schuettei: Murina, 1964a p, 238, fig, 12;

1971: 43, 1973 16; 1974 p. 235; 1977 pp. 191-2,

fig. 128; Stephen & Edmonds 1972 p. 1456, figs

18a and b.

Location of type: Not known; specimen from

Sydney, Australia (coll. by Dr. Schutte in

1876).

Description: Specimens light lo golden brown or

grey, usually long and sub-cylindrical. Trunk 33-

160 mm long (three are longer than 100 mm) and 5-

10 mm wide. Introvert Usually not well differenti-

ated from trunk, 10-25 mm long. Mouth surrounded

by numerous digitiform tentacles which are made to

look complexly afranged by radially projecting

extensions of the tissue from which the tentacles

arise, much as is shown in figs, 191-3 of Théel

(1905), Zone of brown to black, sharply pointed

hooks, lies posterior to tentacles. Hooks either

straight or curved slightly at tip and varying in size

and direction, Anterior hooks usually largest,

projecting almost at right angles to surface; posterior

ones smaller, often just piercing surface. Small

papilla-like swellings usually present near base of

cach hook. Field of hooks may be better developed

On ventral surface and in one specimen whale field

much reduced.

24 REC. 5. AUST, MUS, 18 (1):

FIG. 35, Golfingia

schuettei, (specimen from Port

Jackson, New South Wales),

Trunk papillae may be plumply conical, glabose

or short and cylindrical, the first two sometimes

arising from a narrowed base or stalk, Musculature

of body wal! continuous. Two strong cetractors,

separate for part or most of their length, arise in

middle or posterior half of trunk. Contractile vessel

prominent but without villi. Wing muscle strong

Spindle muscle arises anteriorly from two strong

roots attached to body wall on each side of just

anterior to anus. Roots, separate from wing muscle,

run under rectum and join near caecum to form

spindle muscle (much as in G. margaritacea

adelaidensis and G. herdmant), Spindle muscle free

posteriorly. Three to five fastening muscles. Fl,

arising on left side of nerve cord runs to posterior

oesophagus; F2, F3 and F4 (if present) connect with

penultimate intestinal spiral and F5 with rectum.

Nephridia two, usually tubular (sometimes swollen

anteriorly), long free and opening at about same

level as anus. Brain simple with two eyespots.

Gonads at base of retractors. Twenty to thirty

double intestinal spirals.

Systematic position: Tam naming the specimens as

G, schuettet (Augener) with some reservations,

chiefly because I have not been able to compare

them with the type specimen.

It is possible that the specimens, especially those

from Cockburn Sound, W.A., are the same species

1-74 January, 1980

as those from Cockburn Sound which Fischer (1919

p. 281) called G. pellucida (Keferstein, 1865). The

type locality of the latter species is in the Antilles but

Fischer (1919 p. 281) says that it is “circummiun-

dane”, Unfortunately Fischer gave no information

about his specimens. G. pellucida (especially long

specimens) have heen reported from Indonesia,

Malaysia and the Philippines several times, (1) by

Selenka & de Man (1883) from the Philippines,

Singapore (specimens with trunk 80 mm _ long,

introvert 45 mm and without hooks) and Terres St.

(hooks lacking) and (2) by Augener (1903) from

Amboina (specimens 70 mm long). Selenka says that

he was not able to find any real difference between

specimens from the Antilles and Rio de Janeiro and

from the Philippines etc, and gave all them all the

same name.

After comparison with specimens of G, pellucida

from the Western North Atlantic, which were

named and sent to me by Prof, E, Cutler, and some

from Curacao, collected by C. J. van der Horst

(Zool, Mus., Amsterdam) | am inclined to think that

the Australian specimens are different. They are

much larger, cylindrica! rather than flask shaped,

their tentacles are more complexly arranged, their

wing muscle is strong and the spindle muscle arises

either from a triangular flap or two strong roots.

Concerning size, Cutler (1973 p. 159) found that the

trunk of 834 specimens of G. pellucida fram the

Western North Atlantic was 3-25 mm Jong and 1-3

mm wide (most were 6-10 mm long). Murina (1968

p. 422; 1972 p. 302: 1977 p. 192) also reports small

specimens. For these reasons I am reluctant to Jump

my specimens with those from Curagao and the

North Atlantic. A translation of Keferstein’s

description of G. pellucida is given in this paper (see

p. 25)

Phascolosoma schuettei Augener, 1903 was

described from a single specimen collected at

“Sydney” by Dr. Schiitte in 1876. Augener says that

that the specimen was with some others obtained

from the Gottingen Museum, The type however, is

not in the Museum today and there is no recard that

it was ever lodged there. The length of the trunk is

about 70 mm and Augener figures its club-like

papillae and its introvert hooks. Its tentacular crown

consists of numerous, long, yellowish tentacles

which sutround the mouth in a single row. The

single-rowed nature of the arrangement is obscured

by the radially projecting folds of the tissue from

which the tentacles arise (=‘'Tentakelgrundes”’)

much as itis in Siphonosoma cumanense. The species

possesses two retractors which arise in the posterior

two-thirds of the trunk, three fasteners, 63 intestinal

spirals, a caecum and a spindle muscle not attached

posteriorly, There is no doubt that my Australian

4 REVISLON OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA] 7

specimen and Augener’s description closely corres-

pond and it is unfortunate that the type specimen

cannot be found for a comparison, The main

ditference seems to be in the number of fasteners.

Murina (1964a p. 238) records two specimens from

the Tasman Sea (37° 31°S, 163° 59°F) at 1330 mm

and one from the N-W Pacific at 5397 m and in

Murina (1973 pp. 69-70) several from the Peruvian-

Chilean Trench at depths of up to 7 000 m. The

specimens in my collection, however, were collected

from shallow water, twa from Port Jackson (=

Sydney Harbor).

The trunk of G, (Phascalaides) novaecealandiae

(Benham, 1904) may be lang (up to 310 mm) and is

sub-cylindrical; the species seems closely allied to G,

schuettel. The four specimens so far desctibed

however, all lack introvert hooks, Sluiter (1902 p.

35) considered that his G. subhamata, dredged from

two different localities in Indanésia, although closely

related to G. pellucida was different trom it

Recently 1 re-examined one of Sluiter’s original

Specimens, a small one which I found difficult to

work with, Whether G, subhamata and G, sehuettei

are the same T am unable to say.

G. schuettei differs from the other large Australian

golfingiids (G, margaritacea adelaidensis and G.

herdmani) in that it has only two retractors, while

they have four.

Previous Australian records: New South Wales

(Augener, 1903): Tasman Sea (Murina, 19644).

Distribution: (1) in Australia: Port Jackson

(N.8.W.); Cockburn Sound, Port Gregory and

Dampier Archipelago (W.A.); Tasman Sea.

(2) elsewhere: N.W. Pacific Ocean, Peruvian-

Chilean Trench.

Specimens examined: New South Wales—Port

Jackson (1) AMS W3019 “tinder rocks” and (1)

SAM E1171 (coll. P. Hutchings). Western Aus-

tralia—Cockburn Sound (north of Jervaise Grayne)

SAM 141/76, Cockatoo Is. (1) ‘under stones at low

tide” WAM 149/76; Dampier Archipelago (1) coll.

of WAM.

Subgenus Golfingiella Stephen

Golfingiella Stephen. 1964 p. 459; Stephen &

Edmonds, 1972 p. 118; Cutler & Murina, 1977 p.

180; Murina, 1977 p, 240,

Description: Introvert without hooks but with

tentacles or tentacular lobes. Two pairs of retractor

muscles. Spindle muscle attached posteriorly,

Nephridia single-lobed.

Type species: Phascolosoma procerum Mobius,

Remarks; I am following the lead of Cutler (1977

p. 141) in placing G, murinae murinae in this

subgenus. Cutler (1969 p. 214), however, says that 4

few rows of transparent hooks are present on the

introvert. It is difficult, therefore, to see haw the

species falls in Golfingiella, Murina (1977 p. 238),

with some justification. has placed G. mnrinae in

Mitosiphon. Perhaps the status of G. murinae

murinae (= unilobatae) and G. mutinae bilohatae

needs investigation, Cutler and Murina (1977 p. 181)

claim that, because its mouth lies outside the ring af

tentacles, the species should be transferred to

Fisherana (Phascolosomatidae),

*Golfingia (Golfingiella) murinae murinae Cutler

Golfingia murinae murinae Cutler 1969 p. 214 (as

amended by Cutler): 1973 p, 145; 1977 p, 141-2,

figs 4 and 5; Cutler and Murina, 1977 p, 181,

Location of type: U.S.N.M. no. 38247; type locality,

37° 13’N, 68° 40°W, dredged at 4 540 m,

Description: Trunk 1-5-13 mm long, Introvert

four ta six times length of trunk, with few rows of

transparent hoaks (same of which have an accessory

comb of spinelets at base), Large mammiform

papillae at posterior end of trunk. Four retractars

arising from posterior third of trunk. Nephridia free

and sae-shaped, Anus posterior to nephridiapores

Australian record: Cutler, 1977 p, 142.

Distribution: (1) in Australia: Tasman Sea (off

Port Macquarie, N.S.W.) (31° 27°S, 153° 33'E)

dredged at 4530 m (6) (Cutler, 1977 p, 142).

(2) elsewhere: “A deep water species (1 O00-

4750 m) in the north-western Atlantic and in the

north-western Pacific Ocean (3 950 m)" aceording

to Cutler, 1977, off Kenya (3950 m), east of

Seychelles (3 300m), N-W Philippine Trench

(1 000 m), Sunda Trench (2 810-2 990 m), south of

Bali (780 m), All these records from Cutler, 1977,

Subgenus Thysanocardia Fisher

Thysanocardia Fisher, 1950b p. 551; 1952 p. 400:

Stephen & Edmonds, 1972 p. 120; Cutler &

Murina, 1977 p, 181; Murina, 1977 p. 198.

Description: One pair of retractor muscles.

Contractile vessel with branched or unbranched

tubules. Nephridiopore usually opens in front of

anus, No introvert hooks.

Type species: Phascolasoma procerum Mébius,

1875.

Remarks: The position of G. cortaca (Keferstein)

in this subgenus is very uncertain because it

possesses introvert hooks. Fisher (1950 p. 551; 1952

p. 395) placed it in the subgenus Golfingia s.s,

28 REC. S. AUST. MUS. 18 (1); 1-74

Tore re eti ade HMMM ooh

TC eanretic py PION a:

January, 1980

FIGS. 36-43. Figs. 36-38, Golfingia herdmani; 36, anterior region dissected; 37, introvert hooks; 38, posterior region of trunk. Fig. 39,

Golfingia margaritacea adelaidensis; anterior region dissected. Fig. 40, Golfingia misakiana; 41, anterior region of introvert; 42,

introvert hooks: 43, papillae from trunk.

Keferstein’s fig. 23 and his description ‘‘Contractiler

Schlauch .. . , mit mehreren sehr langen, cylindris-

chen Aussackungen” indicates that long tubules are

present. In which case the position of the species in

Golfingia s.s. is also uncertain.

Specimens of neither of the two species reported

from Australia have seen by me. G. coriacea

possesses hooks, G. semperi does not.

*Golfingia (Thysanocardia) coriacea (Keferstein)

Phascolosoma coriaceum Keferstein, 1865 pp. 432-3,

pl. 32, figs. 23-4; Selenka and de Man, 1883 pp.

34-5, pl, 2, fig, 15, pl. 5, figs. 50-53.

Golfingia coriacea Fisher, 1950b p. 551; Cutler, 1973

pp. 160-161; Cutler and Murina, 1977 p. 181.

Location of type: not known specimen from St.

Thomas, West Indies.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 29

Remarks: Murina’s single specimen from South

Australia is small, cream and pyriform, Trunk 7 mm

long and 5mm wide and introvert 3 mm Jong and

1-5 mm wide. Introvert hooks 0-1 mm tong. Two

retractor muscles present and “fluffy” polian

vesicles,

No specimens of this species are in the collection

of the South Australian Museum. The only ones

from St, Vincent Gulf in the Museum which are

small, possess two retractors, fluffy polian vesicles

and introvert hooks are those of Themiste fusca (see

p. 40). The latter is common.

Cutler (1973 p. 159) places G. coriacea in the

synonymy of G. pellucida (Keferstein) on the

grounds that Keferstein's specimen must have been

aberrant.

Australian record: South Australia, near Adelaide

(Murina, 1972 p, 298),

*Golfingia (Thysanocardia) semperi (Selenka and de

Man)

Phascolosoma semperi Selenka. and de Man, 1883 pp.

37-38, pl. 5, figs. 56-59,

Golfingia semperi Fisher, 1950b p. 551,

Location of type: 7; specimen

Philippines.

from Uhoy,

Description! Trunk 28 mm long and slim. Two

stout retractors. Tentacles filamentous but hooks

Jacking, Contractile vessel with numerous tubules.

Nephridia about half as long as trunk, free and

opening anterior to anus. Spindle muscle not fixed

posteriorly.

Remarks; No specimens seen by the author,

Although it has been reported several times the

species seems not to have been re-described,

Fisher’s 1921 account takes up only three lines.

Australian record; Cape Jaubert, Western Australia

(Fischer, 1921).

Distribution: (1) in Australia: Western Australia.

(2) elsewhere: Philippines, Red Sea, Zanzibar and

Fernanda Po.

Genus Phascolion Théel

Phascolion Théel, 1875 p, 13,

Edmonds, 1972 pp. 164-165,

Stephen and

Description: Specimens usually small, living in

empty shells of gastropods or scaphopods, in tubes

of annelids, in burrows in coral rock or in solitary

corals. Body usually spirally twisted and internal

organs asymmetrically arranged. Only one, usually

the right, nephridium present. Usually only one

gonad. Musculature of bady wall continuous.

Characteristic adhesive or holding papillac usually

present on surface of trunk,

Type species: Phascolion strombi (Montagu, 1804)

KEY TO AUSTRALIAN SPECIES GF PHASCOLION

i. Retractor single forall ormostofitslength =... ... 2

Retractors double for allor most of [heir leneth , = a

2. Retractor divided posteriorly near point of fixation into 3

short roots; caecum placed al a considerable distance [rom

anus (half-third of way from anterior end of trunk)

“P pacificwm (p, 32)

Retractor not divided posteriorly: caecum not reported

P, vollare (p, 29)

3. Some papillae on anterior surface of trunk near base ot

introvert With 2-4 poms P. cronullae n.s.p. (p, 30)

Papillae on trunk with single points. * FP. dentalicolien(p. 32)

Phascolion collare Selenka and de Man

(Fig. 52)

Phascolion collare Selenka and de Man, 1883 pp. 45-

46, pl. 6, figs. 71-74; Fischer, 1922 p. 12; Cutler.

1977 pp. 144-5.

Type locality: Philippines; location of type not

known to author,

Description; This description is based on two

complete and two incomplete specimens, One intact

specimen is spirally coiled but the other not.

Estimated length of coiled specimen 14 mm and

width 2:2 anteriorly: of other specimen 9 * 1-7 mm,

Anterior extremity of trunk of coiled animal formed

into a soft cap which is marked off from trunk by a

rim; it is not an aspidosiphonid shield because it is

not hardened and because introvert arises centrally

and not ventrally to it. No shield-like structure in

other specimen, Posterior region of introvert and

anterior trunk bears flask-shaped papillae. Much of

trunk carries numerous small hemispherical or

rounded holdfasts, capped with a brown-black,.

forked structure of variable size, those on outer

(dorsal?) side being largest, Single-pointed part of

forked structure usually directed anteriorly.

Introvert about three quarters as long as trunk,

bearing anteriorly numerous small, blinted hooks.

One long nephridium. [ntestine short and although

convoluted scems not to be coiled. Retractor single,

arising without bifurcation from posterior extremity

of trunk,

Systematic position: There is no doubt that these

specimens from Heterocyathus are a Phascolion. The

single nephridium, the presence of holdfasts on the

trunk and the absence of shields suppert this

conclusion, The shape of the pointed structure of the

holdfasts and the internal anatomy of the animal

correspond to those af Phascolion collare, a species

previously reported from shells. No three or five

0 REC. §, AUST. MUS, 18 (1)-

pronged structures like those described for P. tridens

Selenka & de Man are present in the Western

Australian specimens. The specimens are closely

related to P. robertsani Stephen & Robertson, 1952,

to P. pacificum Murina, 19574 and especially ta P,

ikedai Sato. 1930, reported from the coral

Stéephanacerls,

Distribution: (1) in Australia: north-west Aus-

tralia-

(2) elsewhere: Philippines (Pangola, Ubhoy,

Bohal) in shells of Stronthus, Cerithiun: and Turbo:

Zanzibar in Dentalium shells; Makassar St. (at 2 DOO

m) and Bali Sea (at 1951 m).

Specimens examined and localities; Western

Austraha—5 specimens in colitary coral, Heleracy-

athus sp. dredged at Norbill Bay, Dampier

Archipelago WAM 36-73; alsa 2 specimens from

Heterocyathus from Phillips Beach, Dampier

Archipelago. AMS W5497. One retained in S.A.

Museum (SAM E11&8).

Phascolion cronullae n.sp.

Location of type: Australian Museum, Sydney;

specimen in shell of Gazameda gunni (Reeves)

off Cronulla, New South Wales,

Description; Trunk spirally twisted, about 9-20

mm long and with maximum width 3-4-5 mm.

Introvert about as long as trunk and 1-5 mm wide.

Body wall of introvert and anterior trunk is thick;

that of part enclosed in Shell of molluse is thin.

transparent and often fragile, except posteriorly

Where it is thicker.

Anterior extremity of introvert slightly bulbous

and with small, scattered hooks 0-05-0-08 mm long.

Amernormost hooks are largest and straightest;

more posterior hooks shorter and more recurved.

Small papilla-like swelling present at hase of hooks,

Introvert and trunk covered with papillae varying

considerably in size and shape. Papillae on introvert

smallest, with height greater than width, and with

pointed and extended tips. Papillae on anterior

trunk largest, 0:10-0:35 mm tall and 0-05-0-15 mm

wide, some being urn-shaped and other more

bulbous. Most terminate in a single tip, making

papillae appear mamillate, but others, especially

most anterior, may possess 2, 3 or 4 fips or prongs,

then resembling to some extent the “four-armed

crass” shown for Phascolion convestirum by Sluiter

(1902: 32). Surface of trunk posterior to region with

pointed papillae covered with numerous. closely

packed, rather flat, pale coloured papillae, elliptical

in shape. Thin walled part of trunk bears prominent.

scattered holding or adhesive papillae about Q-1-0-2

1-74 January, 1980

mm tall and 0-15-0-25 mm wide, brown to dark red-

brown in colour and usually lying with their blunted

tips directed anteriorly. Between holding papillae

and over rest of posterior surface lie numerous flat,

pale, hemispherical to elliptical papillae.

Longitudinal musculature continuous. Two retrac-

tor muscles, one usually stronger than other, arise

separately at base of trunk. Alimentary canal

complexly loaped and eniled, with its oesophageal

section fixed to the thinner retractor. Intestinal

loops held in position by fixing muscles arising from

anterior and posterior body wall. Posterior section

ol intestine may be wound into a loose spiral, Rectal

caecum present and from nearby intestinal wall a

pair of fastening muscles run to body wall

(sometimes two strands may fuse before reaching

body wall), Contractile vessel, without villi, attached

to dorsal surface of oesophagus. No spindle muscle

posteriorly. A single nephridium, fixed for most of

its length.

Systematic position: These specimens resemble P.

convestitum Sluiter, 1902 in possessing some

anteriorly placed trunk papillae which possess 1-4

points. I have, however, compared them with

Sluiter’s type and consider them different. Sluiter’s

specimens, both labelled ‘types’, are pyriform and

bottle shaped and not coiled, and lack holdfast

papillae with horny rims. The four-armed cross-like

papillae are very numerous, more slender and

extended than on the Australian specimens. Sluiter

says that the nephridium of his species lies on the left

side of the animal while in the Cronullan specimens

it is clearly on the right. Sluiter does not mention the

presence of a caecum nor of two fasteners that

connect the rectum near the caecum and the body

wall.

The Australian specimens differ from P, strombt

(Montagu, 1804), in possessing anteriorly placed

trunk papillae with 1-4 points. No such papillae are

mentioned or illustrated by Théel (1905) where he

recleseribes the species, nor by Gerould (1913), nor

in specimens collected by me at Roscoff, France

(SAM F1183) nor in those dredged at 43° 03° N, 65°

30” Wand sent 1a me by Dr, FE. Cutler (Univ. of

Utica, U.S.A,),

P. cronullae differs from P, temporariae Edmonds,

1976 [rom New Zealand in the shape of the introvert

hooks and from P. tortum Edmonds, 1976. also from

New Zealand, in the nature and position of the

“ventral’’ retractor,

Commensal: am indebted to Miss E, Pope of the

Australian Museum, Sydney for the following

information. “A polychaete of the family Syllidae

was found in mast of the shells containing a

sipunculan. The polychaete seemed to be entwined

with the Phascolion with its head near a point at the

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 31

(

49 51

FIGS, 44-52. Figs. 44-45, Golfingia schuettei; 44, anterior region dissected (specimen from Cockburn Sound); 45, introvert hooks. Figs.

46-51, Phascolion cronullae; 46, entire specimen; 47, anterior region dissected; 48, introvert hooks; 49-50, papillae from anterior

trunk; 51, holdfasts from trunk. Fig. 52, Phascolion collare; hardened part of holdfast.

32 REC. $8. AUST. MUS. 18 (1);

base of the introvert of the sipunculan. Of 74

randomly selected shells, 10 (13-3%) contained

living molluscs, 46 (62%) contained Phascolion

worms and 18 (24%) were empty. Syllids were

present in 40 (87%) of the 46 tubes that contained

the Phascolion.” Dr, P. Hutchings of the Australian

Museum has identified the polychaete as Syllis

(Toposyllis) armillaris (Mueller).

Symbiotic associations between species of Phasco-

lion and a number of other animals, including the

syllid Langerhansia cornuta (Rathke) have been

previously reported (Stephen and Edmonds 1972 p.

342),

Distribution: Known only from the type locality,

*Phascolion dentalicolum Sato

Phascolosoma dentalicolum Sato, 1937 p, 165, pl. 4,

figs, 20-21, text figs. 10-14; Cutler, 1977 p, 145,

fig. 8.

Location of type: Saito Ho-on Kai Museum, Sendai,

Japan; type locality, Onagawa Bay, north-east

Honshu, at 23 m.

Description: Found in shells of Dentalium sp,

Introvert 15 mm, trunk 15 mm. Small spines or

hooks on introvert. Holding papillae on trunk low

and laterally compressed. Two retractors. Single

nephridium. Nephridiopore posterior to anus,

Rectal caecum present.

Australian record: Great Australian Bight (off

South Australia) 37° 28’ S, 138°55' Eat 1 360 m (18)

(Cutler, 1977). Specimens from small twisted

gastropod shells.

Distribution: (1) in Australia: off South Australia.

(2) elsewhere: North-east Honshu, Japan.

*Phascolion pacificum Murina

Phascolion pacificum Murina 1957b pp. 1777-1781,

figs. 2a, 2b, 3a-e; 1972 p. 306; Cutler, 1977 p.

146.

Location of type: Not known to author; type locality

Kurile-Kamchatka Trench at 6 156-6 860 m.

Description: Introvert with very small hooks.

Trunk thin, small and grey-yellow, Single

retractor with three roots. Caecum situated half

or third of way from anterior end of trunk.

Nephridia fastened to body wall for whole

length.

Australian record: Great Australian Bight (off

South Australia) 37° 28" S$, 138° 55° B at 1 320-

1 340 m (1) (Cutler, 1977 p. 146); specimen in

empty gastropod shell,

1-74 Junuary, 1980

Distribution; (1) in Australian: off South Aus-

tralia,

(2) elsewhere: North-west Pacific Ocean, Peru-

Chile Trench, Antarctica, Pacific and Indian

Oceans,

Genus Onchnesoma

Onchnesoma Koren and Danielssen, 1875 p, 133:

1877 p. 142; Selenka and de Man, 1883 p. 130.

Description; Small sipunculans, usually pear or

club-shaped, Introvert very long and without hooks

and spines. Anus lies on introvert, well forward near

mouth. Tentacles few or absent, Trunk covered with

papillae of varying size and shape. Longitudinal

musculature continuous, Single retractor arising

from posterior of trunk. One nephridium.

Type species; Onchnesoma steenstrypii Koren &

Danielssen, 1875.

*Onchnesoma steenstrupii Koren & Danielssen

(Fig. 8)

Onchnesoma steenstrupii Koren & Danielssen, 1875

p, 133; 1877 p. 142, pl. 15, figs 28-36; Théel,

1905 pp. 93-96, pl. 10, figs 151-152, pl, 11, figs

157-172, pl. 13, fig. 185; Cutler, 1973 pp. 164-

166.

Location of type: Not known to author; specimen

from Kristiansund, Norway.

Description: Cutler (1973 p. 164) says that the

worms have been aptly referred to as “a tiny football

on a string”, Trunk small, pyriform; length -0-2-5

mm, maximum 4 mm and width 0-8-1:3 mm (2 000

specimens sampled—Cutler, 1973). Introvert 5-10

times length of trunk, lacking tentacles and hooks.

Only one introvert retractor, Anus lies well forward

on introvert just posterior to mouth,

Remarks; The species is well known in the north

Atlantic. It is distinguished from Golfingia

irichocephala (Sluiter) by the position of its anus and

the presence of only one retractor and one

nephridium.

Australian record; New South Wales; off Broken

Bay (33° 34°58, 152° 06'5E) (7) Murina, 1972 pp.

164-166, 304-305.

Distribution: (1) in Australia: New South Wales.

(2) elsewhere: Iceland, Scandinavia, North Sea,

North Atlantic, South-West Africa and Mediterra-

nean,

Genus Themiste

Themiste Gray, 1828 p. 8, pl. 6, figs 4, 4a; Stephen,

1964 p. 458; Rice and Stephen, 1970 p. 66;

Stephen & Edmonds, 1972 p. 193.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) a3

Dendrostumum Grube & Oersted, 1858 p. 118;

Fisher, 1952 p. 404.

Dendrostoma Keferstein, 1865 p. 438; Selenka & de

Man, 1883 p, 83,

Description: Trunk usually pearshaped, some-

times elongate and slender; strangly contracted

specimens may be globose, Tentacles surround

mouth and branching; consisting of 4-8 stems which

divide and sub-divide to form pinnately or palmately

arranged tentacules. Hooks or spines may be present

on introvert, Longitudinal musculature continuous.

Retractor muscles two (rarely four). Spindle muscle

not attached posteriorly. Two or more (usually

three) fastening muscles to intestine, Two free

nephridia, Contractile vessel usually well developed

and bearing few to many villi which may be short or

very long. Nuchal organ prominent in some species.

Type species: Themisre hennahi Gray (= Dendros-

tomum peruvianum Collin).

Remarks: Specimens of this genus have been

collected along the coast of all the States of

Australia. T. lageniformis is found in coral, in

clumps of mussels and under stones, T. cymoadoceae

and T. dehamata are found in mud, in sand and in

the roots of marine angiosperms. T, hutoni, T. fusca

and T. variospinosa are often found in burrows in

caleareous rocks or in cracks.

All the known Australian species have contractile

vessels with numerous short villi. Specimens of T.

lageniformis, T. fusca, T. variospinosa, T, huttoni

and T. cymodoceae are usually pyriform, the last two

and especially T cymodoceae tending to be large

and stout. T. dehamata, however, is more slender

and elongate.

1 am considering as one species the large

specimens of T, dehamata collected in New South

Wales (amongst them the type) and some smaller

ones not uncommon in South Australia and

Victoria. I have found it difficult to separate them

other than on the basis of size, The relationship

between them resembles in some Ways that which

Fisher (1952 p. 418) found between T. dyscrita

(Fisher) and T. zostericola (Chamberlin). Eventually

the larger New South Wales and the smaller

southern forms may prove to be two species.

Subgenera of Themiste

On the basis of the number of retractor muscles

and the length of the contractile tubules or villi the

species of Themisre fall into three groups which I

propose be regarded as sub-genera,

1. Themiste (sensu stricto) n. subg.

Diagnosis: Two retractor muscles. Contractile

tubules or villi long and thread-like.

Type species: Themiste hennahi Gray.

Hooks or spines present

I. alutacea (Griibe) T. dyserita(Fishér)

T blanda (Selenka T, hennahi Gray

and de Man)

Hooks or spines absent

T_ hexadactyla (Sato) T_ lissa (Fisher)

T. petricola (Amor) T_ perimeces (Fisher)

T. pyroides (Chamberlin) T. schrnitti (Fisher)

T. rosacea (Amor) T. zostericola (Visher)

T. spinifera (Sluiter)

(hooks sometimes reported to be

lacking in T. aluracea).

2, Lagenopsis n, subg.

Diagnosis: Two retractor muscles, Contractile

tubules or villi short and finger-like.

Type species: Themiste lageniformis Baird.

Haoks or spines present

T. fusca (Edmonds)

T_ hutfont (Benham)

T. minor (Tkeda)

T_ variuspinasa n- sp,

Huoks of spines absent

T. cythodaceve (Edmonds)

T. dehamata | Resteyen)

T. elhiprica (Saw)

T_ fisheri (Amor)

T_ lageniforms (Baird)

T- fropicva (Salo)

T. robertsoni (Stephen

und Rohertsen)

(in T_ robertsoni the tubules are

deseribed us benig Yonly of

moderate lengrh and fairly

numerous"),

3, Stephensonum mm. subg.

Diagnosis; Four retractor muscles.

Type Dendrastomum

Stephen.

Hooks present

species! stephensoni

Hooks absent

T. pinnifalia (Keferstein) T. slephensoni (Stephen)

Remarks: In Themiste s.s. the contractile tubules

are usually few and in Lagenopsis they are numerous

to very numerous. All Themisre species reported

from Australia are members of the subgenus

Lagenopsis.

KEY TO SPECIES OF THEMISTE (LAGENUPSIS) KNOWN

FROM AUSTRALIA

1, Specimens with introvert hooks ..,... 2

Specimens without introvert hooks...,.--. . ..... 4

2. Hooks of almost uniform size..- _. — 5 P

Hooks of markedly different size, which are Very irregularly

directed ..,, . Themisté variaspinusa no sp. (p 42)

3. Hooks few and scattered; specimens usually small (trunk less

than 15 mm), body wall thin and sometimes nearly

transparent, retractors not staut, intestinal spiral rather

loosely wound Themiste fusca (p. 40)

Hooks many and prominent, extending uver most ol (he

introvert and even posterior to the anus; specimens may be

large and stout, body wall rhick. villi of contractile vessel

crowded, intestinal spiral usually tightly wound

Themiiste huttoni (p. 36)

4. Specimens slender, size medivim 1a very Jong: two dorsal

stems of tentacles usually longer than two ventral stems;

tentacles plumose and flecked at their tips, nuchal organ

usually prominent Themiste dehamata (p- 34)

34 REC. S. AUST. MUS, 18 (1):

1-74

January, 1980

FIGS. 53-54, Themiste dehamata, 53, retracted specimen from New South Wales (previously dissected), 54 specimen from Victoria.

Specimens stout rather than slender and pyriform or even

sub-globular in shape; stems of tentacles about equal

size ae)

5. Specimens small to medium in size; tips of tentacles not

pigmented; usually collected in clumps of mussels or in

coral): is 046 co. 4 SEE Themiste lageniformis (p. 41)

Specimens medium to large tn size; tips of tentacles

pigmented; usually collected in roors of marine angio-

sperms ..,. Themiste cymodoceae (p. 35)

Themiste (Lagenopsis) dehamata (Kesteven)

(Figs. 53-57)

Dendrostoma dehamatum Kesteven, 1903, pp, 69-

73, pl. 7, figs 1-6.

Dendrostomum dehamatum: Edmonds, 1956, p. 296,

pl. 1, fig 1.

Themiste dehamata: Stephen & Edmonds, 1972, p.

198.

Location of type: Australian Museum, Sydney;

specimen from Balmoral, Sydney Harbor, New

South Wales.

Re-examination of three specimens: This rede-

scription is based on three of Kesteven’s original

specimens from Balmoral, New South Wales (SAM

E1210) but not the type material.

Specimens very long and slender. According to

Kesteven (1903 p. 69) they were collected on the

beach ‘‘at a time when there had been much rain and

heavy storms”. One cannot then discard the

possibility that the unusual shape of the specimens is

the result of the changes which take place when

sipunculans are placed in hyposmotic solutions,

Fisher (1952 p. 17) reported that three specimens of

Themiste perimeces, washed up on the beach after a

storm, were “unnaturally lengthened’’.

Body sub-cylindrical, slender, curved almost into

a circle, tapering posteriorly and pointed. Trunk of

largest 170 mm long and 7 mm wide posteriorly, Off-

white to grey in colour with posterior region darker

and almost black in one specimen. Skin smooth but

marked into small rectangular areas by fine

intersecting lines. Small, flat to slightly elevated

papillae with small, white, centrally-placed glandu-

lar pores are present in the middle of the rectangular

areas. Posteriorly placed trunk papillae black in one

specimen,

Introvert, not well differentiated from trunk, with

maximum length estimated as 50 mm; grey to yellow

and without hooks. Tentacles plumose, flecked

purple brown and, according to Kesteven, arranged

“in four stems, two of two and two of three primary

branches. In some specimens, however, the division

between the branches extends right down to the

circumoral muscular ridge, in which case there are

eight branches’. The individual tentacules seem

arranged largely in a pinnate manner along the

extended oral grooves (Kesteven, 1903, pl. 6, fig 2).

Nuchal organ prominent.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 35

Twa very long retractors, arising in posterior half

of trunk and separate for most of their length.

Anteriorly, however, they are rolled to form two

halves of a “tube” containing oesophagus. and

contractile vessel. Alimentary canal very long.

Oesophagus runs to or near to base of retractors,

where it is fastened, then loops up anteriorly and

cails into a very Jong intestine with spirals extending

into posterior third of trunk, Rectum very long and

carrying a caecum, Anus and nephridiopores at

about. same level, although in Kesteven’s fig. 1 the

anus is just anterior, Rectal membrane present but

not a5 prominent as that shown for T. fisheri (Amar,

1964, p, 4, fig, B),

There are three main fastening muscles and a

spindle muscle. Fl, arising. at or just posterior to

base of retractors (= mes’ and mes’! of Kesteven fig.

1), may be single or double; if double the roots may

anse at slightly different levels. Fl is attached to

oesophagus fear posterior extremity of contractile

vessel. F2. arising from left side of and near ta nerve

cord, runs to rectum near caccum, and usually

supplies a well-developed series of connectives to

posterior oesophagus near intestinal spirals (=spm'

of Kestevens fig. 1). F3, arising from right side of

nerve cord (more anteriorly than F2), runs to rectum

at about level of caecum (=spm”™ of Kesteyen’s fig.

1), Spindle muscle arising from rectum just posterior

to caecum, traverses whole length of intestinal spiral

but is not fixed posteriorly, Contractile vessel

prominent, bearing yery numerous, short villi (much

like thase of T, lagenifarms, T. cymodoceae and T

huttoni) clustered along its dorsal surface. Posterior

villi longer and forming short branches, Nephridia

slender, free and about one third as Jong as trunk.

Remarks on some smaller specimens; After

considerable deliberation | am referring a large

collection of smaller specimens of Themiste from

South Australia and Victoria to this species. They

differ from the Sydney specimens most notably in

size.

Trunk of largest 75 mm long but mostly 30-50 mm

long and width 3-4 mm. Colour light grey to pale

FIGS. 55-56. Themiste dehamata, tentacular crown of

two specimens; 5S from Victoria, 56 from Sauth

Australia,

pink but anterior region sometimes yellowish.

Mouth surrounded by four groups or stems of

branching tentacles, two larger stems being placed

dorsally between a prominent nuchal organ and two

smaller stems ventrally. Tentacules often dark

purple, arranged pinnately along stems but at

extremities of smaller branches more palmately.

Introvert without hooks. or spines and often not well

marked off from trunk,

Internally as for T. dehamata. Two points of

difference, however, sometimes noticed; (1) the

recta] membrane 1s much more prominent than in

the four specimens described above and (2) there are

some variations in the points of attachment of the

intestinal fasteners. Fl may consist of a membrane

or three roots of different lengths. F2 may connect

with upward loop of oesophagus more anteriorly and

an additional F2a may connect the posterior

ocsophagus and rectum, F3 seems invariable.

Spindle muscle as in Balmoral specimens. Spherical

eggs 0:15-0-18 mm in diameter in some specimens.

Systematic position; I have not been able to

distinguish satisfactorily between the larger speci-

mens of T. dehamata from New Sauth Wales and

the smaller specimens of Themiste from South

Australia, Victoria and Western Australia other

than on their size. Eventually some other worker

may be able io show that they are different, The

relation between the Jarge and small specimens is

much like that between T. zostericola and T, dyscrita

(Fisher, 1952 p, 418),

Themiste fisheri (Amor, 1964) from Argentina is

closely allied to T. dehamata. Amor (1964 p, 469)

considered that the tentacles of her specimens arose

from six stems. While this may be so I contend that

her fig. 3a could be interpreted to show the presence

of four primary stems, her (1 & 2), 3, 4, her (5 & 6).

The main differences between the two species are

that T, dehamata has a caecum while T. fisheri has

not, that Fl and F2 arise from different positions in

the two and that the spindle muscle of T. fisheri

seems to arise more anteriorly, None of these points

is very Significant taxonomically except for that of

the caecum. Previous Australian recard. New South

Wales (Kesteven, 1903)

Distribution; in Australia: New South Wales,

Victoria, South Australia and Western Australia,

Specimens examined and localities: New South

Wales—Balmoral, Sydney Harbor (4) SAM

E1210—part af Kesteven's collection. Victoria—

Western Port Bay (2) SAM E1211 and (1) SAM

E1212; Port Phillip Bay (4) SAM E1222, South

Australia—chiefly amongst the roots of the marine

angiosperms Amphibolis, Posidonia and Zostera”

Outer Harbor (7) SAM E1214 Aldinga reef (8) SAM

E1216 and SAM E1217; Proper Bay, Port Lincoln

3h REC § AUST. MUS. 18 (1);

(10) SAM Ei21&; Coffin Bay (Eyre Peninsula) (3)

SAM E1219; Cape Jervois (5) SAM 61221; Emu

Bay (Kanyarow Is.,) (6) SAM E1213, Sellick’s reef

‘in Toots. of Amphibolis" (S) SAM E1225 Western

Australisa—Cottesloe (3) SAM E1248,

Themiste (Lagenopsis) huttoni (Benham)

(Figs. 60-64)

Phascolosoma huttoni Benham, 1904, p. 307,

Dendrestomum huttoni: Edmonds, 1960, pp, 164-

16S. pl. 3, text figs, 5-6,

Themiste hunoni: Stephen & Edmonds, 1972, p. 204.

? Dendrostoma signifer (in part) Selenka & de Man,

1883, pp. 86-87; Augener. 1903, p, 337; Fischer,

1914, p, 11; Fischer, 1919a p. 282.

Location of type: Not known by author; specimen

from New Zealand.

Introduction; Selenka & de Man (1883, p. 87)

considered a specimen from Sydney, with back-

wardly directed hooks 0-2 mm long, to be ‘'an armed

variety of Dendrostama signifer’ (= T. lagenifor-

mis), The same term was used by Augener (1903, p.

37) for some specimens from New Zealand, by

Fischer (1914), p. 37) for some specimens from New

Zealand and by Fischer (1919a, p. 282) for

specimens from Albany, Western Australia. None of

these atithors discussed the taxonomic problem of

the presence or absence of introvert hooks, Awati &

Pradhan (1935, p. 10), however, in their account of

“D. stenifer’ from India state that feebly developed

hooks are found on the introvert of the younger

stages but that they ‘‘fall off” tn adults, This does nat

appear to be so far T lageniformis in Australia,

None of the specimens in my collection from

Yeppoon, Dunwich and Darwin possess books,

although they range from young to adult animals.

On the other hand all the armed specimens of

Themisre sent from New South Wales and Western

Australia (including some with a trunk 40 mm long

and others that contain eggs and consequently are

adu ts) are heavily armed. At present I take the view

that since all the specimens collected from any one

lovality are either armed or unarmed and not a

mikx_ure of the two they belong to closely related but

different species. If in the future specimens with and

withaut Hooks are collected from the same habitat

the Jecision would have to be reconsidered, Benham

(1904) possibly took this view (although he did not

expicilly state it) when he called the “armed

variety” from New Zealand a new species, D.

hutroni, Cutler (1973, p, 161) says, with justification,

that the hooks of some species of Galfingia may be

lost While it js possible and even likely that some or

all of the hooks may be lost from a specimen ot

1-74 Jariiary, L980

Themtiste for reasons coneerned with its environment

or method of callectian, my. observatons lead me to

the hypothesis that T. lageniformis is one species and

that the armed form is another.

The specimens of Themuiste from some localities

e.g, South Australia, are small and often almost

transparent and the jntrovert haoks few in number,

small, almost uniform in size and distributed over 4

small area of the introvert, Specimens from New

Zealand, Tasmama, Western Australia and New

South Wales are larger and more robust and the

introvert hooks more numerous and larger but,

while still almost uniform in size, they extend over

much of the introvert, even to a point posterior to

the anus. In another group of specimens from

Queensland the hooks are noticeably different;

some are very large and others almost rudimentary,

they are irregularly arranged and point in different

directions, For these reasons [ regard the armed

specimens of Themiste closely resembling the

unarmed T. lageniformis as falling into three groups

and, because any collection fram one locality

contains only one group and not two or more, |

consider each group on morphological grounds to be

a separate species. The specimens from New

Zealand, Tasmania, Western Australia and New

South Wales are T. hutoni (Benham), those from

South Australia are T. fusea (Edmonds) and those

with vartable spines are a new Species, T.

variospinosa.

Description: The following description ot T-

huttoni is based on specimens from New Zealand,

New South Wales, Queensland, Tasmania and

Western Australia.

(1) Trunk of New Zealand specimens 16-35 mm

long with maximum width about 6mm, grey in

colour, thick walled, smooth and pointed post-

criorly, Estimated length of invaginated introvert

about a quarter that of trunk. Dissected introvert

shows that tentacles arise froma number of stems

and that black, posteriorly directed hooks or spines

are present, Hooks almost uniform in shape and size

and their field does pot extend as far as the anal

aperture.

Two stout retractors arising from posterior half of

trunk and separate for most of their length.

Alimentary canal consisting of descending and

ascending oesophageal loops, a long intestinal spiral

and a rectum with a caecum. Contractile vessel

attached to anterior oesophagus and bearing very

numerous, short, finger-like villi, the most past-

eriorly placed of which often form short branches.

Fixing muscles Fl and F3 well developed but F2

usually reduced, Fl arises from body wall anterior ta

and left of point of attachment of left retractor and is

fastened to oesophagus just posterior to termination

Dee

ET SERS to ate a

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 37

FIG. 57. Themiste dehamata, anterior

region dissected (one of Kesteven’s

specimens), c, Caecum; cv, contrac-

tile vessel: f1-f3, fastening muscles; 1,

introvert; in, intestine: n, nephridia-

pore; o. oesophagus; rm, retractor

muscle: rme, rectal membrane: sm,

spindle muscle,

of contractile vessel. F3 attached to rectum near

rectal caecum. F2 represented by a number of

muscular threads arising from upward loop of

oesophagus which conyerge and join rectum near

caecum. Spindle muscle, arising anteriorly from

rectum, not attached posteriorly. Wing muscle

present. Two free tubular nephridia, opening just

posterior to anus. Two eyespots near brain and a

nuchal organ lying at dorsal extremity of introvert.

Papillae on body wall small, flat with a small white

pore centrally, Eggs spherical, 0:17-0-19 mm. in

diameter. Skin at posterior extremity traversed by

fine lines.

(2) Trunk of Western Australian specimens 10-

28 mm long and with maximum width 2-5-4-4 mm,

curved inwards on the ventral side and with

posterior extremity bluntly pointed. Tentacles as in

New Zealand specimens but pigmented at their

base. Introyert hooks dark brown to black, most

pointing posteriorly, The field of hooks in all seven

specimens. however. extends right round the

introvert and well posterior to the anus (something

very unusual in a sipunculan). Fl fixes oesophagus

to body wall at a point between base of retractors.

F2, F3, caecum, spindle muscle, nephridia and

eyespots like thase of New Zealand material.

(3) Specimens from New South Wales larger:

trunk 12-55 mm long and maximum width 7-11 mm.

Colour predominantly grey but with marked

yellowish tinge. Armed area of introvert large but

not extending posterior to anus as in specimens from

Western Australia. Fl arises between retractors just

to left of nerve cord. F2 not reduced, arising from

body wall anterior to base of left retractor and

running to posterior oesophagus: it may connect

with rectum near the caecum. F3 as in New Zealand

specimens. Hooks, diameter of base 0-15-0-25 mm

and length 0-15-0-22 mm.

Systematic position; This species differs from T.

lageniformis, T. cymodoceae and T, dehaniata in the

possession of introvert hooks. It differs from T,

fusca which is smaller and in which the bady wall is

thinner, the nuchal organ more prominent, and the

intestine shorter and more loosely wound. It differs

from T. variospinosa in which the introvert spines

vary very much in size, shape and orientation.

Previous Australian record: Sydney, New South

Wales (Selenka & de Man, 1883).

Distribution: (1) in Australia: New South Wales,

Tasmania, Western Australia and Queensland.

(2) elsewhere: New Zealand.

Specimens examined and localities: New Zealand

—Stewart Is., (2) SAM E1228; east of Otago (1)

trawled on ‘‘Tacaroa” SAM E1229. Tasmania—

Coles Beach, near Devonport (1) SAM E1234;

38 REC. 8, AUST. MUS. 18 (1):

Jacobs Boat Harbor (1) SAM E1235, Western

Australia—Trige Is., “in limestone reef (7)

SAM E1233 and (2) WAN 242/76; Cockburn Sound

(2) WAM 32/73, New South Wales—Coffs Harbor

(4) SAM E1236; Bottle and Glass Racks (2)

SAM E1237; Long Reef (1) SAM E1238. Queens-

land—Coloundra (6) “on exposed rocky shore”

SAM E1239,

Themiste (Lagenopsis) cymodoceae Edmonds

(Figs 58 and 63)

Dendrostoma cymodoceae Edmonds, 1956 pp. 297-

301, figs 15-16; Akesson, 1958, pp. 147-151 figs 46-

67 and 219-222.

Themiste cvmodoceae Stephen and Edmands, 1972,

pp. 197-198. fig, 102,

Location of type: Australian Museum, Sydney;

specimen from Aldinga Reef, South Australia,

amongst roots of the marine angiosperm Amphibolis

(= Cymodocea) antaretica.

Description: Adults large, robust and elongate to

pyriform in shape; trunk 50-90 mm long with

maximum width of 15-25 mm posteriorly, Colour

light to dark grey with tendency to yellow brown

anteriorly; posterior region usually darker. [Introvert

cylindrical, relatively short, about 5-8 mm_ in

diameter and lacking hooks and spines. Surface. of

introvert just posterior to tentacles smooth, pale and

shiny or sometimes purplish and glossy. A light

brown or yellow band often surrounds introvert near

its middle or towards base. Trunk of fixed specimens

tends to curve inwards slightly on ventral side, Body

wall smooth, although covered with numerous flat,

white papillae, 0-04-0-08 mm in diameter, each of

which bears a white glandular opening at its centre.

Body wall thick, especially posteriorly. Longitudinal

and circular musculature continuous but internal

coelomic wall traversed by oblique muscle striations.

Tentacles arising from four primary stems any of

which may subdivide so that tentacles sometimes

appear to be in 4-8 stems. Subsequent division or

redivision of stems gives rise to small, finger-like,

straw to purple-brown pinnately arranged ten-

tacules. Tip of tentacules always darker than

remainder. Tentacles (= stems and tentacules),

relative to length of trunk seem smaller than those of

T. lageniformis and are more bushy.

Internal anatomy resembling that of T. lagenifor-

mis. Two long, stout retractors fixed in posterior

third of trunk. Oesophagus thin-walled, running

between retractors as far as their base and then

looping sharply upwards to a point near rectum;

intestine Jong and spirally coiled. Rectum short and

carrying a caecum. System of intestinal fasteners

variable. Oesophagus always fastened near base of

4-74 January, 1980

MaTRICY

FIG. 58. Themiste cymodoceaé, (specimen from Aldinga

Bay, South Australia).

retractors by 1-4 short threads (F1), usually arising

between bases of retractors on left side of nerve

cord, F2, arising near left retractor and connecting

with upward loop of oesophagus near the first spiral,

may Or may not be present, F2 may bifurcate or an

additional fastener (F2a), arising close to F2, may

also connect with oesophagus, In a few specimens F2

may connect solely or in part with Jast or

penultimate intestinal spiral. F3, present in all

specimens, is shorter and stouter than F2, arises an

right side of nerve cord in anterior half of trunk and

connects with last intestinal spiral or rectum where,

latter joins the intestine. Spindle. muscle strong,

arising anteriorly from beneath a rectal membrane,

but not attached posteriorly. Contractile vessel

attached to oesophagus as far as base of retractors

and bearing very numerous short sometimes

branching villi, which are larger and very densely

clustered posteriorly. Nephridia thin, brown in

colour, free, about one third as long as trunk and

opening at about level of anus or usually just

posterior to it. Gonads at base of retractors. Nuchal

organ not as prominent as in T, dehamata.

Systematic position: T. cymodoceae is closely

related to T. lageniformis, Although in 1956 I

admitted that the two might be extreme varieties of

the same species J now consider them different, T,

cymodoceae grows to a larger size, its tentacles are

more bushy and plumose and are pigmented at their

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA)

FIG. 59 Themiste fusca, (specimen from South FIG. 60. Themiste huttoni, (specimen from Long Reef,

Australia). New South Wales).

FIG. 61. Themiste lageniformis, (specimen from Curtis FIG. 62. Themiste variospinosa, (specimen from Queensland).

Is., Queensland).

AO. REC, §. AUST, MUS, L8 (1):

tips and there is usually greater variation in the

arrangement of its intestinal fasteners. There is,

however, little difference in the internal anatomy of

the two species. At Aldinga reef, St. Vincent Gulf,

South Australia, the type locality, T. cymodaceae

lives amongst the tangled roots of Amphibolis

(= Cymodacea) antarctica and the sand and debris

contained in them. It lives in a cavity which is

formed as the animal grows in the roots. It seems ta

he almost sedentary; it scarcely moves and shows

little inclination to burrow when it is placed on sand

in an aquarium. Amphibolis normally lives below the

level of inter-tidal reefs. Patches af it occur,

however, on the outer edges of reefs which are well

washed by waves and where there are tidal runs and

drains. J have not found the species in burrows in

rocks. Themiste fusea, however, occurs in burrows

on the same reef at a higher level near the shore.

Although [ have collected about 400 specimens of T,

cymodoceae at this locality | have not found any

which earry introvert hooks, The specimens of T.

lageniformis which T have identified from Australia

have been associated with mussel clumps or coral

rock,

The relation between T, cymodoceae and T.

lageniformis is very much like that between T.

zostericala and T. dyserita as described by Fisher

(1952 p. 418),

Akesson (1958) made a careful study of the

nervous system of T. cymodoceae and Edmonds

(1957a & 1957b) studied its respiration and

excretion. It is a good experimental animal in that it

is very hardy: it Seems reluctant. however, to feed in

the laboratory. Previous Australian record: South

Australian (Edmonds, 1956).

Distribution; Known only from the shores ot St.

Vincent Gulf, South Australia.

Specimens examined and localities: South Aus-

trittia—Aldinga reef (just north of Sellicks Beach),

St. Vincent Gulf, “among roots of the marine

angiosperm Amphibolis (= Cymodacea) anrarctica

(Labill,) Sonders & Ascherson”’, about 50 specimens

comprising SAM E1191, E1193: Cape Jervis (2)

SAM E1104.

Themiste (Lagevopsis) fusca (Edmonds)

(Figs. 59, 65-67)

Dendrostomum fuscum Edmonds, 1960, pp. 145-1067,

hes 7-9, pl. 3.

Themiste fusca: Stephen & Edmonds, 1972, pp. 200-

201,

Lovation of type: Australian Museum, Sydney;

specimen from Proper Bay, near Port Lincoln,

South Australia, Specimens burrowing in a

calcareous, intertidal reef,

1-74 Janhary, T98T

Description; Specimens usually small, Trunk

rarely as Jong as 20 mm and usually less than 15 mm;

white, cream or light brown in colour. Body wall

thin and sometimes. semi-transparent. Introvert

short, Tentacles white and branching, sometimes

flecked with brown and arising from four primary

stems, Introvert hooks black, blunt, posteriorly

directed, 0:05-0:10 mm long, few in number and

scattered. Numerous very small, circular, pale

coloured papillae, Jargest ancl most prominent at

base of introvert and on posterior trunk. Skin

usually smooth but sometimes furrowed into small

square or rectangular areas.

Two slender retractors arising in posterior region

of trunk. Oesophagus fastened to body wall at base

of retractors, then looping anteriorly as in T-

lageniformis and T. hurtoni. Contractile yessel with

fewer and simpler (less branched) villi. Intestinal

spiral Joosely and sometimes irregularly wound

consisting of 6-10 double spirals, Rectal caecum

small, Spindle muscle arising anteriorly fron rectum

but not fixed posteriorly. Two nephridia short, free

and with external openings just posterior to anus.

Reetal membrane present. Fastening muscle FI

cannects base of oesophageal loop ta body wall at

point bear base of left retractor and F3 holds last

spiral of intestine. F2 may be complete, reduced or

absent; if complete, it holds. posterior oesophagus

and if reduced it joins only the posterior oesophagus

and posterior intestinal spiral and not the body wall.

Gonads at base of retractors. Two eyespots, Nuchal

organ present,

Systematic position: This Species is closed allied to

T. huttoni (Benham) and T. minor (Ikeda), the latler

from Japan, It differs from T, hunoriin thatin fusca

(1) the specimens are smaller (2) the body wall is

much thinner and often nearly transparent (3) the

hooks are smaller, fewer in number and more

scattered in their distribution (4) the retractors are

weaker (5) the villi are fewer (6) the intestinal cai) is

shorter and more loosely wound and (7) the nuchal

organ is more prominent.

The smallest specimens are not easily disting-

uished externally from some of T. minor (Ikeda)

kindly sent to me from Matsuyami, Japan by Dr. O.

Ochi (Ehime University), The Japanese specimens,

however, ate smaller, their system of fastening

muscles and their contractile vessel seem simpler.

Both T. cymadoceae and T. fusca are found at

Aldinga and Sellicks Reefs, South Australia, the

former amongst the roots of the angiosperm

Amphibolis on the lowest levels of the reef and

subtidally, and the latter in burrows in the

calcareous rack at higher levels near the shore_

Previous Australian records: South Australia

(Edmonds, 1960).

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 41

Distribution: (1) in Australia: South Australia.

(2) elsewhere: New Zealand.

Specimens examined and localities: South Aus-

tralia—Point Wittelbee, Eyre Peninsula (15)

SAM E1240; Coffin Bay (16) “in calcareous reef

between tide levels’** SAM E1241; Kellidie Bay

(near Coffin Bay) (7) “in limestone reef”; Port

Willunga and Aldinga (both south of Adelaide) (30)

SAM E1243; Encounter Bay (in limestone reef in

front of Yilkie Post Office) (20) SAM E1249; Proper

Bay, near Port Lincoln “burrowing in limestone reef

exposed at mid-tides’’ (8) SAM E1242.

a orl

_ ~sS

Themiste (Lagenopsis) lageniformis Baird

(Figs 61, 70)

Themiste lageniformis Baird, 1868, pp. 98-99, pl. 10,

fig. 3; Rice and Stephen, 1970, pp. 66-67, pl. 3,

figs 12-13; Stephen & Edmonds, 1972, p. 205.

Dendrostoma signifer Selenka & de Man, 1883, pp.

86-87, pl. ti, pl. 11, figs 163-164.

Dendrostomum signifer: Edmonds, 1956, p. 297, pl.

1, fig, 2.

Location of type: British Museum (Nat. Hist.,

London); specimen from Australia.

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FIGS. 63-70. Fig. 63, Themiste cymodoceae; dissected specimen. Fig. 64, Themiste huttoni; introvert hooks, Figs. 65-67, Themiste fusca;

65, introvert hooks: 66, papillae on trunk (figs. 65 & 66 to same sale); 67, dissected specimen. Figs. 68-69. Themiste variospinosa;

68, introvert hooks; 69. dissected specimen, Fig, 70, Themtiste lageniformis; branching contractile villi.

42 REC. §. AUST. MUS, 18 (1);

Description: Size and shape of specimens

depending largely on whether or not they had been

relaxed before fixation, Contracted specimens stant.

pear-shaped or sub-spherical, relaxed ones longer

and less stout. Trunk 5-33 mm long with maximum

width in posterior thicd, Besh-coloured, light to dark

grey or light to dark brown, Most tend to curye

inwards on ventral side, Body wall appearing

smooth although smalj, white, flat glandular pores

present. Surface of unrelaxed specimens often

divided into small rectangular areas by fine lines or

ridges, Posterior extremity rounded or pointed,

Introvert shart, 3-10 mm Jong, often darker in

colour than trunk and often ringed anteriorly with a

dark blue to black band. Tentacles white and not

pigmented or flecked like those of T. dehamata and

T. cymadoceae; arising from 4 primary stems any

one of which may divide into two.or more stems, the

latter branching in turn to form tentacules,

Consequently, if subdivision is near base of primary

stem, number of stems may appear ta be 5-8.

Tentacules finger-like and relative ta body length

longer than those of T, dehamata and T.

cymodoceae, Four primary stems form prooves along

which food moves to mouth, Region of introvert just

posterior to tentacles forming a smooth collar; a

pigmented ring, if present, lying posterior to collar.

All specimens hsted below without introvert hoaks,

although collected at different times by different

people.

Two very Stout retractor muscles arising in

posterior third of trunk. QOesophagus runs to base of

retractars Where it is held in position by a short,

sometimes double, fastener (Fl) and then loops

anteriorly. Intestinal coil a well developed spiral;

rectum short and carrying a caecum, In most

specimens a second fastener (F2), arising near point

of attachment of left retractor, is fixed to

oesophagus near first intestinal spiral. F3, arising

near right retractor, fixed to last or second to Jast

spiral. Stout spindle muscle, arising anteriorly from

beneath a rectal membrane, not fixed posteriorly. A

tubular contractile vesse] with numerous. finger-like

villi extending along the oesophagus to base of

retractors: villi may form short branches near base of

retractors but rarely extend a long upward loop of

oesophagus. Two free, tubular nephridia, about one

third as long as trunk, opening near anus or just

posterior to it, Gonads at base of retractors;

spherical eggs with diameter 0-18-0-20 mm. Nuchal

organ not as prominent or as noticeable as in T,

dehamata and T_ cymodoeceae Brain with two

eyespots.

Systematic position: T. lageniformtis, probably

better known as JT. signifer, isa simall to medium

sized species, pytifarm to sub-globose in shape and

L-74 Jankary, Ls

often curved inwards on its ventral side. It lacks

hooks and its tentacles are not pigmented at their

tips, The body wall is usually stout and thick, A dark

blue-black band may ring the introvert anteriorly,

Previous Australian records: Queensland

(Edmonds, 1956), Cape Yark (Selenka & de Man,

1883). “Varieties” with hooks reported from Sydney

(Selenka & de Man, 1883), from Tasmania (Fischer,

1914) and Albany, Western Australia (Fischer,

1926) are being referred to other species, Baird's

1868 record is from ‘Australia’.

Distribution: (1) in Australia: Queensland,

Northern Territory and Western Australia.

(2) elsewhere: widely in the Indo-Pacific Regian;

West Africa, South Africa, Madagascar, India, Gulf

of Manaar, Amboina, Japan, Philippines, New

Zealand and Hawaii (SAM E1197).

Specimens examined and localities; Queensland

—Dunwich (20) ‘in clumps of mussels, in front of

the cemetry’ SAM E1196 and (12) SAM E1207;

Myora (10) SAM E1200; Cape Cleveland (2) SAM

E1201; Cardwell (1) SAM E1202; Geoffrey Bay,

Magnetic Is. (6) SAM E1203; Wellington Pt.,

Moreton Bay (8) SAM E1205; Wistari Reef,

Capricorn Group (2) call, of Prof. W. Stephenson;

Yeppoon (5) coll. of Prof, R, Kenny; Rat, Is., near

Pt Curtis (7) AMS W2760. Northern Territory

—Lee Point, near Darwin “in coral” (4) SAM E1208

and (4) SAM E1209. Western Australia—Broome

(8) coll, from Hamburg Museum,

Themiste (Lagenopsis) variospinosa n-sp.

(Figs. 62, 68, 69)

Location of type: Australian Museum, Sydney;

specimen from “coral clumps", St. Helena Is.,

Moreton Bay, Queensland.

Description: This species is distinguished from T,

huitoni, which it closely resembles, by the spination

of its intravert hoaks.

Trunk 9-27 mm long and maximum width 4-7 mm

in posterior third, with posterior extremity bluntly

pointed and with surface smaoth, grey in some and

pinkish brown in others, Small flat, circular papillae,

with a very small clear or white pore at centre,

almost uniformly distributed over surface, Tentacles

branching and arising fram 4-5 stems (shown in two

well-relaxed specimens). Nuchal organ present

between two stems of tentacles on dorsal side,

Introvert hooks sharp and black; those most

anteriorly placed largest, 0-15-0-30 mm long, with

most points directed posteriorly, Hooks most

posteriorly placed smallest, 0:02 mm long, with rips

directed anteriorly, posteriorly or at mght angles ta

surface (hence name variospinosa).

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPLINCULA) 43

Internal anatomy much like that of T. hutioni.

Two strong retractors, arising in posterior half of

trunk, Alimentary canal consisting of downward and

upward loops of oesophagus, a spirally coiled

intestine and rectum, Contractile vessel attached to

downward loop of oesophagus and bearing villi not

as densely packed as in T. hutioni, Posteriorly placed

villi forming short. branches and extremities of villi

sometimes expanded into a small knob. Fastening

muscle F1 arising to left of base of left retractor (as.it

does sometimes in T. huttoni) and connecting with

oesophagus just posterior to extremity of the

contractile vessel, F2 arising more anteriorly than Fl

and connecting with posterior oesophagus and

rectum. F3 short and strong. Spindle muscle nat

fixed posteriorly, Rectal caecum present. Nephridia

twa, free, tubular and arising just posterior to anus,

Systematic position: In the retracted state these

specimens look very much like a Golfingia. Some

expanded specimens, however, show that the

tentacles arise from at least four stems each of which

divides and subdivides to form tentacules. The

species is closely related to T. huttaniand T, fusea. It

differs from both in the nature and arrangement of

its introvert hooks.

No previous Australian record,

Distribution: in Australia; only from the type

locality (St. Helena Is., Moreton Bay).

Specimens examined and localities: Queens-

land—St. Helena Is., Moreton Bay “in coral

clumps” SAM E1245 and same locality (5) SAM

E1246.

Family Aspidosiphonidae and key ta genera

Aspidosiphanidae Baird, 1868, p. 100: Stephen

and Edmonds, 1972. pp. 215-216

Description: Rather small sipunculans with either

a hardened, boroy, usually brown shield or a white.

caleareous cap clearly marked off on anterior region

of trunk, A caudal shield may be present at posterior

extremity of trunk. Introvert lies ventral to shield or

cap in all genera, except Cloeasiphon, where it arises

from centre of cap, Longitudinal musculature

continuous or in bundles, the latter sometimes

anastamosing considerably. Two nephridia.

Type genus: Aspidosiphon Diesing. 1851.

KEY TQ GENERA OF ASPIDOS([PHONIDAE KNOWN

FROM AUSTRALIA

1, Amerior (anal) shield or cap present and posterinr (caudal)

shield as Well 20... oe, pisses Pate pt annaee

Anterior (anal) shield or cap present but no posterior

(calidal) shield-- -.... -.... een, 8

2. Longitudinal musculature of trunk wall forming bundles-

which may anastomose considerably

Paraspidosiphon (p. 49)

Longitudinal musculature of trunk wall continuans

Aspidosiphan (p. 43)

3. Longitudmal musculature of trunk wall continuous (anterior

tap rounded, caleareous and pineapple-like). Introvert

from middle of cap ---,.,,,,. -. , - Cloeosiphon (p. 54)

Longitudinal musculature of trunk wall forms bundles, which

thay anastomose. Anlerior cop has shape of # truncated

cone. Introvert arising ventral ro cap Lithacrosiphon (p, 54)

Remarks; Stephen (1964, p. 457) split off from

Aspidosiphon a new genus, Paraspidosiphon, in

which the longitudinal muscles are grouped into

bundles, sometimes with much anastomosation.

Stephen regarded the condition of the longitudinal

muscles of considerable taxonomic importance in

the phylum. Stephen and Edmonds (1972) followed

Stephen's lead. Neither Cutler (1973, p. 173) nor

Murina (1975b, p. 1748, 1752), however, considers

that Paraspidosiphon is justified at the generic level,

each preferring to regard. Aspidosiphon as consisting

of two subgenera. Aspidosiphon (sensu stricta) and

Paraspidosiphon,

Centrosiphon (Shipley, 1903), which Stephen and

Edmonds (1972) included in this family, is being

omitted because it now seems to me that C.

herdmani, the type of the genus, is a galfingiid, The

taxonomic position of the species 1s, however, still

uncertain.

The tentacular arrangement in Aspidosiphon and

Paraspidosiphon as stated by Stephen and Edmonds

(1972, p, 217) is naw known to be wrong, Gibbs

(1977) pointing out that the tentacles lie in a near

ring dorsal to the mouth, as in Phascalosoma (see

remarks on p. 8), | am not certain how the tentacles

are arranged in Cloeosiphon and Lithacrosiphon,

Genus Aspidosiphon Diesing

Aspidesiphon Diesing, 1851, p. 67 (in part)s

Stephen, 1964, p. 457,

Description: Anal and caudal shields usually

prominent and longitudinal muscles of body wall

continuous and not separated into bundles, Anal

shield may be circular, elliptical or horseshoe

shaped, darker in colour than the trunk, with its

surface sometimes furrowed, groaved or covered

with hard, irregolarly arranged granules. Caudal

shield may be flat, hemispherical, conical or

truncate; it may or may not be furrowed or grooved.

Sometimes the caudal shield is not strongly

developed, Introvert arises from the yentral side of

the anterior shield, Tentacles digitiform and

arranged in a horseshoe-shaped ring dorsal to mouth

(see foot-note on p. 8). One or two retractors. Two

nephridia. Contractile vessel simple-

Type species: Aspidesiphon muelleri Diesing,

1851.

44 REC. 8. AUST, MUS, 18 (1):

KEY TO SPECIES OF ASPIDOSIPHON FOUND IN

AUSTRALIA

1. Introvert hooks—all with single points ., Be

Introvert hooks—some or all with two points tevernrt ned

2. Hooks with long, bent points . TA. ealrigustus Cp: 4b)

Hooks without long, bent points - a Le ay ee.

3. A number of knobby papillae on anterior pate of anal shield

modified io form spine-like structure , A. inquilinus(p 47)

No spine-like structures on anal shield; tendency forcircular

musculature of body wall to be striated . A, gracilis(p, 46)

4, Surtace of anterior shield furrowed ar nbbed --- 0 ..,,..,,5

Surface of anterior shield not furrowed

A. elegans elegans (p. 44)

. Anterior shield with about 12 furrows, posterior Shield with

wenker striations (well known inhabitant of solitary

corals) —. ‘ _ A. Jukesin(p. 49)

Antenor shield with: 5.6 inrzasws, posterior shield with 24-

2b TEE OCEETE Pio hem ae iat A. harmeyeri (p. 47)

A. elegans elegans and A- gracilis inhabit coral

reefs and A, hartmeyeri limestone reefs. A. Jukesit

lives in the solitary coral Heteropsammia and A,

inquilinus inhabits the shells of Dentalium and other

molluscs,

Aspidosiphon elegans elegans (Chamissa and

Eysenhardt)

(Figs. 71-76)

Sternapsis elegans Chamissa and Eysenhardt, 1821,

p. 351 pl. 24, figs. 5a-e.

Aspidosiphon elegans Griibe 1868a, pp. 645-647, pl.

8, fig. 5-Sa-b; Selenka and de Man, 1883, pp.

124-6, pl. 1. figs. 10-10a, pl, 14, figs. 205-208;

Sato, 1935, p. 316; Sato, 1939, p. 426-7

Aspidosiphon exilis Sluiter, 1886, p, 497, pl. 3, figs.

11-12.

Location of type: Not known to author; type

locality, “small islands of the Pacific Ocean”.

Description; Slender, delicate, pale to straw

coloured and coral dwelling. Trunk more or less

cylindrical, 8-14 mm long and 1:7-2:8 mm wide, thin

walled and fragile and sometimes slightly curved so

that ventral is slightly shorter than dorsal side.

Longitudinal musculature continuous,

Anal shield prominent, elliptical, dark brown

(although golden around edges) and usually warty,

consists of brown polygonal plates placed closely

together and usually lacking furrows (although edges

may be ridged to some extent), Caudal shield more

weakly developed and sometimes almosi insignific-

ant, usually hemispherical with chitinous structures

around its periphery; furrows, if present are only

marginal.

Papillae on trunk flat and usually restricted to (1)

a narrow region just pasterior to anal shield and (2)

a narrow region at posterior of trunk, consisting of

1-74 January, 1980

ageregates of flat polygonal plates grouped around a

small circular opening. Caudal shield of sorne

specimens consists of structures resembling closely

set papillae.

Introvert of fixed specimens about as long as or a

little longer than trunk. slender, thin walled, arising

ventrally to anal shield and bearing anteriorly a few

tentacular-like structures; armed with 15-30 rows of

small dark hooks with two points, Clear area inside

hook usually possesses a small, backwardly directed

tongue or extension (slight variations of the angle at

which the hagk lies on a shde can cause considerable

differences in its appearance), Hooks in most

posterior rows of some specimens are smaller and

lack internal markings—whether they are develop-

ing hooks or not is not known.

Dark, single pointed, closely set spines of varying

size and shape he on introvert posterior ta haaks.

their field extending to its base; spines (1) are larger

and more closely set an dorsal surface of introveri

and (2) beconie larger the more posteriorly they are

placed. In same specimens, where spination is

heavy, spines arise from strang bases or rooting

Processes and clearly possess a central, tubular

canal,

Retractor muscle, arising [rom body wall well in

front of caudal shield, consists of two short roots,

Intestine loosely wound and frail. Rectum long and

lacking a caecum. No intestinal fasteners observed,

Two thin and delicate nephridia, about half as long

as trunk, are fixed to body wall for most of their

length, Spindle muscle fixed posteriorly.

Systematics and discussion; There is a group of

aspidosiphonids in which the longitudinal muscula-

ture is continuaus, which possess introvert hooks

with two points, well developed introvert spines wilh

Single points (often with strong rooting processes)

and which bear small truncated papillae on the

introvert. They possess few or no papillae on the

trunk except near the anal and caudal shields and the

papillae are flat and consist of polygonal plates.

They possess a single retractor muscle that arises

from two roots which are fastened to the body wall

well anterior to the caudal shield, Their nephridia

are about half as Jong as the trunk and are fixed to

the body wall for most of their length. Some of these

species are Aspidosiphon spinalis Ikeda, 1904 (from

Japan and Jaluit {s.), A. spinosus Sluiter, 1902 (fram

Indonesia), A, homoniyarius Johnson, 1964 (from

west coast of India), A. exilis Sluiter, 1886 (from

Indonesia), A. carolinus Sato, 1935 (from West

Caroline Ts,), A. hrocki Augener, 1903 (fram

Malaya), A. elegans elegans (Chamisso and

Eysenhardt, 1821 (from Pacific ls., Philippines and

the red Sea) and A. elegans yapensis Sato, 1935

(from West Caroline Is.).

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 45

—\ Ui

~y \ Wy 8

88 WWF 86

FIGS. 71-88, Figs. 71-79, Aspidosiphon elegans; 71, dissected specimen; 72, introvert; 73, introvert hooks (from posterior region); 74-

75, introvert hooks with two points; 76, hook from Aspidosiphon exilis Sluiter; 77, spines from introvert; 78, larger spine from

posterior region of introvert; 79, papilla fron anterior region of trunk (Figs. 74. 75 & 77 to same scale). Fig, 80, Aspidosiphon

hartmeveri; introvert hook (after Fischer). Figs. 80-86, Aspidosiphon inquilinus; 81, entire animal; 82, anal shield: 83, small papilla

from introvert, 84, single pointed hook from introvert; 85. spiny papilla from anal shield; 86, dissected specimen, Figs. 87-88,

Clogosiphon aspergillus; 87, anterior cap; 88, introvert hook.

ah REC, §. AUST, MUS, I8 (1):

The first of these to be described was A. elegans,

The description of Chamisso and Eysenhardt is very

brief and deals with external features only, Their

figures, however, show that their specimens arc

those of an Aspidosiphon with an anal and caudal

shield. an introvert which arises ventral to the trunk,

that the introvert is armed wth hooks and spines, the

latter increasing in size the more posteriorly they lie.

Their specimens were collected ‘on the low-lying or

small (= humiles) islands of the Pacific Ocean, in

places that are washed by the tides, and boring in

coral rocks, apparently by means of their shield (=

scutum)”, Griibe’s (1868) description of A. elegans

from specimens from the Red Sea is fuller. The

anterior shield is furrowed to some extent, the spines

on the introvert are largest posteriorly and the

papillae on the trunk are more prominent must

posterior to the anal shield, His drawing of the

bilobed hooks, however, is difficult to follow, The

most detailed description of the species is that of

Selenka and de Man 1883, their specimens having

been collected from the Philippines and the Red

Sea, the latter specimens showing some differences

from those from the Philippines.

The specimens which I am describing from the

Pacific Ocean (Hawaii, Solomons, New Hebrides

and the Great Barrier Reef) resemble A, elegans.in

the follawing respects (1) the presence of double-

pointed hooks with internal marking like those af A.

elegans (2) flat papillae composed of polygonal

plates (3) the spination of the introvert posterior to

the hooks, the spines being largest on the dorsal

surface and at the base of the introvert (4) the

presence of a single retractor which stems from two

roots attached to the posterior third of the body wall

(5) the presence of two nephridia attached for most

of their length.

Selenka's specimens, however possessed a caecum

and a fastening muscle. Nevertheless, | am regarding

my specimens. as A. elegans,

An examination of Sluiter’s specimens af A. exilis

from (1) Siboga Stn. 172 (2 spec.) labelled '‘type”’

and (2) Diuzend Ejl (Tausend Is.) (1 spec.) shows

that they are identical with what I am regarding as

A. elegans. The hooks, spines and papillac

cortespond in all respects. Consequently T have

placed A. exilis Sluiter, 1886 in the synonymy of A,

elegans, A- homomyarius Johnson, 1964, according

to his description, is also very closely related to A,

elegans.

A, elegans seem to have been found only io

formations of coral,

No previous Australian record,

Distribution: (1) in Australia: Queensland at

Three Is. and Lizard Is. (Gt. Barrier Reef).

1-74 January, 1980

(2) elsewhere: Pacific Is. (Chamisso and

Eysenhardt, 1821), West Caroline Is. (Sato, 1935),

Japan (Sato, 1939); Hawaii (in the present paper);

Funafuti (Shipley, 1898): Indonesia (Sluiter, 1886);

Philippines (Selenka & de Man, 1883); Gulf of

Manaar (Gravely, 1927), Mauritius (Wesenberg-

Lund, 1959b); Red Sea (Griibe, 1868, Herubel,

1904, Fischer, 1914. Wesenberg-Lund, 1957)

Specimens examined and localities; Queensland

—Great Barrier Reef at Three Is. (4) coll. Dr. P. E.

Gibbs SAM E1265, at Lizard Is. (5) coll, P. Wearne,

at Heron Is. (1) coll, R- Reichelt.

The description of the species given abave was

based on these specimens and about 20 more from

Hawaii.

*Aspidosiphon exhaustus Sluiter

Aspidosiphon exhaustus Sluiter 1912, pp, 20-21, pl.

1, fig. 11,; Murina, 1972, pp. 295-298, 1977, p.

42-43, fig. 98,

Location of type; ? Zoological Museum, Amster-

dam; specimen from off Morocco (36° 42'N, 8°

40’ 3C’W at 310-749 m).

Description (based on Sluiter 1912): Trunk 17 mm

long and 2mm wide, Introvert about as long as

trunk and armed with numerous rows of very small,

transparent hooks with long bent points. Anal shield

aval and marked posteriorly with 12 radial furrows,

Caudal shield with 10 furrows. Two retractor

muscles, attached to caudal shield, join to form a

single muscle. Two long nephridia attached to trunk

for most of their length.

Murina (1972) gives the length of her Australian

specimen as 15 mm and reports the presence of 20

furrows on the anterior shield,

Australian record: off coast of N,S.W_, (33° 34’ 5”

S, 152° 06’ S$” E) at depth of 425 m. 1 specimen;

Murina 1972, pp. 295-298.

Distribution: (1) in Australia: off coast of New

South Wales.

(2) elsewhere; off Morocco (Sluiter, 1912); Pacific

Ocean at 27° 13" 8, 109° 25" W and Atlantic Ocean at

5° §, 11° E (Murina, 1972).

Aspisosiphon gracilis Baird

(Figs 94-964)

Pseudaspidosiphon gracile Baird 1868, p. 103, pl. 10,

figs 1, la,

Aspidasiphon gracilis: Selenka and de Man 1883, pp.

122-123, fig. 22, 209-213; Rice and Stephen

1970, p, 69,

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 47

Location of type: Natural History Museum,

London, reg. no, 43,5,15, 58a-b; specimens

from Philippine Is.

Description: Specimen long and slender, Trunk

about 45 mm long and 2:5-3:0 mm wide, curved to

resemble a horse-shoe, grey and bearing dark

brown, hemispherical papillae which contrast with

lighter coloured background of trunk, Because the

darker papillae and the lighter areas of the trunk

tend to lie in short longitudinal rows the trunk

appears moitled.

Anal shield with 10 almost complete longitudinal

furrows, Caudal shield sharply conical and with

about 28 complete and a few incomplete radial

furrows, Brown papillae. consisting of numeraus

closely set polygonal plates, are 0:10-0:22 mm in

diameter and at their summit there is a white pore,

Similar but smaller papillae lie on posterior surface

of introvert and on body wall between radial furrows

of caudal shield.

Introvert, light yellow in colour, about 40 min

long and 1-4mm wide arising ventrally to anal

shield; armed with small single pointed hooks, with

width of base greater than height. Small conical

papillae also present on surface, Prominent spines

(fig, O00) scattered over much of introvert. A

tendency for the musculature to be striated circularly

is evident in the dissected specimen, Baird

comments on a similar condition in his specimens,

“Corpus gracile, — - - —striis circularibus cinctum,

Two retractors arise fram body wall near posterior

extremity of trunk, Alimentary canal very long and

much. coiled. Contractile vessel poorly developed.

Neither an intestinal fastener nor caecum found,

Two very long, free, almost black nephridia, nearly

as long as trunk, arise at about same level as anus,

Systematic position: This specimen is one of the

few which have been recorded and described,

No previous Australian record.

Distribution: (1) in Australia: Queensland at Low

Ts,

(2) elsewhere; Philippine Is.

Specimen examined and locality: Queensland

—Low Is. (1) coll, Dr. P, E, Gibbs, SAM E1254.

Aspidosiphon hartmeyeri Fischer

(Fig. 80)

Aspidosiphon hartmeyeri Fischer, 1919a, p. 277:

1927, p. 204, Edmonds, 1956 p. 306, fig. 18;

Cutler, 1977, pp, 147-148.

Location of type; Not known to author; type

locality, Shark Bay, Western Australia,

Description; Trunk as long as 20 mm. Introvert

about same length. Shields darker than trunk and

composed of thick polygonal plates. Anterior shield

with 5-6 furrows, posterior with 24-25, Introvert with

hooks of two points and more posteriorly irregularly

arranged spines. Tentacles 6-8. One retractor muscle

with two long roots fixed near caudal shield. Strong

spindle muscle fixed posteriorly, Nephridia more

than half length of trunk, free for most of their

length and opening just posterior to anal aperature.

Longitudinal musculature continuous} circular mus-

cles tend to form anastomosing bundles, most

readily noticed in dissected animals. This probably

accounts for the ‘“Querstreifung” noticed by Fischer

(1919), Two eyespots. Edmonds (1956) reports an

intestinal fastener to last intestinal whorl and a

caecum,

Systematic position; Although described from

Australia the species is not well Known in this

country, The hooks of Edmonds’ (1956) specimens

are slightly different from those of Fischer, The

Species needs redescription from specimens col-

lected at the type locality, T have used Fischer's

figure of the hook.

Previous Australian records: Western Aus-

tralla—Shark Bay (Fischer, 1919a); Rottnest Is,

(Edmonds, 1956), South Australta—Great Austra-

lian Bight (37° 18" S, 138° 43" E ar 795 m) (Cutler,

1977),

Distribution; (1) in Australia: Western Australia

at Shark Bay and Rottnest Is,; off coast of South

Australia,

(2) elsewhere; West African coast (Wesenberg-

Lund, 1959a); Cuba (Murina, 1967).

Specimen examined and locality: Western Aus-

tralia—Rottnest Is., (1) SAM E1250,

Aspidosiphon inquilinus Sluiter

(Figs. 81-87, 89)

Aspidosiphon inquilinus Shuiter, 1902, p. 29-30, pl. 2,

figs 21-22.

Location of type: Zoological Museum, Amsterdam;

specimen from Siboga Stn. 282 (8° 25-2” S, 127°

18” E, 27-54 m). In shell of Dentalium sp.

Description: Description based on three speci-

mens found in shells of Denralium sp. and one from

shell of a nassariid mollusc. Specimens from

Dentalium small, slender, sub-cylindrical becomine

more slender posteriorly, Trunk 4-8 mm long, 0:7-

1-0 mm wide anteriorly and 0-5-0-7 mm posteriorly

Colour off-white to pale yellow. Specimen from

gastropod coiled.

48 REC. 8. AUST. MUS. 18 (1):

Anal and caudal shields strongly differentiated

from trunk and brown to dark brown in colour. Anal

shield subtriangular with about 14-18 longitudinal

furrows, surmounted by hard, warty and knobby

papillae which are usually arranged so as to form

approximately 3-5 transverse furrows. A feature of

the shield is that 6-15 of the most anterior placed

papillae are modified to form spine-like structures,

Caudal shield conical with about 18 complete and

incomplete furrows.

Introvert (not completely evaginated in any

specimen) about as long as trunk or little longer,

slender and about 0-5 mm wide. No tentacles

observed in a dissected specimen but this point

needs checking. Anteriorly introvert bears 25 ar

more rows of very small single-pointed hooks about.

0:015-0:020 mm tall and wide. Between rows of

hooks and over most of surface of introvert are

numerous, small, tubular papillae or glands about

0-02 mm tall, Spines are also distributed over surface

of introvert.

Papillae or skin bodies on trunk mostly elliptical in

shape. Longitudinal musculature continuous, Intro-

vert retractor single for over half its length but

arising from two shorter roots attached to or near

caudal shield. Oesophagus emerges from between

roots of retractor and loops up; single fastening

muscle attached to jt. Spindle muscle fixed

posteriorly and a wing muscle present, Rectum with

small conical caecum. Two free nephridia, about

half as long as trunk, attached to body wall at about

level of anus.

Systematic position; I have compared these with

the holotype of A. inquilinus Sluiter, described from

a single specimen found in a Dentalium shell in the

Timor Sea (8° 25" S, 127° 18’ B) and consider them to

be the same. The holotype is cylindrical and its

shields are well defined. The anterior shield,

however, is obliquely inclined to the longitudinal

axis of the trunk but in the opposite way to the usual,

its anterior region is tucked in and slopes down to

the introvert (the effect is produced by the

contraction of the longitudinal musculature nearby

in the trunk), One important point, not mentioned

by Sluiter, is that a few of the knob-like papillae

which comprise the anterior part of the anal shield

are modified to form sharply pointed hooks or spines

which are obscured to some extent in the holotype

by the introvert, The introvert retractor of the

holotype (a dissected specimen) is single for well

over half its length and arises from two shorter roots

fixed to the trunk near the caudal shield. The

specimens from Moreton. Bay are smaller and so are

their hooks, They possess a caecum and an intestinal

fastener, neither of which is mention in the type

description. Unfortunately, owing to the frail and

1-74 January, 1980

damaged condition of the gut, it is difficult to say

whether the structures are now present in the

holotype or ever were.

FIG. 89. Aspidosiphon

from Moreton Bay, Queensland; one in shell of

Pentalium).

inguilinus, (twa specimens

The Australian specimens are also closely related

to Aspidosiphon kovaleskii Murina, 1964a collected

from the shells of Denzalium sp. in the Adriatic and

Aegean Seas. They differ, however, from A.

kovaleskii in that (1) externally they possess spines

and tubular papillae on the introvert and (2)

internally they possess a caecum, only one fastening

muscle and nephridia which are free and not

attached,

Murina, 1972, p. 295 reported A. exhaustus

Sluiter, 1912 living in the dead shells of gastropods

(Mitra sp.) from the coasts of New South Wales and

Western Australia. A. exhaustus, however lacks the

spine-like structures on its anal shield and is

consequently thought to be. different.

No previous Australian record.

Distribution:

Queensland.

(1) in Australia: Moreton Bay,

(2) elsewhere: Timor Sea (Siboga Stn, 282),

Specimens examined and localities: Three speci-

mens from shells of Dentalium sp. collected during

Moreton Bay Survey, Queensland by Stephen Cook

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 49

(Dept. of Zoology, Univ. of Queensland) at Stn. 28

in March, 1973. Two were dissected. One specimen

also collected from shell of a nassariid at same

locality; depth 10m. SAM E1266,

Aspidosiphon jukesii Baird

(Figs, 97-99)

Aspidosiphon jukesti Baird, 1873, p. 97; Rice and

Stephen, 1970, pp, 68-69

Aspidosiphon corallicola Sluiter, 1902, pp. 19-25,

Location of type: Nat. Hist. Museum, London: reg.

na, 1965.25.3; specimen from Lee Sandbanks

(Great Barrier Reef), Queensland,

Description: All the specimens had been removed

from the coral and fixed and the introvert was

retracted except in two, where it was partly

extended. Trunk either curved in a circle or coiled.

with am estimated maximum length of 28 mm and

maximum width of 5-8 mm. Anterior shield almost

horseshoe-shaped, very dark in colour and warty,

with radial furtows well developed on dorsal half.

Furrows on ventral side of shield less strongly

marked and run almost at right.angles to radial ones,

Anus. prominent dorsally, just below anal shield,

Posterior shield conical to hemispherical, light

yellow and with radial furrows. Both shields

correspond very closely to those of A. jukesii shawn

in figs. 21-23 of Rice and Stephen (1970).

Introvert (in retracted state) with length about

thcee quarters that of trunk and bearing (1) small,

doubly pointed hooks, (2) spines of variable size and

(3) small, slender, sub-conical papillae which arise

from hemispherical base. Papillae well developed on

anterior and posterior fourth of trunk, A strong

introvert retractor, dividing posteriorly into two

stout roots, attached to body wall near caudal shield,

Longitudinal musculature cantinuous, Rectum

long; without caecum and fastening muscle, Spindle

muscle attached posteriorly. Nephridia slender,

about half to three quarters as long as trunk and

attached for about half of their length, Elliptical ta

spherical eggs 0-12-0-16 mm in diameter,

Systematies and remarks; These specimens from

Queensland correspond very closely to Baird's A

Jukesii as redescribed by Rice and Stephen (1970)

and A. coerallicola Sluiter 1902. Baird's specimens

were obtained from Lee Sandbanks (Great Barrier

Reef) and Sluiter's from solitary corals, including

Heteropsammia michelini from Indonesia, Through

the kindness of the authorities of the Zoolapical

Museum of Amsterdam | have been able to compare

my Specimens with those of Sluiter (reg. no.

V.Si.7.). The examination confirms the conclusion

of Rice and Stephen (1970), that A. jukesii and A.

corallicala are synonymous. The species is well-

illustrated in Rice and Stephen, 1970.

The association between the salitary corals

Heteropsammtia and. Heterocyathus and the sipuncu-

lan A, jukesii was first reported by Edwards and

Haime, 1848a and 1848b and has been redescribed

by a number of writers, listed by Rice (1976). In a

discussion on the matter Rice says, “The

Aspidosiphon inhabits a spiral cavity in the base of

the coral and, through an opening of the cavity on

the under surface of the coral, the sipunculan

extends its introvert into the surrounding substratum

pulling the coral about as it probes and feeds in the

sand. Through this association, the sipunculan is

provided with a protective habitat and, by

movements of the sipunculan, the coral is

maintained in an upright position on the surface of

the substratum and transported about to different

feeding areas.” The developmental history of the

association has also been carefully studied, Accord-

ing to Rice, “The juvenile Aspidosiphon, when

| mm or less in length, enters an empty pastrapod

shell, usually a small Cerithium. A coral planula

settles on the shell, overgrowing and eventually

enclosing it. Only those planulae settling on shells

occupied by sipunculans have any chance of survival.

Growth of the sipunculan and the coral are well

synehronized, the sipunculan actively enlarging its

cavity as 4 spiral tube in the base of the growing

coral while maintaining an opening of the tabe on

the underside of the coral”.

Preyious Australian record: Queensland: Baird

(1873), Goreau and Yonge (1968).

Distributian: (1) in Australia: Queensland at

Heron Is. and Lizard Is.

(2) elsewhere: Malaysia, Gulf of Manaar, Zan-

zibar and Madagascar,

Specimens examined and localities: Queensland

—Lizard Js., (20) in the solitary coral Heterapsam-

mia michelini; coll, D, Fisk (Dept, of Botany, Univ.

of Queensland) SAM E1255 and E1256; Wistari

Reef, Heron IJs.. ‘tin salitary coral’ AMS W5620-

Genus Paraspidosiphon Stephen

Paraspidosiphon Stephen, 1964, p, 457; Stephen &

Edmonds, 1972, p. 237,

Description; Sipunculans with anal and caudal

shields and with characters similar to those of

Aspidesiphon Diesing but differing in that the

longitudinal musculature of body wal) is grouped

into bands. which may anastomose, sometimes

considerably, Intravert, arising from ventral side of

trunk, may be armed with haoks or spines, often

SU REC. §, AUST. MUS. 18 (1);

both. Tentacles as in Aspidosiphon, Introvert

retractors one or two short roots. Spindle muscle

attached posteriorly.

Type species: Paraspidosiphen steenstrupii (Diesing,.

1839),

Remarks; Stephen considered that the condition

of the longitudinal musculature warranted the

separation of Paraspidosiphon from Aspidosiphon

and Fisherana from Phascolosoma. Not everybody

agrees with this. Both Cutler (1973) and Murina

(1975b) consider that Paraspidosiphon warrants the

status anly of a subgenus, The genus contains about

25-30 species, most of which liye in burrows. in

limestone or coral reefs.

Specimens of Paraspidosiphon klunzingeri and P.

steenstrupii reported from Low Is, by Edmonds

(1956) are now being referred to other species, the

former to P. johnstoni n.sp. and the latter to P,

formasanus (Sato),

KEY TO AUSTRALIAN SPECIES GF PARASPIDOSIPHON

Ll. Introvert hooks with asingle point...) 6.5... peepee 2

Introvert hooks with Two paints... ...- baal caees ad

. Numerous rectal appendages present... .. P. cumingit (p.50)

Numerous rectal appendages absent - P. jahnstonia.sp. (p.5!)

3. Introvert hooks with a thin, posteriorly directed, tongue-like

extension of cleatarea ... -... .... P. steensirupit (p.51)

Introvert without tongue-like extension of clear area

P. formosatus (p.50)

*Paraspidosiphon cumingii (Baird)

Aspidosiphon cumingii Baird, 1868, p, 102; Selenka

& de Man, 1883, pp. 113-115, pl. 13, figs 183-

186; Rice & Stephen, 1970, p, 67,

Paraspidosiphon cumirgii; Stephen & Edmonds,

1972, pp. 243-244.

Location of type: Nat. Hist. Museum, London:

specimen from Philippines.

Remarks: This species is often reported from

tropical and near tropical localities and is allied to P,

kKlunzingeri. The introvert, however, is armed with

single-pointed hooks and spines. The anterior shield

has 12-13 furrows around its margin and the caudal

has 30 of which only about 12-13 reach the centre of

the structure. There are 27-28 longitudinal muscles

antériorly and 32-34 posteriorly. According to

Selenka the posterior region of the tectum is thickly

bordered with long, tufted villi (‘Das letzte Ende

des Mastdarmes ist dicht mit langen zottenartigen

Gebilden besetzt’”), There is one broad retractor

which spans about 19 muscle bands; sometimes. the

retractor may split into as many as four roots. Two

nephridia, attached for about two-fifths of their

length extend to the posterior of the trunk. Selenka

states that it has a strong fixing muscle.

1-74 January, L980

Australian record: Low Is., Queensland (Monro,

1931),

Distribution: (1) in Australia; Low Is,, Queens-

land.

(2) elsewhere; West Indies, Red Sea, Philippines,

Zanzibar, Malaysia,

Paraspidosiphon formosanns (Sato)

(Figs 90, 92, 93)

Aspidosiphon formosanus Sato, 1939, pp, 421-424,

text figs 55-57, pl, 21, fig. 23.

Paraspidosiphon formosanus: Stephen & Edmonds,

1972, p. 245; Edmonds, 1971, pp, 144-145,

Aspidosiphon steenstrupii: Edmonds, 1956, p. 307,

fig 19.

Location of type: Not known to author; specimen

from Sinko, Taiwan (Formosa).

Description: Specimens yellow to pink and tend to

be slender. Trunk of largest 30-32 mm long and 2:5-

4-0 mm wide and tending to curve inwards on

ventral side. Anal shield elliptical, dark brown to

black and composed of small knobby wartlike bodies

to which white calcareous material may be attached;

without furrows, although its periphery may be

ridged, Caudal shield light brown, rounded, or

conical and sometimes only weakly developed; if

conical, weak radial furrows may be present but if

rounded they do not appear. Papillae on trunk most

prominent on surface adjacent to anal shield and are

composed of small, closely packed, polygonal to

rounded platelets; diameter of largest papillae

0-45 mm, Papillae near anal shield usually separated

by grooves, which themselves are covered with small

polygonal platelets, Papillae on mmid-trunk region

usually smaller, more rounded and scatteied or even

absent in some specimens, Papillae on posterior

trunk larger and more prominent,

Introvert of retracted specimens about half as Jong

as trunk; of expanded ones nearly twice as long. A

few stoutly digitiform tentacles protrude adjacent to

mouth in one specimen; surface of introvert just

posterior to mouth is smooth and _ callarlike.

Introvert armed with many rows of hooks with two

paints and spines with one point. Small truncated to

cylindrical papillae lie between rows of hooks and

among spines. Secondary or minor tooth of hooks

tends to curve more strongly than primary ane.

Internally, clear area which at base of hook is halt to

three quarters as wide as hook, gradually tapers

towards apex of hook, None of the mounted hooks

show the posteriorly directed, tongue-like extension

of the clear area like that shown for P. steenstrupii in

fig. 192 of Selenka & de Man (1883) nor the

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 5!

FIG. 90, Paraspidosiphon formosanus,

from Queensland),

(Specimen

indentation on the right side of the clear area shown

in fig. 59 of Sato (1939), Although all spines are

single pointed, their shape varies considerably; some

closely resemble single pointed hooks, some are

more tetrahedral and others more slender and

pointed, The variation is much like that shown for P.

grandis in fig. 49 of Sata (1939).

Two introvert retractors, arising from muscles 2-5,

2-6, or 3-7 a short distance in front of caudal shield,

remain separate for about one to two thirds of length

of trunk before fusing to form a stout, single

retractor, Oesophagus, carrying a simple contractile

vessel, is attached to single retractor. Intestine with

about 22 double spirals and rectum with a caecum.

Spindle muscle strong and fixed posteriorly. No

fixing muscle. Nephridia long (extending to

posterior extremity of trunk), fixed for about half

their length and opening at about same level or just

posterior to that of anus. Brain simple with two

eyespots.

Sytematics: Although the internal anatomy is very

much like that of P, steenstrupii (Diesing), the hooks

of these specimens lack the thin tongue-like

extension of the clear area shown in fig. 192 of

Selenka & de Man. (1883) and in pl. | fig. 4 of

Augener (1903), The shape of the hooks is close to

that of P, amthonensis (Augener, 1903), The shields

of the latter, however, are furrowed. The specimens

are also close to P. makoensis (Sato, 1939) and P.

formosanus (Sato, 1939), two species which

themselves are allied. The shape of the hooks of the

Australian specimens corresponds to those of the

latter.

Edmonds (1956, p. 307 and fig. 19) described

some specimens from Heron Is,, Queensland as P.

sieenstrupit. | now think that this identification was

wrong. The hooks, as shown in fig. 19, are like thase

of the specimens which 1 am now describing from

Magnetic Is, and lack the tongue-like extension of

the clear areas as shown by Selenka & de Man and

Augener. Consequently the record of 1956 is now

placed in the synonymy of P. formosanus,

No previous Australian record.

Distribution; (1) in Australia: Queensland.

(2) elsewhere: Formosa, Guam,

Specimens examined and localities: Queensland

—Low Is,, (2) SAM E1275; Magnetic Is. (2) SAM

E1277.

*Paraspidosiphon steenstrupii (Diesing)

Aspidosiphon steenstrupii Diesing, 1859, p. 767, pl.

2, figs 1-6; Selenka & de Man, 1883, pp, 116-

118, pl. 1, figs 12-13, pl. 13, figs 190-192.

Paraspidosiphon steenstrupii: Stephen & Edmonds,

1972, p, 254.

Location of type: Not known by author; specimen

from St. Thomas.

Remarks: This species differs from P. formosanus

in possessing introvert hooks in which the clear area

carries a thin tongue-like extension, much like that

of Aspidosiphon elegans, Edmonds (1956, p. 307,

fig. 19) described some specimens from Low Is, as P.

steenstrupil. | now think that they should have been

called P. formosanus and am transferring the 1956

record to the synonymy of P, formosanus. P.

Steenstrupii has often been recorded from tropical

and subtropical seas.

Australian record: Low Is,, Queensland (Monro,

1931).

Distribution; (1) in Australia: Low Is., Queens-

land.

(2) elsewhere: West Indies; Brazil; Mauritius:

Red Sea; Laccadive Is,; Japan; Philippines; New

Hebrides; Indo China; Loyalty Is., and Cuba.

Paraspidosiphon johnstoni n. sp.

(Figs. 91, 100-102)

Aspidosiphon klunzingeri: Monro, 1931, p, 34;

Edmonds, 1956, pp. 308-9, fig. 20, pl, 3, fig. 1.

52 REC. S. AUST. MUS. 18 (1):

Location of type: Australian Museum, Sydney;

specimen fram coral in Moreton Bay, Queens-

land (coll. Prof. T. H. Johnston).

Description: Trunk usually cylindrical, sometimes

curved ventrally, 20-45 mm long and 3-5 mm wide.

Introvert, arising ventrally from trunk, is in fixed

specimens about as long as or a little longer than

trunk and armed anteriorly with very numerous

(165) rows of single pointed hooks with a stout base

and a curved tip. Hooks in posterior rows noticeably

smaller than those in anterior rows. Posterior to

rows of hooks is a narrow zone of scattered, single

pointed “hooks” or ‘‘spines”, closely resembling

those more anteriorly, but which are much smaller

and lack fine internal markings, Posterior part of

introvert with out spines. Small, truncated ta sub-

conical papillae lie between rows of hooks and on

surface of trunk. Tentacles short, few and finger-like

lying in a part ring dorsal to mouth as in

Phascolosoma,

FIG, 91

Paraspidasiphon johnstoni, (specimen from

Queensland).

Shields dark gold to red brown in colour, wartlike

in texture and appearance and prominent. Anal

shield fan-like, lying obliquely to trunk and

possessing 10-16 complete and incomplete furrows

which radiate dorso-ventrally like ribs of a fan.

Posterior shield conical, often carrying a rim at

junction with trunk and possessing 22-28 complete

and incomplete radial furrows. Furrows of both

|-74 fanuary, 1980

shields may extend over lateral margins of shield,

thus making surface of trunk joining shields appear

furrowed. Furrows correspond approximately to

tissue between longitudinal muscles.

Body wall thick and longitudinal musculature in

25-32 anastomosing bundles. Circular musculature

tends to form fascicles, Retractor muscle very

strong, arising from two short roots each of which

spans muscles 1-8, 2-7, 2-6 and 1-9 well in front of

caudal shield. Oesophagus attached to retractor

muscle tor part length of latter. Intestinal spirals

numerous and extend to posterior of trunk,

Nephridia tubular, about three: quarters as long as

trunk and fixed for anterior quarter. No intestinal

fastener and no rectal caecum. Wing muscle strong;

in addition two strong muscles are attached to

rectum near anus and run transversally across body

wall to muscle 1 or 2 on each side of nerve cord.

Spindle muscle stout, arising anteriorly from body

wall under rectum, and fixed posteriorly, A strand of

fine tissue connects rectum and body wall in some

specimens, Contractile vessel without villi.

Systematic position; The specimens closely

resemble P. klunzingeri (Selenka & de Man) in the

shape and marking of the introvert hooks and in

internal anatomy. They differ from it, however, in

that (1) no rectal sac bearing numerous lobes on

both sides is present (‘am Rectum sitz ein grosser,

an beiden Seiten vielfach gelappter Blindsack”’) (2)

they lack a well defined region of introvert spines

and to a less extent (3) the spindle muscle arises

from the body wall under the rectum and not in front

of the anus (“‘Ein Spindelmuskel nimmt unmittelbar

vor dem After seinen Antang’’),

The Australian specimens also resemble P.

pachydermatus (Wesenberg & Lund, 1937), P.

grandis (Sato, 1939) and P. cumingit (Baird, 1868),

They differ in that the last three species possess

rectal appendages as clearly shown in Wesenberg-

Lund 1937, fig. 9, Sato, 1939, fig. 50 and Selenka &

de Man, 1883, fig. 186b. They also differ from (1) P,

angulatus (Ikeda, 1904) in the internal markings of

the hooks (2) P. speciosus (Gerould, 1913) in the

shape of the hooks and the number of longitudinal

muscles (3) P, schnehageni (Fischer, 1913) in which

the shields are not furrowed, the longitudinal

muscles 10-14 and in which there is. no fastening

muscle (Ditadi, 1975). and (4) P. tenuis (Sluiter,

1886) in which the hooks are long and slender,

Edmonds (1956, pp, 308-9) identified two

specimens from Low Is. as Aspidosiphon klunzin-

geri, remarking at the time that they lacked rectal

appendages. | now think that [ was wrong in so

doing. I have re-examined them and consider that

they should be placed in the synonymy of P.

Johnstoni.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 53

Specimens examined and localities: Queensland Raine Is, (11° 35’ S, 144° 1’ E) coll. 1. M. Thomas,

—Low Is., (2) coll. Dr. P. E. Gibbs, SAM 1274; SAM E1272; Moreton Bay, in coral (8) coll. T. H.

Two Is., (4) coll. Dr. P. E. Gibbs, SAM E1273; Johnston SAM E1270,

FIGS, 92-102, Figs. 92-93, Paraspidosiphon formosanus: 92, dissected specimen; 93, introvert hooks, spines and papillae. Figs. 94-96,

Aspidosiphon gracilis; 94, dissected specimen; 95, introvert hooks; 96, spine fram introvert. Figs. 97-99, Aspidosiphon jukesii: 97,

introvert hook; 98, Introvert spine (same scale as 97); 99, trunk papilla. Figs. 100-102, Paraspidosiphon johnstoni; 100, dissected

specimen; 101, posterior shield; 102, introvert hooks.

54 REC. S. AUST. MUS, 18 (1):

Genus Cloeosiphon Grube

Cloeosiphon Grube, 1868b, p. 48; Selenka & de

Man, 1883, p. 126; Stephen & Edmonds, 1972,

p. 267.

Description: Trunk surmounted by a raunded to

pineapple shaped cap, consisting of numerous.

small, white, calcareous plates. Introvert arises from

centre of cap. Rows of two-pointed hooks present on

introvert, No cap or shield at posterior of trunk,

Longitudinal muscles continuous, Spindle muscle

fixed posteriorly. Two nephridia.

Type species: C, aspergillus. (Quatrefages, 1865),

Remarks; Although four species have been

described in this genus most workers consider them

to be synonymous. C. aspergillus is commonly found

in coral formations in most tropical seas,

Cloeosiphon aspergillus (Quatrefages)

(Figs. 15, 87-88)

Loxosiphon aspergillus Quatrefages, 1865, p. 605, pl.

20, fig. 20.

Clogosiphon aspergillus; Selenka & de Man, 1883,

pp. 126-7, pl, 2, figs 23-24, pl. 14, figs 214-216;

Edmonds, 1956, pp. 309-310, fig 21, pl. 3, fig 2;

Stephen & Edmonds, 1972, pp, 267-268.

Location of type: Not known by author; type

locality: 7?

Description: Trunk cylindrical, 10-45 mm long

and 2.5-4-9 mm wide, bearing anteriorly a white,

calcareous knob or cap, 2-5 mm in diameter and

composed of small, white, polygonal plates of 4-4

sides, arranged in spiral rows. Each plate possesses a

small brown pore, No cap, shield or knob at

posterior extremity. Introvert slender, about as long

as trunk, arises from centre of anterior cap and

armed with double-pointed hooks. No spines on

introvert,

Longitudinal musculature continuous. Trunk

smooth although covered with glands, largest and

most closely placed on anterior and posterior

surfaces, Single retractor arising from twa short

roots. One fastening muscle. No intestinal caecum;

spindle muscle fastened posteriorly. Nephridia dark,

long (extending to base of retactor) and fixed for

their entire length.

Systematic position: The identification of these

specimens depends on the presence of the calcareous

cap or knob and the shape of the introvert hooks.

The cap is sometimes rounded, sometimes shaped

like a pineapple or more flattened. The internal

marking of the hook also varies ta some extent, A

number of authors consider that C, japonicus Ikeda,

1-74 January, 1980

1904, C. javonicus Stluiter, 1886 and C, mollis

Selenka & de Man are synonymous. The matter is

discussed in Stephen & Edmonds, 1972, p, 268.

Previous Australian records: Queensland: Munro,

1931; Edmonds, 1956,

Distribution; (1) in Australias Queensland (in

coral formations); Low Is., Heron Is., Lodestone

Reef, One Tree Is.

(2) elsewhere: widely in the Indo-Pacific region.

Specimens examined and localities: Queens-

land—Heron, Is., SAM E1292 (8), E1293 (1), E1294

(2), Low Is,, SAM E1301 (4): Townsville SAM

E1295 (1).

Genus Lithacrosiphon Shipley

Lithacrosiphon Shipley, 1902, p, 139; Fischer, 1919b,

p. 289, Fischer, 1922, pp. 26-28; Stephen and

Edmonds, 1972, p. 259; Cutler and Jurczak,

1975, p, 243,

Description: Anterior end of trunk surmounted by

a hard, calcareous, cone-shaped cap borne ona pad

of skin; longitudinal muscles fit into pad_ Introvert

arises On ventral side of trunk or cap and bears

numerous rows of hooks with one or two points.

Longitudinal muscles form bands which may

anastomose freely. Two retractors and two nep-

hridia. Spindle muscle fixed posteriorly.

Type-species: Lithacrosiphon maldivensis Shipley,

1902.

Remarks; All the species so far described have

been found in formations of coral, Cutler and

Jurezak (1975) have critically reviewed the genus

and reduced the number of species from 9 to 3. The

valid species. are now L. maldivensis Shipley (the

type), L, cristatus (Sluiter) and L, wniscistatus

(Ikeda). Only L. ecristatus has been found in

Australia,

Lithacrosiphon cristatus (Sluiter)

(Fig. 16)

Aspidosiphon. cristatus Sluiter, 1902, p. 26, pl. 2, figs.

15-16,

Lithacrosiphon cristatus Fischer, 1922, p. 26; Cutler

and Jurczak, 1975, p. 243.

Location of type: Zoological Museum, Amsterdam;

specimen from Malaysia (Siboga Stn. 53,

Waingapu),

Description; Specimens small; trunk of Jargest

9 mm long and 3 mm wide, Anterior region of trunk

with form of a truncated cone, the surface of which is

marked with a number of almost longitudinal

furrows. Posterior to furrowed area trunk bears

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 55

prominent, hemispherical swellings or papillae.

Introvert estimated to be about as long as trunk,

Longitudinal muscles in 17-22 anastomosing bands.

Two introvert retractors, Two nephridia about half

as long as trunk. Both uni- and bidentate hooks

present on intravert, those most distally being

bidentate.

The identification of these specimens depends on

the facts that (1) the anal shield ts furrowed and (2)

the distally placed hooks have two teeth while the

proximal ones have one. No previous Australian

record,

Distribution: (1) in Australia: Lizard Is., Queens-

land,

(2) elsewhere; ‘wide tropical distribution”

(Cutler); Malaysia, Timor, Saipan, Gilbert Is.,

Hawaii, Panama and the Caribbean,

Specimens examined and localities: Queensland-

—Lizatd Is, (8) coll, P, Weate AMS and (3) SAM

£1289; Yonge Reef, east of Lizard Is., (1) AMS

W 10542.

Family Phascolosomatidae and key to genera

Phascolosomatidae Stephen and Edmonds, 1972 p.

269.

Description; Specimens bottle- or flask-shaped,

sometimes sub-cylindrical, Finger-like tentacles

lying in a near circle dorsal to mouth and

surrounding nuchal organ. Longitudinal muscles

grouped in bands, except in one small genus.

Papillae conical to hemispherical in shape, often

mamillate and darkly pigmented and usually covered

with small platelets; most prominent and most

densely grouped at anterior and _ posterior

extremities of trunk, Retractor muscles four, very

rarely two,

Type genus: Phaseclosoma Leuckart, 1828,

KEY TO GENERA OF PHASCOLOSOMATIDAE

1. Species in which longitudinal musculature forms bands

Phascolosoma (p, 55)

Species in which longitudinal musculature is continuous

Fisheranat

Genus. Phascolosoma Leuckart

Description; Trunk spindle-lke sub-cylindrical,

flash-shaped or fusiform. Longitudinal muscles

always in bands, which may anastamose. Muscle

bands not always visible externally but readily seen

on dissection. Tentacles lie in a horse-shoe dorsal ta

mouth and enclose nuchal organ. Introvert and

* Specimens of this small genus have not yet been found in

Australia, Cutler & Murina (1977, p. 183) consider Fisherana

better placed in the family Gollingjidae

trunk usually covered with prominent, conical ta

hemispherical papillae, often brown to dark red-

brown in colour. Trunk papillae usually largest and

most densely distributed on anterior and posterior

surfaces (especially on dorsal side), Retractors four,

very rarely wo. Contractile vessel single and usually

simple.

Type species; Phascolosoma granulatum Leuck-

art, 1828.

SUBGENERA OF PHASCOLOSOMA AND KEY TO

SUBGENERA

Stephen & Edmonds (1972, p, 271) divided the

genus inta four subgenera. Of these, two,

Phascolosoma s,s, and Satonus, are known in

Australia.

1, One pair of retractormuscles,.........,,... Rueppellisorna

Two pairs of retractor muscles .... _... piig ca Bibs

. Contractile yessel with villior tubules,..,...... Anmillesorna

Contractile vessel withour yillior tubules... ___, , ee

3. Spindle muscle fixed to posterior of trunk

Phascalosama s.s (p, 56)

Spindle muscle not fixed to bady wall posteriorly

Satonus (p, 61)

Remarks: Specimens of Phascolosoma ate the

most commonly collected intertidal sipunculans in

Australia. They are found in limestone and coral

reefs (some species bore into the rack; Rice, 1969,

1976), in masses of tubiculous polychaetes, in

clumps of mussels, in rock crevices, under stones

and in mangrove flats, Most appear to be detritus

feeders (Rice, 1976). The larval development af

some species has also been studied by Rice (1970). A

list of the species reported from Australia is given on

p.7 of the present paper.

Identification of species: I have not been able to

construct a simple and satisfactory key to the

Australian phascolosomatids.

P. nigritorquatum, however, differs from all the

other species because (1) its spindle muscle,

according to the type description, is not fixed

posteriorly (2) its mouth is surrounded by a row of

very dark granular swellings and (3) the introvert

bears four rows of hooks. It is not a well-known

species.

The shape and markings of the introvert hooks

and the structure of the papillae are two characters

most commonly used in the identification of species,

Consequently L have figured the hooks of all the

species,

The hook of P. noduliferum has a comparatively

wide base, a tendency to narrow near its mid length

and a fine clear streak running from the apex to the

mid-base. The papillae of P. noduliferum are usually

hemispherical and carry few platelets; they are not

56 REC. 8. AUST. MUS. 18 (1):

heavily pigmented and tend to be uniformly

distributed. Pigmented bands on the introvert are

lacking,

P. arcuatum differs from other species in (1) the

arrangement of its retractor muscles (2) the

presence of sub-epidermal coelomic extensions

(3) the tendency of its circular musculature to form

small bundles and (4) its lack of fixing muscles. The

clear area of the hook is very wide basally. The

species has becn found only in association with

mangroves,

The papillae of P. anulatum are covered with

large contiguous, polygonal plates which usually are

darker in colour at the base of the papillae.

Polygonal plates like those on the papillae are

scattered on the body wal] between the papillae, The

hoak of P. annulatum is “scolops-ugassizi"'-like and

consequently not a very useful distinguishing

character.

The hook of P. albolineatum is stout; its pointed

part is strangly bent and lies almost parallel] to the

base of the hook. The central clear “streak” is wide

and slopes sharply to a basal corner of the hook.

There is a well developed clear, triangular area on

the side of the streak away from the tip of the hook.

The papillae of P. pacificum tend to be sharply

conical so that specimens may feel ‘“bristle-like”.

The introvert hook is sharply bent, its clear streak

widens basally and it possesses a clear triangular

ared, The nephridia extend almost to the extremity

of the trunk and are fixed to the body wall for most

of their length. Specimens are sometimes mottled in

colour,

The hook of P, stephensoni has a clear triangular

area on one side of the clear streak that runs from

the apex to the base and on the other side a clear

crescentic area. The latter is characteristic,

The hooks af P. rottnesti and P. scolops resemble

each other closely except that the clear streak of P

rottnesti may sometimes be slightly swollen neae its

middle. A rectal caecum, reported to be absent in P-

scolops, is present in P. rotmesn, The papillae of P.

rommesti consist of more numerous and smaller

platelets than those of P. scalops.

In P. perlucens a number of elaw-like papillae or

spines are present at the base of the introvert

(especially on the dorsal side) and on the posterior

region of the trunk. The hook is bent rather sharply.

The clear streak of the hook of P nigrescens is

swollen slightly in its upper half and very much

expanded posteriorly, The basal part of the

expansion is indented slightly so as to form a bulge

or a tongue Which is directed towards the tip of the

hook.

1-74 January, 1980

| now consider that P.. heronis Edmonds, 1956 and

P, dunwichi Edmonds, 1956 were incorrectly named;

the former should have been called P, stephensoni

(Stephen, 1942) and the latter P. scolops (Selenka

and de Man, 1883), The reasons for the changes are

given in the text,

Phascolosoma (Phascolosoma) albolineatum Baird

(Fig. 106)

Phascolosoma albolineatum Baird, 1868, pp. 91-92;

Rice and Stephen, 1970, p. 59; Stephen and

Edmonds, 1970, p. 293.

Phymosoma albolineatunt Selenka and de Man,

1883, pp, 71-74, pl. 9, figs. 128-129,

Location of type: Nat. Hist. Museum, London, reg,

no. 1925.25.1; specimen from Philippine fs,

Description: This. description is based on the four

specimens mentioned below and 16 others from the

Solomon Is. Trunk off-white to pale straw in colour

except at junction with introvert where it is dark

brown; length 9-34 mm and width (in mid-region of

trunk) 3-8 mm. Straw coloured introvert, about half

to three quarters as long as trunk, with a number of

black bands of varying size on its dorsal side. An

almost complete ting of short, stubby, light tentacles

(26 in one specimen) lies dorsal to mouth,

Immediately posterior to mouth there is a smooth

eollar-like region. Posterior to collar, iutravert ts

atmed with rows (28 in one specimen) of dark

coloured stout hooks, with a strongly bent tip.

Central, clear region af hook wide and slopes

sharply to basal corner, A triangular shaped, clear

area lies basally on one side of central clear area.

Papillae on introvert brown, pointed, sub-conical

or mamillate, becoming larger and more densely

placed basally. Papillae on trunk largest anteriorly

and (-2-0-3 mm in diameter; in mid-trunk region

they are smaller and more scattered and about

0-15 mm in diameter and posteriorly slightly Jarger

and closer together,

Longitudinal musculature in about 20 anastomos-

ing bundles. Four retractors, arising in posterior half

of trunk: a stout ventral pair from muscles 2-6 or 2-7

and a more slender, dorsal pair more anteriorly from

bands 4-6 or 5-7. Oesophagus fastened to retractors

by thin mesenteries, A strong fastener arising from

muscle 1 on left side near base of darsal retractor

runs towards junction of oesophagus and intestinal

spiral, giving off one branch to posterior oesophagus

and another branch to last spiral of intestine. No

rectal caecum. Wing muscle strong. Spindle muscle

attached near anus and at posterior extremity of

trunk. Contractile vessel simple. Nephridia, about

half as long as trunk, fixed for about half their length

and attached at same level as anus.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) »

Remarks: The shape of the hooks is a very useful

specific character of this species.

No previous Australian record.

Distribution: (1) in Australia: Great Barrier Reef,

Queensland.

(2) elsewhere: an Indo-Pacific species: Natal,

Aboina, Java, Indo-China, Philippines, Japan,

Tokara Is., Palau, West Caroline Is.

Specimens examined and localities: Queensland

—Great Barrier Reef at Three Is,, (2) SAM E1315

and at Casuarina Beach, Lizard Is., (2) WAM

220/76.

Phascolosoma (Phascolosoma) annulatum Hutton

(Figs. 103, 107-8)

Phascolosoma annulatum: Hutton, 1879, p. 278;

Benham, 1903,-p. 174; Edmonds, 1960, pp. 160-

2, text figs. 1-2; Stephen and Edmonds, 1972,

pp. 296-297; Cutler, 1977, p. 151.

Physcosoma scolops var. mossambicense Augener,

1903, p. 339 (in part).

Physcosoma scolops var. tasmaniense Fischer, 1914,

p. 3, pl. 1, figs. 4-6.

Physcosoma scolops Wheeler, 1938, p. 346,

Phascolosoma tasmaniense Edmonds, 1956, pp. 285-

286, figs. 5-6.

Location of type: Not known by author; specimen

from Campbell Is,, New Zealand.

Description: Trunk 25-49 mm long and 4-7 mm

maximum width, Introvert about as long as trunk (in

fixed specimens), with 20-30 short finger-like

tentacles lying in a near ring dorsal to mouth.

Numerous rows of dark coloured introvert hooks,

resembling those of P, scolops (Selenka and de Man)

and P. agassizii Keferstein. The hook possesses (1) a

clear streak (sometimes swollen near its middle)

running down through the ‘centre’ of the hook and

(2) a clear triangular area on one side of the streak.

Specimens brown but anterior and posterior of

trunk made dark- to red-brown or almost black by

pigmentation of papillae. Papillae at base of

introvert dark red, conical and closely packed,

particularly on dorsal surface; those on anterior and

posterior surface of trunk largest, most prominent,

conical or hemispherical, about 0-2-0-4 mm _ in

diameter and densely packed. Papillae may coalesce

or run into one another; composed of polygonal

plates set closely together. At base of papillae there

1s usually an almost complete ring of what appears to

be darker plates, much as is shown for Phascolosoma

japonicum (Griibe;) by Sato (1939, fig 29), Papillae

on mid-region of trunk smaller and more scattered,

A feature of the species is that dark-brown

polygonal plates are scattered over the surface

between papillae. Whether they are detached plates

of the larger papillae is not certain, A number of

pigmented bands, usually best developed on dorsal

side, partly surround anterior surface of introvert

Pigmented patches sometimes appear on trunk,

FIG. 103. Phascalosoma annulatim, (specimen trom

South. Australia).

Longitudinal muscles in 18-25 anastomosing

bands, usually visible externally. Two stout ventral

retractors arising from muscles 2-5, 2-6, 2-7, 3-5 or 3-

6 and two more slender dorsal retractors from 5-6, 4-

6 or 4-7 are attached to posterior half of trunk, One

fastening muscle, connecting muscle 1 on left side ta

last spiral of intestine, may bifurcate giving a second

root to oesophagus. Contractile vessel without villi.

Spindle muscle arises anteriorly near anus and is

fixed posteriorly, Nephridia, usually about half

length of trunk fixed to body wall for about half to

two thirds of their length and arising near anus.

Gonads at base of ventral retractors. Largest sub-

elliptical eggs 0:11-0:13 mm long. Rectal caecum

present in some specimens from South Australia

(SAM 1323), rudimentary or absent in others but

present in those from New Zealand (SAM 1324 and

1325).

Sytematic position: P. annulatum differs from all

other Australian phascolosomatids in the structure

58 REC. 8. AUST. MUS. 18 (1);

of its papillae and in the presence of platelets on the

body wall between the papillae. The platelets are

polygonal and appear ta be detached platelets. from

the papillae. It differs from P, agassizii Keferstein in

the structure of the papillae and from P. japenicurn

Griibe which lacks polygonal platelets on its surface.

In spite of the fact that the rectal caecum is bettet

developed in New Zealand specimens I regard the

specimens from southern Australia and New

Zealand as the same species. According to Cutler

(1977, p, 181), Wesenberg-Lund in an unpublished

work disagreed with this. Although T have not

examined Augener's, Fischer's nor Wheeler's

specimens | have placed them in the synonymy of P.

annulatum, Figs, 4 and 5 of Fischer (1914) are the

papillae and platelets of P. annulatum and the Sir

Joseph Banks Group of islands, where Wheeler's

material was collected, lie off Port Lincoln, South

Austraha, a locality where P. annulatum is

commonly collected,

Previous Australian records: Augener (1903),

Fischer (1914), Wheeler (1938) and Edmonds

(1956),

Distribution: (1) in Australia: along coast of South

Australia irom Baird Bay (Eyre Peninsula) to

Kileunda; Victoria; along northern and eastern

coasts aft Tasmania.

(2) elsewhere: New Zealand; Campbell and

Stewart Islands.

Specimens examined and localities: South Aus-

tralia—Nuyt's Archipelago (2) SAM E1529, Baird

Bay (5) SAM E1231; Pearson Is, (12) SAM E1528;

Grindel fs. (5) SAM E1334; Port Lincoln ‘in

calcareous reef’ SAM E1322; Cape Donnington (2)

SAM E1333; West Is, (1) SAM E1329; Corny Pt. (8)

SAM E1329; Pondalowie Bay (5) SAM E1396;

Beachport (3) SAM E1527; Emu Bay. Kangaroo Is,,

(2) SAM 61537; Stokes Bay, Kangaroo Is., (4) SAM

E1532; Elliston (10) SAM E1540. Vic-

toria—Shoreham (5) SAM E1327 and (1) NMV

G1218; Portland (1) coli, NMV; Cape Patterson (1)

coll. NMV; Kilcunda (5) call. NMV. Tas-

mania—Eaglehawk Neck (3) SAM E1331 and (1)

TMH £101-1553; Port Latta (1) coll, NMV; Dunally

(5) TMH 166; Fossil Is. (1) TMH K82-i4482,;

Taroona Beach (1) TMH K206; Bicheno (2) coll,

NMV;, Erith Ts. (1) AMS W6562.

Phascolosoma (Phascolosoma) arcuatum

1828)

(Figs. 104, 109-111)

Sipunculus arcuatus Gray, 1828, p. 8,

(Gray,

Phascolosoma arcuatum: Baird, 1868, p. 88%; Rice

and Stephen, 1970, pp. 50-52, pl. 1-

174 January, L980

Phymasoma lurco Selenka and ce Man, 1883, pp. 61-

63, pl. 1, fig 5, pl. 8, figs 103-110.

Physcosoma lurce; Fischer, 1895, p. 12.

Phascolosoma lurco: Edmonds, 1956, pp, 290-291,

text fig 10.

Location of type: Nat, Hist, Museum London;

specimen from India.

Description: Trunk stout, almost uniformly

cylindrical, 15-120 mm long and 6-10 mm wide; body

wall light-brown to brown except at anterior and

posterior of trunk Where it is usually dark-brown to

brown-black. Posterior extremity of trunk may be

ipvaginated slightly,

Introvert of fixed specimens with length varying

from half to same as that of trunk, Horseshoe-

shaped ring of tentacles lies dorsal to mouth. More

posteriorly as. many as 75 complete or partially

complete rows of dark coloured hooks, with a large

expansion basally of clear streak which runs from

apex to base. Papillae between raws of hooks small,

about 0-02 mm in diameter, those at extremities of

trunk larger, 0-5-0-:6 mm in diameter, prominent

and very dark, consisting of numerous polygonal

plates, Their structure is.clearly shown in pl, 1, fig. 2

af Rice and Stephen (1970).

Longitudinal muscles in 16-23 broad, anastomas-

ing bands not always visible externally. Circular

muscles tend to form fascicles, as shown in fig. 103 of

Selenka and de Man (1883). Rice and Stephen

(1970, p. 50) point out that small coelomic pockets

extend into spaces between Jongitudinal and circular

musculature. Four retractors fixed ta body wall in

posterior third of trunk, a strong ventral pair to

muscles 1-2, 2-3, or 1-3 and a more slender dorsal

pair more anteriorly to 1-2 or 2-3. Retractors fuse for

most of their length into one, larger flat muscle,

Thin walled oesophagus attached to fused part of

retractor for most of length of latter. Spindle muscle

fastened anteriorly near anus and posteriorly to

body-wall. Rectum short and without a caecum. No

fastening muscle, Wing muscle strong, Nephridia

about halt to three quarters as long as trunk and

fixed for about three quarters of their length.

Contractile vessel without villi. Gonads at base of

ventral retractors, Brain, a white swelling with

pigmented eyespots,

Systematic position and remarks: This species is

distinguished from other Australian phas-

colosomatids by (1) the structure of the introvert

hooks (2) the arrangement of the retractors and (3)

the presence of subepidermal, coelomic extensions.

Tt is associated with mangroves and was for a long

time known as P. lurea. Harms and Dragendarf

(1933) and Green and Dunn (1976) have studied the

ionic and osmotic balance of the species.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 59

FIG. 104. Phascolosoma arcuatum,

from Rockhampton, Queensland),

(specimen

P. arcuatum, according ta Green and Dunn, lives

from the highest to the lowest levels of mangrove

swamps along the west coast of peninsular Malaysia

and consequently is subject to extremely variable

salinities. According to their researches P. arcuatum

is physiologically an osmoconformer. Green (1975)

has studied the reproductive cycle of Australian

specimens.

Previous Australian records; Baird (1868), Fischer

(1895), Edmonds (1956).

Distribution: (1) in Australia: in mangrove flats in

Queensland at Sandgate, Townsville and Rock-

hampton and in Western Australia at Derby,

(2) elsewhere: Malaysia, Indo-China, Philippines

and Java.

Specimens examined and localities: Queensland

—mouth of Ross River “in mangroves” (8) SAM

F1335; Sandgate (5) SAM E1336 and (3) AMS

W3605; Calliope River (3) “amongst mangrove

seedlings” SAM E1526; Townsville “in mangroves”

(10) SAM 1337, Western Australia—Derby (King

Sound) “in mangroves on tidal flat’ (2) SAM E1388

and E1389.

Phascolosoma (Phascolosoma) nigrescens Keterstein

(Fig. 112)

Phascolosoma nigrescens Keferstein, 1865, p, 424,

pl. 32, fig. 14-15: Edmonds, 1956, p. 289, text

fig, 9; Stephen and Edmonds, 1972, p. 315.

Phymosoma nigresecens Selenka and de Man, 1583,

pp. 73-74, pl. 9, figs. 130-137,

Location of type: Not known to author; specimen

from Fiji.

Description: Trunk 15-45 mm long and 2:5-6 mm

wide, light to dark-brown in colour, Introvert of

fixed specimens may be as long as trunk but usually

shorter, with group of short tentacles dorsal to

mouth, Smooth collar present on introvert and more

posteriorly 18-35 rings of light to dark-brown hooks

(fig. 112). Although the clear area that runs from the

apex of the hook towards its base is always expanded

slightly in the middle, the shape of the clear area

basally varies noticeably; in specimens {rom Broome

there is a small hump but in those from Low Is, and

Myora the hump is enlarged to a tongue-like

protrusion. Introvert with stoutly conical papillae

(0:05-0:07 mm in diameter) between rows of hooks,

and usually with a number of pigmented bands on

dorsal surface.

Papillae on trunk and introvert numerous, pointed

and more uniform in shape and distribution than in

most Phascolosoma, At posterior extremity of trunk

papillae concial to mamillate, about 0-2 mm tall and

0-2 mm broad, with a pore at apex and consisting of

dark coloured, elongate, radiating plates. A number

of very dark papillae may be scattered randomly on

trunk and introvert of specimens.

Longitudinal muscles in about 20 anastomosing

bundles. Two stout ventral retractors arise in mid-

third of trunk from muscles 1-7, 2-7, 2-8, or 3-9 and

two slenderer dorsal retractors more anteriorly from

6-8, 5-8 or 7-9, the dorsal and ventral muscle on

same side of nerve cord soon fusing, Strong spindle

muscle attached near anus and at posterior extremity

of trunk. Rectum and spindle muscle may fuse for

part of length, A fastening muscle to posterior

intestine may give a branch to oesophagus. Strong

wing muscle and rectal caecum present. Contractile

vessel simple, Nephridia fixed for half or third of

their length, about half as long as trunk and arising

at about level of anus.

Systematic position: This species is identified

largely by the structure of its hooks as shown in figs.

130 and 135 of Selenka (1883) and in the present

material. Edmonds (1956) mentioned that the

contractile vessel of his specimens possessed very

REC. S. AUST. MUS. 18 (1):

1-74 January, 1980

FIGS. 105-115. Fig. 105, anterior region of introvert of a Phascolosama, showing position of tentacles, mouth and nuchal organ (after

Theel). Fig. 106, Phascolosoma albolineatum: introvert hook, Figs. 107-108, Phascolosoma annulatum: 107, introvert hooks: 108,

trunk papilla, Figs, 109-111, Phascolosoma arcuatum: LO9. dissected specimen; 110. intravert hook; 111. trunk papilla. Fig, 112.

Phascolosoma nigrescens; introvert hooks. Fig, 113, Phascolosoma nigritorquatum; introyert hooks (from one of Sluiter’s

specimens), Figs. 114-115, Phascolosoma naduliferum; 114, introvert hook; 115, trunk papilla.

small villi. Although I have not been able to re-

examine the same specimens I now think that I

mistook some very small wrinkles or crenations of

the muscular wall of the vessel to be villi.

It is difficult to find a satisfactory character which

can be used to distinguish P. nigrescens from P.

puntarenae Griibe. The relationship between the

two is discussed by Fisher (1952, p, 430-432). If they

are the same, P. puntarenae Griibe, 1859 has

priority. The species is often associated with coral

formations,

Previous Australian records: Queensland—Low

Is,; (Munro, 1931); Heron Is. (Edmonds, 1956).

Western Australia—Cape Jaubert (Fischer, 1921);

Shark Bay (Fischer, 1919a), Cape Dennison

(Edmonds, 1956).

Distribution: (1) in Australia—Queensland;

northern New South Wales; north west Australia.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (S/PLUINCULA) pl

(2) elsewhere; widely in Indo-Pacific region; West

Afnica.

Specimens examined and localities: Queensland

—Low Is,, “in coral” (2) SAM E1344; Magnetic Is,,

(1) SAM E1345; One Tree Is,, (1) SAM E1346;

Three fs., (1) SAM E1347; Cockle Bay (2) SAM

E1350; Myora (1) SAM E1351; Wistan Reet

(Capricorn Group) (2) SAM E1352. New South

Wales—Woolgaoga (1) coll, NMV: Lennox Head (3)

AMS W10545 and (6) AMS W10544. Western

Australia—Cable Beach (near Broome) (3) SAM

E1349; Broome (Pt, Gantheaume) (3) WAM 10922,

Dampier Archipelago (1) WAM 132/76 and (1)

WAM 222/76.

*Phascolosoma (Satonus) nigritorquatum (Sluiter)

(Fig, 113)

Phymosoma nigritorquatum Sluiter, 1882, pp, 151-

152, pl. i, figs 3, 8, 11; Selenka and de Man,

1883. pp. 68-69.

Physcasema_ nigritorquatum; Sluiter, 1902, p. 13;

Fischer, 1919a, p, 280; 1921, p, 4-5, figs 1-2;

1927, p. 416,

Phascolosoma nigritorquatum: Stephen & Edmonds,

1972, p. 186.

Location of type: Zoological Museum, Amsterdam:

specimen from Batavia ‘trom coral islands".

Description; T have not examined any Australian

specimens of the species. It has, however, been

reported twice from Western Australia by Fischer

(19194, 1921). According to Sluiter the length af the

body is 2-5-3 times its width. Posterior region

pointed. Papillae very numerous, especially at

anterior and posterior extremities of trunk. Longitu-

dinal muscles in 20-22 bands and circular muscula-

ture continuous. Introvert about a quarter as long as

body, Two anterior retractors weaker than two

placed more posteriorly, A narrow papillae-tree

zone at base of introvert. A row of black granulated

bodies? (‘‘Karner'') and swellings? (‘‘Wiilste”) lie in

a half ring around margin of mouth, The pigmented

bands extend in successive rows around anterior part

of introvert. Four rows of small, slender introvert

hooks. Tentacles arising from two folds. Intestine

with only a few, nine, spirals, No spindle muscle.

Two anteriorly swollen nephridia, Colour of animals

dark yellow to brown. Length without introvert

13mm,

In a ‘discussion’ that follows the description,

Sluiter says that the black bands? or ridges?

("*Leiste’’) on the introvert are characteristic of the

species and that sometimes they anastomose. The

first of the black bands is never completely

invaginated and remains visible even if the introvert

is retracted.

The account of Selenka & de Man is only a

Tepetition of Sluiter’s. Fischer (1921. p. 5) records

that a specimen fromm Western Australia ig 12 mm

long and its introvert 8-9 mm. Skin yellow-white.

Introvert hooks sickle-shaped and in 5-6 tows.

Longitudinal muscles in 20-22 bundles. Two broad

ventral retractors arise in posterior third from

muscles. 2-7, Intestine of 10-12 double spirals.

Fischer says that a spindle muscle is present and that

it is fixed posteriorly.

Systematic position: Recently T have re-exarnined

two of Sluiter’s specimens (V 51, 80, Baai v.

Batavia) from the Zoological Museum, Amsterdam

The tube cantaining them bears the label “type”;

one was partly dissected. Body small, trunk 5-8 mim

long and 25-4mm maximum width; pointed

posteriorly. Introvert of each specimen almost

completely retracted. Longitudinal muscles in 19-23

bundles visible externally; some anastomeasdtion.

Papillae most prominent on anterior and posterior

surfaces of trunk; often sharply pointed and spine-

like, Surface of papillae carries small rounded-

polygonal platelets (much like those of P. seolops)

and underlying there seem to be rather long radially

arranged plates. In the specimen that I dissected a

number of red-brown bands were found on the

introvert. especially the dorsal surface. 1 was not

able to find Sluiter’s “intens schwarzer Kérner and

Wilste”. A few rows of clear, unpigmented

introvert hooks were present (fig. 113); they do not

correspond very well with those drawn by Sluiter

and Fischer and resemble closely those of P. scalops

and P. rottnestti, Introvert retractors four, the dorsal

and ventral pair on each side almost spanning the

same number of muscles. The gut of the specirneni

Whieh I dissected was thin, frail and not in gaod

condition and that of the other (1 extended the cut

already present a little further anterjorly and

posteriorly) was surrounded by coagulated fnaterial

which was difficult to remove, In neither was [ able

ta find @ spindle muscle anteriorly nor the point at

attachment of such a musele near the anus. No sign

of a posteriorly placed spindle muscle was present ii

ane specimen but in the other a stout muscle

emerges from the last intestinal spiral and is attached

to the bady wall. Consequently I have not been able

ta check conclusively whether a spindle musele is

present or absent in P. nigritorquatum. The answer is

important because in the subgenus Satonus a spindle

muscle is either absent or not attached posteriorly.

P. nigritorquatum is the type of the subgenus, If P-

nigritorquatuim possesses a spindle muscle Satonus is

invalid and another subgenus must be erected and a

new type found, No information about the presence

of a caecum or fastening muscle can be given. The

two nephridia are free, Some sub-elliptical-shaped

eggs (0-10-0-13 mm) lang and (0-07-0-075 mm) wide

a2 REC. S. AUST, MUS, 18 (1);

with a nucleus about 0-025 mm in diameter were

present.

Previous Australian records: Fischer, 19194; 1921.

Distribution: (1) in Australia—Western Aus-

tralia, Cape Jauberr and Shark Bay.

(2) elsewhere: Bay of Bantam, Indonesia.

Phascolosoma (Phascolosoma) noduliferum Stimpson

(Figs 114-115)

Phascolasoma noduliferum Stimpson, 1855, p. 390;

Keferstein, 1865, p. 423, p. 32, figs 16-17;

Edmonds, 1956, pp. 286-288, text figs 7-8;

Cutler, 1977, p, 152,

Physcosoma grayi Baird, 1868, pp. 88; Rice &

Stephen, 1970, p, 52,

Location of type: Not known to author; specimen

from Port Jackson, New South Wales,

Deseription; Trunk 12-65 mm long and 3-7 mm

wide, brown or light brown in colour with only a

slight darkening at its extemities, Introvert of fixed

specimens about as long as trunk, lacking pigmented

bands but possessing (1) about 20-25 short,

digitiform tentacles and (2) a varying number of

dark brown hooks (fig. 114). Shape of hook

distinctive, with broad base and a thin clear streak,

running from apex to base of hook,

Papillae usually hemispherical and more uni-

formly distributed on trunk than in P. annulatum,

largest 0:3-0-4 mm in diameter. consisting of

numerous small plates and scattered granules.

Longitudinal musculature in 20-26 bands, not always

visible externally because body wall is thick. Four

retractors, a stout ventral pair attached to muscles 2-

6, 2-7 or 1-6 in posterior half of trunk and a weaker

dorsal pair more anteriorly to muscles 4-6, 5-7 or 6-

7, Oesophagus fixed to retractors by two short

mesenteries. Spindle muscle fixed anteriorly near

anus and posteriorly to body wall, Wing muscle well-

developed, A fastening muscle arising from muscle 1

or 2 near base of left ventral retractor runs to last or

penultimate intestinal spiral; sometimes supplying a

branch to oesophagus.

Nephridia half to three quarters as long as trunk

and fixed for about three quarters of their length,

Contractile vessel simple. No rectal caecum. Brain

appears as slight swelling in some specimens} two

eyespots. Elliptical eggs about 0-12 mm in diamter.

Systematic position; Edmonds (1956, p, 288)

discussed at some length the status. of this sipunculan

commonly collected along the eastern and southern

coasts af Australia and which has often been called

P. japonicum (Selenka and de Man, 1883, p. 77;

Dakin, 1952, p, 157 and Whitlegge, 1899, p. 11).

This. probably stems from the identification of two

specimens from New South Wales as P. japonicum

1-74 January, 1980

by Selenka and de Man, 1883. The two specimens,

tegistered as 85.12.3.30 and 85.11.3.30, are in the

British Museum (Natural History, London). They

possess the hooks and papillae of P. noduliferum and

not P. japonicum.

There is considerable overlap in the distribution of

P, noduliferam and P. annulatunr in Victoria and

Tasmania, Specimens of both have been recorded

fram Eaglehawk Neck, Dunally and Bicheno (in

Tasmania} and from Kilcunda and the Nobbies (in

Victoria). The differences in their habitats at these

places is not known, P. noduliferum occurs in New

South Wales, Victoria, Tasmania and South

Australia. Two specimens are also known from the

South coast of Western. Australia.

P, noduliferum differs from allied Australian

species in the structure of its hooks and papiilae.

Pigmented bands on the intravert have not been

described for the species.

Previous Australian records: Stimpson (1885);

Edmonds (1956); Cutler (1977) (7 specimens

dredged at 85-1340 m off South Australian coast),

Distribution: (1) in Australia—New South Wales,

Victoria, Tasmania, South Australia and south coast

ot Western Australia.

(2) elsewhere—Malaeca Sts, (at 1140 mm),

Mindanao (at 22 m), Pt. Moresby (New Guinea) (at

8 m), Campbell [s. (off south coast of New

Zealand). All of these records are from Cutler

(1977, p, 152),

Specimens examined and localities: New South

Wales—Newport (6) SAM E1353; Long Reef (3)

SAM E1354 and (1) AMS W10548; near Sydney (6)

SAM E1356; Harbord (3) AMS coll,; Botany Bay

(2) AMS W548. Victoria—the Nobbies, Western

Port (8) SAM E1355 and (1) NMV GI129: Phillip Is.

(2) SAM B1357; Godfrey reef near Larne (1) NMV

coll.; Cape Bridgewater (1) NMV coll.; Portsea Pier

(1) NMV coll.; Malacoota (2) AMS W10547,

Tasmania—Bicheno (2) NMV coll.; Jacobs Boat

Harbor (1) TMH coll.; Dunally (2) TMH K/166;

Eaglehawk Neck (8) TMH coll.; and (2) TMH

K102/15532 and (5) SAM E1397; Erith Is, (1) AMS

W10569, South Australia—Cape Northumberland

(1) SAM E1390. Western Australia—10 miles cast

of Hopetown (1) SAM E1400; off south coast (35°

15' S, 126° 22' E) (1) WAM 145/76.

Phascolosoma (Phascolosoma) pacificum Keferstein

(Figs, 120-121)

Phascolosoma pacificum Keferstein, 1866, pp. 8-9;

1867, pp. 49-50, pl. 6, Figs 1-2, Edmonds, 1956,

pp. 291-292, text fig. 11, Stephen and Edmonds,

1972, pp, 317-318.

A REVISION OF THE SYSTEMATICS GF AUSTRALIAN SIPUNCULANS (SIPUINCULA) 63

Phymosoma pacificum: Selenka and de Man, 1883,

pp. 63-65, pl. 2. fig 6, pl. 7, figs 111-112.

Location of type; Not known to author; specimen

from Gilbert and Tarawa Is. (Kingsmill Group,

Pacific Ocean).

Description: Trunk of largest specimen 102 mm

long and maximum width (in posterior third of

trunk) about & mm. Light brown to brown in colour

with a darker mottling in some specimens. Mottling

on anterior surface of trunk may be less irregular

and more band-like, especially on dorsal surface.

Body wall thick and banding of longitudinal muscles

not usually visible externally.

Introvert in fixed specimens about as long as

trunk, with a group of digitiform tentacles dorsal ta

mouth and with humerous complete and incomplete

rows of large, dark-brown hooks, 0-07-0-10 mm

wide basally and 0-07-0-09 mm tall.

Papillae on introvert and trunk numerous, closely

packed, sharpely pointed and conical so that animal

feels rough and bristle-like (a fact mentioned by

Selenka and de Man), On mid-region. of introvert

papillae 0-20-0-25 tm tall, 0-15-0-2 mm wide; on

posterior of trunk becoming 0-3 mm tall, 0-3 mm

wide. Papillae composed of very small granules with

a pore at apex,

Longitudinal muscles in anastomosing bundles,

ahout 20 anteriorly and 35-40 posteriorly, Four

retractors, a stout ventral pair arisingin mid-third of

trunk and attached to muscles 2-7 or 2-8, and a

dorsal pair more anteriorly and attached to same six

or seven muscles. Retractors fuse anteriorly,

Intestinal contents of coarse sand and coral

fragments. One fastener to last or second to last

spiral or intestine, Spindle muscle stout and attached

posteriorly. No caecum in two dissected specimens

Contractile vessel simple. Nephridia pigmented

brown ar brown-black, very long (extending almost

to extremity of trunk) and fixed to trunk wall for

their whole length,

Systematic position; This species is identified by

the size and shape of its introvert hooks and the

presence of nephridia which extend almost to the

posterior extremity of the trunk and which are fixed

ta the body wall for their whale length-

Previous Australian records: Queensland—Low

ls. (Monro, 1931); Heron Is. (Edmonds, 1956),

Distribution: (1) in Australia—Great Barrier

Reef, Queensland; Northern Territory.

(2) elsewhere: Indo-Pacific, Madagascar, Zan-

zibar, Mauritius, Red Sea, Ambaina, Indo-China,

Japan, Philippines, Gaum. New Britain, West

Carolines,

Specimens examined and localities: Quceens-

land—Heron Is, (1) SAM E1358; Low Is. (1) SAM

E1359; Turtle Is, (2) SAM £1360; Dunwich (1)

SAM E1364; Michaelmas Reef, Cairns (2) AMS

coll. Northern Territory—Fannie Bay Rocks,

Darwin (1) AMS coll.

Phascolosoma (Phascolosoma) perlucens Baird

(Figs, 117-119)

Phascolosoma perlucens Baird, 1868, p. 90, pl. 10,

fig. 2; Rice and Stephen, 1970, pp, 63-64,

Phyimosoma dentigerum Selenka and de Man, 1883,

pp. 67-68, pl, 1, fig, 7, pl, 9, figs. 118-123,

Phascolosoma dentigerum: Fisher, 1952, pp. 432-

434, pl. 36, figs. 4-7.

Location of type: Nat. Hist. Museum, London. reg.

no. 1847,12,30,11; speeimen from Jamaica (in

coral rock).

Descriptian; Specimens tend ta be slender and

straw coloured in alcohol, Trunk of largest 45 mm

long and 4-5 mm wide. Body wall thin and almost

transparent in some regions, Introyert about half to

three quarters as long as trunk (in fixed specimens)

with (1) a number of very dark-brown bands on

dorsal surface (2) prominent, dark-red-brown,

spine-like papillae or tubercles on posterior dorsal

surface and (3) rows of light- to dark-brown hooks

anteriorly, Spine- or claw-like papillae mostly point

backwards and vary in size, the largest being about

0-6-0-7 min tall and about 0-5 mm wide basally (fig

00). A fine tube runs from tip of spine to its base, the

diameter of tube increasing basally, Anterior half or

third of hook strongly bent and lying almost parallel

to base of hook, central clear area expanded near its

middle and a clear triangular area present basally

Papillae on posterior trunk darker than on trunk and

tend to be claw-like} largest about 0-3 mm tall and

0-25 mm wide at base. Posteriorly placed claw-like

papillae of two Australian specimens not as

prominent as those shown in Fisher, 1952, pl. 39,

fig, 5,

Longitudinal muscles in about 22 anastomosing

bands, Retractor muscles four, a ventral pair

attached fo muscles 2-7 or 2-6 in mid-third of trunk

and a dorsal pair more anteriorly to 5-8, Contractile

vessel slender and without villi. One fastening

muscle ta last spical of intestine, Caecum not

abseryed (Fisher, 1952 claims that one is present),

Nephridia fixed to trunk for about one third of their

length and about one third as Jong as trunk

Systematic position and remarks: The shape of the

introvert hooks and the presence of large claw-like

structures on the dorsal surface of the introvert

make this one of the easier species of Phascolosoma

64 REC. S. AUST. MUS. 18 (1):

to identify, Fisher (1952) redescribed and figured it.

The species is probably better known in the

literature as P. dentigerum, The Australian speci-

mens were collected in coral rock, Rice (1976)

describes the feeding behaviour of the species.

Previous Australian record; Low Is., Queensland

(Monro, 1931).

Distribution; (1)

Reef, Queensland,

in Australia—Great Barrier

(2) elsewhere—Baja California, Gulf of Panama,

Rotuma, Funfuti, Marshall Is., Eniwetok Atol,

Philippines, Laccadive Is., Batavia, Madagascar,

and the West Indies,

Specimens examined and localities: Queens-

land—on Great Barrier Reef; Three ts., (1) SAM

E1365; Tree [s,; (2) SAM E1366; One Tree Is,, (1)

AMS W5525, Lizard Is., "in coral” (3) SAM E1395.

Phascolosoma (Phascolosoma) rottnesti Edmonds

(Figs, 116, 124-126)

Phascolosoma rottnesti Edmonds, 1956, pp. 282-284,

text figs. 1-4.

Physcosoma agassizii Fischer, 1919a, p. 277;

Fischer, 1922, p. 7, pl. 1, figs. 3, 5 and 6.

Location of type: Australian Museum, Sydney;

specimen from Rottnest Is., Western Austraha,

“from burrows in calcareous rock’

Description; Trunk 10-52 mm long and maximum

width (in posterior third) 3-5 mm, Expanded

introvert of fixed specimens about as long as trunk.

Freshly collected animals pale to pink-brown in

colour except at extremities of trunk which are dark-

to red-brown. Red-brown bands, varying in number

and width, usually present on dorsal surface of

introvert, Trunk of few specimens is mottled by a

number of red-brown spots. Partial ring of tentacles

lies dorsal to mouth, From 15-45 rows of light ta

dark brown hooks, resembling those of Phas-

colosoma scolops (Selenka and de Man 1883, fig.

143) and Phascolosoma agassizii Keferstein, 1867,

pl. 6, fig. 4 but not fig, 8, present on introvert

anteriorly, A clear triangular area is always present

on one side of the clear streak, which runs fram the

apex of hook to its base.

Papillae between rows of hooks very small and

forming complete transverse rings; towards base of

introvert they are larger, about 0-1 mm in diameter,

pointed or conical and darker in colour. On anterior

surface of trunk papillae larger, especially on dorsal

side, 0-2-0:35 mm in diameter, more hemispherical,

more densely packed and dark brown in colour. In

mid-trunk region papillae smaller, rounded, more

scattered and less prominent, On posterior surface

1-74 January, [980

FIG.

Rottnest Island, Western Australia),

116. Phascalosoma rattnestt, (specimen from

they are large, 0-2-0-45 mm in diameter, closely

packed and dark brown, All papillae consist of small

plates of almost uniform size.

Longitudinal muscles in 20-24 anastomosing

bands, not always visible externally. Four retractor

muscles, a strong ventral pair arising in posterior

half of trunk from muscles 2-5, 2-6, 2-7 or 3-7 anda

weaker dorsal pair more anteriorly from 4-6, 5-6, 5-7

or 6. Anterior oesophagus attached to dorsal

retractors by thin mesenteries, Spindle muscle

fastened near anus and at posterior extremity of

trunk, Contractile vessel simple, One fastener

arising from muscle 1 on left side joins last or

penultimate spiral of intestine, sometimes giving off

a branch to posterior oesophagus. Rectal caecum

present, Nephridia, attached to body wall for about

half their length, about two thirds as long as trunk

and arising at about same level as anus, Brain

sometimes observed as slight swelling with two

eyespots, Gonads at base of ventral retractors.

Systematic position: The identity of the specimens

was discussed by Edmonds (1956, p, 284), They

resemble most closely Phascolasoma scalops

(Selenka and de Man) and to a less extent

Phascolosoma agassizii Keferstein, two very vari-

able species. Edmonds (1956) considered that they

differed from P, scolops (1) in the structure of the

papillae (they haye more uniformly distributed,

A REVISION OF TRE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 63

more closely set and rounded platelets) and (2) in

the possession of a rectal caecum, a structure neither

mentioned or figured by Selenka in the type

description nor by Sato (1930) nor Wesenberg-Lund

(1957) in their descriptions of specimens from Japan

and Red Sea, There are two specimens of P. scolops

in the Nat, Hist. Museum, London, both named and

presented by Selenku. No caecum is present in the

dissected specimen of the two, Just haw important is

the presence of absence of a rectal caecum in the

systematics of Phascolosoma I am not sure. In the

circumstances [| prefer ta specify (with some

reservations) the specimens [rom Western Australia

as P. rottnestij and not P. scolaps,

From figs. 3 and § of Fischer, 1922 it seems likely

his P. agassi2ii from Shark Bay and Rottnest Is, are

P) rotmesi. | have not, however, been able to find

his Specimens

The species is commonly found in limestone

rocks. coral and lithothamnion and coralline mats.

Previous Australian record; Western Australia

(Edmonds, 1956), 7 Fischer, 19194 and 1922,

Distribution. (1) in Australia—Western Australia;

as far north as Point Cloates and as far south as

Albany, Whether these localities are at the

extremities of its distribution is not known.

(2) elsewhere—no records,

Specimens examined and localities: Western

Austraha—Fremantle (3) “in calcareous rocks near

railway bridge’’ SAM E1367; Rottnest Is. (Wilson

Bay) (4) WAM 174/76, Rottnest Is. (Green Is,) (20)

WAM 169/76 and (8) SAM E1368; Cottesloe (4)

SAM E1369; Pt. Peron (2) WAM 180/76; Garden Is,

(1) WAM 234/76; Abrolhos Group at Beacon Is, (2)

WAM 237/76 and North Is., ‘in coral’ (2) WAM

241/76; Yallingup Beach (2) AMS W9246; Cheyne

Beach, Albany (2) WAM 239/76; Eagle Bay, Cape

Naturaliste (2) WAM 174/76; Warroora (1) AMS

5470; Point Cloates (3) SAM E1370.

Phascolosoma scolops (Selenka & de Man)

(Figs 122-3)

Phiviniosama scolaps Sclenka & de Man, 1883, pp.

75-7. figs 17, 138-144) Cutler. 1977, p. 152,

Phascolosoma dunwichi Edmonds, 1956, pp, 292-3,

text figs 12-13: Stephen & Edmonds, 1972. pp.

300-401.

Location of type, Not known to author; specimen

from Philippines.

Description: Trunk 15-34 mm long and maximum

width (in posterior third) 3-6 mm light brown except

at anterior and posterior extermities which ate dark-

brown, Introvert of fixed specimens. about as long as

trunk with @ variable number of dark-brown bands

on dorsal surface. Tentacles 15-20 and digitiform,

lying in a near ring dorsal to mouth. Surface of

introvert just pasterior to mouth smooth and collar-

like,

Introvert hooks in [5-45 complete and incomplete

rows, closely resembling those of Phascolosoma

scolops as shown in fig 139 of Selenka & de Man

(1883), Size of hook may vary in different

specimens, smallest being 0-045 mm tall and largest

0-064 mm. Clear streak (running from apex to base

of hook) (1) does not always form a uniform are but

may be bent in its middle (much like an Australian

boomerang) and (2) is sometimes expanded at or

just below bend. Most hooks with a small

“secondary tooth’, something not shown in

Selenka's fig 139. A clear triangular area present on

one side of clear streak at base af hook but lacking

the clear crescentric area as found in P, stephensoni,

Papillae between hooks small, flat, circular to

elliptical with diameter 0:02-0:03 mm. Papillae at

base of introvert darker, cancical (sometimes

sharply) to hemispherical, consisting of an apical

pore surrounded by almast contiguous, polygonal

plates af about equal size. Papillae on trunk largest

and most closely packed at anterior and posterior

extremities, particularly on dorsal side; 0.2-0-35 mm

tall and 0-:2-0:3 mm wide. Papillae in nmd-trunk

region smaller, more scattered and less pointed. In

Some specimens. number of papillae on middle and

posterior surfaces may be reduced

Longitudinal muscles in 20-25 anastomosing bands

often clearly visible through body wall, Four

retractors, a Stout ventral pair atrached to muscles 2-

7, 1-6 or 2-6 in posterior half of irunk and a more

slender dorsal pair more anteriorly to muscles 6-7, 5-

7 or 6-8, Retractors on each side fuse anteriarly.

One fastener runs from muscle 1 near base of left

dorsal retractor to last spiral of intestine, sometimes

giving a root to posterior oesophagus. Wing muscle

well-developed, Spindle muscle fastened anteriorly

near anus and posteriorly to body wall. Rectal

caecum not present. Contractile vessel simple.

Nephridia brown, about quarter to half as long as

trunk, fixed for about half their length and opening

at about same level as anus. Two cyespots present.

Systematie position: Edmonds (1956, p, 292)

identified some specimens from Dunwich as P.

dunwichi a new species allied to P. scolops, The main

differences were that P. dunwichi possessed a

caecum and that there were some differences in the

size and distribution of the platelets of the papillae,

More recently I have re-examined the specimens and

consider that a caccum is not present. TI must have

mistaken a small outpocketing of the gut wall

66 REC, S, AUST. MUS. 18 (1):

containing some hard food particles for a caecum. In

addition the size and distribution of the platelets of

some of the newer specimens in my collection

approach more closely those of the two specimens of

P. scolops (named by Selenka) now in the Natural

History Museum, London, The differences then

between P. scolops and P. dunwichi then become

insignificant and the latter becomes a junior

synonym of the former.

What the distribution of the species is in Australia

is not known. Very few phascolosomatids from

northern and north-western Australia are available

for study. However, all the specimens of P, rottnestii

from south-west Australia that I have examined do

possess a well-developed caecum,

ATI

oi,

Ov?

——— a-O5

1-74 January, 1980

Superficially P. scolops and P. stephensoni are

almost indistinguishable. The crescentic-shaped

clear area present on the hook of P. stephensoni,

however, is not present in P. scolops.

Previous Australian records: Queensland—Ed-

monds (1956); Monro (1931). Cutler (1977) reports

one specimen off South Australia.

Distribution: (1) in Australia—Queensland.

(2) elsewhere—widely reported from the Indo-

Pacific.

Specimens examined and localities: Queensland

—Dunwich (on Stradbroke Is.) (1) in clumps of

mussels SAM E1339; (3) SAM E1340: (6)

SAM E1341.

FIGS. 117-129, Figs. 117-119, Phascolosoma perlucens; 117, complete specimen: 118, spine-like papillae from base of introvert (Figs.

117 & 118 after Fisher); 119, introvert hook, Figs. 120-121, Phascolosama pacificum; 120, introvert hook; 121, papilla from

introvert. Figs, 122-123, Phascolosoma scolopss 122, introvert hook: 123, trunk papilla. Figs. 124-126, Phascolosonia rotmesti: 124,

dissected specimen, 125, introvert hook; 126, trunk papilla (same seale as Fig. 123), Figs. 127-129, Phascolosonia stephensoni; 127,

intravert hook; 128, trunk papilla; 129, hook from one of Stephen's specimens.

4 REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 67

Phascolosoma (Pascolosama) stephensoni (Stephen)

(Figs. 127-129)

Physcosoma stephensoni Stephen, 1942, p. 25(), pl.

11, figs 3-5.

Phascolosoma stephensont; Wesenberg-Lund. 1963:

pp. 121-126, text figs 7-9.

Phascolosoma heronis Edmonds, 1956, pp, 293-295,

fig. 14,

Physcosoma scalops: Monro, 1931, p, 31.

? Physcosoma agassizti: Fischer, 1922, p 7, pl. 1,

fig. 4,

Location of type: Royal Scottish Museum, Edin-

burgh; specimen from Natal, South Africa.

Deseriptian: This account is based oan the

specimens below and a collection from Hawati

(SAM E1176). Trunk slender, 1.5-3.5 mm long and

pale-straw to flesh coloured. Introvert of fixed

animals about as long as. trunk, with 3-9 brown ta

red-black bands of variable width on its dorsal

surface. Closely packed pigmenied papillae make

base of introvert darker than surface of trunk, Rust

coloured patches present in some places on trunk,

Trunk of some Hawaiian specimens uniformly pale

although some large papillae present posteriorly.

Introvert with almost complete ring of 20-24

digitiform tentacles lying dorsal to mouth and with a

smooth, collar-like region just posterior to mouth.

Introvert hooks in many complete and incomplete

rows. may vary in size even when taken trom same

animal, those more posteriorly placed being smaller,

Hook always possesses a clear streak (running from

apex to base), a clear erescentic area on ane side of

streak and a clear triangular area on other, a small

secondary “tooth” and 4-6 bars hasi-laterally, Rings

of very small, subconical papillae, 0:02-0-04 men tall

and 0-025-0-05 mm wide basally, lie between rows of

hooks.

Trunk papillae al posterior extremity of trunk

usually conical, 0-3-0-5 mm tall and 0-25-0-4 mm

wide basally, but sometimes a more rounded form is

present as Well. Conical form covered with small,,

brown polygonal plates but rounded form lacking

small plates and possessing a number of curved

petal-like plates radiating from an apical pore, It ts

possible that the rounded type is. the result of the loss

of the outer polygonal plates from the conical type,

Papillae on dorsal surface largest. Mid-trunk

papillae smaller, hemispherical, Jess pigmented and

more scattered.

Longitudinal muscles in 20-25 anastamosing

bands. Four retractors. a stouter ventral pair

attached to muscles 1-5, 2-6, 2-7 or 2-8 near middle

of trunk and a more slender dorsal pair more

anteriorly to 5-8, 4-7 or 6-8, One fastening muscle,

attached to muscle 1 near base ol left dorsal

retractor, runs to last spiral of intestine, sometimes

giving off a small reat to posterior oesophagus

Rectal caecum present (dissected specimens

SAM E1373 and E1531). Stephen in the type

description did not mention the presence or absence

of the structure, Wesenberg-Lund (1963. p. 124)

states with emphasis that no rectal caecum i$

present. Dr. G. Smaldon (Royal Scottish Museum),

however, informs me (personal communication) that

a caecim is present in Stephen’s dissected holotype.

Cantractile vessel without vill. Nephridia, about a

third as long us trunk and fixed for about half their

length, arise at about same level as anus.

Systematic position: PF, heronis was deserihed by

Edmonds (1956) from some Specimens collected at

Heron Is. in the Capricorn Group, Queensland.

Wesenberg-Lund (1963, p, 126) pointed out that the

hooks of P, heranis closely resembled those of P-

Stephensoni (Stephen. 1942) from South Africa.

Through the kindness of the Trustees of the Royal

Scottish Museum | have been able to examine three

of Stephen's specimens from Inhaca, Mozambique

(1960.48.14), one a dissected specimen. J now

consider that the two species are conspecific,

Stephen's having priority, The hooks of the South

African and Australian specimens are the same.

although slightly different in size, and the papillae of

Stephen's specimens are covered with fine plates,

something not mentioned by him but reported for

her South African specimens by Wesenberg-Lund

(1963), Tam grateful to the late Dr. Wesenberg

Lund tor her observations. Phascolosoma heronis

must be sunk.

It seems most likely that Fischer's (1922) record of

P, agassizti from “Port Jackson near Sydney” is that

of P. stephensoni; the drawing of the hook (pl. 1, fiz

6) matches that of the latter species. Fischer's fig 6,

however, in my opinion, differs from the books of P.

agassizji as shown in plates 37 and 38 of Fisher,

1952. What I am now calling P. stephensomti is. widely

distributed in the Pacific Ocean, I have specimens

from Hawaii, New Hebrides, Solomon Is.,.and Fiji.

It js puzzling to know what name has been given to

such specimens previously, ] do not think that they

are P. scolops because 4 re-examination of two

specimens named by Selenka himself and now

lodged in the British Museum (Nat. Hist) shows thal

the hooks Jack the clear, crescentic area which is so

prominent in P. srephensoni. The introvert hook of

P, fasciatum Baird, 1868 as shown in fig 17 of Rice &

Stephen (1970) also possesses a clear crescentic area,

The current name of P. fasciatum is P. granulatun

(Leuckart), I consider, however, that the hook of

68 REC. 8. AUST. MUS. 18 (1):

these Australian specimens does not match that of P.

granulatum as shown in Selenka & de Man (1883, fig

147).

Stephen considered that the distribution of P.

stephensoni was localised along the coast of Natal

(Wesenberg-Lund, 1963 p. 126). The occurrence of

the species in Australia, Norfolk Is., Hawaii, etc.

shows that its range is Indo-Pacific. What is difficult

to account for is that it has not yet appeared

anywhere else in the Indian Ocean nor on the

western coast of Australia. In Australia, Hawaii and

the Solomon Is, it is usually associated with coral

formations.

The species differs from closely related species in

the markings on its hooks.

Previous Australian record: Queensland—Heron

Is., Low Is. (Edmonds, 1956).

1 SS. indicus °

2 S. robustus e

3° § titubans r?)

4 X,mundanus +

§ S.cumanense Vv

6 S. vastum v

7 S. australe 2)

8 S. boholense @®

9 S.novaepommeraniaea

10 §. rotumanum &

1-74 January, 1980

Distribution: (1) in Australia—Queensland; New

South Wales; Norfolk Is., Lord Howe Is.

(2) elsewhere—South Africa; Hawaii (SAM

E1376-7); Solomon Is,

Specimens examined and localities: Queensland

—Caloundra (5) SAM E1372 and (1) AMS W19550;

Low Is. (3) SAM E1374; Albany Passage (North

Queensland) (2) AMS G11224; Heron Is. (3) AMS

W3597 (includes type of P. heronis) and (2) WAM

150/76; Norfolk Is. (2) SAM E1378. New South

Wales—off Coff’s Harbor (10) SAM E1373; Minnie

Waters (2) AMS W9247; Lennox Head (6) AMS

W10552; Arrawarra (2) AMS W10555; Lord Howe

Is. (2) SAM E1375; Lord Howe Is. (Salmon Beach)

(6) SAM E1531.

SIPUNCULIDAE

130

FIG. 130. Distribution of Sipunculidae in Australia.

.Margaritacea adelaidensis ©

.misakiana

. Schuettei

QDNHADDADRDADHAD

.trichocephala-a

AA ADHD VN VD

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 69

.coriacea y

_herdmani e fa

improvisa ° MO

minuta °

murinae murinae

ohlini

pellucida

semperi

.VulgariS queens. A

. collare >

. cronullae + GOLFINGIIDAE

dentalicolum v

pacificum x T. huttoni oO ©

cymodoceae 9% T. lageniformis ° Q

.dehamata 9 T. variospinosa co)

. fusca v O. steenstrupii &

FIG. 131. Distribution of Golfingiidae in Australia.

70 REC. S. AUST. MUS. 18 (1): 1-74 January, 1980

A,elegans

A,exhaustus

A. inquilinus

A.hartmeyeri

A. gracilis

A. jukesii

P, cumingi

P, formosanus

OMAN DOO FW p=

q~4a-PePP0gmgQ0 @O0O

P. steenstrupii

10 P johnstoni 4 ASPIDOSIPHONIDAE

11 C.aspergillus

12 L.cristatus

132

FIG. 132. Distribution of Aspidosiphonidae in Australia.

A REVISION OF THE SYSTEMATICS OF AUSTRALIAN SIPUNCULANS (SIPUNCULA) 7

,albolineatum

| annulatum

arcuatum

nigrescens

. nigritorqatum

noduliferum

. pacificum

NOonf® WN =

ua UU DU

> @e@ooese Ce

> @peg

v *O

ev o

le t

B P perlucens A

9 Prottnesti v

10 PR, scolops v

11 Pp stephensoni 0)

133

FIG, 133. Distribution of Phascolosomatidae in Australia.

REFERENCES

AKESSON, B, 1958. A study of the mervous system of the

Sipunculoideae, with some remarks on the development of

the two species Phascolion strombi (Montagu) and Golfingia

minuta (Keferstein). (In) Undersdkningar éver Oresund, 38

Lund, C. W. K. Gleerup, 249 pp.

AMOR, A, 1964. El genero Dendrostomum (Stpunculida) en Ja

Argentina. Physis, 24 (68); 457-470, 5 figs.

AUGENER, H. 1903, Beitrage zur Kenntnis der Gephyreen

nach Untersuchung der im Géttinger zoologischen Museum

befindlichen Sipunculiden und Echiuren, Arch. Naturgesch.

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RECORDS oF THE

SOUTH AUSTRALIAN

MUSEUM

VOLUME 18 NUMBERS 2—6 May, 1980

No. 2. THE DISTRIBUTION IN AUSTRALIA OF THE GRASS BUGS OF THE

TRIBE STENODEMINI (HETEROPTERA-MIRIDAE-MIRINAE)

by JOSE C. M. CARVALHO and GORDON F. GROSS

No. 3 TWO NEW GENERA OF THE SUBFAMILY PENTATOMINAE (HETER-

OPTERA: PENTATOMIDAE) FROM THE AUSTRALIAN REGION

by IMTIAZ AHMAD and NASEER AHMAD KHAN

No. 4 SOME REMARKS ON THE BREEDING DISTRIBUTION AND TAXON-

OMY OF THE PRIONS (PROCELLARIIDAE : PACHYPTILA)

by J. B. COX

No. 5 EVOLUTIONARY SYSTEMATICS OF XENYLLA: XI. SPECIES FROM THE

AUSTRALIAN REGION (INSECTA : COLLEMBOLA)

by MARIA MANUELA DA GAMA

No. 6 MACROPODID SKELETONS, INCLUDING SIMOSTHENURUS, FROM AN

UNUSUAL “DROWNED CAVE” DEPOSIT IN THE SOUTHEAST OF

SOUTH AUSTRALIA

by N. S. PLEDGE

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

THE DISTRIBUTION IN AUSTRALIA OF THE GRASS BUGS OF THE

TRIBE STENODEMINI (HETEROPTERA-MIRIDAE-MIRINAE)

BY JOSE C. M. CARVALHO AND GORDON F’. GROSS

Summary

The Stenodemini are represented in Australia by the three genera and species, Dolichomiris linearis

Reuter, 1882, Chaetedus longiceps Eyles, 1974 and Trigonotylus doddi (Distant, 1904).

A key is provided to separate these three species and figures and a brief diagnosis of each are given

to further assist in their recognition. Their known distribution in Australia is plotted on a map, and

in the case of the two less common species Dolichomiris linearis and Chaetodus longiceps, new

records to hand since the Eyles (1975) and Eyles & Carvalho (1975) revisions of the group are

given in detail.

THE DISTRIBUTION IN AUSTRALIA OF THE GRASS BUGS OF THE TRIBE STENODEMINI

(HETEROPTERA-MIRIDAE-MIRINAE)

JOSE C. M. CARVALHO! AND GORDON F. GROSS’

1. Museu Nacional. Rio de Janeiro, Brazil.

2. South Australian Museum, Adelaide, South Australia 5000-

ABSTRACT

CARVALHO, J. C. M, & GROSS, G, F., 1978, The distribution

in Australia of the grass bugs ot the tribe Stenodemini

(Heleroptera-Miridae-Mirinae) Rec, 5 Aust, Mus: 18(2); 75-42.

The Stenoderaini are represented in Australia by

the three genera and species, Dolichomiris linearis

Reuter. 1882, Chaeledus longiceps Eyles, 1974 und

Trigonotylus doddi (Distant, 1904).

A key is provided to separate these three species

and figures and a brief diagnosis of each are given to

further assist in their recognition. Their known

distribution ia Australia is plotted on a map, and in

the case of the two less common species Dolichamiris

linearis and Chaetodus longiceps, new records to

hand since the Eyles (1975) and Fyles & Carvalho

(1975) revisions of the group ure given in detail,

INTRODUCTION

The authors have in preparation a series ol papers

ia Which we shall attempt to give as complete an

account of the species of Miridae and their

distribution in Australia as is possible on the

material to hand in collections, with the exception of

the Bryocorinae and the non-mimetic Phylini which

have been entrusted to other authors.

In the paper which immediately preceded this

(Rec. S, Aust. Mus 17 (30): 000-000), though

covering the tribe Hyalopeplini of the subfamily

Miridae on a world basis, we were able to include all

revords and specimens of Australian material of that

group.

This paper deals with the allied tribe Stenodemini

in Australia, but it has been purposely kept short as

there have been two other recent contributions (vide

infra) on the taxonomy and distribution of the

Stenodemini in this general region.

Tribe Stenodemini

A small group of Miridae which was raised to

tribal rank in the subfamily Mirinae by China (1943),

This placement and ranking was accepted by

Carvalho (1952 and subsequent papers). The group

has subsequently been lowered to an unstated rank

in the Mirtnae by Schuh (1976) as a consequence of

his demotion of the subfamily Mirinae sensu

May, 1980

Carvalho to a tribe of an enlarged sublamily

Mirinaé. However, we propose fo retain the group

at its former rank for the time being.

The principal characters which taken together

distinguish this group from other Mirinae (sensu

Carvalho, or Mirini sensu Sehuh) are a rather

delicate appearance with the first segment of hind

tarsi longer than. (or as long as) the second and third

together, the antennae and legs relatively long and

the pronetal collar incomplete.

The group are grassfeeders, their elongate and

delicate shape presumably playing a role in

camouflage. Their small size and delicate stature, as

with a number of orher Heteroptera of similar size

and shape (c.f, Nabis species), appear to have

helped them ta be very vagile. They are known from

all googeographie regions with a number of genera

(e.g. Acelrapis Fieber, Collaria Provancher,

Dolichomiris Reuter, Leptapterna Fieber,

Mevaloceracu Fieber. Stenvdeme Laporte de Castel-

nau, Teratocoris Fieber and Trigonotylus Fieber) and

species (e.g. Dolichomiris linearis Reuter, Leptup-

rerma dolobrara (Linnaeus), Megaloceroed recticornis

Geotfroy, Stenodema [rixpingsumt Reuter.

Stenodema laevigatuim (Linnaeus), Stenodema virens

(Linnaeus), Teratocoris saundersi Douglas & Scott,

Teratocoris. viridis Douglas & Scott, Trigonotylus

doddi (Distant) and Trigonotylus ruficornis (Geot-

froy)) widespread in two or more zoogeographic

regions. The group is represented in Australia by

three genera and species, two of the species and

their genera are widespread. the third species is

endemic to Australia and New Zealand and belongs

to a genus (Chaetedus) known only from the

Australian, Papuan and New Zeuland regions, As

Eyles (1975) and Eyles and Carvalho (1975) have

recently reviewed these genera and species we give

here only a Summary account of the Australian

species. However, the distribution of the three

speci¢s which occur in Australia is much more

extensive than the above authors realised, so we

have mapped all locality records now available. bul

have cited in detail further records only of the two

less common species.

The three Australian species may be separated as

follows:—

76 REC. S, AUST. MUS_ 18(2); 75-82

Key to Australian Stenodemini

1. First antennal segment and base of second covered by long

erect pubescence, hairs as long as or longer than, width of

Segments, first antennal segment as long as {rom its

insertion to hase of pronotum

Dolichomiris linearis Reuter, 1882.

All antennal segments with only short pubescence, hairs

shorter than half width of segments; first antennal segment

much shorter than distance from its insertion to base of

ead; second

antennal segment with dark, bristlelike hairs near base;

ratio of length: width pronotum at base about 45:60; body

léngth in range 5-2-8-7 mm

Chaetedus longiceps Eyles, 1974

First antennal segment about as long as head; second

antennal segment without dark, bristle-like hairs: ratio

length: width pronotum about 20:33, body length in range

3-7-6-6mm) 0.55, Trigonotylus daddi (Distant, 1904)

Dolichomiris Reuter, 1882

Dolichomiris Reuter, 1882, p. 29. Additional

references and synonymy by Eyles & Carvalho,

1975, pp. 258, 267,

Type-species: Dolichomiris linearis Reuter, 1882.

For the type-species of synonymous genera see

Eyles & Carvalho opp.cit.

Body elongate and slender, smooth or very finely

punctured. Head horizontal, clypeus large and

visible from above, vertex sulcalte between eyes

which touch or nearly touch pronotum. Antennae

May, 1980

long, segments IT and II (at least basally) with long,

fine, erect, hairs which are at least as long as half

width of segment.

Pronotum with calli flat, smooth or very finely

punctured, collar depressed and weakly marked,

lateral margins carinate. Scutellum flat, smooth or

very finely punctate, Hemelytra smooth or very

finely rugose; cuneus distinctly longer than wide,

Legs long, hind tibiae covered with long, erect hairs.

Aedeagus with two spiculi,

Remarks: Only the type-species

Australia.

occurs in

Dolichomiris linearis Reuter, 1882

(Figs. 1-7, 16)

Dolichomiris linearis Reuter, 1882, p. 29. Additional

references and synonymy by Eyles and

Carvalho, 1975, pp. 258, 260-2 and by Carvalho

and Wallerstein, 1976, pp. 511-515.

Location of type: Lectotype ¢ in the Zoological

Museum of the University, Helsinki,

Colour: A faint, iridescent, pale brown with a

dark, reddish longitudinal stripe on head and a pale,

median longitudinal stripe on pronotum and

scutellum which is margined with light red, Eyes

grey, some red longitudinal marks laterally on head

and thorax, hind femora tending reddish brown,

Figs. 1-7—Dolichomiris linearis Reuter: Fig. 1—dorsal aspect:

Figs. 2-7—male genitalia; Fig, 2—vesica; Fig, 3—small

spiculum; Fig. 4-5—two aspects of left paramere; Figs, 6-

7—two aspects of right paramere,

GRASS BUGS OF THE TRIBE STESODEMINI V7

Structure: Very elongate, membrane considerably

surpassing apex of abdomen, hind femora not as

long as hemelytra. Long and erect hairs on antennae

restricted to first segment and base of second;

second and third antennal segments very long,

Male genitalia: Pygophore with a tooth on lett

hand side near upper corner, vesica (fig. 2) with two

spiculi, each with small sclerotized teeth (fig. 3).

Left paramere (figs. 4-5) falciform, tapering to apex.

Right paramere (figs. 6-7) smaller, constricted

medially, apex curved and pointed.

Length: 7:3-9-0 mm.

Distribution: Almost cosmopolitan, ranging from

southern Europe through east and west Africa, the

Canary Islands, Madeira, North, Central and South

America, the Galapagos Islands, India, Ceylon,

Burma, Java, Sumba, New Guinea, Queenstand, the

New Hebrides, New Caledonia (new record) and

Fiji.

Specimens examined: AUSTRALIA-Queensland

2¢¢ Redlynch, 31.viii. 1938, ex E,P. Van Duzee

collection May 1934, A, R, Brimblecombe; 2d,

Running Creek, 15.iv. 1941, A. W. Smith, all in

DPIO; Id. Rockhampton, 13.vii, L940, C. N.

Smithers, AM; 1¢, 14 km (9 mi.) n of Dayboro,

23.iv.1966, Z, Liepa, ANIC. These new localities

together with those of Eyles and Carvalho (1975)

have been platted on fig. 16.

tn addition we have examined material from the

following new localities outside of Australia. NEW

CALEDONIA—I1¢é 12, Mt, Quen Toro,

19.ix.1962, G. F, Gross; 19, by sweeping roadside

vegetation, 3 km. 5 of Touho, 10.x,1962, G. PF.

Gross, all in SAM. NEW HEBRIDES-BEfate, 124,

19, at light, Vila, 11 and 12.vii.1971, G. F. Gross on

Royal Society-Percy Sladen Expedition,

Erromanga, 1d 12 Ipotak, 5.viii.1971, G, F. Gross

on Royal Society-Percy Sladen Expedition, all in

SAM. FIJI Viti Levu, 12, sweeping and beating

herbs and shrubs, Nandarivatu, 800 m (2700 ft.),

10.xi.1964, N. McFarland; 19, Nagatosota Village,

1.1x.1976, J, A. Herridge SAM,

Remarks: According to Eyles and Carvalho

(1975) Dolichomiris linearis is more slender and

more delicate looking than other species of

Dolichomiris and has only a very finely punctate

pronofum,

Chaetedus Eyles, 1975

Chaeledus Eyles, 1975, p. 155-6; Eyles & Carvalho,

1975, p. 268.

Type-species:

1878. O.D.

Megalocervea reuteriana White,

Body elongate and slender, smooth with pro-

notuim dorsally finely punctured. Head horizontal,

clypeus large and visible from above; frons striate,

vertex sulcate between eyes which do not touch

anterior margin of pronotum. Antennae long,

segments I and IT (at least basally) with short,

bristle-like, dark hairs which are not as long as width

of segment,

Pronotum with calli flat, finely punctate except on

calli, collar depressed and weakly marked, lateral

margins thickly carinate except anteriorly. Scutellum

nearly flat. Hemelytra smooth or very finely rugase,

cuneus longer than wide. One of two spiculi of

aedeagus hooked, serrate or plumose.

Remarks: Chaetedus Eyles has four species,

reulerianus (White) is restricted to New Zealand,

longiceps Eyles occurs in beth Australia and New

Zealand, plumalis Eyles in Norfolk and the Raoul

Islands and rutilans Eyles in New Guinea,

Chactedus longiceps Eyles, 1975

(Figs, 8-16)

Chaetedus longiceps Eyles, 1975, pp, 156-7, 162,

Location of type: Holotype &

Entomology Division, DSIR,

Zealand

and allotype 2 in

Auckland. New

Colour: Fresh specimens green but later fading to

stramineous. Eyes grey, Antennal segments Il to 1V

red or suffused with reddish. pronotum with four

longitudinal dull stripes between inner stripes pale.

Scutellum with two longitudinal. dull stripes

between these paler. Apices of tibiae and tarsi red or

sulfused with reddish.

Structure; Second and third antennal segments

equal in length and each about three times as long as

first, fourth equal in length to first. Scutellum faintly

strigose on either side of midline. Hemlytra longer

than hind femora, membrane considerably surpas-

sing apex of abdomen.

Male genitalia: Pygophore (figs. LO-LL) with a

comical process on left side; vesica (fig. 12). both

spiculi sclerotized with the usual plumosity on one

reduced to minute teeth (fig. 13), Left paramere

(fig. 14) falciform and tapering to apex pointed and

curved apically. Right paramere (fig. 15) pointed

anically.

Length: 68-8 T mm.

Remarks: Species of this genus were formerly

thought to be all Megalocveroea reuteriana White

until Eyles (1975) showed that four species were

present in our general region and belonged to a new

genus which ditferes from Megaloceroea Fieber in,

inter alia. having the first antennal segment distinctly

78 REC. S$. AUST. MUS. 18(2): 75-82

May, 1980

Figs. 8-9—Chaetedus longiceps Eyles: Fig. 8—dorsal aspect; Fig.

9—lateral view of head and thorax.

shorter than the head and pronotum together and

one of the spiculi of the aedeagus, hooked, serrate

or feathery. Chaetedus reuterianus and C. longiceps

are green in life whereas the two species with a more

tropical distribution, plumalis and rutilans, are

reddish. C. reuterianus has a prominent, red, lateral

abdominal stripe which is absent in longiceps.

Distribution: New Zealand and Southern Aus-

tralia, including Tasmania.

Specimens examined: AUSTRALIA: New South

Wales, 1d, Oxford; 31.xii.1962, D. K. McAlpine;

Id,12, Mt. Gibraltar National Park, 24.17.1965, D.

K. McAlpine, all in AM; ¢ Rydalmere, at Mercury

vapour light, 15.x.1971; Broadwater NSWDA,

A.C.T. 19, Canberra, 27.iii.1959, G. E. Chadwick,

NSWDA. Victoria 12, Glen Waverley, sweeping

Gramineae and Campositae in garden,

19-31.xii.1976, J, J. H. Szent-Ivany. South Aus-

tralia, 32 2, Mylor, Nov. 1949, G. F. Gross, SAM.

Western Australia, 19, Walpole-Nornalup National

Park (35° 00'S, 116° 49’E) at The Knoll, 11.xi.1969,

E. B. Britton, ANIC. These localities and those of

Byles (1975) for Australia are plotted on fig. 16.

GRASS BUGS OF THE TRIBE STENODEMINI 79

|co-

Figs. 10-15—Chaetedus lonpiceps Eyles, male genitalia: Fig

[(}—ventral aspect of pygophore; Fig, | 1-—lateral aspect of

same; Fig, 12—vesica; Fig. 13—sclerotized spiculum, Fig.

14—left paramere; Fig, 15——-right paramere.

Trigonotylus Fieber, 1858

Trigonotylus Fieber, 1858, p.302. Other references

and synonymy in Eyles, 1975, pp. 153, 162.

Type-species: (of Trigonorylus) Miris ruficornis

Fallen, 1807 (=Cimex ruficarnis Geottroy in

Fourcroy, 1785) O.D,

Smallish and elongate, pronotum punctate. Head

horizontal, clyqeus large and visible from above,

frons usually prolonged over base of clypeus, vertex

suleate medially between eyes, eyes touching

pronotum, Antennae long, first segment not as long

as head and pronotum together, without black

bristle-like hairs.

Pronotum depressed laterally, collar depressed

and weakly marked, calli flat, lateral margins

carinate, Scutellum flat and smooth. Hemelytra

smooth and glabrous, cuneus distinctly longer than

wide. Legs relatively jong. Acdeagus with a single

spiculum, or spiculi lacking,

Remarks: Only the one cosmopolitan species is

represented in Australia. The genus was last revised

by Carvalho and Wagner 1957.

Trigonotylus doddi (Distant, 1904)

(Figs. 16-24)

Megaloceroea doddi Distant, 1904, p. 269; other

references and synonymy in Eyles 1975, pp.

162-4.

Location of type: Lectotype ¢ in British Muscum

(Natural History).

Colour: Fresh specimens green but later fading to

stramineous. Eyes grey. Antennae suffused with

pink, Pronotum and scutellum with a pale median

stripe bordered by a pale brown or red stripe on each

side, on pronotum on each side are additional fainter

brown or red stripes between the median stripes and

lateral margin. Apex of hind tibia, first two

tarsomeres and base of third of hind leg red, claw

and apex of last tarsomere of hind leg black, Corium

paler between veins R and M and costal margin,

Beneath lateral margins sometimes reddish.

Structiire: Second antennal segment a little longer

than third and both about three times as long as first.

fourth a little shorter than first. Hemelytra longer

than hind femora, membrane considerably surpas-

sing apex of abdomen,

Male genitalia: Pygophore (figs. 54-55) with a

conical spinous process on left side; aedeagus (fig.

56) with membranous lobes and one sclerotized

spiculum. Left paramere (figs. 57-58) falciform with

apex tapering and sinuate, right paramere (fig, 59)

small with a spinous apical process.

Length: 4-2-5-7 mm.

Remarks: This is much the most common and

widespread of the three Stenodemini species in

Australia and differs from Dolichomiris linearis and

Chaetedus longiceps in ats much smaller size and very

much shorter first antennal segment,

Distribution: Widely distributed mainly in tropical

and subtropical areas but in New Zealand and

Australia extending into the temperate zones and tn

the latter also into the arid regions, Trigonotylus

doddi has not been recorded from Tasmania, New

Caledonia, New Hebrides, Polynesia east of the

Gilbert and Kermadec Islands, Solomon Islands,

New Guinea or Indonesia. We have seen material

from the Cocos, Keeling and Fijian Islands from

which there were no previous records.

Specimens examined: 7 ex Department of Primary

Industries, Queensland; 9 ex QM) 10 ex ANIC; 4ex

AM: 15 ex New South Wales Department of

Agriculture; 2 ex NM; 69 ex SAM; 1 ex Western

Australian Department of Agriculture; 1 ex

Zoological Museum of the University, Helsinki.

As this species is now known to be widespread in

Australia we have not listed all the localities in detail

but have plotted the Australian records of Eyles

1975 and all the additional material we have

examined on the map on figure 16.

80 REC. S. AUST. MUS. 18(2): 75-82 May, 1980

DARWI

@+Hall’s Creek

\Q - iS ene

\ j e

val

PERTH

Albany

Dolichomirus linearis

4 records by Eyles & Carvalho

A new records

Chaetedus longiceps

0 records by Eyles

@ new records

Trigonotylus doddi

@ new records

Fig. 16—Distribution in Australia of Dolichomiris linearis Reuter,

Chaetedus longiceps Eyles and Trigonotylus doddi (Distant).

ane

oi ey

ADELAIDE op

‘ ’

Cairns

©Normanton

Mt. Morgan * 4

fe) Ao® "

j fo) , _7 BRISBANE A+O

—~, @ OR .

Cunnamulla _ 2S —4+0(Tambourine)

=a 5

[eR eo 5 yee ot us

*: e

? ss

SYDNEY gi @+ (Rydalmere)

~ a

A ak re

MELBOURNE: t

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HOBART,

GRASS BUGS OF THE TRIBE STENODEMINI 81

Figs. 17-18—Trigonotylus doddi (Distant): Fig. 17—dorsal aspect;

Fig. 18—lateral view of head and thorax.

82 REC. S. AUST. MUS. 18(2): 75-82

Figs. 19-24—Trigonotylus doddi (Distant) male genitalia: Fig.

19—ventral aspect of pygophore; Fig. 20—lateral aspect of

pygophore; Fig. 21—aedeagus and vesica; Figs. 22-23—1wo

aspects of left paramere; Fig. 24—right paramere.

May, 1980

REFERENCES

CARVALHO, J. C. M. 1952. On the major classification of the

Miridae (Hemiptera), (With keys to subfamilies and tribes

and a catalogue of the world genera.) Ann. Acad. bras. Sci.

24 (1): 31-110.

CARVALHO, J. C, M. 1978. The tribe Hyalopeplini of the

world. The Australian fauna in collaboration with Gordon F,

Gross. Rec. S. Aust. Mus. 17 (30): 429-531.

CARVALHO, J. C. M. and WAGNER, E. 1957. A world

reyision of the genus Trigonotylus Fieber. (Hemiptera-

Heteroptera, Miridae.) Arg. Mus. nac. Rio de J, 43: 121-155.

CARVALHO, J. C. M. and WALLERSTEIN, P, 1976. Revision

of the genera Anthropophagiotes, Nesosylphas and Notostira

pacifica Kirkaldy, 1908 (Hemiptera, Miridae). Rev. brasil.

Biol. 36 (2): 511-515.

CHINA, W. E. 1943, The generic names of British Insects—Part

8, The generic names of the British Hemiptera-Heteroptera,

with a check list of the British species."’ (Royal Ent. Soc.;

London.)

DISTANT, W. L. 1904. Rhynchotal notes—XXIH. Heteroptera

from North Queensland. Ann. Mag. nat. Hist., 13: (7): 263-

276.

EYLES, A. C. 1975, Further new genera and other new

combinations for species previously assigned to Megaloceroea

(Heteroptera: Miridae: Stenodemini), J, nat. Hist., 9 (2):

153-167.

EYLES, A. C. and CARVALHO, J.C. M. 1975. Revision of the

genus Dolichomiris, with a revised key to the genera of

Stenodemini (Heteroptera: Miridae). J. nat. Hist., 9 (3): 257-

269.

FIEBER, F. X. 1858. Criterien zur generischen Theilung der

Phytocoriden (Capsini auct). Wiener Ent. Monatsch. 2: 289-

327.

REUTER, O. M. 1882. Ad cognitionem Heteropterorum Africae

Occidentalis, Ofv. F. Ver. Soc. Férh. 25: 1-43.

SCHUH, R. T. 1976. Pretarsal structure in the Miridae

(Hemiptera) with a cladistic analysis of relationships within

the family. Amer. Mus. Novit, 2601; 1-39.

TWO NEW GENERA OF THE SUBFAMILY PENTATOMINAE

(HETEROPTERA: PENTATOMIDAE) FROM THE AUSTRALIAN REGION

BY IMTIAZ AHMAD AND NASEER AHMAD KHAN

Summary

Two new genera, Grossiana and Knightiella, are described to accommodate Strachia persignata

Walker and Stenozygum flavifrons Distant known from Queensland and New Caledonia

respectively. The above species are redescribed, the characters of their metathoracic scent gland

ostioles and male and female genitalia are described for the first time and their position in the

subfamily Pentatominae is briefly discussed.

TWO NEW GENERA OF THE SUBFAMILY PENTATOMINAE (HETEROPTERA: PENTATOMIDAE)

FROM THE AUSTRALIAN REGION!

IMTIAZ AHMAD and NASEER AHMAD KHAN

Department of Zoology, University of Karachn, Karachi, Pakistan

ABSTRACT

AHMAD, L., & KHAN, N. A. 1979. Two new genera of the

subfamily Pentatominae (Heteroptera!Pentatomidae) trom the

Australian region. Rec, S, Austr, Mus. 18(3); 83-90,

Two new genera, Grossiana and Knightiella, are

described to accommodate Strachia persignata

Walker and Stenozyguin flavifrons Distant known

from Queensland and New Caledonia respectively,

The above species are redescribed, the characters of

their metathoracie scent gland ostioles and male and

female genitalia are described for the first time and

their position in the subfamily Pentatominae is

briefly discussed.

INTRODUCTION

Strachia persignata Walker, 1867 and Stenozyguin

flavifrons Distant, 1914 were described trom

Queensland and New Caledonia respectively in the

Australian region, During a revision of Srenozygum

Fieber, 186) from the Oriental and Australian

regions by the present authors the above species

were compared with the type-species of the genus

and with all the Oriental and Australian species

described under the genera Strachia Hahn, 1833 and

Stenozygum and were found to be very different,

especially in the characters of metathoracic scent

gland ostioles and male and female genitalia, These

two species are redescribed here with special

reference to the above-mentioned characters and

placed each in a new genus, Grossiana and

Knightiella respectively. Their position in the family

is briefly discussed. For the dissection of male and

female genitalia, for measurements and diagrams

the conventional procedures, especially those of the

present authors (in press) have generally been

followed.

Genus GROSSIANA New Genus

Body brilliantly patterned.

Head: Eyes stalked; basal antennal segments

uniformly thick and slightly inwardly curved,

distinctly passing beyond apex of head, labium

reaching posterior coxae,

Thorax; Metathoracic scent gland ostioles without

peritreme and with ill-defined evaporatoria,

' Financially supported by an ARC-USDA Research Project

A17-ENT-37, (FG-Pa-181),

Male genitalia; Pygophore with latero-posterior

lobes distinct but not detached or marked from the

rest of the pygophore;, parameres simple, sickle-

shaped; inflated aedeagus with vesica distinctly

shorter than penial lobes and apically semi-

sclerotized, dorsal membranous conjunctival appen-

dage, and ventral conjunctival appendage fully

membranous.

Female genitalia: Female terminalia with triangu-

lin visible in between Ist gonocoxae, with ends

pointed and reaching to inner anterior margin of Ist

gonocoxae, spermatheca without processes on

spermathecal bulb.

Comparative note: Grossiana is somewhat closely

related to Strachia in having the basal antennal

segment long and surpassing apex of head and

second antennal segment subequal to third, but it

can easily be separated from all the species of that

genus in having a simple unilobed pronotum as

opposed to distinctly laterally bilobed in the specics

of Strachia and in other characters noted under

“Systematic positions”.

The genus is named in honour of Dr, G. F, Gross.

of South Australian Museum, Adelaide, Australia to

acknowledge his work on Heteroptera.

Type-species: Strachia persignata Walker, 1867.

Grossiana persignata (Walk.)

(Figs. 1-10)

Strachia persignata Walker, 1867, p, 347.

Stenozygum persignatum Distant. 1881, p. 213.

Coloration; Body shining black with prominent

yellow markings (fig. 1), uniformly punctured except

head and anterior portion of pronotum, ventrally

yellow with a small black patch in the middle on

either side along the margins of bucculae, a small

black spot at the base of antenniferous tubercles on

either inner side, broad black longitudinal lines on

outer portion of pro-, meso-, and metapleuron, on

either side, black punctures on the rest of pro-.

meso-, and metapleuron, dark brown 2nd, 3rd and

4th segments of labium, 4 black transverse patch

posterior to spiracles on 2nd-7th abdominal

segments and a black transverse patch on outer

portion along the lines between 2nd-7th abdominal

segments on either side,

Rd REC. S. AUST, MUS, 183): 83-90 May, 1980

Fig. 1—Grossiana persignata (Walker) female, dorsal view.

TWO NEW GENERA OF THE SUBFAMILY PENTATOMINAE 84

Head: Anteocular portion equal to the posterior

portion of head including eyes, length of anteocular

portion 0:75 mm, length of posterior portion of head

including eyes 0-75 mm (0:65-0-80 mm); width

including eyes 2:30 mm (2-20-2:30 mm); interocular

distance 1-1] mm; interocellar distance 0:80 mm

(0-80-0:90 mm); paraclypei with tips slightly

separated from each other. terminating above and

subequal to clypeus, basal antennal segments. well

passing the apex of head, 3rd segments slightly

longer than the 2nd antennal segments; length ot

segments 1, 0:50 mm (0-50-0-55 mm), H, 1-10 mm

(0:-00-L-1 mm), Hf, 1:20 mm (1-L0-1-20 mm), IV,

140 mm, V, mutilated, antennal formula

1<2<3<4; Labium reaching posterior coxae, Ist

segment slightly extending beyond bucculac, latter

nal reaching to the base of head; length of segments

1, 0-80 mm (0°70-0-80 mm), IT, 0-95 am ((°835-0-95

mm), UI, 0-60 mm (0-55-0-66 frm), IV, 0-70 mm

(0-65-0:70 mm), labial formula 3<4<1<2,

Tharax: Pronotim 25x wider than long, length

1-70 mm (1:60-1:80 mm), width 4:30 mm (3-70-4-30

mm); scutellar length about 1 x the length of

pronotum, scutellar length 3-10 mm, width 2:70 mm

(2:65-2:70 mr); metathoracic scent gland ostioles

small elliptical, without pentreme and with ill-

defined evaporatoria; distance from base of

scutellum to apex of clayus 2-15 mm, apex of

scutellum to apex of clavus 0-985 mm: apex of

scutellum to apex of abdomen (including mem-

brane) 3-10 mm (2:6-3-10 mm),

Abdomen: Anterior margin of 7th abdominal

sternum acute in male and medially convex in

female, posterior margin medially broadly concave

in male and medially bilobed in female, connexiva

slightly exposed at repose. Total length ¢ 7-60 mm,

2 9-40 mm (9:05-9:40 mm).

Male genitalia: Pygophore (figs. 2-4) nearly 2x

wider than long, ventro-posterior margin medially

concave with a pair of quadrate proecsses consisting

of a group of hairs al their apices on either inner side

adjacent to median concavity, procter sclerotized;

parameres (figs. 5 and 6) with comparatively short

straight stem and elongated tapering curved blade

and spine-like inner process at junction of stem and

balde; inflated aedeagus (figs. 7 and 8) with a more

or Jess cuboidal theca. broader posteriorly than

anteriorly and with minute lateral thecal appen-

dages, conjunctiva along with vesica enclosed in a

cup-shaped structure with a hexagonal ventral

surface and more or less trapezoidal dorsal surface,

dorsal membranous conjunctival appendage taper-

ingly bilobed and semisclerotized apically, ventral

membranous conjunctival appendage bilobed and

shorter than dorsal membranous conjunctival

appendage, penial lobes fused, enclosing a slightly

longer vesica.

Female genilalia: (figs, 9 and 10) Posterior margin

of triangulin slightly convex; exposed part of Ist

gonocoxae 3x broader than long; 9th paratergites

with apices truncate with apical angles distinctly

touching the posterior margin af 6th paratergites:

spermatheca (fig. 10) with spermathecal median

dilation divided into a posterior large and anterior

small pouch by a constriction, spertnatheca with

distal duct slightly dilated, bulb without processes,

Material exantined: Wolotype, Austraha,

"'Strachia persignata (Walker) “57-130 “96" in

British Museum (Natural History) Lovdon, | ¢.2 2

Australia: Queensland, Rockhampton; [ ¢d Aus-

tralia: Brisbane, Queensland Mus, R. Hamlin Harris

1914-202; “647; 1 @ Australia: Goodna, 10-11-24,

R. Backer 163" ‘Australia, British Museum, 1926-

241° - Australia: Peak Downs”, Distant Coll. 1911-

“3R3°-9 2,2 2 Australia, in B.M.(N,H.);bd,2 4%,

Australia, Queensland: Bundaberg, Gayndah,

Rockhampton, A.M. Lea det.A.Musgrave in South

Australian Museum, Adelaide: Australia.

Genus KNIGHTIELLA New Genus

Body predominantly green and less brilliantly

patterned than other genera of Strachiini,

Head: Eyes sessile; basal antennal segments

uniformly slender, straight, distinctly surpassing

apex of head; lahium reaching to 4th abdominal

sepment

Thorax: Metathoracic scent gland ostioles with

elongated well developed peritreme and well

defined evaporatorja-

Abdomen: Strigose vittae present on 2nd, 3rd and

4th abdominal segments,

Male genitalia’ Pygophore without distinct latero-

posterior lobes; parameres F-shaped, inflated

aedeagus with vesica distinctly longer than penial

lobes and dorsal membranous conjunctival appen-

dage.

Female genitalia: Female terminalia with triangu-

lin visible in between Ist gonocoxac, large,

rectangular-shaped and reaching to middle of inner

margins of Ist gonocaxae; spermatheea with bulb

having one lang and one short processes.

Comparative note: Knightiella is isolated in the

entire tribe Strachiini in the characters of predomin-

antly green and less patterned body, uniformly

slender basal antennal segments passing distinctly

beyond apex of head, and in the characters o!

metathoracie scent gland osticles and male and

female genitalia noted under systematic positions

& REC. S. AUST. MUS. 18(3): 83-90

Mey, Take)

EN Nic Snneay met

eon

0.60mm

Figs, 2-10—Grossiana persignata (Walker),

vy"

‘

(

‘i

0.60mm

Pygophore; Fip

2—Dorsil view: Fig 3—Ventral view: Fig, 4—Horizantal

view, paramere: Fig. 5—Dersyl view: hig. h—Vontral view,

aedeagus; Fig. 7-Dorsal view: Pig. 8—Ventral view. Fig.

Y—Femule terminulia, ventral yiew: Fig. \0—Spermatheva,

ventral view,

The genus Knightiella is named in the honour of

Dr. W. J, Knight who is in charge of the Hemiptera

section, Britsh Museum (Natural History) London

to recognise his work on Hemiptera.

Type-species: Stenozyguin flavifrans Distant, 1914.

Knightiella flayvifrons (Dist, )

(Figs. 11-21)

Stenozygum flavifrons Distant, 1914, p. 375.

Coloration; Body green with prominent yellow

and light green markings, uniformly punctured

except head (fig. 11); ventrally green with head,

thoracic sternum, posterior coxae, lateral margins of

propleuron, small outer portion of mesopleuron,

postenior margin of metapleuron yellow, 4th and

basal portion of Sth labial segments brown, and apex

of labium black, portion around ostioles and

peritreme white, lateral und a little outer posterior

margin of each abdominal segment and broad

median protion of 2nd-6th abdominal segments

yellow,

Head: Anteacular portion equal to the posterior

portion of head including eyes, length of anteecular

portion 0-65 mm (0:65-0-70 mm), length of posterior

TWO NEW GENERA OF THE SUBFAMILY PENTATOMINAE 87

portion of head including eves 0-65 mm (0-65-0:70

mim); width including eyes 1.75 mm (1-75-1-85 mm);

interovwlar distance (9S mm (0-95-L-05 mr),

interocellar distan¢e 0-40 mm (0-60-0-65 mm);

paraclypei With tips separated and subequal to the

clypeus; basal antennal segments. uniformly slender,

straight, distinetly passing the apex of head, 3rd

segments slightly longer than ['/2 x the length of 2nd

antennal segments; length of segments T. 0-45 mm

(0:45-0:50 mm), 11, 0:60 mm, 11, 1-00mm, 1V, & V.

mutilited, labium teaching to 4th abdominal

segment, Ist sogment distinctly extending beyond

bucculae, latter not reaching to the base of head,

length of segments 1, 0-95 mm, 11, 1-10 mm, HT, 0-60

mm, LV, 0-65 mm, labial formula 3<4<1<2

Thorax: Pronotum slightly more than 2-5 x wider

than long, length 1-30 mm (1-30-1-40 mm), width

3-40 mm (3-40-3-80 mm); scutellar length about L's x

the Jength of pronotum, scutellar length 2-35 mm

(2:35-2-55 mm), width 2-20 mm (2-20-2-40 mm);

metathoracie scent yland ostioles (fig, 12) with

075mm

Fig. Ll—Knightiella flavifrony (Dyst ) male dorsal viene

88 REC. 8. AUST

elongated crescent-shaped peritreme and well

defined evaporatoria; distance base of scutellum to

apex of clayus 1:60 mm (1:60-1:80 mm); apex of

scutellum to apex of clavus 0-75 mm; apex of

scutellum fo apex of abdomen (including mem-

brane) 2-05 mm (2-05-2-20 mm).

Abdomen: Anterior margin of 7th abdominal

sternum medially broadly and smoothly convex and

posterior margin medially broadly and smoothly

concave in male and female; connexiva nat exposed.

Total length ¢ 7-0 mm, 2 7-55 mm,

Male genitalia: Pygophore (figs. 13-15) nearly 3x

broader than long in middle, ventroposterior margin

medially broadly concave with a pair of elongated

lobe-like appendages on either side of median

concavity (visible in posterior view only); proctiger

sclerotized posteriorly and laterally while remaining

portion thin and membranous; parameres (figs. 16

and 17) with a short foot-like stem and a long

itregularly-shaped blade with outer three bulges,

large inner tooth and distal curved part rounded

apically, inner tooth and distal part of parameres

MUS. 18(3); 83-90 May, 1980

imparting a more or less F-shaped appearance ta this

structure; inflated aedeagus (figs. 18 and 19) with

elongated theca tapering posteriorly without distinct

lateral thecal appendages, dorsal membranous,

conjunctival appendage bilobed, equal to penial

lobes, latter fused anteriorly and terminating into

truncate apices,

Female genitalia: (figs. 20-21) Posterior margin of

triangulin with an obtuse angle in middle; exposed

part of Ist gonocoxae slightly broader than long; 9th

paratergites with apices rounded, with apical angles

not touching the posterior margin of 8th parater-

gites; spermatheca (fig. 21) with elongated median

dilaton, without any constriction and with uniform

distal duct and two processes on the bulb, one

thicker and larger as compared to other.

Material examined: 2 3, 1° New Caledonia: Ba

Bay. 5-VI[-19174, P. D. Montague 1918-87 "New

Caledonia Exped", Stenozygum flavifrons Dist. "C,

H, Lyal det. 1977 British Museum (Natural History)

London,

Figs, 12-21—Knightiella flavifrons (Dist.); Fig. 12—Metathoracic

scent gland ostioles, pygophore; Fig, 13—Dorsal view; Fig.

I4—Ventral view; Fig. 15—Horizontal view, paramere; Fig.

16—Dorsal view: Fig. 17—Ventral view; uedeagus; Fig.

18—Dorsal view; Fig, 19—Ventral view: Fig. 20—Female

terminalia, ventral view; Fig, 2)—Spermatheca, ventral

view.

TWO NEW GENERA OF THE SUBFAMILY PENTATOMINAP 89

SYSTEMATIC POSITIONS

The Genus Knightiella is not only very different

fram all the species of the genus Stenozygum but is

completely isolated in the entire tribe Strachiini Stal,

1876 (Strachia group Gross, 1976) on the basis of

metathoracic scent gland ostioles with well

developed peritreme and completely defined

evaporatoria, 4edeagus with vesica distinetly longer

than penial lobes, parameres F-shaped and sper-

mathecal bulb with processes, It has stvigose vittae

on UI, Il and JV abdominal segments laterally

similar to those in the members of Mecideini

Distant, 1902 and of Diemenia group Gross, 1976.

The members of the former group are gencrally

elangated and slender having shorter peritreme in

the metathoriacic scent gland ostioles and have lobe-

like short and rounder blade without inner lobe in

parameres, acdeagus with elongated and conical

lateral conjunctival lobes and the spermatheca is

without pracesses on the bulb, In the latter group

the paraclypei generally surpass the clypeus and may

be reflexed laterally and often produced into small

lobes or processes in front of the eyes, the aedeagus

has prominent bilobed conjunctiva with or without

addijional median and ventral lobes aod the vesiva Is

short and robust (Sailer 1952, Abbasi 1978, Gross

1976).

On the other hand F-shaped parameres ure found

in the species of Carpovorin Stal, 1876 (Curpaceris

group Gross, 1976) including the speeics of

Avonoscelis Spinglu, 1837 for whieh Sal (1876)

formed his division Agenoseelaria and this was also

follawed by Atkinson (1888). Similarly V-shaped

parameres are found in many species of Halyi Stal,

i872 (Halys group Gross. 1976) as reported by

Ahmad and Abbasi (1974), Abbasi and Alimad

(1975), Gross (1976) and Ahmad and Afzal (in

manuscript). Some of the species af Diemenia wroup

also have rather blade-shaped parameres which

could look vaguely P-shaped (Gross. personal

communication), But in none of these groups a

combination of charaeters as listed in the generic

description are found. Either it should have its own

group within the subtamily Pentatominae or its

position remain unascertained until # world wide

revision of the subfamily is completed.

On the contrary the genus Grossiana certainly

belongs to the tribe Strachiini in the characters of

metathoracic scent gland ostioles without peritreme

and with ill-defined evaporatoria, brilliantly col-

oured body, somewhat stylate eyes, simple sickle-

shaped parameres. in the male and_ bilobed

spermathecal median dilation without processes on

the bulk of the females. However Grossiana differs

remarkably from all the species of the genus

Stenozygum in the characters of the long, uniformly

swollen and slightly inwardly curved basal antennal

segment surpassing apex of head (in S. laelibile

(Walker) the basal antennal segment distinctly

surpasses the apex of head but the second is

distinctly shorter than the third) and the second

antennal segment subequal to third, The latter is

also found in the species of Sirachia bul the

prominent laterally bilobed pronotum, the simple

hook-like parameres, and the medially deeply

concave posterior margin af the seventh female

abdominal sternum in all the species of Strachia

(Ahmad & Kamuluddin 1978) clearly separate the

two gentra,

ACKNOWLENGEMENTS

The present wuthors are deeply indebted to Dr,

W. J. Knight. in-charge of Hemiptera Seetion, to

Dr P. Freeman Keeper. Department of Ento-

mology aud the trustees of the British) Museum

(Natural History) Londen for their kind permission

for the first author fo examine the genitaha of the

holotype of Srruchia persignata and other identified

specimens of the above species and Stenozyguin

flavifrons Distant laged at their Museum during his

receur visit (1977-78) to their Museum, Dr. G F,

Gress of South Australian Museum, Adelaide,

Australia is sincerely acknowledged for lnaning

three identified Specimens of persignaium (Lo .2 2)

and for critically reading the manuscript

REFERENCES

ABBAS. G. A. 1974 A new speaes of a palacarctic penus

Holeosiethus Weber (Pentatominae; Carpocorni) fron

Pakistan Karn Univ f Se, § (1 & 2), 81-84,

ABBASL, O.A. and AHMAD, |. 1975. A new species of the

genus Hayly Faber (Pentatomidae! Halyini) from East Bengal

With notes on jls genitalia dnd there bearing on classification,

Kar, Univ. Se. 4 (1-2); 35-41

AHMAD. Ll. and ABBASI. Q. A. 1974, a new genus and species

of Halyini Stal with phylagenetic considerations. Kar. Liniv,

J. Se. 3 (1-2): 61-65.

AHMAD, land KAMALUDDIN. S, 1978. Two new species of

the genus Sirachia Hahn (Pentatominae; Strachiini) trom

Bangladesh with a taxonamic note on the Tribal name. Kar,

Uni. J. Se. 6 (i): 35-41.

ATKINSON, FE. T. i888, Notes on Indian Rhynchota;

Heteroptera. /. Asiat. Soc, Beng, 23 66-69,

DISTANT, W. L. L881. Rhynchota from the Australian and

Pacific regions. Trans. ent. Soe Lond. 188: 211-218.

DISTANT, W. L. 1902, The fauna of British India, including

Ceylon and Burma, Rhynchota. 1. Taylor and Francis,

London,

DISTANT, W. L. 1914. Rhynchota from New Caledonia and the

surrounding islands, In Sarasin. F, and Roux, J. Nowa

Caledonia, Zaologie 1: 369-390,

FIEBER. F. X, 1861, Die Europaischen Hemiptera Halbfliigler

(Rhynchota: Heteroptera): 113-444, Carl Gerold’s Sohn,

Wien.

90 REC. S. AUST. MUS. 18(3): 83-90

GROSS, G. F. 1976. Plant feeding and other bugs (Hemiptera) of

South Australia. Heteroptera. Part II. Government Printer,

South Australia.

HARN, C. W. 1833. Die Wanzenartigen Insecten (I-III). C. H.

Zeh’schen, Niirnberg.

SAILER, R. I. 1952. A review of the stink bugs of the genus

Mecidea. Proc. U.S. Nat. Mus. 102: 47-505.

May, 1980

STAL, C. 1872. Enumeratio Hemipterorum. K. svensk. Vetensk.

Akad. Handl. 3: 31-40.

STAL, C. 1876. Enumeratio Hemipterorum (1). Bidrag till on

fortechning of ver alla hittills Kanda Hemiptera Janta

Systentiska Meddentandon and Stockholm. K. svenska

Vetensk. Akad. Férh. 1: 1-232.

WALKER, F. 1867. Catalogue of the specimens of Heterop-

terous—Hemiptera in the Collection of the British Museum,

Pt. 2. British Museum, London.

SOME REMARKS ON THE BREEDING DISTRIBUTION AND TAXONOMY

OF THE PRIONS (PROCELLARITIDAE: PACHYPTILA)

BY J. B. COX

Summary

The breeding distributions of prions are reviewed and the characters used to differentiate Pachyptila

vittata, P. salvini, P. desolata, P. belcheri, P. turtur and P. crassirostris are critically assessed.

Feeding habits and the concept that differing breeding times isolate these prions genetically are also

discussed. Evaluation of the evidence indicates that their previous classification into three genera or

sub-genera is invalid because only two groups are clearly separable by morphological

characteristics: the Fairy Prions (turtur and crassirostris) and the Whale-Birds (belcheri, desolata,

salvini and vittata). The former are considered to be one polytypic species and the whale-birds to be

one monotypic and one polytypic species.

SOME REMARKS ON THE BREEDING DISTRIBUTION AND TAXONOMY OF THE

PRIONS

(PROCELLARIIDAE; PACH YPTILA)

J. B. COX

2 Rockbourne Street, Elizabeth North, South Australia 5113

ABSTRACT IV. Discussion .,,...., ceTeesaDEereerttertve. 2

COX, J. B, 1979. Some remarks on the breeding distribution and V. Acknowledgements ....)......-5. sense = BO

taxonomy of the Prions (Procellariidae: Pachyptila). Rec,S.Aus!. WY]. References............0....2.22-2 2221 30

Mus. 18(4); 91-121,

The breeding distributions of prions are reviewed

and the characters used to differentiate Pachyptila

vittata, P, salvini, P, desolata, P. beleheri, P. turtur

and P. crassirostris are critically assessed. Feeding

habits and the concept that differing breeding times

isolate these prions genetically are also discussed,

Evaluation of the evidence Indicates that their

previous classification into three genera or sub-

genera is invalid because only two groups are clearly

separable by morphological characteristics; the Fairy

Prions (turtur and crassirostris) and the Whale-Birds

(belcheri, desolata, salvini and vittata), The former

are considered to be one polytypic species and the

whale-hirds to be one monotypic and one polytypic

species.

Characters described as differentiating turtur and

crasstrostris jatergrade through many island popu-

lations north-south and west-east.. However, the

sharp boundary between each form in the Chatham

[s, allows their recognition as subspecies, although

elsewhere delimitations are arbitrary. The character-

istics of Whale-hirds also intergrade, but in their case

two species can be recognised, P, belchert 1s here

treated as specifically distinet from the desolata-

salvini-viltala complex, ullhough they have a zune of

overlap and hybridization in the subantaretie Indian

Ocean, Southern desolata and northern viltata are in

all probability conspecific and evidently salvinris an

imermediate form of hybrid origin, [In two southern

oceanic regions (South Atlantic and New Zealand)

there are no intermediates between desolata and

viltata and both are clearly distinguishable; but in

the southern Indian Ocean region their differences

are less pronounced nd intergrade through salvini

populations.

CONTENTS

‘U

ADSIHTACE occ ere eee stele ewelacetnee!s

1, Introduction . 2.0.00... ee te “

LL, Methods. 20. 0 cc cee cee ee ee

Ill. Observations ..-, 02.20 62.4 eee eee

(a) Breeding Distribution... , r

(b) Comparative Morphology .......,

(d) Differences in Breeding Times ...,

(e) Feeding Habits .... ATT Yeere et

Sow NM ee

MW MM

1. INTRODUCTION

The small southern oceanic petrels known as

prions or whale-birds were al different times

classified in several genera, a varying number of

species and many subspecies by Mathews (1912,

1932, 1937, 1938), Murphy (1936) disagreed wath

Mathews’ changeable concepts and believed there

were only four species in the single genus Pachyptla.

His elucidation of the group was welcomed by

Fleming (1939) who defined a fifth species. Ina later

systematic review Falla (1940) recognised five

polytypic and one monotypic species of three sub

genera, and Fleming (1941) hypothesized their

phylogeny, Some of these species had minor

differences in plumage coloration, but all six were

distinguishable only by the size, shape and structure

of their bills, occasionally in conjunction with

relative wing dimensions, [n this respect systematists

have emphasised the distinctiveness of prion bills.

Murphy (1936) wrote that the bill of each species

“stands out clearly as a distinct entity, withoul

exhibiting a degree of variation which might be

interpreted as interpradation,’’ Fleming (1941)

considered that "the bill characters by themselves

are diagnostic,”

Contrary to these views there is increasing

awareness that the bill characters of each species

vary greatly. Clancy (1965) and Watson (1975) listed

salvini afd vittata as conspecific. Furthermore,

beach-washed specimens of unknown origin are

frequently not identifiable (Cox 1976). Neverthe-

less, most authors still follow Falla's (1940)

classification and difficulties in identification have

been explained in various ways. Fullagar (1972)

wrote: ‘Since identification depends so much on the

form of the bill, not only its size, but also an

appreciation of its shape, many of the subtle

characteristics of each species can be recognised only

with experience.’ Serventy et al. (1971) noted that

some immatures have smaller bills and may not be

recognisable, and Kinsky and Harper (1968) showed

that through shrinkage the bill-width of dried

specimens may be less than in fresh or live

specimens. However, under the current classitica-

tion of Species these points do not appear realistic

92 REC. S$. AUST, MUS, 18(4); 91-121

when, as is often the case, two otherwise

indistinguishable specimens are allocated to diffe-

rent species simply because their bill-widths differ by

one or two millimetres,

Il. METHODS

In an attempt to resolve the problems of

identification, specimens of adults from most known

breeding places were obtained on loan to the South

Australian Museum from eight other museum

collections. Specimens collected at sea were alsa

borrowed, and use was made of numerous beach-

washed birds collected by myself, and fram

Australian museum collections.

Falla (1940) wrote "the older preoccupation with

bill measurements for specific diagnosis is mislead-

ing” and the “importance of emphasis on certain bill

characters was first clearly stressed by Mathews".

However, bill characters can be adequately specified

only by illustration and most authors have shawn

only one supposedly typical bill of cach species

(Murphy 1936; Fleming 1941; Slater 1970; Serventy

eral. 1971; Fallagar 1972; Macdonald 1973, Watson

1975). This aspect has obscured the existence of

great vanation in the bills of the species recognised.

Por these reasons, 4 range of prion bills are here

illisteated. The drawings were all made al twice

actual size on graph paper of lwo-millimetre squares

Uirect from specimens, traced onto cartridge paper

and reduced to getual size for publication, The

museum registration numbers of each specimen are

piven, and each drawing is also consecutively

numbered and prefixed A (Fairy Prions) or

(Whale-Birds) for cross references.

Measuremenis were all caken from dry skins m

accordance with merhous deseribed by Disney e/ al

(1974) bul, because many authors have emphasised

that the bill is the ovajor identification feature of

pnians. fhe methods of measuring bills are detailed:

culmen—tltom the juncture of the forehead feathers

and nasal tubes taken as a chord to the bill tip; bill

width—taken at the widest point of the bill, usually

but not always at the base; bill depth—a vertical line

from the point immedistely in front of the nostrils ta

the underside of the lower mandible on a properly

closed bill. MeEvey (1957) showed that measure-

ments taken from the same specimens by different

persons sometimes vary; (herefore, there may be

small differences between the dimensions | give and

those given by other wrirers, Ail dimensions cited

are tn millimetres.

Registration mumbers of specimens are abbrevi-

uted; AMNIT—Americsn Museum of Natural

History. New York; AMS—Australian Muscum,

Sydney, BMNH—British Museum (Natural His-

tory), Pring; CMC—Canterbury Museum, Christ-

church; CSTRO—C.S.1.R.O. Division of Wildlife

May. 1980

Research, Canberra; DMW—Dominion (now

National) Museum, Wellington; MNHN—Muséum

National d'Histoire Naturelle, Paris; NMV—

National Museum of Victoria, Melbourne; SAFM—

South Africa Museum, Capetown; SAM—South

Australian Museum, Adelaide,

Various Geographical regions and zones are

considered. Murphy (1936) and Servyenty et al,

(1971) have described the surface water zones of the

Southern Ocean, the Antarctic and Subtropical

Convergences are shown in Figure 1, Definitions of

the Antarctic and Subantarctic Zones follow

Roberts (1964), and the Subtropical Zone js

considered here to be the seas north of the

Subtropical Convergence. Different sectors of the

Southern Ocean are also discussed: the southeru

Atlantic region (from southern South American and

Antarctic Peninsulas and outlying islands, east to

South Africa, approximately 18°E); the southern

Indian Ocean region (from 18"E to the Western

Australian coastline, approximately 115°E): and the

Australian-New Zealand region (from 11S°E to

165°W), Prions breed at many localities in each of

these sectors, but are not known to breed in the

southern Pacific Ocean sector between 165°W and

75" W

lil, OBSERVATIONS

(a) RREEDING DISTRIBUTION

Prions occur over the uintaretic, subantaretic and

neighbouring subtropical seas, and breed on oceanic

and offshore islands and the mainlands of Antaretica

and Souch [,, N.Z,, The breeding grounds of ¢ach

recognised species have been frequently listed

inemrectly and require clarification. They are shown

In Figure | und listed below, but some remin

indefinite.

Falla (1940) did mot clearly define the breeding

distribution of PF. tarrur fallai Oliver, 1930, With the

exception of this subspecies, all subspecies he

recognised are listed, but with later evidence und

amendments included.

P, turtur (Kuhl, 1820): Islands of Bass Strait. off

Tasmania (Wood Jones 1937; Condon 1975). and off

New Zealand fram the Poor Knights Is, squth to

Cook and Foveaux Straits (Falla et al. 1966, Condon

1975); Snares Is. (sec below); Big and Little

Mangere Js of the Chatham Croup (Fleming 1939,

Falla 1940); Marion 1. (van Zinderen Bakker 1971)

and Beauchene I., Falkland Group (Strange 1968),

Also, probably on fle de VEst of Archipel Crozet

(Despin et al, 1972), possibly Antipades Is. (see

below), and two specimens were collected on

Macquarie I. (Keith and Hines 1958).

I agree with Condon (1975) who wrote that no

subspecies ‘can be recognised at this time.” Falla

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 93

KNTIC

5S. GEORGIA

geht

Van

SS =@ S oFKneY Is,

S. SHETLAND IS.

SCOTT |.

BOUNTY

CHATHAM IS.

x D,

POOR KNIGHTS

@ turtur i belcheri

O POSSIBLE

g Crassirostris

O POSSIBLE

Fig 1.—Breeding distribution of prions.

(1940) listed the breeding distribution of fallai as:

‘Breeding: Stewart Island, Akaroa, Chatham

Islands, ? Bass Straits, ? South Indian Ocean or

West Australia.’ He also listed Bass Strait,

Chatham Is. and ? Stewart I. in the breeding

distribution of nominate turtur. Falla et al. (1966)

said Antipodes Is. turtur are a small subantarctic race

(P. turtur subantarctica Oliver, 1955), although,

when naming the subspecies, Oliver (1955) sug-

gested they are a larger form because he wrote that a

specimen “‘thus resembles P. crassirostris more than

P.t.turtur.”’.

PR @ stan DA CUNHA

(@) Goucn |

S$, SANDWICH IS.

ANTIPODES |S.

‘ pM rucxvano Is.

Moy,

“Nay

ik ARCHIPEL

CROZET

CAPE DENISON

MACQUARIE I.

NARES IS

STEWART 47

TASMANIA

@ desolata

I salvini

@ vittata

Snares Is.: Falla (1940) said ‘Oliver’s suggestion

(1930: 115) that the stout-billed form occurs at the

Snares is not supported by a specimen, in the Otago

Museum, which has the bill form of typical

P.t.turtur.’ Oliver later (1955) listed this specimen as

P.t.turtur. Fleming (1948) wrote that on the main

Snares Island ‘‘Fairy Prions (Pachytila turtur) nested

deep in rock crevices among boulders piled along the

shores,”” Warham (1967) also reported turtur

‘nesting solely in rock crevices and talus debris” on

the main island, but after a fourteen-month stay on

this island Horning Jr. and Horning (1974) only

94 REC. §, AUST, MUS. 18(4): 91-121

reported ““Prions, probably both Fairy (Pachyptila

turtur) and Broad-billed, were seen flying near and

vver the main island during mast months,” without

mention of breeding. Fleming and Baker (1973),

reporting on birds of the Western Chain of islands in

the Snares Group, wrote ‘A new record from the

Snares is the Fulmar Prion (Pachyptila crassirostris)

which we found perching on ledges and nesting in

crevices’. Sagar (1977a) said the “Fulmat Prion”

was found on the Western Chain and that ‘On Rima

nests were found in the NW and NE areas of the

islet. Here large rocks formed 4 jumble over the

solid rock foundations Nests were deep under these

rocks”. Surprisingly, Edgar (1977) wrote that Sugar

had found the “Fairy Prion” on “Snares Islands”

and Sagar’s report (1977b) of the birds found during

his expedition to the Snares included no mention of

Fairy or Pulmar Prions, J, Warham (pers. comm.

1976) said that he is not aware of any specimens. of

crassirosiris taken from the Western Chain of the

Snares. Therefore it ts difficult to assess whether

typical crassirostris do in fact breed there,

Tt is known that prions of the Western Chain and

the main Shares Island have similar nesting habitats

und preferences, and that they probably breed at the

same time Sagar (19774) reported that Fulmar

Prions on the Western Chain had eggs on 21

November, and Warham (1967) reported that Fairy

Prions had chicks on the main island in January-

February (see below —''Differences in Breeding

Times"). Therefore the only difterences between the

birds must be morphological. Photographs of nesting

Fairy Prions trom Mownau I. and from Snares 1.

published by Serventy et al. (1971: Figs 62 and 139)

shaw that birds from both localines have very similar

bills, Furthermore, another photograph of two

Fulmar Prions at the Western Chain of the Snares,

published by Fleming and Baker (1973), depicts a

bird with a bill that is certainly too slender for a

typical crassirostris, albiel rather unclear. Because

Motunau J. prions vary, and some are not separable

fram crassirostris, While others are indistinguishable

from turtur (see below—'Comparative Morphol-

ony”), the evidence strangly suggests that the Snares

Group alsa contains one variable form of Fairy

Prion, ft is further jnteresting that there are no

teports of two different forms breeding during single

expeditions to the Snures, or to the Antipodes Is.

Where two forms have also been reported to breed

(see below); this aspect suggests that the prions have

been indiseriminately identified as either furtur or

CPASSIFOSITIS,

P.. crassirostris eatant (Mathews, 1912): Heard T,

(Falla 1937; Downes et al 1959) and Auckland Is,

(Falla et al. 1946), Also, possibly on the Antipodes

Is. (see below).

May, 1980

Although Condon (1975) listed fles Kerguelen as

a breeding ground, only one bird (the type of eatoni)

is known from this locality (Derenne e7 al, 1974) and

there ig no evidence of breeding.

P.c.crassirostris. (Mathews, 1912): Bounty Is.

(Falla 1940),

P. crassirostris pyramidalis Fleming, 1939:

Pyramid Rock, possibly Forty Fours islets and

probably The Sisters islets of the Chatham Group

(Fleming 1939: Falla 1940; Dawson 1955).

Dawson (1955) found that burrows on the eastern

Sister islet contained ‘two almost fully-fledged young

of the Chatham fulmar prion’, Whether these birds

Were crassirostris or turtur is open to doubt. They

were found in ‘burrows’ at the end of January, which

indicates, if one followed Fleming’s assertions (1941)

that turtur breeds earlier than crassirosiis, that they

were furmr which came from egys laid in mid-

October (sec below—'Differences in Breeding.

Times’). Also, crassirastris, at all other localities

Where it is known to breed, nests in rock crevices,

undec boulders, or in similar sites, and is not known

to nest in ‘burrows’. Hawever, the identilication of

many prions breeding in the Chatham proup is

questionable, because A, Wright stated (pers.

comm, 1975) that during extensive work on these

istands he found many prions that were not

distinguishable as mirtur or ¢rassirostris,

Antipodes Is.: Whether turiur or crassiroytris or

both breed at the Antipodes Is_ is unconfirmed,

Many writers have stated that one or hoth forms

breed on these islands, but their souree of

information is. obsture. Both are little known from

this locality and Dr J. Warham (pers. comm. 1976)

Stated that he knows of no specimens actually taken

on land Dr F. C. Kinsky (pers comm. 1976) agreed

that none Was known to have been taken on hind,

bul suggested that turtyr does breed there, because

twa specimens, taken On board ship anchored close

io the rocky shoreline in 1950, were in breeding

condition and had ‘naked brood patches,’

Although Oliver (1955) said of Bounty Is. birds

(erassirustris) that ‘this subspecies was found

breeding at Antipades Island in 1902,' it seems that

his statement Stems trom two specimens that are in

the Canterbury Museum. Waite (1909) mentioned

these specimens, and Falla (1940) gave their

registration numbers as 1200.1. Two crassirasiris

labelled *Antipodes Is.’ are in the Canterbury

Museum. Both are dated 1949, but this date

probably is when their registration numbers were

changed ta CMC AVI208 and CMC AV1247,

because they appear to be very old and were once

mounted; doubtless, they are the same two

specimens mentioned by Waite and Falla. Interest-

ingly, Wajte also said that a specimen of ~Prion

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 45

desolatus’ was obtained at the Antipodes Is. in

February 1907; but this specimen was probably

either turtur or crassirastris (its present location is mot

known) because Waite said “Captain Bollens tells me

thal this Prion breeds in crevices in racks at the

Bounty Islands’ (which implies it was not found

breeding at the Antipodes [s.) and he listed the

desolata of Auckland Is. as ‘Prion banksii,

Oliver (1955) listed specimens of turiur taken at

the Antipodes Is. on 31 July 1924 and in November

7950. Ti is unlikely that the July specimen was

hreeding, for that month is outside the normal

breeding season, and the 1950 birds are those which

were taken aboard ship. Of two specimens

examined, one (DMW 17222) is lahelled ‘off

Antipodes Is,’ and dated 21 November 1972, and the

other (DMW DMS5615) is a bird listed by Oliver,

labelled ‘Antipodes Is.” and dated 6 November

1950—one of the birds taken aboard ship.

From these data ali one can say is that both farms

have been collected cither on or near the Antipodes

Is. There is therefore a possibility they do breed at

this locality, but Warham added (lve. cit.) that a

recent thorough search of the islands during the

breeding season failed to find any prions. Sumimuriz-

ing, it seems that all reports of these prions at the

Antipodes Is. stem from two crassirastris taken, not

necessarily on land, in 1902, a doubrfully identified

specimen taken in 1907, a single rurtur taken wutside

the breeding season in 1924, and a few other /urtur

taken at Jater dates over the nearby sea.

Consequently there are no grounds to suggest both

torms breed sympatrically at the Antipodes Ls.

P. belcheri (Mathews, 1912); Falkland Is.

(Cawkell and Hamilton 1961), Tle de PEst of

Archipel Crozet (Despin et al. 1972) and Hes

Kerguelen (Falla 1937, Derenne et al. 1974). Also.

possibly on Staten 1. and offshore islets of Tierra del

Fuego (Harper 1972), and the remains of birds were

found in the nests of the Great Skua Stercorarius

skua on South Georgia (Watson 1975),

Serventy er al. (1971) and Condon (1975) listed

Bouversya as a breeding ground of helehert, but

Fleming (1941) and Tickell (1962) suggested that

“nly desolara breeds there. Holgersen (1960) did not

list prions in his review of the birds of Bouvetoya,

and Jaler papers by Solyanik (1964) and Holdgate er

al. (1968) are probably significant in that they do not

mention prions breeding or seen On Bouvetwya.

Most desolata breed on islands in the Antarctic

Zone, and Murphy (1936) said, without evidence but

merely because Bouvetpya is in the Antarctic Zone.

that desolata presumably breed on Bouvetoya. Later

confusion has doubtless resulted from such conjec-

ture.

P. d. desolara (Gmelin, 1789): Tles Kerguelen

(Tickell 1962).

P. desoluta banksi Smith, 1840: South Georgia. 5,

Orkney [s. and Heard J, (Tickell 1962).

P. desolata alter (Mathews, 1912): Auckland Is.

and Macquarie J, (Tickell, 1962),

P.desolata subsp.; Tle de l'Est of Archipel Croze|

(Despin ef al. 1972); 8, Sandwich Is,, §.Shetland Ts.,

Scott 1, (Harper 1972, Watson 1975) and Cape

Denison of Antarctica (Falja 1937).

Falla (1937) listed three specimens collected at

Cape Denison in 1913, and wrote that five birds

were seen and three eggs found. In January 1931, he

sighted. one bird flying towards land in the evening,

There are no further reports of desolata at this

locality, Watson (1975) wrote that the Cape Deniscm

population is “possibly extirpated.”

P.s.salvini (Mathews, 1912): Marion and Prince

Edward fs, (van Zinderen Bakker 1971).

P.salvini crozeri (Mathews, 1932); Archipel

Crozet (Despin et al, 1972).

P.vitrata nraceillivrayi (Mathews 1912): Me Saint-

Paul and fle Anssterdam (Mathews 1912; Falla

1940). According to Watson (1975) the last

population 1s “probably extirpated” and subject to

offshore stacks being searched, only a few birds may:

still breed on fle Saint-Paul,

P v.villata (Forster, 1777): The Tristan da Cunha

Group and Gough 1. (Murphy 1936, Swales 1965);

the mainland coasts of Faoveaux Strait and south

western fiords of South I., New Zealand. islets off

Stewart I. Snares Is. and the Chatham Group

(Fleming 1939, Richdale 1965, Falla er al. 1966).

Also, two vittata were collected in the Falkland fs,

(Cawkell and Hamilton 1961).

Condon (1975) followed Clancy (1965) in listing

South Georgia as a breeding ground of vittala

However, Murphy (1936) considered that desolata

of South Georgia had been confused With virtara and

that this mistake in identfication was subsequently

carric¢d on by others through quotation. But both

Murphy and Clancy listed vi/rara as breeding on the

Auckland Is, This locality was not listed by Falla

(1940), Falla er al. (1966), Serventy er al. (1971),

Condon (1975) and Watson (1975). According to

Tickell (1962) desolata breeds on South Georgia and

the Auckland Is.

(b) COMPARATIVE MORPHOLOGY

All prions have white underparts, a pale

superciliary, bluc-grey ear coverts and upperparts

with a dark ‘'M" acrass the wings and rump, a black

terminal tail patch, bright blue feet with paler webs,

96 REC. S. AUST. MUS. 18(4): 91-121

and blue-grey or dark grey bills. Their bills are

diverse in shape and have a row of comb-like

lamellae inside the cutting edges of the upper

mandibles. These lamellae are well-developed in

broader-billed forms but are often present only as

rudimentary ridges in narrower-billed forms.

Two groups of prions are readily recognisable: the

fairy prions (turtur and crassirostris) and the whale-

birds (belcheri, desolata, salvini and vittata). The

usage of these vernaculars follows Murphy (1936).

In many cases prions of each group are separable

by bill characteristics only. Compared with whale-

birds, most fairy prions have stouter (shorter and

turtur

oad

SAM B20293

LAWRENCE ROCKS. aust

DMW 14479

POOR KNIGHTS 15,,NZ

MNHN C.G. 1956 No. 127!

SLES KERGUELEN

belcheri

BMNH 1932.7 2.17

FALKLAND IS,

belcheri

BMNH 1932.72. 2)

FALKLAND 1S.

belcheri

AMNH 792675

FALKLAND IS.

belcheri

May, 1980

deeper) bills with a proportionately larger and more

bulbous dertrum. The exceptions are some turtur

that have bills closely resembling those of some

belcheri (Fig. 2: cf Al, A2, B1), or, in external

appearances only, even those of some desolata.

Birds of both groups are more easily recognisable by

marked differences in the amount of black on their

tails and slightly different head plumage colorations.

PLUMAGE PATTERNS

There are three plumage patterns evident in the

prions: (1) fairy prions have a large area of black on

the tail and a fairly uniform pale head and facial

pattern; (2) whale-birds have a small area of black

56 :

—=a(

sae =

Fig. 2.—Bills of some fairy prions and whale-birds compared to

show that some birds of each prion group have similar bills.

In this and subsequent illustrations of prion bills, the

drawings were all made direct from specimens. The bills of

some specimens were not set properly in preparation or,

since they have dried, shrinkage of the bill plates has caused

distortion, These aspects were copied to achieve a true

representation of the specimens. In life the nasal tubes of

prions are soft and, consequently, after death they gain a

variety of shapes which are not indicative of distinctions.

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 97

on the tail; of the latter belcheri has a prominent

white superciliary and facial area contrasting with a

dark blue-grey crown and ear coverts, and a pale

grey back; desolata, salvini and vittata usually have a

darker facial area, back and sides of breast.

Examples of these plumage patterns are shown in

Figures 3 and 4. These patterns are usually constant,

but some desolata have plumage indistinguishable

from some belcheri.

Tail Patterning

SAM B30285 and SAM B30288 (Fig. 3) are beach-

washed specimens of turtur and belcheri respectively,

prepared with spread tail feathers to show the

differences in uppertail patterning between the two

prion groups. Watson (1975) incorrectly showed in

illustrations that fairy prions differ from whale-birds

by having black extending in a broad band across the

outer rectrices. All prions have pale grey outer

rectrices.

The black terminal tail patch of fairy prions

extends for about half the tail length on the upper

surfaces of the inner rectrices. Specimens examined

from all populations have the outer rectrices wholly

pale grey and the second outer pair have a trace of

black on the ends of the inner vanes only. The inner

pairs have progressively larger areas of black with

that of the central feathers underlying the ends of

the uppertail coverts, the extreme tips of which are

also sometimes blackish.

Fig. 3—P. turtur SAM B30285 with typical tail pattern of fairy

prions (left) and P. belcheri SAM B30288 with typical tail

pattern of whale-birds (right). Note the grey outer rectrices

in each species and that the black of the tail extends to the

uppertail coverts.

98 REC, S. AUST. MUS. 18(4): 91-121

In whale-birds the terminal tail patch occupies less

than a third of the tail length on the upper surfaces

of the inner rectrices. The two outer pairs of

rectrices are wholly pale grey and the inner rectrices

have progressively larger areas of black, with that of

the central pair extending to the tips of the uppertail

coverts.

Head and Body Coloration

Fairy prions have paler, more uniformly coloured

heads than whale-birds, but like belcheri have a pale

May, 1980

grey back (fig. 4). The fore part of the crown of fairy

prions is usually slightly, but distinctly mottled, and

the dark coloration of their ear coverts, while

variable, is usually not extensive. The sides of their

breasts, flanks and undertail coverts are lightly

coloured blue-grey, and on some specimens this

colour extends over the abdominal feathers; this last

feature being particularly noticeable on some

Falklands Is. and Pyramid Rock (Chathams Group)

specimens.

Fig. 4—Head and back coloration of P. belcheri (SAM B30288,

left), P. turtur (SAM B30285, middle) and P. vittata subsp.

(SAM B29293, right). Note: these are selected examples to

illustrate basic differences; some vittata have paler plumage

than the specimen shown.

Harper (1972) correctly illustrated belcheri as

having white lores and a brvuad superciliary, which,

in all specimens I examined, contrasts with a

uniform dark blue-grey crown and ear coverts. He

also stated that desolata differs from belcheri by

having darker breast markings and ‘‘M”’ pattern

across the wings and rump, mottled grey lores and a

much smaller pale superciliary.

Specimens of belcheri and desolata that I

examined from populations studied by Harper

usually showed the differences he describe; but some

desolata had intermediate plumage colorations.

Tickell (1962) published photographs of desolata

showing a broad white superciliary, and fles

Kerguelen desolata are often indistinguishable from

belcheri by plumage. For example, fles Kerguelen

desolata’ BMNH_ 80.11, 18.687a and BMNH

80.11.18.647g and Falkland Island belcheri BMNH

1932.7.2.21 and BMNH 1932.7.2.17 were critically

examined and could not be differentiated by

plumage coloration. Therefore, the two forms are

not so clearly identifiable as Harper believed, and

extreme caution must be used in separating them by

plumage features alone.

Fleming (1941) stated that salvini and vittata have

darker plumages than desolata, but Harper and

Kinsky (1974) did not differentiate them by plumage

coloration. There was little difference between these

forms in specimens examined, except for a tendency

for larger-billed birds to be darker; but there is much

variation, For example, of two vittata collected on

fle Saint-Paul by the same person, one (MNHN

RREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 99

C.G. 1875 No. 112) has white lores slightly mottled

grey and a prominent white superciliary, while the

other (MNHN C.G. 1875 No. 115) has almost

wholly grey lores and very little white above the cye,

making the head appear almost uniformly grey.

The six examples cited above are typical of many

specimens examined. Most fles Kerguelen desolora

are indistinguishable by plumage from_ belcheri,

whereas desolata from other populations usually are

separable. Other whale-birds, from one or different

populations, vary in depth of plumage colour like

most desolata,

THE FAIRY PRION GROUP

Many island populations of these prions have been

characterised by bill distinctions. Consequently two

species are generally recognised: northern slender-

billed turtur and southern stout-billed crassirosrris.

Following descriptions of Falla (1937), Fleming

(1941) and Fullagar (1972) 1 previously stated (Cox

1976) that crassirostris is distinguishable from turtur

by having a dertrum lacking an angular ridge,

bulbous latericorns and a bill appearing bowed

outwardly when viewed from above. Subsequent

examination of specimens from most known

breeding localities of both forms showed that this

definition was incorrect. [ndividuals of cach farm

can have dertra the same size and shape, and bowed

or straight bill outlines when viewed dorsally. Only

smaller-billed examples of turtur have an angular

ridge along the top of the dertrum, An immature

from Marion Island (SAFM 559546) has this ridge.

but an adult olf smaller hill size (SAFM 21881) and

an adult of larger bill size (SAFM 55954a) from the

same island have larger, evenly rounded dertra,

Because many immatures (mainly beach-washed

specimens) possess this angular ridge and most

adults from all breeding localities have rounded

dertra, it seems that the character is usually a sign of

immaturity which may persist in some narrow-billed

adults. Fleming (1941) stated that) width and

bulbosity of prion bills develops with age.

There has been much confusion in differentiating

lurtur and crasstrostris, particularly in beach-washed

specimens. The often quoted first Australian record

of crassirostris (Learmonth 1957) was questioned by

Robinson (1972), with whom | agree that it was an

incorrect identification, This specimen (NMV

B6840) and CSIRO 14670 (which is also alleged to

be an Australian crassirosiris (MeKean and Lewis

1971)) are illustrated in Figure 5 for comparison with

drawings of specimens from breeding grounds.

- MMV Baadd

— ~‘,

(e=

Portland, Victoria

CHRO M670

40m E Moruya Heads,

NSW

(f=

Fig. S—Australian

specimens

wer ss —_ ul

of jurtur that have been

(erroneously?) identified as crassirostris. The lower specimen

was prepared with its bill open at the gape, giving itu stouler

appearance, which possibly resulted in it being wrongly

identified, The drawing was made with it clamped shut,

Which reduced the artificial stoutness,

NMV B6840) more resembles Australian or northern

New Zealand turtitr than other forms, and CSIRO

14670 resembles some specimens from Heard Is.

currently classified as crassirostris, but often

indistinguishable from some turtur, Other specimens

alleged to be crassirostris from Australia that were

examined are also incorrectly identified or are not

recognisable, including one described by myself

(Cox 1976).

Identifications of specimens from breeding col-

onies have also been doubted. When describing

Falkland Is. turtur, Strange (1968) said Dr. R. C,

Murphy and Dr, D. Amadon had confirmed his

identification and furthermore had found the

specimens to be inseparable from smaller-billed

turtur of New Zealand; but Watson (1975), when

citing Strange, said either turtur or crassirosiris

breeds on the Falkland Is.

100

Harper and Kinsky (1978) listed both turtur and

crassirostris as breeding on the Falkland Is. Harper

said (pers. comm., January 1979) their statements

were based on specimens in AMNH and birds seen

at sea. However, only one colony is known to exist

on the Falkland Is, and specimens from it (detailed

below and almost certainly those examined by

Harper) are morphological intermediates that have

the alleged characters of both taxa. It would be easy

to identify individual specimens as either form; but it

FALKLAND IS

turtur?

HEARO »

REC, S$. AUST. MUS, 18(4): 91-121

AMNH 808833

FALKLAND 1S

turtur ?

AMNH 786638

NMV B2898

crassirostris

May, 1980

would not be easy to believe that such specimens

could occur in one mixed colony as separate entities,

Bill Characters

Most published descriptions of the bills of turtur

and crassirostris are subjective. Their classification as

two species was largely based on such descriptions.

The purpose here, in illustrating many of their bills,

is to reduce this subjectivity by showing actual bill

shapes.

<a ae

7 DMW 14902 i

| MOrUNAL |. NZ .

Pr} turtur

1 i

(a :

fo DMW. 17222

/ ANTIPODLS 1s .

ut turtur ee

Po ae ‘ ee ee ‘

ara gts soe Me;

= Ts

—_——

. CMC AVI241

Lo MANGERE ) CHATHAMS:

/ turtur we

cS =a

_ it \ Er | A8

[< }——— ———

Fig. 6—The bills of two Falkland Is. turtur compared with other

turtur specimens and a Heard J. crassirastris.

Two Falkland Is. specimens (AMNH 808833 and

AMNH 786638) have bills similar to some Motunau

I. (New Zealand) turtur (Fig. 6: cf. A3, A4, A6).

They also closely resemble some Antipodes Is.,

Mangere Is. (Chathams) and Cook Strait turtur, and

some Heard I. crassirostris (Figs. 6 and 7: cf. AS, A7,

Al0, All).

Variation in the bills of Heard IJ. (Figs. 6 and 8:

AS, A16, A19) and Motunau I. birds (Figs 6 and 9:

A6, A15, A23), and in other populations of which

examples are illustrated, shows that it is extremely

doubtful whether all turtur and crassirostris are

distinguishable. Some Mangere Is., Antipodes Is.

and Motunau I. turtur closely resemble some Heard

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 101

DMW 18454

PYRAMID ROCK, CHATHAMS

crassirostris

NMV BIl438

LADY JULIA PERCY | AUST

turtur

DMW 9296 _

LIGHTHOUSE BROTHERS |,

COOK STRAIT Nz

turtur

DMW 18091

MANGERE |, CHATHAMS

turtur

DMw 18090

MANGERE |, CHATHAMS

turtur

DMW 18454

PYRAMID ROCK, CHATHAMS

crassirostris

Fig. 7—Two Mangere Is. rurtur compared with an Australian and

a New Zealand turfur, and two Pyramid rock crassirastris,

Mangere Is, specimens have stouter bills than many other

turtur, and, in bill characteristics, approach the form of

Pyramid Rock crassirosiris. The aciual bill lengths of

Mangere Is. and Pyramid rock specimens are similar.

and Auckland Is. crassirostris (Figs. 6 and 8: cf. AS,

Ao, A7, A8, Al5, Al6, A17, Al8, A19). Some

specimens of crassirostris from the Bounty Is.,

Pyramid Rock (Chathams) and Antipodes Is. are

very similar to each other (Fig. 10: cf. A29, A30,

A31), although Falla (1940) recognised each of their

populations as a different subspecies. Birds of these

populations sometimes also closely resemble Heard

and Auckland Is. crassirostris (Figs. 8 and 10: cf.

A19, A20, A27, A28).

While the bill shapes of turtur and crassirostris

cannot be used to distinguish many specimens, of

greatest critical interest are the characters of

differing populations that breed in close proximity,

Fleming (1939) described the largest form of

crassirostris from the Chatham Group breeding on

Pyramid Rock, approximately 17 km from turtur,

breeding on Mangere Is.

Like most southern New Zealand turtur, some

Mangere Is. birds closely resemble some crassirostris

and differ from Australian and northern New

Zealand turtur by usually having stouter bills with

more rounded dertra (Figs. 2, 6 and 7: cf. Al, A2,

A8, All, Al2). They therefore have somewhat

intermediate bill characteristics.

Pyramid Rock crassirostris differ from Mangere Is.

turtur by having broader and deeper bills with more

bulbous dertra. But the bills of both vary and

differences in the width and depth correspond to the

size of their dertra. The actual bill lengths of both

forms are similar, as illustrated in Figure 7 (A11,

102 REC. S, AUST. MUS. 18(4): 91-121

A1l2, Al3, Al4), which also shows that their bill

differences are not always significant.

Dimensions of Fairy Prions

The dimensions of specimens examined and those

listed by Buddle (1941), Despin et al. (1972), Falla

(1937, 1940), Fleming (1939), Richdale (1944) and

Wood-Jones (1937) are compared in Figure 11,

which shows mean and range of measurements.

Although few specimens are known from some

breeding grounds, the dimensions of birds from

some separate localities clearly differ. Notably,

Pyramid Rock crassirostris have absolutely longer

wings and wider and deeper bills than Mangere Is.

turtur. Also, Bounty Is. crassirostris is not a small

maTUNAU 1 Ne

turtur

NMV B4667

HEARD

DMW 14908

May, 1980

form as stated by Condon (1975), because, though

smaller than Pyramid Rock birds in mean bill and

wing measurements, its dimensions are all greater

than those of Heard I. and Auckland Is. crassirostris,

Differences between the mean dimensions of 15

Pyramid Rock and seven Bounty Is. crassirostris (0-5

bill-depth, 0-9 bill-width, 0-9 culmen, 5-5 wing

length) are so small that they are unlikely to have

much taxonomic significance (especially because the

data are from only 22 specimens) and _ surely

constitute insufficient evidence for subspecific

differentiation as Fleming (1939) proposed by

naming Pyramid Rock birds P. c. pyramidalis after

examining only 14 specimens, Furthermore,

although there is no evidence that the specimens

represent random samples of each population, the

ae Ns)

crassirostris

CMC AVI242

MANGLRL |, CHATHAMS

furtur

/ DMW. 17498

/ AUCKLAND 15

crassirostris

NMV B6284

HEAR |

crassirostris

} DMW 18426

oo BOUNTY thy

/ crassirostris Pai

pe =—{ A2O

eo Be

Fig. 8—Heard I. crassirostris (A16) have bills very similar to

Motunau I. and Mangere I. turtur, or (A19), very similar to

Auckland Is. and Bounty Is. crassirostris.

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 103

fo SAFM 21881

turtur 1

—. :

ra iar —

/ SAFM 559544

marion |

turtur 22

(-E \

ie DMW (4906

MOTUNAUL | rie

turtur

ao =

a SK

DMW DM5615

ANENOBES. 15

turtur

gs ~ —

NMY BIlg64

MACOUAHIE |

turtur

——_s

MW 14457

POOR KNIGHTS 15 Na

turtur

Fig. 9—Examples of turtur bills from various populations. The

similarity of these specimens suggests that races cannot be

distinguished by bill characteristics.

means of the bill width, bill depth, culmen and wing

measurements of four crassirostris from Pyramid

Rock and seven from Bounty Is. were tested on the

hypothesis that for each character the means of the

two populations would be equal. This was done with

a t-test after assuming that each of the four

characters is normally distributed in both popu-

lations and also after having tested the hypothesis,

again for each character, that the variances from

both populations would be equal (Table 1). At the

conventional 95% level, bill width was the only

character for which the means were found to

significantly differ. However, because this character

is only indicative of a very small difference between

the populations, and the samples were extremely

small, with the added problems such as measuring

the bills so finely, it is felt that it would be premature

at this stage to attach any nomenclatural significance

to the statistical result.

The two specimens of crassirostris alleged to be

from the Antipodes Is. are also better classified with

Pyramid Rock and Bounty Is. birds, for their bills

have very similar proportions but are slightly

narrower, suggesting immaturity, Also, one (CMC

AV1247) is otherwise doubtfully distinguishable

from some Bounty Is. specimens, while the other

(CMC AV1208, Fig. 10: A29) only differs from

specimens of both populations by having shorter

wings (by 4:0). However, their age when collected

cannot be established, because the collecting dates

are unknown; only one (male) is sexed, without

mention on its label of gonadal condition, and,

judging from. plumage condition, they are old and

have been handled frequently; consequently, plum-

age wear is difficult to ascertain. If they are

immature (therefore non-breeding) the shorter

wings of one have little bearing on their

classification.

104 REC. S. AUST. MUS. 18(4): 91-121 May, 1980

TABLE 1

eee

Pyramid Rock Bounty Islands

n= n=7

Mean (Range) Mean (Range) Variances F ty Significance

Billdepth ............-..... 8-58 8-17 Equal 1-64 2-156 0-1>P>0.05

(8-2-8-7) (7-8-8-8)

Bill width ................., 12-45 11-07 Equal 3-33 3-00 0:02>P>0-01

(11-8-12-9) (10-3-12-6)

Culitiert. atittaactiegr¢etaa 23-35 22:79 Equal 1-87 1-44 P>0-1

(22:7-24-3) (21-6-23)

WYER oi ee fates eA EE 192-5 187-57 Equal 8-99 1-63 P>0-1

(190-194) (180-197)

DMW 17500

AUCKLAND IS,

crassirostris

DMW 0M5604

BOUNTY (S

crassirostris é

CMC AVI208

ANTIPODES iS.

crassirostris |

DMW 17286

BOUNTY JS,

crassirostris

DMW 18455

PYRAMID ROCK, CHATHAMS

crassirostris

NMV BII963

MACQUARIE |

turtur

Fig. 10—Examples of crassirostris that have similar bills, and the

dimensions numbered on A32 correspond to the measure-

ments given in Table 2.

Most Pyramid Rock, Bounty Is. and Antipodes Is. because they appear to be intermediates. Many have

crassirostris can be separated from all other fairy stout bills, thus closely resembling crassirostris or

prions by having a bill depth of over 8-0; but Heard some turtur, but the dimensions of specimens are

I, and Auckland Is. birds present a difficult problem closer to those of turtur (Fig. 11).

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS

BILL-DEPTH & WIDTH

FALKLANDS

MARION

cROZETS”

AUSTRALIA

POOR KNIGHTS(NZ) +

COOK STR. IS. }

MANGERE (Chatham)

turtur

ANTIPODES”

JACQUES LEES(NZ)

WHERO 1, (NZ)

MACQUARIE”

MOTUNAU 1.(NZ)

HEARD i.

AUCKLANDS

ANTIPODES*

BOUNTY IS.

crassirostris

PYRAMID ROCK

(Chathams}

¥ - Breeding uncertain

105

SPECIMENS

200

CULMEN WING

180 190

—s ok

BOO @wWWNM

—_—

aw nn © ~ PM

Fig. 11—Dimensions of fairy prions (mean and range).

Falkland Is., Marion 1., Australian and New

Zealand furtur specimens have slight dimensional

differences, but more material is needed before

assessments of their relationships can be made. Falla

et al, 1966 said Antipodes Is, furtur are a small!

subantarclic race, but the few specimens of

uncertain natal origin are similar to many other

turtur (Figs. & and 9; A7, A24) and cannot be

recognised as distinct. The type of fallai, taken at

sea near South J., New Zealand, is a young bird just

out of nestling down (Falla 1940) and also cannot be

recognised,

THE WHALE-BIRDS

Although it is extremely difficult, if not

impossible, to distinguish many desolata from

salvini, Falla (1940) and Fleming (1941) followed

Mathews (1912) in listing belcheri and desolata in a

separate subgenus from salvini and vittata, Fleming

gave the subgeneric characters of belcheri and

desolata as: ‘Long bill with straight outlines, viewed

from above, Palatal lamellae become obsolete some

distance behind the dertrum and are invisible when

the bill is closed. Plumage pale grey, and head

pattern moderately distinct,’ and those of salvini

and vittata: ““Large very broad bill with bowed

outlines, Palatal lamellae extending all the way from

gape to dertrum and visible even when bill is closed.

Plumage dark and head pattern well defined.”’ He

also said, “the reality of the subgeneric grouping can

be emphasised by drawing the bill of one species in a

network of Cartesian co-ordinates, and by Tepre-

senting the same co-ordinates on similar drawings of

other species,"’ By this method he suggested that

helcheri and desolata have bills based on a different

“plan” to those of salvini and vittata.

However, the plumage characters (detailed

previously) of desolata and salvini are not different

and the bill characters of both intergrade, indicating

that the subgeneric prouping is wrong, Furthermore,

Fleming’s representation of the bill of desolata in

Cartesian co-ordinates is not true of most desolala,

because only a minority (typically fles Kerguelen

birds) have straight cutting edges to their upper

mandibles as he depicted. Their bills in fact usually

have curved cutting edges (beneath which their

lamellae are often visible) and are very similar to

those of salvini (Figs, 14 and 15: ef. BIS, B17, B19,

B20, B21. B23, B24), Many desolata also have

bowed bill outlines viewed dorsally (Figs. 12, 14 and

15: B13, B16, B19, B21, B22) and their lamellae are

often visible when the bill is closed (also see Tickell

1962) and usually extend inside the bill as far as the

dertrum. The lamellae in desolata, salvini and vittata

are large at the gape and progressively diminish in

size towards the dertrum, as described by Murphy

(1936) for vittata, but in narrower-billed desolata

specimens they are usually present only as

rudimentary ridges towards the dertrum and closely

resemble those of belcheri, also described by

Murphy. Irrespective of natal origin, adult speci-

mens of desolata, salvini and vittata with wider and

more bowed bills also tend to have larger lamellae.

106

and

In whale-birds the major problems are

recognising belcheri from some desolata,

recognising desolata and vittata from salvini.

Bill Characters of Narrower-billed Whale-Birds

Although bill measurements of allopatric popu-

lations of belcheri and desolata overlap slightly, both

forms are clearly separable except in their

populations of fles Kerguelen and fle de l'Est,

Archipel Crozet.

Falla (1937) identified fles Kerguelen belcheri and

desolata by bill shape and said specimens of each

form with ‘“‘exactly the same”’ bill dimensions could

NMV 83133

ILES KENGULLEN

beicheri

ILES KERGUELEN

belcheri

NMV B2902

ILES KERGUELEN

belcheri

NMV B4674

ILES KERGUELEN

desolata?

NMV B4673

ILES KERGUELEN

desolata?

NMV B4676

desolata

NMV B467!

LES KERGUELEN

desolata

(LES KERGUELEN

desolata

REC. 8. AUST. MUS. 18(4): 91-121

MNHN CG. 1969 No 925

ILES KERGUELEN

May, 1980

only be recognised by the ‘‘compression of the

maxillae in P.belcheri and consequent — slight

concavity of outline when viewed dorsally."’ He

described two desolata populations on_ the

archipelago: A—a wide-billed form, and B—a

narrow-billed form. Of form B he said that “At the

time of collecting I regarded these birds as a new

form and a possible hybrid between P.desolata and

P.belcheri, but subsequent examination of the

specimens shows that they possess none of the

distinctive characters of P.belcheri, and are certainly

referable to P.desolata.”’ Tickell (1962) used Falla’s

measurements from these two forms and concluded

that fles Kerguelen desolata are a narrow-billed

subspecies.

BMNH 80.1! 18.687

B12

fn BMNH 80.1..18, 647 A

ae ES KE! =

wae _ — “dasolata <2 B13

Fig. 12—The intergradation of bill characteristics of belcheri and

desolata of fles Kerguelen.

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS

The bill dimensions of all three forms from fles

Kerguelen and belcheri from the Falkland Is. are

plotted separately in Figure 13, which also shows the

means and range in bill size of other desolata (from

Tickell 1962), and the known bill size variation of

belcheri. There is little difference between fles

Kerguelen form A and desolata of other breeding

localities in the mean or variation of bill size. Form

B is intermediate between form A and belcheri (i.e.

intermediate between desolata and belcheri).

24 25 26 27

CULMEN LENGTH

A —FORM A

107

The bill shapes of fles Kerguelen belcheri and

desolata are also variable, and this variation is shown

by drawings of their bills in Figure 12 (B5-B13).

Some desolata have bowed outlines to their bills

(B12, B13) while others have straight outlines (B10,

B11). Some belcheri do not show a slight concavity in

their dorsally viewed bill outline; NMV B3133 (BS)

with a bill width of 8-7 has a slightly bowed outline.

Falla (1937) did not elaborate on “the distinctive

characters of P.belcheri’’ beyond their bill shape, and

28 29 30 31 mm

B—FORM -} Pdesolata, Iles Kerguelen (Falla 1937) o — specimens examined.

a — P belcheri, Iles Kerguelen; O — Falkland Is.

e@—disputed specimens

Pdesolata- mean bill sizes: Py —Heard I, He =S. Georgia, Ay —Macquarie I, r= 5. Orkney Is..

hr) —Auckland Is. (Tickell 1962)

Fig. 13—Bil! dimensions of belcheri and desolata. Solid lines

illustrate the known size variation of each form, and keyed

symbols the size of individual specimens. Note the bill size of

fles Kerguelen form B is intermediate between form A and

belcheri, and that form A is similar in mean bill size to other

desolata populations.

10s

it seems that he only found concavity io the bill

outlines of his specimens once they had dried, Also,

live and fresh beach-washed specimens I have

collected had no indication of concave bill outlines

but, when dried, concavity was noticeable. Concav-

ity in bill outline can be caused by shrinkage and

consequent inward cantrachion of the latericorns on

drying specimens, BMNH 1932.7.2.17 (fig. 2: B2)

has its bill shghtly swollen at the base and slighdy

concave towards the dertrum; 4 conditian probably

caused by shrinkage.

Differences between Tles Kerguelen belcheri and

desolata are ill-defined, and it is not surprising that

the identifications of some specimens have been

disputed, NMV B4673, NMV 84674 and NMYV

B4676 are labelled desolata from Tles Kerguelen (by

the collector?) but also written on their labels is P.

©, Harper's opinion that they are beleheri, whieh is

indicated as correct by the plots of the specimens in

Figure 13. However, the collector also ohtained

other desolaia al the same Jocality on the

archipelago (NMV 64671, NMW 4672, NMV

B4675 and NMV B467&), and because the drying

hills of some specimens shrink more than others, the

question remains whether Wes Kerguelen birds can

be wWenttied when living. Kinsky and Harper (1968)

stated that under controlled drying conditions the

bill width of adult belcheri shrinks by 5 to 16 per cent

(average |3 per cent). This wide variation in

shrinkape suggests that al least one of the disputed

specimens (NMV B4673) could have been quite

justifiably identified as desolai when collected: it

does nol have concave bill outlines viewed from

ubove, and on the dried skin its bill width measures

116. which 1s within the range of variation of both

torn.

Tickell (1962) found Tles Kerguelen desolata to be

narrower in bill width than other desolata because he

combined the dimensions of forms A and B in his

calculations. But form A closely resembles other

desolara, whereas form B is intermediate between

helchert and desolala- and other specimens from the

archipelago show there is intergradatian of the

characters of bath forms, This evidence suggests

hybridization and. consequently, Tles Kerguelen

deselata should not be classified ay a separate

subspecies,

The measurements of two belcheri from ile de

Est given by Despin ef al. (1972) are: bill width 10-2

and IL-8, culmen Jength 25 und 27) Two desolete

fram the same colony measure 12-5 anc 14-3, and 26

dod 28. Despin ef al said of the helcheri

colonyCette calonie est de trés petites dimensions

-elle ne compte que quelques nids—et trés

nettement délimitee’ (This colony is very small—

only a few nests—and extending over a very limited

area). No other belehert were found on the istand.

REC. 5, AUST. MUS. 18{(4}: 91-121

May, 1980

Furthermore, they said; ‘Le Prion de la Désolation

est apparemment trés peu abondant aL'Ule de |'Est

ou une seule colonic a élé observée, la méme

dailleurs que celle du Prion de Belcher mentionée

précédemment” (The Desolate Prion is apparently

very scarce on East Island where only one colony hus

been observed, the same colony moreover as that of

Belcher’s Prion mentioned above), Because of the

Scunty data few conclusions can be formed,

Nevertheless, the culmen lengths of the two forms

overlap, and the difference in bill width between the

largest belcheri and the smallest desolata 1s only (7.

When the specimens were examined, the two forms

could not be separated by plumage coloratian,

though plumage wear varied. and it was noted that

their dorsally viewed bill outlines intergraded. The

smallest beleheri (MNHN C,G. 1974 No. 1823) has

very shghtly cancave outlines, while the Jargest

(MNHN C.G. 1974 No. 1824) has straight though

slightly swollen outlines. The smallest desolata

(MNHN ©.G. 1974 No, 1816) has very slightly

bowed outlines, and the largest (MNHN C.G. 1974

No, 1815) has obviously bowed outlines. It is

interesting that the former two are both females, and

that the latter are both males. [n bill shape the two

desolata are intermediate between the beleheri and

salvini (MNHWN C,G. 1974 Nos. 1808 and 1809) that

are also fram Jle de VEst. Therefore, as on Tles

Kerguelen, the bill characters of Te de l'Est beleheri

and deselara are inconsistent.

Bill Characters of Wider-billed Whale-Birds

Bill dimensions of 245 specimens are shown in

Figure 16, which is a reduced continuation of Figure

13, [nh must cases smaller-billed deselare were

excluded if their identification from belcheri was

doubtful. The known dimensional limits of desolata,

salvuini and vitrata, derived from specimens and

mejsurements given by Despin er al. (1972), Falla

(1937, 1940), Mathews (1912, 1938), Murphy (1936).

Serventy ef al. (1971), Swales (1965) and Tickell

(1962) are superimposed around the plots of the

individual specmens, and the mean bill dimensions

of separate breeding populations ure given except

those of desolata, which are shown in Figure 13.

Figure 16 shows that (a) there is wide overlap in bil]

dimensions of the three forms; (6) many specimens

have intermediate dimensions; (c) there is a strong

tendency tar birds with a longer culmen to have a

wider bill; and also that saluini is an intermediate

form between desolata and vittata,

Falla (1940) and Fleming (1941) revognised two

subspecies of salvia? (nominate salvin? of Marion T-

and crezeli af Archipel Crozet) because the

specimens available to them suggested that Marion

1. birds have larger bills. However, Despin ef al,

(1972) Jater showed thar Tle de Est salpini have

larger bills (culmen 29-34 (av. 31-6), and bill width

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS

AMS 0.22083

ADELIE LAND, ANTARCTICA

desolata

BMNH 1958.12.10

SOUTH ORKNEY IS.

desolata

salvini

NMV B2891

HEARD |

desolata

MNHN C.G.1974 No. 1807

ARCHIPEL CROZET

109

AMS 0.22082

MACQUARIE 1

desolata

AMS 0.22084

ADELIE LAND, ANTARCTICA

desolata

AMS 0.22085

MACQUARIE |

desolata

—

B20

Fig. 14—Typical desolata specimens from various populations

compared with a salvini specimen. Note that the cutting edge

to the upper mandible of desolata is usually curved upwards

at the base, in profile view, as in salvini, and that desolata

frequently have bowed bill outlines, viewed dorsally.

15-5-20-5 (18-4) on fresh specimens). It is possible

that vittata macgillivrayi is intermediate between

salvini and vittata (Elliott 1957; Clancy 1965), but in

bill shape two specimens from fle Saint-Paul more

resemble vittata. Their measurements and those of

two others given by Mathews (1938) and Falla (1940)

are: culmen 30-31 and bill width 17-18. Fifteen

presumed beach derelicts measured by Serventy et

al. (1971) are 29-5-34-5 (av. 32-3 ad 16-2-21-5 (18),

and appear smaller than vittata which they measured

as 32-37 (34) and 18-8-22-6 (20-6), but the overlap in

size indicates that many specimens are not

separable. The bulge in outline of dorsally viewed

vittata bills is variable, with some being more evenly

bowed (Fig. 15: cf. B27, B28), and although

macgillivrayi also have a similar bill shape (B25,

B26) the bulge in bill outline is also present in some

salvini and even some desolata (Figs. 14 and 15:

B16, B22). Differences in bill profile between salvini

and vittata suggested by Fullagar (1972) are

inconsistent (Fig. 15). The dertra of salvini and

vittata do appear “weak” (Falla 1940) in proportion

to bill size compared with those of belcheri and

desolata, but in these whale-birds it appears that

differences have evolved through spreading

latericorns and palates without a corresponding

enlargement of the dertra and nasal tubes. In all the

wider-billed whale-birds the dertra and nasal tubes

are of similar size, but with individual variation

(Figs. 14 and 15).

REC. S. AUST. MUS. 18(4): 91-121

BMNH 1958 12.12

SOUTH ORKNEY IS

desolata B21

NMV B2892

HEAM |

desolata

MHNP. C G.1974 No 1776

ARCHIREL CROZET

salvini B23

BMNH 1949.5) 5

MARION |

Salvini

INHINC.G B75 112

(LL saint —eaun

vittata

MNHN C.G.1875. (5

ILE SAINT~PAUL

vittata

DMW 13540

CHATHAM 1p

vittata

: B27

Fig. 1S—A comparison of the bills of desolata, salvini and vittata.

Although variation is wide, the bills of each form are

fundamentally similar. B28 has a bill that is larger than that

May, 1980

of average sized wittata.

A further bill characteristic used to separate

salvini and vittata is colour. Watson (1975) said the

bill colour of salvini is “wholly blue” and that of

vittata is “steel gray”’, a difference accepted by most

writers. However, Murphy (1936) said the bill of

vittata is “light blue or bluish grey” and two males

collected on Gough I. in April have their bill colours

labelled as “‘blue-grey’”” (BMNH 1956.36.20) and

“blue (BMNH 1956.36.19). Falla (1937) said New

Zealand vittata have the upper bill ‘glistening iron

grey, with an occasional bluish patch appearing as an

abnormal marking.” Although there is variation, it

seems possible that vittata of the southern Atlantic

region differ slightly in bill colour from New Zealand

birds. But confusion could stem from the fact that

after death the bills of both forms rapidly become

blackish, Specimens of desolata and salvini that |

have collected on beaches have had entirely blackish

bills and eyes not yet shrunken. Narrower-billed

whale-birds on the other hand, have bills which do

not usually turn blackish until they are dry.

Although Falla (1940) and Fleming (1941) said

salvini is more closely related to vittata than to

desolata, and Clancy (1965) and Watson (1975)

listed salvini and vittata as conspecific, there appears

to be no reason, by way of bill size or structure (their

criteria) to support these views. It seems that salvini

consists of an assemblage of slightly differing

populations which are intermediate between desol-

ata and vittata (Fig. 13).

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS Ih

24) 254 26) 27 28 29 30 31

32) 33 34) 35] 36) 37] 38ymm

a) -5 plus specimens, e- 4, e-3, 4-2,+- I.

salvini: P - POSSESSION; M— MARION; E- ILE DE L’EST.

vittata: S— SAINT-PAUL; C- CHATHAMS; Z-NZ; G- GOUGH.

Fig. 16—Dimensions of the bills of desolata, salvini and vittata,

The known size variation of each form is superimposed

around the plots of individual specimens. the mean bill sizes

of salviniand vittata populations are shown; those of desolata

are shown in Fig. 13. Note the wide overlap in the bill

dimensions of each form, and that many specimens are

intermediate. Unlike

desolam and

vittata, bill sizes of

different salvint populations are inconsistent

Other Characters of Whale-Birds

Bill, tarsus, tail and wing measurements of

specimens from breeding localities are shown in

Figure 17, the data derived from specimens

examined and measurements given by Despin ei al.

(1972), Falla (1937, 1940), Mathews (1912, 1938),

Murphy (1936), Rand (1954), Swales (1965) and

Tickell (1962). The dimensions of each recognized

species overlap with those of one or more of the

other three, and differences are hard to define.

There is a tendency for larger-billed birds to have

their other dimensions larger, but this generalization

does not always hold true.

Some belcheri have the shortest wings but their

measurements widely overlap those of desolata.

However, belcheri usually have a longer tarsus than

those desolata which have only slightly larger bills;

but two specimens of each form collected on fle de

l'Est by Despin et al. (1972) differ from most other

belcheri and desolata in having a longer tarsus. In

REC. S. AUST. MUS. 18(4): 91-121 May, 1980

112

ore

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——

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Abs

its

SZ oz St

HLOIM-1118

SWVHIVHS 8 ZN

2YELIA

HuIAleS

e}ej0sap

14984919q

HONOD

TAVd-LNIVS

41S4.1 30 311

NOISS4SSOd

NOIWVW

GuvaH

AANYYO'S

VIDYO39D ‘S

VOILOYVLNY

SONVTNONV

AIYVNOOVW

1831 30 311

v- N373nN9Yay

g@-N314aNDY3ay

SONV1N1V4

N31S5NDY4ay

1S3q 30 311

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 113

tarsus measurement the Tle de Est birds more

resemble each other than other populations of each

form, Furthermore, both forms from this small

colony have similar wing lengths, and in all othe

given dimensions the desolata are intermediate

between the belcheri and salvini of the same island,

Therefore, there appears to be little convincing

evidence that at the two localities (fle de l'Est and

fles Kerguelen) where belcheri and desolata both

breed they are morphologically distinct; although

their allopatric populations show fairly clear

interspecific differences. At the Falkland Is,, where

beleheri is the only breeding form, they are distinct

from desolata by their bill width, which is not known

to exceed 11-5, Most desolata can be distinguished

hy having a wider bill and a shorter tarsus,

Tickell (1962) classified South Georgia, S. Orkney

Is, and Heard I. desolata as a different subspecies

from Macquarie I, and Auckland Is. birds because

they have differences in wing and tail lengths,

Nevertheless, these differences are slight and are

probably not indicative of subspecific distinction,

because Macquarie [. desolata are intermediate in

wing and tail length between Auckland Is. and Cape

Denison (Antarctica) desolata, and the latter are

intermediate between Macquarie 1. and Heard 1.

birds. Moreover, Macquarie |. birds more resemble

Heard L. desolata in wing length than birds of the

Auckland Is. Therefore only a minority of

individuals are likely to be identifiable, and Tickell’s

classification of subspecies should be treated with

caution,

The longest-winged birds are vithate. but their

wing measurements Overlap those of salvini by 12

and desolata by 19. Most vita are nevertheless

distinguishable by the combination of long wings and

large biil, because mast longer-winged desolata have

measurably smaller bills. There is little difference in

the bill sizes of Tristan da Cunha, Gough 1. and New

Zealand vittata (Falla 1940, Elliott 1957) and, except

for forms of the southern Indian Ocean region. all

desolata populations haye similar mean bill sizes

(Tickell 1962, Fig. 13). On the other hand, the mean

hill sizes of salviai populations vary, and some

closely resemble desolata while others more

resemble vitrata, The mean bill size of desolata is

approximately 14-5 x 27:5 and vitrata 21 x 34; the

desolata specimens in Figure 14 are typical examples

of their respective populations, and in Figure 15

vitlata B27 represents average size. The similarity of

both forms is shown in Figure 15 (B22, B26, and

others). Murphy (1936) illustrated typical specimens

of cach, but Fleming (1941), followed by Serventy ef

al. (!971), diagrammatically illustrated their bill

characters and showed a viltata with an abnormally

wide bill compared with a desolata of ayerage bill

size, but with an unusually wide dertrum, which

gives the impression that they are vastly different

birds.

At all their proven breeding grounds populations

of iurtur atid erassirostris are allopatric. The different

whale-birds are also allopatric breeders except that

both belcheri and desolata breed on tes Kerguelen

and Despin et al. (1972) reported that belcheri,

desolata and salvini breed on fle de Est, Archipel

Crozet.

The Intergrading Characters of Populations of Fairy

Prions

The fairy prions were classified as two specics

mainly because of differences in the stoutness of

their bills. To ascertain the comparative stautness of

the bills of different populations, a graded series of

six bill dimensions was scored according to index

values from 0 ta 4 (Table 2), A true hybrid index

could not he devised because the measurements are

very small and individuals of the most different

forms someltinies share characteristics, and, conse-

quently, it is very difficult to define phenotypes. Use

of wider scales of dimensions produces less

differences in the mean index values of the various

populations, and they would be barely distunguish-

able.

The measurements required for the bill stoutness

index could only be taken from a rather small sample

of specimens. These measurements (illustrated in

Fig. 10: A32) were: (1) bill width at base, (2) the

width of the maxillary unguis, (3) bill depth from the

base of the nasal tubes, (4) bill depth from the point

in frone of the nostrils, (5) bill depth trom (he top of

the maxillary unguis to the lower angle of the

mandibular unguis, (6) the actual bill length (i.c. mar

culmen Tength). Pertaining fo stoutness, higher

index values were given for greater width and depth

ineasurements, but lesser scores were afforded to

prealer bill length measurements,

The index value of each specimen equals the mean

score of the six dimensions, and the mean index and

per cent value of bill stoutness of populations was

calculated (Table 3). Figure 18 diagrammatically

illustrates geographical variation in the bills of these

prions: cach population is represented by a circle,

which, aecording to its per cent value of bill

stoutness, is partly black. A greater portion of black

indicates populations with stouter bills.

114 REC. S, AUST. MUS. 18(4): 91-121 May, 1980

TABLE 2

Index values according to stoutness of fairy prion bills. Specimens with bills of greater width and depth but lesser length are afforded

higher index values.

Width Depth

Bill Index

from base in front at unguis Length Value

at base of max. of nasal of nostrils

(1) unguis (2) tubes (3) (4) (5) (6)

—10 —4 —9 —7 —7°5 20-6— 0

10:1—10-5 4.]—4.3 9-1 —9-5 7-1—7:5 7-6—8 20:1—20-5 1

10-6—11 4:4—4-6 9-6—10 7-6—8 8:1—8-5 19-6—20 2

11+1—11-5 4-749 10-1—10-5 §-1—8:5 8-6—9 19-1—19.5 3

11-6— s— 10-6— 8-6— 9-1— —19 4

TABLE 3

The comparative bill stoutnesses of fairy prion populations.

No, of birds with index value: Mean Per Cent

Locality 0-1 1-1-2 2-1-3 3-1-4 Index Value

turtur:

Australias wmetenussitseugtohe «shitit edctotencstits see tta tala 7 1 0-66 16-5

Marion hs ss: partecsssactisusittvettoy;titvtes 2 0:71 17:75

Poor Knights Is, -.,.-....-- : eS wen 1 seh et re! 2 2 1-08 27

NESS od OT 2 4 1:27 31-75

Cook Strait Is... _- 2 1-41 35-25

WHETOAG 5 6 och ye seteens 0 she a peigele oncdat po eclebien debate wedi, o's 1 3 | 1:43 35-75

Falk lara Ts. by edscsse 444 oe tose dt eh 2 1-49 37°25

Matunau T,-o SAS so Ca oe ET hes eecee cle 3 I 1:7 4255

crassirostris:

Heatd-£, . 0s yoce bat ssebenreosri: Sse 2 2 2-03 50:75

ACK Nd Ts. 4 oe lyste eset par tela be ce tosdo ne easiegte eta gy shade 1 l 2-33 58-25

(Antipodes Is.) ........ 1 1 2:99 74:75

Bounty Is, ........, Rie FARES sells ecrenn ac ede al 2 2 3-24 81

Pyramid Rock ..........00 00.00 cece eee eee 1 3 3-49 87-25

AUSTRALIA —

KNIGHTS 15S,

——____,»

MANGERE IS.

FALKLAND IS, RYRAMID

ROCK

q << _—_—_ MARION t.

WHERO |.

BOUNTY [S.

HEARD }. 4S. 3)

Fig. 18—Geographical variation in the bills of fairy prions. The

comparative bill stoutness of each population is indicated by

the percentage of black in the circle representing each.

Stouter-billed forms are indicated by a greater percentage of

black, and by arrows pointing from more slender-billed

forms. Although the Antipodes Is. are not a confirmed

breeding ground, the bill stoutness of two crassirostris said to

be from this locality is indicated to show that they are

intermediate between Auckland Is. and Bounty Is. birds.

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS

Subjective judgments of bill stoutness compliment

the index. Falkland Is. birds have neither a stout nor

a slender bill (fig, 6: A3, A4) and in many respects

appear to be intermediate between turtur and

crassirostris, Stouter-billed birds breed on Heard [.,

and Auckland Is, birds seem intermediate in bill

bulbosity between the birds of Heard |. and

Antipodes or Bounty Is. Fleming (1939) pointed out

that there is a progression towards greater bill

bulbosity from birds of Antipodes Is., to Bounty Is.,

and to Pyramid Rock in the Chatham Group.

Marion I. and southern New Zealand turtur are

different intermediates between the stout-billed

forms and slender-billed Australian and northern

New Zealand iurtur.

The distributions of tur/ur and crassirostris cannot

be correlated with the surface water zones of the

Southern Ocean as was believed by Fleming (1941),

for both breed in the Subantarctic Zone (Fig. 1).

Throughout their respective ranges, furtur and

crassirostris are connected by populations with

intermediate characteristics, suggesting that both are

heavily affected by gene-flow. This intergradation is

probably indicative of adaptations to local environ-

ments rather than of hybridization. However, the

Mangere Is. and Pyramid Rock populations (both in

the Chatham Group) probably each stem from

different parental stock; the former closely resem-

bies New Zealand turtur and the latter Bounty Is.

crassirostris. These taxa are nevertheless allopatric,

and their characteristics, being mainly differences in

size, are not greatly divergent from their hypotheti-

cal common ancestor (see Fig. 7).

The Intergradation of Different Whale-Birds

To some extent the differing bill characters of

whale-birds can be related to latitudinal distribution.

Small-billed desolara breeds in the Antarctic and

Subantarctic Zones; intermediate salvini breeds on

islands in the Subantarctic; and large-billed vittata

breeds on islands close to the Subiropical Converg-

ence. This distribution indicates that the three forms

replace each other geographically, but belcheri,

which has the smallest bill of the whale-birds, does

not fit this pattern, and is known to breed only in the

western Subantarctic Zone. Therefore these prions

do not differ in characteristics entirely according to

their latitudinal distribution or to the surface water

zones.

In Figure 19 the three desolata population groups

of the Antarctic Zone (Scotia Sea, Heard |. and

Cape Denison) are diagrammatically illustrated on

the same plane, and eastern Subantarctic desolatu

(Macquarie I, and Auckland Is.) are connected by a

vertical line in the east to indicate their geographic

position. Arrows point from Scotia Sea and Cape

Denison populations to Heard I. desolata and then

to Subantarctic Indian Ocean salvini, Subtropical

Indian Ocean vitiata (macgillivrayi) and vittata of

the southern Atlantic and New Zealand regions to

indicate their progressively preater bill sizes.

Falkland [s. belcheri are shown isolated NW of

Scotia Sea desolata and connected by an arrow to the

Subantarctic Indian Ocean islands where intermedi-

ates between belcherj and other whale-birds breed.

Figure 19 indicates that two taxa (desolata and

NORTH

O-—_w—_—

©—-@-—

ATLANTIC

INDIAN

Fig. 19—Geographical variation in bill size of whale-hirds.

Arrows point to larger-billed forms. D=desolala, S=salvini.

V=viltai Arrow from B=belcheri points to H=hybrid

belcheri x desolata—salvini. Intergradation of bill size occurs

in thé southern Indian Ocean region.

116 REC. S. AUST. MUS. 18(4): 91-121

vittata) are allopatric in the southern Atlantic and

New Zealand regions, but are connected by

intermediate forms (salvini) in the southern Indian

Ocean region.

The intergrading characteristics of desolata and

vittata can also be demonstrated by their bill

dimensions. In the southern Atlantic and New

Zealand regions, where there are no intermediate

salvini populations, the differences in their mean bill

widths are 7-1 and 7-3 respectively. But in the

southern Indian Ocean region where intermediate

salvini populations also breed, the difference

between desolata (excluding fles Kerguelen and fle

de I‘Est forms which hybridize with belcheri—see

below) and vittata in mean bill width is only 3-3.

Heard I. desolata have a slightly larger bill than

other desolata (Fig. 13) and fle Saint-Paul vittata

have a smaller bill than other vittata (Fig. 16). The

salvini populations, having varying mean bill sizes,

are probably of hybrid origin but have not

themselves become a new specific entity because in

their range of variability (bill width from 14-0 to

20-5) they include the characteristics of both

parental phenotypes. Gene-flow through the appa-

rent hybrid swarms to the parental populations

indicates that desolata and vittata are conspecific.

Hybridization of Thin-billed and Broad-billed Whale-

Birds

Differences between belcheri and members of the

desolata-salvini-vittata complex indicate that paren-

tal populations of both were isolated for a prolonged

period and that the intermediate forms of the

southern Indian Ocean region result from hybridiza-

tion.

At fles Kerguelen belcheri is much less numerous

than desolata (Falla 1937) and although specimens

have been taken at different localities in the

archipelago, Derenne et al. (1974) mapped only one

belcheri colony as compared to thirteen desolata

colonies. The characteristics of birds from only three

colonies are known, but of these one consists of

birds which closely resemble belcheri parental

phenotypes; another apparent desolata parental

phenotypes; and a third consists of birds with

intermediate characteristics (fig. 13). Also, speci-

mens taken elsewhere in the archipelago show that

the characteristics of both forms intergrade (fig. 12:

BS5-B13). It is therefore highly probable that many

Tles Kerguelen prions are hybrid belcheri x desolata.

Nevertheless, because prions usually return to their

natal colonies for breeding (Fleming 1941) each

colony may consist of very similar birds that differ

slightly from birds of other colonies. The fles

Kerguelen prion assemblage probably therefore

consists of locally different parental and hybrid

populations.

May, 1980

Despin et al. (1972) said that on fle de Il’Est salvini

are abundant and breed over most of the island

wherever suitable nesting sites are available. They

also reported a very small isolated colony containing

both belcheri and desolata, but the characteristics of

these birds appear analogous to the intergrading

characters of belcheri and desolata of fles Kerguelen.

However, the similarity in tarsus and wing lengths of

these fle de l’Est belcheri and desolata indicates they

are more closely related to each other than to any

other populations of either form. The data suggest

they were only separated by bill size, and this

distinction appears quite artificial, The alleged

desolata appear to be hybrid belcheri x desolata, but

they may well be hybrid belcheri x salvini because in

bill, tarsus and wing dimensions they are intermedi-

ate between belcheri and salvini of the same island.

A hypothetical hybrid belcheri x salvini would

probably closely resemble true desolata. The

apparent isolation of these probable hybrids and

belcheri in one small colony suggests they are

unlikely to extensively and freely interbreed with

salvini from outside the colony.

Cawkell and Hamilton (1961) said belcheri is very

numerous in the Falkland Is. and is greatly

increasing its numbers and colonizing new islands.

This increase suggests that elsewhere belcheri may

have encroached upon the breeding range of other

whale-birds.

(d) DIFFERENCES IN BREEDING TIMES

Prions of each island population have probably

developed a rather stable breeding cycle adapted to

the local environment. Fleming (1941) reasoned that

differences in breeding dates of the species he

recognised were effective isolating mechanisms.

However, similar forms breed at different times that

correspond to latitude and climate. Birds breeding

further south in a cooler climate usually have later

egg-laying dates than those breeding in northern

warmer latitudes. These facts (detailed below), and

that belcheri and desolata hybridize although their

parental populations usually breed at different

times, suggest that differences in breeding cycles

have not yet become effective isolating mechanisms.

Fairy Prions

Fleming (1941) said that turtur usually lay eggs in

mid-October while crassirostris, which he recognised

as of a cooler zone, lay in mid-November. But all

turtur populations do not breed synchronously.

Buddle (1941) said birds of the northern Poor

Knights Is. lay eggs about 15 October, Crockett

(1954) suggested the more southerly Montunau I,

birds breed slightly later, and Richdale (1965) said

the peak egg-laying date of a southern New Zealand

BREEDING DISTRIBUTION AND TAXONOMY OF THE FPRIONS 117

population is about 16 November, Subantarctic

Marion J. turiur also probably lay eggs in mid-

November, for van Zinderen Bakker (1971) said

that; “ar the end of February 1966 two nests were

found both containing nearly fully fledged chicks

which were ready (o depart to sea,”’ The incubation

period of this species is about 56 days and the

fledging petiod 44-55 days (Serventy et al. 1971),

Snares Is. turtur also probably lay in mid-November.

judging from dates on which chicks were found (see

above— Breeding Diswibution), and two chicks

found on eastern Sister islet in the more northerly

Chatham Group, listed by Dawsan (1955) as the

“Chatham fulmar prion’, must have come from eggs

laid in October; the normal laying time of northern

turtur. Therefore the breeding times of many /urtur

and crassirostris are similar.

Whale-Birds

The whale-birds also difter in breeding times

according to latitude and climate. On Tristan da

Cunha vitiaia usually completes egg-laying by the

end of August (Eliott. 1957). whereas further south

on Gough |, eggs are laid in nid-September (Swales

1965). The peak egg-laying date of southern New

Zealand vitlata is about 6 September (Richdale

1944). Subantaretic salvini lay about mid-November

(Rand 1954+) and antarctic desolata lay eggs

throughaut December (Tickell 1962). Like sulvini

sub-antaretie belcheri also lay their eges mainly in

mid-November (Falla 1937). Some of these prions

have flexible breeding times which indicate that

different forms may meet when jna similar breeding

condition, On Gough 1. a fresh vittara egg was found

in mid-November (Swales [965) antl Falla (1937)

found a desalata on Tes Kerguelen (parental

phenotype, form A) in a burrow with a fresh egg on

19 November. Tickell (1962) said of antarctic

desolata that “eggs are laid throughout December,

the earliest record being 5 December” and that some

eggs are laid as late as January if snow blocks nesting

holes.

There is little consistency in the breeding tintes of

hybridizing populutions, At Tles Kerguelen heleleri

lay exes in November and most desolara lay in

December (though an egg is known from mid-

November), but form B oof probable mixed

parentage have ua Garber eyele than other desolate

(Falla 1937), Despin et al, (1972) said that on fle de

(Est beichert and salbini breed at the sume time. but

that desolara breeds later. However, they presumed

heleher? hasan incubation period 1) days longer than

these alleged deselata when calculating laying dates

fram We belche chieks and an unspecified mumber

of alleged desolata eggs, While incubation periods

do vary. [here is no reason bo suppose the periods of

these birds chffer, Tp is also odd thar they listed the

dimensions of fle de l'Est belcheri eggs, but did not

mention finding any, whereas they did not give

dimensions of the alleged desolata eggs, Accepting

that they assigned the eggs to the right (doubtfully

separable) adults and that their judgments of lapsed

egg incubation times were correct, the data of

Despin ev al. indicate in fact that two belcheri chicks

came from eggs laid 15 days apart, and that the

alleged desolata eggs were laid 23 days later, While

the laying dates of the last eggs are not known, it is

reasonable to presume that some eggs were laid

earlicr than others. Therefore in this small colony

there is wide variation in laying dates, a situalion

very similar to the different breeding times of Mes

Kerguelen birds. Because different breeding times

do not separate forms on fles Kerguelen it is

doubtful that they act as isolating mechanisms on fle

de Est. [t must also be considered that the alleged

desolata are probably hybrid and their different

laying times may be the result of some selective

factor that enables the parental forms to retain their

awn identity.

(e) FEEDING HABITS

The feeding habs of Australian ¢irar are

probably similar to those of belchert, but both differ

in feeding behaviour from other whale-birds,

Robinsan (1961) observed mtur diving from che

surface of Port Phillip Bay. Victoria, to feed on

small whitish crustacea. On surtacing they ran acrass

the sea with wings held horizontally before alighting

at another spot preparatory to diving again, | have

seen surtur feeding in a similar manner, but off the

South Australian coast have also seen them leeding

in large flocks, picking from the surface while in

flight. pattering with their feet and fluttering

erratically over the sea. Serventy eral (1971) listed

the food of utr as “Crustacea” and thal of

crassirosiris as “Pretopeds and orher molluses.

amphipods, small fish’. but Watson (1975) also

listed “Small crustaceans” in the diet of crassirosiris

and “Small squid’ in the diet of natar

Harper (1972) said heleheri largely feed on the

amphipod Pararhemisto gaudichaudil and less atten

on small cephalopods, and observed them at night

“dashing helter-skelter over [he water feeding

voraciously on surface plankton like storm petrels

-oceasionally patting the water with the feet to

obtain extra momentum’. On the other hand he

found that desalata feed mainly in the day in @

manner described by Murphy (1936), who said “the

birds worked along with an odd creeping matin,

resting their bodies lightly upon the surface but

holding the wings just above it. the feet furtshing all

of the motive pawer.” The major food of dewlant,

salrin? and Litrdta are €rustaceans, particularly the

Krill EBupluiisia superba of southern waters.

118 REC. S. AUST, MUS. 18(4): 91-121

although cephalopods, pteropods and small fish are

also taken (Murphy 1936; Serventy ef al. 1971;

Harper 1972; Watson 1975),

IV. DISCUSSION

There is agreement that vittata evolved from thin-

billed ancestors (Murphy 1936; Falla 1940; Fleming

1941), but Fleming (1941) also hypothesized that

crassirostris has retained many features of the

ancestor of all prions. This latter hypothesis means

that thin-billed forms evolved from ancestors

resembling modern crassirostris, which is antithetic

because the phyletic lineage of thin-billed forms

would probably have included transitional forms

resembling modern turtur, yet Fleming believed that

turtur and crassirostris only recently diverged from a

single common ancestor. However, there is evidence

indicating that the common ancestor of prions had

an elongated bill with a large but narrow dertrum,

like the bills of Australian and northern New

Zealand turtur. This evidence is gained by comparing

prions with close relatives,

Monotypic Halobaena is very closely-related to

Pachyptila. It is also close to Pterodroma but differs

in having a prion-like palate (Alexander et al. 1965),

Contrary to Alexander et al. (1965), Harper (1978)

said, when summarizing results of plasma protein

studies, that it is “probable” that ‘“‘Pachyptila and

Halobaena are only very distantly related; the prions

evolved from a small fulmarine ancestor, whereas

the Pterodroma group gave rise to the blue petrel.””

Nevertheless, Harper did not have plasma samples

of Halobaena and he aligned it with Pterodroma and

not Pachyptila because the former two show “little

tumescent development of the latericorns” (p,529).

However, it is generally agreed that primitive prions

had narrow bills and that broad-billed forms evolved

later (Murphy 1936, Falla 1940, Fleming 1941), and

some modern prions lack broad latericorns. Harper

also compared the long nasal tubes of fulmarines to

the short tubes of Pterodroma, but in respect of these

Pachyptila and Halobaena have consistently short

tubes. Although Halobaena has similar plumage

characters (except crown and forehead) to those of

prions and these may have resulted from converg-

ence (Harper loc. cit.), it also differs from

Pterodroma in lacking an entirely black bill (Cox

1976). The question of the relationships of these

genera must remain open because little evidence

supports the hypothesis that Pachyptila is more

closely-related to fulmarines than to Halobaena, or

the hypothesis that Halobaena is more closely-

related to Pterodroma than to Pachyptila. Therefore

there is little reason as yet to dispute the opinions of

Alexander et al. (1965) that Halobaena is closely-

related to Pachyptila and Pterodroma. The bill of the

May, 1980

Blue Petrel Halobaena caerulea greatly resembles

that of belcheri although viewed in profile its

dertrum is larger and more strongly hooked,

resembling some slender-billed turtur. Viewed

dorsally the dertra of Halobaena and _ small

Pterodroma, particularly of the subgenus Cookilaria,

are narrow and similar to those of turtur and belcheri

but unlike that of crassirostris. Slender-billed turtur

and belcheri are the only forms of each prion group

with similar bills, and the only prions which

resemble in bill characteristics birds of closely-

related genera.

Salient aspects of modern prion distributions also

indicate that turtur is an ancient form. In the

Australian and New Zealand region its populations

are large and abundant, but in the southern Atlantic

and Indian Ocean regions, where whale-birds are

numerous, it is only known from a few isolated

colonies. Together with the fact that turtur has bill

characteristics like those of belcheri only in the

Australian and northern New Zealand region

(Marion I, rurtur have thin, but short bills) where the

breeding ranges of the two prion groups do not

overlap, these aspects suggest ancestral turtur were

widespread but that only remnant populations

survived competition with ancestral whale-birds in

the western parts of their range. In areas of

overlapping range turtur has a stouter bill, indicating

that both prion groups were able to co-exist only

after ecological adaptations evolved,

It is evident that populations of the common

ancestor of whale-birds became isolated. Some, the

progenitors of belcheri, did not diverge in morphol-

ogy to any great extent, but others appear to have

utilised the teeming crustacea of the Southern Ocean

as their major food source and through selection

developed a different bill structure. These birds,

having adapted to a food supply largely unexploited

by other prions, successfully spread and, eventually,

their populations became isolated. The northern

birds developed larger bills (conforming to Allen's

Rule that warm-blooded animals have greater

surface areas in warm regions, and have a reduced

surface area—volume ratio in cooler regions as an

adaptation to prevent heat loss—Mayr 1963) and in

two regions where they are not connected by zonally

intermediate populations to southern smaller-billed

forms they appear quite distinct. However, their

different bill sizes may be purely adaptive

characteristics which are not necessarily indicative of

fundamental changes in genetic composition,

because where intermediate forms occur the

differences in nearby parental forms are less

pronounced. The early isolate from common

ancestral whale-bird stock, belcheri, presumably

increased in numbers and range and, where it

colonised islands already occupied by wide-billed

whale-birds, interbreeding occurred.

BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 119

Classification

Fundamental to a valid classification is am

understanding of the relationships of the different

prions, and recognition of their dynamic evoluno-

nary trends, Previous classifications of prions have,

to say the least, artificially categorised populations

by doubttul morphological characteristics. The

synonymy is vast and confusing. Condon (1975)

listed 58 names, of which 34 were bestawed by G.

M. Mathews, Hellmayr and Conover (1948) said

Mathews had “within a years time, based Iwo

different names on the very same specimen in the

British Museum”. A major cause of this confusion ts

the inter- and intra-specific variation in prion bills.

Beach-washed specimens far from their unknown

natal home were sometimes named as distinct forms

because of bill differences (Mathews 1912), but

substantial collections in the Australian Muselm

and the South Australian Museum show complete

intergradation in the characteristics previously used

to distinguish species. Intermediates are numerous.

Smaller samples from many breeding localities

demonstrate similar inconsistencies.

The characteristics of tirtir and crassirostris

intergrade over a wide area, indicating canspecific-

ity, Many island populations appear to have slightly

diverged in different directions and a dimensional

unalysis of specimens is required to clarify

geographic variation. Both forms have been subject

to recent pene-flow but their respective ranges

appear ta be clearly demarcated in the Chatham

Group although delimitations elsewhere are unclear,

As iin interim measure to indicate the relaionships

of populations, Heard I. and Auckland Is. birds are

here classified as u phenotypie tarm that ts

intermediate between crassirosinis of Pyramid Rock

and the Bounty Ts, (including the twa specimens

alleged to he from Antipodes Is.), and winur of the

Falkland Is.. Marion |., Australia and New Zealand.

Differences in morphology and feeding habits

indicate that belchen’ is evolving on a different

evolutionary line from other whale-birds. Parental

populations of both are thriving and in their zone of

overlap and hybridisation it ts probable, at one

locality, that hybrids are disadvantaged.

The other whale-birds consist of two taxa

connected by intermediate populations in a wide

area that contains few, if any, parental phenotypes.

Because this zone of intergradation is between two

regions where both taxa are clearly demarcated in

allopatric breeding distributions, it is probable that

intergradation occurred through hybridization. By

virtue of the island distribution of these probable

hybrid populations, and to illustrate clearly their

relationships, they are here tentatively classified as a

phenotypic form. Other than desolata, salvini and

vittatg, Subspecific recognition of other slightly

different papulatians within the species is arbitrary,

Tickell’s (1962) recognition of banksi and alter =

macquariensis is not followed here because the

characteristics of cach vary over a wide area,

indicative of slight geographic variation, and few

individuals would be identifiable. In view of the

occurrence on files Kerguelen of both desolata and

desolata x belcheri, the identity of the lost holotype

of desolaia may be questioned. In the absence of

evidence to the contrary, [ here consider the type of

desolata to have come from the ‘desolara phenotype

colonies rather than from one of the hybrid colonies.

If the type of desolata (Gmelin, 1789) is found ta be

hybrid, banksi Smith, 1840 should be resurrected as

the name for this southern subspecies of P. vittara-

The following tentative classification is proffered

until sufficient evidence suggesting otherwise is

forthcoming. In the hope of preventing: possible

future misinterpretations of “possible” or “‘prob-

able’ breeding grounds. only proven breeding

loculities of each taxon are listed. In both subspecies

and species the taxa of presumed oldest lineage are

listed first. Condon (1975) listed synonyms of the

scientific nomenclature used. and the English names

proposed by the Royal Australasian Ornithologists

Union (1978) are followed; one is of common usage

and two are descriptive.

Genus Pachyptila [Ihger. 1511,

Pachiyplila turtur (Xuhl, 1820) FAIRY PRION

Pachyprtila turtue turtur (Xuhl, 1820)

Breeds: Beauchene |, Falkland Group: Manan

L.; islands in Bass Strait and off Tasmania,

islands off New Zealand irom the Snares Is.

North to the Poor Knights 1s.; Big and Little

Mangere Is., Chatham Group.

Pachyptila turtur eatoni (Mathews, 1912)

Breeds: Heard I. and the Auckland Is,

Pachyptila turtur crassirasiris (Mathews, 1912)

Breeds: Bounty [y., and Pyramid Rock of the

Chatham Group.

Pachyptila belcheri (Mathews, 1912) SLENDER-

BILLED PRION

Breeds: Falkland fs., Tle de VEst of Archipel

Crozet, and fles Kerguelen.

N.B. Interbreeds with the following species where

both occupy the same islands.

Pachyptila vitrata (Forster, 1777) BROAD-BILLED

PRION

Pachyptila vittata desolata (Gmelin, 1789)

Breeds: S. Shetland Is., S. Orkney Is., &,

Sandwich Is., S. Georgia, fles Kerguelen,

Heard J,, Macquarie I,, Auckland Is., Scott [-

and Cape Denison of Antarctica,

120

Pachyptila vittata salvini (Mathews, 1912)

Breeds: Marion and Prince Edward Is., and

Archipel Crozet.

Pachyptila vittata vittata (Forster, 1777)

Breeds: Tristan da Cunha, Gough I., fle Saint-

Paul, fle Amsterdam, Snares Is., Chatham Is.

and the mainland coasts of Foveaux Strait and

south-western fiords of South I., New Zealand

and islets off Stewart I.

V ACKNOWLEDGEMENTS

I am indebted to the Director and staff of the

South Australian Museum for providing the facilities

necessary to undertake this work, particularly to Mr.

S. A, Parker (Curator of Birds) who gave much time

to discussion and constructive criticism. For

providing facilities in the Australian Museum I

thank Mr. H. J. De S. Disney and Mr. W. Boles. I

also thank the Directors and staff of other museums

for the loan of specimens, Dr. F. C. Kinsky, Mr. A.

R. McEvey and Mr. J. L. McKean for assistance in

locating material, Dr. D. H. Close for translating

references, Mr. L. Joseph for analysing dimensions,

and Mr. R. Ruehle for taking the photographs.

Special thanks are due to Mr. R. F. Brown, Dr. G.

W. Johnstone, Mr. W. B. Hitchcock and Mr. S. A.

Parker for their valuable comments and criticisms of

the manuscript.

VI REFERENCES

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BUDDLE, G. A. 1941. Birds of the Poor Knights. Emu 41: 56-

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CAWKELL, E. M. and HAMILTON, J. E. 1961. The Birds of

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DERENNE, P., LUFBERY, J. X. and TOLLU, B. 1974.

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FALLA, R. A., SIBSON, R. B. and TURBOTT, E. G. 1966. ‘A

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BREEDING DISTRIBUTION AND TAXONOMY OF THE PRIONS 121

RICHDALE, L. E. 1965, Breeding behaviour of the Narrow-

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EVOLUTIONARY SYSTEMATICS OF XENYLLA.

XI. SPECIES FROM THE AUSTRALIAN REGION

(INSECTA: COLLEMBOLA)

BY MARIA MANUELA DA GAMA

Summary

In some material from all states of Australia the author found twelve Xenylla species, one of which

— X. victoriana, is new. In addition the chaetotaxy of X. littoralis from Australia and Japan was

studied. The systematics and evolution of some of these taxa are considered.

EVOLUTIONARY SYSTEMATICS OF XENYLLA. XI, SPECIES FROM THE AUSTRALIAN REGION

(INSECTA: COLLEMBOLA)

MARIA MANUELA DA GAMA

Zoological Museum of The University of Coimbra, Portugal.

ABSTRACT

GAMA, M.M.da, 1979, Evolutionary systematics of Xenylla. XI,

Species from the Australian region (Insecta: Collembola). Rec. S.

Aust. Mus, 14(5); 123-129.

In some material from all states of Australia the

author found twelve

which—X. victoriana, is new. In addition the

chaetotaxy of X, litroralis from Australia and Japan

was studied. The systemati¢s and evolution of some

of these taxa are considered.

INTRODUCTION

Following my studies on some Xenylla species

from Australia, principally from South Australia

(Gama 1974), Mrs. Penelope Greenslade of the

South Australian Museum, Adelaide, kindly sepa-

rated out for me the Nenylla species from 62 samples

which she had assembled from all States of

Australia, the Torres Strait Is., Tasmania and New

Caledonia. Most of the material had been collected

by Mrs. Greenslade herself.

The material contained 12 species, of which one ts

new, but the most interesting find is, in my opinion,

that of X, littoralis, which also lives in the littoral

zone in Japan.

Mrs, P. Greenslade also sent me the H,

Womersley collection of examples of this genus

which has been very useful for my study, especially

when considering X. liftoralis,

The material is deposited in the Institutions cited

below, whose names have been abbreviated in the

text as follaws:—

SAM— South Australian Museum, Adelaide

ANIC— National Insect Collection, c/o C.S.1.R0.,

Division of Entomology, Canberra

MG— Muséum d'Histoire naturelle de Geneve

Mo Museu Zooldgico da Universidade de

Coimbra

SYSTEMATICS AND EVOLUTION

SPECIES

OF THE

1, Xenylla welchi Folsom, 1916

Xenylla welchi Folsom, 1916, p. 497.

Material ¢xamined—Launceston, Tasmama, in

horse manure, garden, very numerous specimens,

9.v.1977. Some specimens in alcohol (SAM); some

specimens, in alcohol (ANIC) and some specimens,

in ulcohol (MC),

Xenylla species, one of

2. Xenylla littoralis Womersley, 1934

Xenylla littoralis Womersley, 1934, p. 56.

Description—Body length, 1.2-2.1 mm, Blue.

Cutaneous granules coarse and third pleurite

possessing a lateral conical projection.

Chaetotaxy consisting of supernumerary setac,

sometimes very numerous, principally in those

specimens with a long body, and often very

irregular. Therefore I found it very difficult to

determine the chaetotaxie formula of this species.

On the drawing of dorsal chaetotaxy (fig. 1), the

dotted line setae represent the most frequent

supernumerary setae, but most specimens have

many more. Moreover in this species one can

distinguish a differentiation into macrochaetae and

microchaetae, which is unusual in the genus Xenylla.

Dorsal chaetotaxy (fig. 1):

Head: all

(character [),

Th.1-I[l; all setae present and central setae

arranged in three rows; there are 2 §.s. on each side,

one of which in position P,.

Abd. I-UL; S.s.=P,; ps present.

Abd. IV: §.5.=Ps; py absent (character n). I am

not sure whether aj is, or is not, present.

Abd. V: all setae present.

Ventral chaetotaxy:

Head: I think that one may consider that all setae

are present, although p, or m,; are absent on some

specimens (fig. 1 in Gama 1969: 4).

Th.U-Uk: without setae (character t).

Abd. [[: p, and p, absent (character vy) and p, also

(character w),

setae present; L, longer than L,

Abd. IIT: without medial setae nor median seta.

Abd. [YV: all setae present,

Antenna IV with four sensillae, of Which the twa

most external of the three dorso-external are very

long, the ratio between the length of the other two

sensillae and the length of these ones being more or

less 3:4,

5+5 eyes.

Unguis with inner tooth, Tibiotarsus with two

dorsal tenent hairs, Tenaculum with 2+2 barbs,

Mucro very long, distinctly separated from dens,

which has two setac, tapering to a fine point and

124 REC. S. AUST. MUS. 18(5): 123-129 May, 1980

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)

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Fig. 1—Xenylla littoralis. Dorsal chaetotaxy of head, th. I-II and

abd. I, IV, V, VI.

EVOLUTIONARY SYSTEMATICS OF XENYLLA 125

possessing one inner narrow lamella. Mucro more or

less equal in length to dens, a little longer or a little

shorter (fig. 41-E in Womersley 1939: 93).

Anal spines and papillae very well developed.

Systematics and evolution—When Prof. Yosii, in

1972, sent me one sample of Xenylla from Japan

(see material examined) he thought that it perhaps

belonged to X. littoralis, but added that Australia

was quite a distance from Japan!

Several times I looked again at this halophilous

species, without arriving at any conclusion.

When only some months ago, I had the chance to

compare this material and the material cited below

with the Womersley collection, and was able to

confirm that the Japanese species is identical with X.

littoralis from Australia, not only in the adaptative

characters but also in the chaetotaxic characters (fig.

1).

The examination of three paratypes of X. arenosa

Uchida and Tamura, 1967, a Japanese species which

also lives in the seashore, reveals that it can be

separated from X. littoralis by some fundamental

nonadaptative characters, namely the character b

(cephalic setae p,p,; absent) although the central

setae on th. II-III are arranged in three rows

(absence of characters h, and hj) as in X. littoralis.

I think that the principal adaptative characters by

which the Womersley species is separated from X.

arenosa are that the former species has two tenent

hairs on its tibiotarsus while the latter has only one;

furthermore X. littoralis possesses anal spines but X.

arenosa does not; finally X. littoralis has a less

tapered mucro than X. arenosa.

Material examined—South Australia: Yorke

Peninsula, Jolly’s Beach, 21 mi. E. Sleaford Bay,

under stones between tides, 7 specimens, leg. P.

Greenslade, 3.ix.1975—1 specimen, on_ slide

(SAM); 1 specimen, on slide (ANIC); 1 specimen,

on slide (MG); 4 specimens, on slide (MC); Yorke

Peninsula, Royston Head, on beach between tides, 8

specimens, leg. P. Greenslade, 27.1.1976—2 speci-

mens, in alcohol (SAM); 2 specimens, in alcohol

(ANIC); 2 specimens, in alcohol (MG); 2

specimens, in alcohol (MC); Christies Beach, 4

specimens, 25.i.1932 (identified by Womersley).

Western Australia: Rottnest Island, 1 cotype,

31.1.1931 (identified by Womersley); Marino Rocks,

2 specimens, 25. XII. 1939 (identified by Womers-

ley). Japan: Cape Tachimachi, near Hakodate,

Hokkaido, among the debris thrown on the sea

shore, 9 specimens, leg. R. Yosii, 5.viti.1971. These

specimens are on a slide and in alcohol (MC).

3. Xenylla thibaudi thibaudi Massoud, 1965

Xenylla thibaudi Massoud, 1965, p. 374.

Observations—This is the first time that X.

thibaudi thibaudi has been found in Australia, but it

has been already found in New Guinea, the Solomon

Islands and the Bismarck Archipelago.

Material examined—Queensland: Cooloola, rain

forest litter, 7 specimens, leg. P. Greenslade,

23.ii.1977—in alcohol (MC); Idem, 3 ad. + 3 juv.,

16.i1.1977—in alcohol (ANIC); Cooloola, rain forest

pitfalls, 2 specimens, leg. P. Greenslade, i1.1977—in

alcohol (MG); Cooloola, Warrawonga pitfalls, 2

specimens, leg. P. Greenslade, 11.1977—in alcohol

(SAM); Cooloola, Warrawonga leaf litter, 8

specimens, leg. P. Greenslade, 16.i1i.1977—in

alcohol (MG); Idem, 12 specimens, 23.ii.1977—in

alcohol (ANIC); Idem, 40° specimens,

29.ii1.1977—in alcohol (MC); Cooloola, Birwillah

litter, 12 specimens, leg. P. Greenslade,

16.ii.1977—in alcohol (SAM). New Caledonia:

Metzdorf Valley, litter, 5 juv., leg. Gross,

5.1.1966.—in alcohol (MC).

4, Xenylla yucatana Mills, 1938

Xenylla yucatana Mills, 1938, p. 183.

Material examined—Torres Strait Islands: Murray

Is., litter from vine, bamboo forest, about twenty

specimens, leg. Cameron, 17.viil974—in alcohol

(MC); Murray Is., leaf litter, about fifteen

specimens, leg. Cameron, 17.vii.1974—in alcohol

(SAM).

5. Xenylla womersleyi Gama, 1974

Xenylla womersleyi Gama, 1974, p. 81.

Material examined—South Australia: Nuyts

Archipelago, Maselon Island, litter from Westringia

sp., 5 specimens in poor condition, leg. P.

Greenslade, 17.i1.1977—in alcohol (MC).

6. Xenylla australiensis australiensis Gama, 1974

Xenylla australiensis Gama, 1974, p.75.

Synonym—Xenylla mucronata Womersley, 1939

non Axelson, 1903.

Systematics—The Womersley collection includes

other specimens, mounted in slides, named as X.

mucronata, coming from the You Yang Mountains,

Wartook, and also Kenwick (see Womersley 1939:

92).

However, only the specimens cited below could be

studied in this work, because the other specimens

are not in good condition.

Material examined—Western Australia: Ragged

Mountain, 70 mi S Balladonia, near Israelite Bay,

litter under Petrophile sp, 1 specimen, leg. P.

Greenslade, 24.v.1977—in alcohol (SAM).

Victoria: Coranderrk Reserve, Healesville, about

fifteen specimens, 27.1.1976—in alcohol (ANIC);

Beaufort, Boxer Cutting, in grass, 1 specimen, leg.

126 REC, 5. AUST, MUS, 18(5): 123-129

P, Greenslade, 12,vi.1975—in alcohol (MG); You

Yang Mountains, forest Reserve, fram log, 1

specimen, leg, P, Greenslade. 9.vi.1975—in alcohol

(MC); You Yang Mountains, 2. specimens,

23,ix,1931 (identified by Womersley as X. miuc-

ronata); Fish Falls, Wartoak, 3 specimens, 30. XII.

1939 (identified by Womersley as X, mucronata).

South Australia: Innes National Park, Yorke

Peninsula. litter, 35 specumens, leg. P. Greenslade,

2.0.1974—in aleohol (SAM); Yorke Peninsula,

south central, mallee litter, 15 specimens, leg. P,

Greenslade, 2.11.1974—in alcahol (MG); 7 km NW

Morgan, Casuarina litter, 1 juv., leg. P. Greenslade,

17.xti,.1976—~in alcohol (ANIC); 7 km NW Morgan,

pitfalls, 22 specimens, leg. P. Greenslade, 15-

(7.xii,.1976—in aleohal (MC); near Loxion, mallee

leaf litter, 2 specimens, leg. P. Greenslade,

25,1x,1974—in alcohol (SAM); Nuyts Archipelago,

Maselon Island, litter under bush, 1 specimen, leg.

P, Greenslade, 171.1977—in alcohol (ANIC);

Belair, 9 specimens, Jeg P, Greenslade, 28

iv,1971—1n_~ alcohol (MG); Coorong, Coolatoo,

disturbance pitfalls, clay pan, in grass, 10 specimens.

leg. P Greenslade, 13.x.1975—in alcohol (MC):

Coorong. Coolatoo, disturbance pitfalls, clay pan, in

bushes, 6 specimens, leg. P. Greenslade, 28.ix.-

13,x.1975—in alcohol (SAM); Mt. Bold, Finniss

Raver, rotten log, about thirty juv., leg. P.

Greenslade, 4.¥.1975—in alcohol (ANIC); Glenelg

River Reserve, Evicalypius leaf litter, 12 juy.. leg. P.

Greenslade, 18,v.1975—n alcohol (MG): 12 km N

of Mt. Gambier, needles under Pinus radiata

planted 1929, 3 juy,, leg. P, Greenslade,

19,y.1975—in aleahol (SAM),

7. Xenylla greensladeae Gama, 1974

Xenylla greensladeae Gama, 1974, p. 72.

Systematics—Among the Womersley material

there are 4 specimens, on slides, labelled as X. prisca

i.sp., Which as far as I know, has never been

described.

l have been able to study two of these specimens

which were found to be X. greensladeae.

Material examined—Queensland: 15 mi E Killar-

ney, rain forest near Queen Mary Falls, leaf litter, 5

specimens, leg. P, Greenslade, 16.v,1974—in

alcohal (SAM); Emu Creek, 20 mi E Warwick, leaf

jitter, about fifty specimens, leg. P. Greenslade,

14.v.1974—in alcohol (ANIC). Western Australia:

Ashburton River, stony flood plain, 5 specimens,

leg. P. Greenslade, 22.vi.1975—in alcohol (MG),

Australian Capital Territory: Brindabella Range nor.

Picadilly Circus, Eucalyptus forest leaf letter.

iumerous specimens, leg. P, Greenslade, 26. vii-

May, 1980

i.1972—in alcohol (MC), New South Wales: Dorrigo

Run, pitfalls, 1 specimen, leg. Weir, 1.1974—in

alcohol (SAM). South Australia: Yorke Peninsula,

Innes National Park, tall mallee, 9 specimens, leg, P,

Greenslade, 14.x.1974—in alcohol (ANIC); Ferries-

McDonald Reserve, 3 specimens, leg. E. G.

Matthews, [3.x.1977—in aleohol (MG): Belair, in

moss, | specimen (identified by Womersley as X,

prisca nesp.) 26,ix,1943; 7 km NW Morgan,

Casuarina litier, 4 specimens, leg. P. Greenslade,

17.xi1.1976—in aleohol (MC); Idem, 1 juv.—in

aleohol (ANIC): Morgan, soil samples. 2 specimens,

leg. Butler, 1974—in alcohol (SAM); 7 km NW

Morgan, pitfalls, 12 specimens, Jeg. P. Greenslade,

15-17.xii. 1976—in alecohal (MC): Morgan, under

mallee, 35 specimens, leg. Hutson, x,1974—1n

alcohol (SAM); Coorong, Coolatoo, disturbance

traps, clay pan, in grass, 5 specimens, leg. P.

Greenslade, 13,x.1975—in alcohol (MC); Coorong,

Banff Transect, 35 specimens, leg, P, Greenslade.

¥.1975—an alcohal (MG), Muston, (Kangaran

Island), in moss, 1 specimen (identified by

Womersley as X. prisca i.sp.), 23.vin.1943.

Victoria: Tarra Falls Valley, in Brass, | specimen.

leg. P, Greenslade, 11_vi.1975—in alcohol (ANIC),

8, Nenylla meritima Tullberg, 1869

Xenylla maritima Tullberg, 1869, p.11.

Material examined—South Australia: 75 mi N Mt.

Gambier, Acacia/Casuarina litter, SAM 325. 16

specimens in poor condition, leg. P. Greenslade,

19.v,1975—in alcohol (SAM); Kiutpo Forest, on

dead Pinus radiata branches, 10 specimens, leg. P-

Greenslade, 20.x.1974—in alcohol (ANIC); Glen

Osmond, 3 specimens, iii.1935 (identified by

Womersley). Western Australia: Perth, 1 specimen,

23.v.1931 (identified by Womersley). Victoria

Studley Park, 1 specimen, viii,1931 (identified by

Womersley},

9. Xenylla vicfonana n.sp.

Holotype and Paratypes: Victoria—Otway

Ranges, fern forest litter, 45 paratypes, leg, P.

Greenslade, 8.vi.1975. Holotype, on slide (SAM); 5

paratypes, an slide (MG); 7 paratypes, on slide. and

32 paratypes, in alcohol (MC), Erskin River,

Eucalyptus woodland, litter and logs, 20 paratypes,

leg. P. Greenslade, 8.vi.1975, 16 paratypes, in

alcohol (SAM) and 4 paratypes, on slide (MC);

Toorangi State Forest, litter and logs, 4 paratypes,

leg. P_ Greenslade, 1f,vi,1975—in alcohol (ANIC);

Tara Valley Reserve, falls leaf litter, 3 paratypes,

leg. P. Greenslade, 11.vi.1975—in alcohol (MG);

Tarra Falls, Wall Valley, in grass, 1 paratype, leg. P

Greenslade, 11.v1.1975—on slide (ANIC).

EVOLUTIONARY SYSTEMATICS OF NENYLLA 27

Description

Body length 0-62-0-92 mm. Blue, Cutaneous

granules fine. Dorsal chaetotaxy revealing the

following characters:

Head: p, is absent (character b); L, longer than Ly

(character f).

Th.ILIL; all setae present, central setae arranged

in five rows (characters h, and h,) and with 2 8.8, on

each side, one of each in position Py,

Abd. I-Il: S.s.=P,: ps present.

Abd. IV: S.s.=Ps;; a; present.

Abd. V: S.s.=P3; a) absent (character q).

The characters of ventral chaetotaxy are as

follows:

Head: all setae present,

Th. II-III: one seta on each side.

Abd. II: two medial setae.

Abd. Ill: without medial setae nor median seta,

Abd. 1V: m, is absent (character a,); m, is absent

(character a,)—This is the first species of Nerrylla in

which I have found character as,

Antenna IV with four cylindrical similar sensillae.

5+5 eyes.

Unguis without visible inner tooth, Tibiotarsus

with two dorsal tenent hairs. Tenaculum with 3+3

barbs, Mucro with curved apex, possessing an inner

lamella and separated from the dens, which has two

setae (fig. 2). Length of mucro about one half the

length of dens; mucro shorter than claw, the ral

between these two structures is between 3:5 and 3:4.

Anal spines on anal papillae with usual conforma-

tion.

Fig. 2—X. victwrlana asp, Mucrodens in profile.

Systematics and evolution—The new species is

genealogically very near to the group maritima,

brevisimilis and uniseta, being distinguished essen-

tially from these species by the presence of a, on

tergite abd. IV and by the absence of m, on sternite

abd. TV (character as—see phylogenetic tree—Fig.

34 in Gama 1969). Moreover, it is the only species |

know in which this seta is absent.

X, victoriana o.sp. is further separated from the

group maritima by the structure of mucrodens,

mucro being separated from the dens in the new

species, While in the species of this group mucro and

dens are fused, Thus, it seems that X. vietoriana

nsp. is anagenetically more primitive than the

species of yroup Miaritinia,

10. Xenylla grisea Axelson, 1900

Xenylla yrisea Axelson, 1900, p. 108,

Material exumined—Queensland: Emu Creek, 20

km E Warwick, rotten log, 3 specimens, leg. P.

Greenslade, 14.v.1974—in alcohol (MC), South

Australias Adelaide, 4 specimens, vy.1933 (identified

by Womersley as X. maritima),

Tl. Xenylla stachi wolffi Gama, 1967

Xenylla siachi wolffi Gama, 1967, p. 15.

Observations—This subspecies is found in the

Solomon Islands and in the New Hebrides. This is

the first time it has been found in Australia.

Material cxamined—Queensland: Cooloola, rain

forest litter, 2 specimens, leg. P. Greenslade,

23.1,,.1977—in alcohol (MC): Cooloola, War-

rawonga pitfalls. J juv., leg, P. Greenslade.

i.1977—1n aleahol (SAM); Cooloola, Warrawonga

litier, 7 specimen, teg, P. Greenslade,

29 ji1.1977—in aleohol (MC); Cooloola. Chalumbar

pilfalls, 2 specimens, leg. P. Greenslade, i, 1977—in

alcohol (MC); Cooloola, Chalambar litter, about

thirty specimens, leg. P. Greenslade, 23.i1,1977—in

(ANIC); Cooloola, Birwillah litter, 2

specimens, leg. P. Greenslade, 16.i1.1977—in

alcohol (SAM): Cooloola, Pertaringa litter, about

fifteen specimens, leg. P. Greenslade, 23.11.1977—in

alcohol (MG); Idem, about thirty specimens.

29.111,1977, 23 specimens. in alcohol (SAM): 7

specimens, on slide (MC). Northern Territory: Alice

Springs. Kunoth Paddock, hills litter, 2 specimens,

lez. P. Greenslade, 17.ii.1975—on slide (MC).

Torres Strait Islands: Murray Is., litter from vine,

bamboo forest, about twenty specimens, leg.

Cameron, 17.vii. 1974, 8 specimens, on slide and the

other specimens, in alcohol (MC).

alcohol

12. Nenylla cf. obscura Imms, 1912

Xenylla obscura Imms, 1912, p. 84.

Systematics and evolution—The populations cited

below from Australia and New Caledonia are

128

distinguished from the populations of X. obscura

from Sikkim, Nepal and India (for a redescription

see Gama 1969: 43-46) in the following features:—

1, The tip of the mucro is wider than on the

drawing represented in fig. 28.

2. The cephalic setae L, and L, are more or less

similar in length (absence of character f).

3. The cephalic seta ao is present in some

populations, but is absent in others, although within

a population some variability of this character, a,

may be observed.

4, Setae are in general blunt and not sharp as in

the other populations of X. obscura which I have

previously studied (Gama 1969 and 1971).

These populations that present the character, a,

could possibly be considered as X. hawaiiensis

Gama, 1969, because they are placed in the same

cladogenetic level as this species (fig. 34 in Gama

1969). However, in typical specimens of the species

from the Hawaii Islands the mucro is different in

form (fig. 30: 51 in Gama 1969) from the mucro

found in populations from Australian regions.

In 1971 (Gama 1971: 152-153) I verified, after

studying some populations of X. obscura from

Nepal, that ‘ta greater length of the body and a

greater development of anal spines seems to be

correlated with the absence of a; on abd. IV and the

greater length of L, in comparison with L;. A similar

correlation appears to occur between a shorter body

and a lesser development of anal spines and the

presence of a; on abd, IV and the near similarity of

length of L, and L,’’. (The above is English

rendering of the original French).

However, these populations from Australia and

New Caledonia do not have the seta a; on tergite

abd. IV and the anal spines are relatively poorly

developed. In body length, specimens from New

Caledonia are very long whilst those from Australia

are relatively short.

I have also recently examined further specimens

from Nepal which Prof. Cassagnau has kindly sent

me. In the four populations of X. obscura that I have

studied, the mucro presents the characteristic form

of the species, the setae are sharp, L, is longer than

L, (character f), the cephalic seta ay is present

(absence of character a), and the seta a, on tergite

abd. IV is absent. Anal spines are only well

developed in sample n° 65.

Material examined—Queensland: Cooloola, Kab-

ali East litter, pitfall traps, 2 specimens, leg. P.

Greenslade, i1.1977—on slide (MC); Cooloola,

Chalambar pitfalls, 1 specimen, leg. P. Greenslade,

ii.1977—in alcohol (MC). Northern Territory: Alice

REC. S, AUST. MUS. 18(5): 123-129

May, 1980

Springs, Kunoth Paddock, ungroved Mulga litter, 2

ad. + 4 juv., leg. P. Greenslade, 7.ii.1975—in

alcohol (ANIC); Idem, about fifty specimens,

15.ii.1975—in alcohol (SAM); Idem, 5 specimens,

15.1i.1975—in alcohol (MG); Alice Springs, Kunoth

Paddock, grove Mulga litter, 8 specimens, leg. P.

Greenslade, 15.ii.1975—on slide (MC); Alice

Springs, Kunoth Paddock, hills litter, numerous

specimens, leg. P. Greenslade, x.1974—on slides

and in alcohol (MC); Idem, 40 specimens, leg. P.

Greenslade, 17.ii. 1975, 34 specimens, in alcohol

(ANIC), 6 specimens, on slide (MC). South

Australia: Monarto South, Ferries-McDonald

Reserve, in Lepidosperma, 15 specimens, leg. P.

Greenslade, 2.x.1977—in alcohol (MG); near

Loxton, mallee leaf litter, 25 specimens, leg. P.

Greenslade, 25.ix.1974—in alcohol (SAM). New

Caledonia: fle Nou, on the surface of a puddle of

water, very numerous specimens, leg. G. Fabres,

28.x11.1977. sent by Prof. P. Cassagnau—on slide

and in alcohol (MC),

ACKNOWLEDGEMENTS

This work was supported by a research grant from

my country’s National Institute of Scientific

Research (Grant CB/2—Portuguese Ministry of

Education and Scientific Research).

The sample from Japan containing X. littoralis was

sent to me by Prof. R. Yosii, whom I thank very

much. My thanks are also extended to Prof. P.

Cassagnau, who forwarded the material from Nepal

and some of the material from New Caledonia.

I also thank sincerely Prof. H. Uchida, Prof. S.

Chiba and Dr. H. Tamura, who kindly provided

some types of X. arenosa Uchida and Tamura, 1967,

the study of which led me to conclude that this

species from the Japanese littoral is not X. littoralis.

Finally I am particularly grateful to Mrs. P.

Greenslade for giving me the opportunity to get to

know the Australian Xenylla species.

I should like to thank Dr. M. Tully for help in the

translation of this article.

REFERENCES

AXELSON, W. M., 1900. Vorlaiifige Mittheilung liber einige

neue Collembolen Formen aus Finnland. Medd. Soc. Faun.

et Flora fenn. 26: 105-123.

FOLSOM, J. W, 1916. North American Collembolous Insects of

the subfamilies Achorutinae, Neanurinae, and Podurinae.

Proc. U.S. nat. Mus. 50: 477-525,

GAMA, M. M. da, 1967. Collemboles du genre Xenylla trouves

dans les fles Salomon e dans |’Archipel de Bismarck (Noona

Dan Papers No. 39). Il. Mems Estud. Mus. zool, Univ.

Coimbra 300: 1-21.

GAMA, M. M. da, 1969. Notes taxonomiques et lignées

généalogiques de quarante deux espéces et sous-espéces du

genre Xenylla. II. Mems Estud. Mus. zool. Univ. Coimbra

308: 1-61.

EVOLUTIONARY SYSTEMATICS OF XENYLLA 129

GAMA, M. M. da, 1971. Nouvelle contribution 4 la connaissance

du genre Xenylla. 1V. Khumbu Himal 4: 152-155,

GAMA, M. M. da, 197la. Application de la méthode de la

“systématique idéale” A quelques espéces du genre Xenylla.

V. Revue Ecol. Biol. Sol 8(1): 189-193.

GAMA, M. M. da, 1974. Systématique évolutive des Xenylla.

VII. Espéces provenant d’Australie. Ciénc. Biol. 1(3): 71-83.

GAMA, M. M. da, 1979. ye évolutif d’une septantaine

d’espéces et sous-espéces de Xenylla provenant de tous les

continents. Revue Ecol. Biol. Sol (in press).

IMMS, A.D., 1912. On some Collembola from India, Burma and

Ceylon, with a catalogue of the Oriental species of the order.

Proc. Zool. Soc. Lond.: 80-125.

MASSOUD, Z., 1965, Les Collemboles Poduromorphes de

Nouvelle-Guinée. Annls Soc. ent. Fr. n.s.1(2): 373-391.

MILLS, N.B., 1938. Collembola from Yucatan Caves. Carnegie

Inst. Washington Publ. no. 491: 183-190.

TULLBERG, T., 1869. Om_ skadinaviska Poduriden af

Underfamiljen Lipurinae. Akad. Afhandl. Uppsala: 1-20.

UCHIDA, H., and Tamura, H., 1967. Eine neue litorale

Collembolen-Art von Hokkaido. J. Fac. Sci. Hokkaido

Univ. s. VI., Zoology, 16: 234-237.

WOMERSLEY, H., 1933. A preliminary account of the

Collembola-Arthropleona of Australia, I Poduroidea. Trans.

R. Soc. S. Aust. 57: 48-71.

WOMERSLEY, H., 1939. Primitive Insects of South Australia.

Silverfish, springtails and their allies. 322 pp. Govt.

Printer, Adelaide.

MACROPODID SKELETONS, INCLUDING: SIMOSTHENURUS

TEDFORD, FROM AN UNUSUAL “DROWNED CAVE” DEPOSIT

IN THE SOUTH EAST OF SOUTH AUSTRALIA

BY NEVILLE S. PLEDGE

Summary

A drowned cave in the South East of South Australia has yielded exceptionally well-preserved

vertebrate fossils, particularly of the Pleistocene macropodid Simosthenurus Tedford, to scuba

divers. Three distinct groups of Simosthenurus have been recognised, including S. gilli (Merrilees).

One (Species II) may be referable to S. orientalis (Tedford) while the third is a new species. Post-

cranial remains associated with some skulls offer, at last, definite knowledge about the anatomy of

these extinct kangaroos.

MACROPODID SKELETONS, INCLUDING: SIMOSTHENURUS TEDFORD, FROM AN UNUSUAL

“DROWNED CAVE” DEPOSIT

IN THE SOUTH EAST OF SOUTH AUSTRALIA

NEVILLE S. PLEDGE

South Australian Museum. Adelaide. South Australia S000

ABSTRACT

PLEDGE, N. S. 1980. Macropodid skeletons, including

Simosthenurus Tedford, from a unusual “drowned caye"' deposit

in the sacl East of South Australia, Ree. S. Aust. Mus 18(6):

A drowned cave in the South East of South

Australia has yielded exceptionally well-preserved

vertebrate fossils, particularly of the Pleistocene

macropodid Simosthenurus Tedford. to scuba divers.

Three distinct groups of Simosthenurus have been

recognised, including S. gilli (Merrilees). One

(Species II) may be referable to S. orientalis

(Tedford) while the third is a new species. Post-

cranial remains associated with some skulls offer, at

last, definite knowledge about the anatomy of these

extinct kangaroos.

The geometry of the cave is described briefly and

supports an hypothesis about the accumulation of

some of the bones, It is proposed that in drought

periods during the latter stages of the last glacial

phase, when caves were the only local source of free

water, animals entered this cave to drink and some,

weukened or over-extended, fell into the under-

ground pool and could not get out. Their carcasses

sank to the bottom to leave near-complete skeletons

that were protected by the post-Glacial rising

groundwater.

INTRODUCTION

In 1964, scuba diver G. McKenzie passed to Mr.

F. W. Aslin a small quantity of bones he had

collected in a drowned cave 24 km from Mt Gambier

in the South East of South Australia. The bones

were subsequently examined by Dr. B. Daily, who

confirmed Aslin’s identification of possum bones

and a palate of the extinct macropodid Sthenurus

(s.l.). The importance of the site was therefore

apparent, and Aslin undertook to coordinate

activities in the cave so that any future finds would

he brought to the attention of the South Australian

Museum.

This paper describes the cave site and contents in

general terms and presents a possible mode of

accumulation,

In 1965, some more bones were collected and

examined by Daily who recognised kangaroo

(Muacropus sp.) and pig (Sus) remains. This material

had apparently been found near the entrance on the

talus cone where rubbish had been dumped.

The next finds were made in January, 1968, by

divers B. Brawley and G. McKenzie. Six sets of

skulls and jaws as well as other elements of

Sthenurus were collected jn three dives, most bones

being in excellent condition, The divers reported

they had located “a veritable graveyard”, with the

bones “laid out in a semi-articulated state’. The

bones occurred in little groups, each apparently

representing a single animal, Aslin quickly sought

advice from Daily on the best way of retrieving the

fragile bones. In February, brief mention of the

fossil deposit was made in an article in the local

newspaper (“South-East has Tourist Attraction

Underground”; The Border Watch, Tuesday, 27

February, 1968).

A further diving expedition in November, 1968

yielded several more jaws and partial skulls, and it

was noticeable that cranial material was becoming

less obvious.

The next expedition was in January, 1969 with

Brian Brawley diving again. Only eight dives were

possible in the two days available, but a considerable

quantity of bone, mainly post-cranial, was retrieved,

including several short sequences of articulated

vertebrae in partial skeletons of Macropus and

Sthenuris.

In May, 1968, Daily encountered a specimen from

the locality in the Australian Museum, Sydney,

where it had been taken unceremoniously by some

divers, whose account of their trip later appeared in

192 RRO S

a somewhat sensationalised article in Pix (15

February, 1969) under the title “In Deep, Blue

Water—Kangaroos Who Lived 12 000 Years Ago!”

The sinkhole named (‘'The Pines”) is nat the same

as the subject of this paper, but many of the features

mentioned are suggestive of it, and the inset

photograph was, according to Brawley, taken in the

fossil cave. In a subsequent issue (Pix, 22 March,

1969) Dr Alex Ritchie of the Australian Museum

made a plea to divers not to disturb the fossil

remains,

A final collecting expedition took place in June,

1974, to salvage any remaining obvious fossils,

following evidence that unauthorised divers had

been active in the cave, and would be returning.

Again, Brawley did the underwater work but had

trouble in finding much of significance apart from

some jaws of Simosthenurus gilli. A quantity of

scallered post-cranial material was also recovered

before the water silted up to the extent that nothing

could be seen,

METHODS

The very situation of the fossils caused special

methods to be devised for their collection, and for

plotting their relative positions.

As the fossils occurred in a flooded cave, at depths

as much as 15 m, and in total darkness, special

urrangements had to be made. Initially, two divers

operated together on the ‘‘buddy system”. Later, a

diver worked alone, connected to the surface by a

safely tine which doubled as a communication

system by way of certain signals,

Bones were exceedingly. delicate and soft when

found und were carefully transferred underwater

into a neutral buoyancy basket lined with cotton-

wool for their removal to the surface, and a shiny tag

put in their place. The specimens were carefully

washed ta remove remnants of silt and clay, dried

slowly, then treated with a dilute solution of

“Glypolin” in acetone, applied several times, to

harden the bone. Later collections were treated with

“Bedacryl” in methyl-ethy] ketone.

Determining truc relative positions of the bone

assemblages proved difficult. The positions of the

first finds were not noted but were indicated by tags

placed by divers on a later expedition, working from

memory. A grid reference system of surveying was

initially considered for fixing the position of fossils,

but was decided against because of the uneven

topography of the rock pile. Instead, the safety line

was employed as a survey instrument to determine

specimen positions, Al each paint where the diver

stopped for several minutes to pick up specimens, a

lag was attached at the surface end of the line, level

with a fixed datum point, At the same time, a

AUST, Muy, I(t: 141-14)

May, 1980

compass reading was taken on the bearing of the

line. When the rope was hauled in at the end of a

dive, the distance indicated by the tag was

measured, This system gave only marginal accuracy,

as it could not quantify the variations in slope, nor

the detours the diver might take on his descent.

More accurate on a smaller scale were the sketches

and notes made underwater by the diver, but again

distances and directions were only estinnites. lt was

found that determinations made on the same points

varied considerably on different occasions, and

correlation was exceedingly difficult. Shiny metal

tags were left at each collecting point by the divers,

ou their second and subsequent expeditions-

However, in between trips, other unauthorised but

tidy-minded divers removed most of them, and it

was subsequently found that the collectors’

memories were not sufficient to relocate these

points. Consequently, the collecting sites marked in

Figure | are mostly only tentative.

Another major problem was that of rapidly

Increasing turbidity of the water, caused by the

divers’ movements near the bottom slirring up fine

silt. It was found that two dives a day for wo days

were all that was feasible, and that the water then

took several days to clear. In fact, a 30 mitute dive

was the longest possible at any time, and this

duration rapidly diminished over a two-day period as

remnant turbidity built up,

It is likely that many bones that are missing from

the skeletal assemblages have slipped down between

rocks on the slope. However, search for them was

not attempted because of the twin-problems_ of

nonvisibility and safety.

Tooth notation is that now advocated by Archer

(197%) based on embryological evidence for three

premolars and five molars, the first of which is

replaced by P3,

OBSERVATIONS

The Cave Deposit

The South East province of South Australia js a

vast limestone shelf of Oligo-Miocene age, capped

by superficial Quaternary calcareniles (Bridgewater

Formation) in the form of stranded beach ridges

(Boutukoff 1963). Extensive cave development

occurs, und centres mainly on two areas: Naracoorte

and Mt Gambier (See Marker, 1975) In the farmer,

the caves are mostly dry or with only minor pools of

standing water, They tend to be labyrinthine,

following joint-controlled weaknesses linking larger

collapse chambers, In the Mt Gambier area, many

caves have standing water and quite a number cqn be

said to be “Drowned” following the post-placial rise

in sea level and water table, Many are designated as

sink-holes, ice. large flooded chambers breaching

MACROPODID SKELETONS, INCLUDING SIMOSTHENURUS TEDFORD 133

the land surface in small apical openings (e.g, The

Shaft), while others are typical cenotes (e.g. Little

Blue Lake). Cave formation may have occurred as

long ago as the late Miocene (Moriarty, pers.

comm.) following regional regression of the sea,

with later rejuvenation during the Pleistocene,

I consider the subject of this paper to be a

drowned caye rather than a sink-hole. The fossil

cave is essentially a large elangate chamberenlarged

by roof collapse, in which the collapse talus cone has

penetrated the breached ceiling, and full of water to

within 4m of the land surface outside, The entrance

doline is roughly oval, about 12m x 20m, aligned on

a NW-SE axis, Emergent rocks occupy the floor but

aré surrounded on nearly all sides by visible water

with free access limited to the south-eastern

quadrant. Mr Aslin has kept records of the water

table, which has fluctuated by less than Im between

winter and summer but as much as 1-8m over several

years, as the result of rainfall variations. At the low

water level, part of the ceiling of the cave is exposed,

for a distance of about 12m, A plan and longitudinal

section of the cave, as cornpiled from divers’ notes

are shown in Figure |. There is an apparent step-like

profile of the floor which may have important

implications regarding the mode of accumulation of

the fossils.

While bones were scattered over the upper floor

of the cave, apparently derived by surface

movement from the apical talus cone, the

immediately obvious feature, seen by the first divers

in the far recesses of the cave, were the discrete piles

of bones, cach of which appeared to represent a

single individual.

The Faunal Assemblage

(1) Aves

Numerous bones, mainly wing bones, indicate the

presence of small birds, but have not yet been

studied.

(2) Marsupialia

Dasyurts maculatus (P17254)—A well preserved

skull represents this taxon, undistinguishable from

the madern species, Area (Fi,

Thylacinus cynocephalus (P17257)—A fragment

of mandible with canine, premolars and first molar is

referable to this extinct species. Between Areas @

and @,

Trichosurus vulpeculus is represented by a

mandible, This may be a more recent addition, as

fossil deposits in dry caves in the region seem to be

devoid of Trichosurus while itis abundant in modern

accumulatians, Area &)

Vombatus ursinus is represented by a well-

preserved skull and jaws. It is possibly a more recent

addition to the assemblage and its exact position is

unknown.

Macropus cf. giganteus—The prey kangaroa is

represented by several specimens including an

immature skull and jaws (P17256), a mature skull

and jaws (P17457) with partial skeleton from area D

another partial skeleton (P17523) with a jaw

(P17522) but no skull from area @ footbones

(P18291) from area @. Tibiae and footbones from

between areas @and @, a partial skeleton (P17518)

from area [5](no skull or jaws).

M, rufogriseus is represented by a skull (P17255)

and isolated bones and skull fragments,

Sthenurinue

In his monograph in this family, Tedford (1966)

erected a new subgenus (Simosthenurus) for those

species of Sthenurus having particularly brachy-

cephalic skulls. He also cited a marked difference of

elevation between palatal and basicranial planes,

However, since a skull of the dolichocephalic

Sthenurus (Sthenurus) cf, atlas from Naracoorte

(Henschkes Fossil Cave) also shows this feature,

while S. gilli does not, it is felt that the character may

be more of specific importance than generic. | have

no hesitation, in view of the excellent material fram

this cave, in raising Simosthenurus to full generic

rank, and including S. gilli in i,

Inspection ceveals three distinct species of

Simosthenurus. present in this deposit, of which only.

one seems to be described:

Graup I Simoasthenurus gilli: Specimens P17248

(skull and jaws), P17265 (left maxilla), P18326

(ankylosed jaws). These are closely similar to the

Haystall Cave (Naracoorte) specimens referred by

Merrilees (1965) ta this taxon, See Table 1; Figure

2A,

Group 2 Simasthenurus Species 1: Specimens

P17246 (skull and jaws), P17247 (skull and jaws),

P1747 (partial skull lacking cranium), P17252 (left

and right jaws), P17263 (right and half of Ieft

ankylosed jaws). (Table 2 and Figure 2B). The teeth

of this taxon are superficially similar to §. gilli;

molars have the same width, but are relatively a little

longer, and have a weaker midlink with fine

crenulatians; the upper premolar is relatively

narrower and almost rectangular compared to S. gilli

where it is rather triangular due to the broader

posterior moiety: lower premolars are much

narrower than in S, gilli, The obvious. difference

however, is in the skull, Even mature specimens of

S. gilli, as estimated by dental wear, have arelatively

small, low skull far Simosthenurus (though shorter

than one of Sthenurus of equivalent breadth). In

134

contrast, even relatively young specimens of this

new taxon have a large skull, roughly equivalent in

size and shape to Simosthenurus occidentalis, The

nasals are Jang, painted and down-curved, The

zygomae are heavy and deep with a prominent

downward process opposite the first permanent

molar M*. On S_ gilli the nasals are shorter and less

curved, and there is no distinet process on the

zygoma,

Group 3 Simosthenurus Species It specimens

P17244 (skull), P17245 (skull and jaws), P17253

(skuJ] and jaws), P17250 (left and right jaws),

P17251 (juvenile left jaw). (Table 3 and Figure 3).

These Simosthenurus are tentatively referred to S.

orientalis which previously has been known only

from a few lower jaws, They have been compared

with published data (Tedford 1966; Merrilees 1965,

1967, Bartholomai 1963; and Marcus. 1976) of

Simosthenurus occidentalis, 8. oreas, 8, pales, S.

antiquus, S. orientalis, S. brownei and the unnamed

large species from Strathdownie and Haystall Cave

(Mertrilees 1965), but only S. orientalis approximates

them in lower dentition, The skulls are large, deep,

broad and shortfaced: one specimen (P17245) is sa

well preserved that even the delicate postpalatine

bar is retained intact.

One of the exciting aspects of this fossil discovery

is the presence of more-or-less complete skeletons,

associated with identifiable skulls, Several partial

skeletons referable to Sthenurus s.1, have been

recovered, aud are being studied. As a preliminary

guide, however, certain bones may be distinguished

from Muecropus thus:

femur cobust, relatively greater shaft diameter,

shaft eurved convex forwards,

tibia robust, relatively shorter length and greater

shaft diameter. slight sinuous curve when viewed

anteriorly.

scapula: shart and broad blade, very strong and

long acromion process of spine, anterior moiety of

blade reduced to distal carner only.

pelois less triangular and mare ovate obturator

foramen than in Macropus; ihac blades broad.

flarings ventral (pubes) suture relatively shorter.

Epipubic bone ts 4 large broad blade, more than

Iwice as long and wide asin Macropys of same pelvic

SIZE.

Cervical vertebrae very short, with the centrum

about 19 mm long fora transverse diameter of up to

50 mm.

At least one of the skulls has a known associated

partial skeleton, This is P17471, Simosthenurus sp, 1,

from area @, and the remains consist of the pelvis

and sacral vertebrae, right epipubie, both femora,

left tibia and fibula, both arms, most cervical and

dorsal vertebrae (only Sth cervical and 12th thoracic

REC. S. AUST. MUS

IR(®)> Lally] May, 1980

vertebrae are missing), some proximal caudal

vertebrae, and a broken series of ribs. Other partial

skeletons are less complete, at the moment, and

there is some doubt abour their associated skulls

because they were not always collected during the

same dive, or even the same expedition.

DISCUSSION

The deposit is so unusual and contains such well

preserved material, that it readily lends itself to

speculation concerning the mode of accumulation,

(t is almost certain that the cave contains an

assemblage of mixed ages, even without the modern

rubbish dumped there, Material being added at the

top of the rock pile must inevitably move by sheet

wash, surface creep or other disturbances further

down the slope. In doing so, however, it alsa would

become dissociated and contribute to the extensive

collection of isolated bones, This process has

continued at Jeast as long as the situation has

approximated that of the present iime, with the

water level close to the surface, indeed, within easy

reach of thirsty animals.

However, a Sarge proportion of specimens ace af

extinct species, the bones often associated or even

semi-articulated as in several instances of vertebrae.

This implies that there have been no distutbances

since the eadavers came to rest, and also the

skeletons hayeé an age of at least 10 000 years, The

actual date of extinction of the Pleistocene

megafauna is not known, but is more likely to be

related to climatic/environmental changes associated

with the end of the last glacial period than with any

other simple cause, Martin (1447), Merrilees (1968)

and others notwithstanding; and the present

(Holocene) mild climatic regime is generally

considered to have started 10 000 years ago (Bowler

et al. 1976).

During the glacial stages of the Pleistocene. much

seawater was removed as huge continental ice

sheets, with the result that sea Jevels were

universally depressed, Consequently, so too was the

water-table in the Mount Gambier area. Aceord-

ingly, the water level in the cave would be lower,

und possibly some considerable distance inside. even

heyond the “twilight zone”. It is quite possible that

at the glacial maximum when sea levels were

possibly as much as L30m Jower than at present

(Flint 1971) the cave was totally dry, and an animal

could conceivably wander to the bottom and die ona

rock and be undisturbed until the divers came. In

reaching this position in the dark, however, it would

scatter and break many of the remains already there.

Yet the evidence is for numerous complete or partial

skeletons resting undisturbed on rock slabs.

MACROPODID SKELETONS, INCLUDING SIMOSTHENURUS TEDFORD

1 am suggesting here, that the cave had a

substantial lake when the “skeleton” individual

entered it. This is based on an idea by B- Brawley,

who noted that some distance from the entrance,

there is a steplike break in the slope, about 7m

below present water level (see fig. 1), If the water

has reached that point, there would be a steep,

possibly insuperable bank ta the lake. In this

scenario, any animal seeking water during a drought

period, such as in the Late Pleistocene (Bowler et al,

1976), would have to stretch in order to drink, and if

it stretched too far and fell in, it might not be able to

clamber out, but would struggle until it drowned or

died of exhaustion. The cadaver might float for a

while, disintegrating slowly, or sink immediately to

leave a beautifully laid-out skeleton on the bottom.

The situation is somewhat like the Morwell fire-hole

deposit of kangaroos excavated by Rich (1977),

except that there has been no areal concentration of

the skeletons by prevailing winds. The alternative

hypothesis that the 'skeletan’’ kangaroos entered a

dry cave is not acceptable, since it requires some

unknown form of protection far the skeletons until

the rising groundwater had covered them,

Using hydrological data gathered in recent years,

Mr. Aslin (pers. comm, 1979) has calculated that a

water level at the break-in-slope, about 7 m below

the present level, would correspond (at a first

approximation) to a sea level drop of about 20m,

Applying this amount to the sea level record curves

figured by Chappell (1978: fig. 1:10) gives an age of

8 000-9. 000 years, which is quite outside expecta-

tions based on fossil evidence elsewhere in

Australia. No dated Holocene site has yielded ia situ

Sthenurus 8,1, This suggests that the relationship

between water table and sealevel in this area is not

as simple as has been supposed, The age of the

specimens must be sought by more direct methods,

With this in mind, an attempt was made in 1974 to

abtain material for dating, Bottom sediments proved

to be useless calcite flakes. There was hope of

finding pollen in the fine silt filling one skull but it

proved to be too oxidized. The age, therefore, will

remain unknown until or unless associated charcoal

or wood is found. However, circumstantial palaeo-

climatological evidence, provided by work in

western Victoria (Bowler et al. 1976) and elsewhere,

suggests that the associated skeletal specimens might

date from the cold arid period of the last glacial

phase, approximately 17 000 to 14 000 years ago, if

the scenario proposed above is correct.

145

ACKNOWLEDGEMENTS

The author owes a debt of gratitude to Mr F. W.

Aslin of Mt Gambier for his untiring interest and

enthusiasm in organising the collection of the

specimens described herein, and to Dr B. Daily for

making it available for study, Also to be thanked are

the several divers, Messrs G. McKenzie, R. Pulford,

and particularly B. Brawley, who made most of the

collections. The manuscript was typed by D.

Scipioni and J, Casaretto and Figure 1 drawn by J,

Thurmer,

REFERENCES

ARCHER, M_ 1978. The nature of the molar—premojar

boundary jn marsupials and 2 reinterpretation of the

homalogy ot marsupial cheekteeth, Mem, Od Mus. 18 (2):

157-164.

BOUTAKOFF, N. 1963. The geology and geamorphology of the

Se age area Geol. Surv. Vict. Mem. 22: 1-117, Sappend., 6

tab.

BOWLER, J, M.. HOPE, G_S_.JENNINGS, J, N.; Singh, G, &

Walker D, 1976, Lale Qualernary climates of Australia and

New Guiner, Quaternary Researeh 6: 359-394.

CHAPPELL, J. 1978. Chronological methods and the ranges and

rates of Quaternary physical changes in Biology. anil

Quaternary Environments (ed. D, Walker & J, G. Guppy)

Aust, Acad, Sei., Canberra, 1-34,

FLINT, R, F. 197]. Glacial and Quatemary Geology, Wiley, New

York, XIE + 892pp..

MARKER, M. E, 1975. The Lower Southeast of South Australia:

a karst province. Occas. Pap. 13. Dept Geogr, and Envir,

Studies, Univ. Witwatersrand, Johannesburg, South Africa,

MARTIN. P. S. 1967. Prehistoric overkill. in ‘Pleistocene

Extinctions, The Search for a Cause,’ (ed, PS. Martio, & H,

E. Wright, Jr.). Yale Univ, Press, New Haven.

MERRILEES, D. 1965. Two species of the extinct genus

Sthenurus Owen (Marsupiala, Macropodidae) from south-

eastern Australia. including Sthenurus gillisp. nov. J. R. Soc

W. Aust. 48 (1): 22-32.

MERRILEES, D, 1967. South-western Australian occurrences of

Sthenurus (Marsupiala, Macropodidae) including Srhenurus

brownei sp. nov. J. R. Soc. W. Aust. 50 (3): 65-79.

MERRILEES, D- 1968. Man the Destroyer: Late Quaternary

changes in the Australian marsupial fauna. J. R, Soc. W

Ausi, 51 (1); 1-24.

RICH, T. H_ V. 1976, Recent fossil discoveries in Victoria. Vict

Nat, 93: 198-206,

TEDFORD, R. H. 1966, A review of the Macropodid genus

Sthenurus, Univ. Calif. Publs in Geol. Sciences 87: 1-72.

NOTE ADDED IN PROOF

The large but microdont species Sfmosthenurus sp,

1 of this paper has recently been recognized in cave

deposits at Naracoorte. where it has been named

Sthenurus maddocki. (Wells, R. & Murray, P., 1979:

A uew Sthenurine (Marsuptaha, Macropodidac)

from southeastern Sourl Australia. Trams. R. Soc,

S. Anst 103 (8); 213-219.)

May, 1980

TABLE 1. SIMOSTHENURUS GILLI, TOOTH DIMENSIONS (LENGTH x MAXIMUM WIDTH)

131-141

REC. S. AUST. MUS. 18(6):

136

g=8

gSs

22s

Ee?

gen

15:5 x 113

9-5 x 10-1

10:5 x 10-4

10-8 x 10-8

10-8 x 10-9

Fonte)

Bist

——

11:3 x 10-4

14-5x 9-9

14:4 x 10-0

w own =

S Ao ao

“ ad eK

wy aD =m

eltolen

cS tun io seo,

S$) 34

ann

11-5 x 10-0

11-4x 9-8

11-6 x 10-0

11-4x 10-1

mare

‘Tx 93

Ox 9-9

10-

“Data from Merrilees (1965); means taken to one decimal place

TABLE 2. SIMOSTHENURUS SP. 1. TOOTH DIMENSIONS (LENGTH x MAXIMUM WIDTH)

ios

95x 81

16-5 x 11-3

Sx 95

10-5 x 10-5

10-7 x 10-4

10-6 x 10-8

10-6 x 10-6

Lat

11:3 x 10-7

11:3 x 10-8

oo oo

“x

oe!

PAA

tm oo

wy ies

me O

aed

00 06

awn

ene

2 90

=3S

aes

oO GO

Pui

“S

ack

90 0

ne)

| nie

o} a]

an o an

| «| lw] |

So] ml] &

= “4 i)

ham = heal

me faq lad

nO coo coo

Wafer [es

| nial mre ae

ae ee -——

ao |or |e

an oo oo

Cad mK mr

Mae | E> | eft

aA ae ae

ae tae c-

wa low [or

aAn!| oo of

Cad Cad

AN | mer

sa lan|as

ore Sn ol

REIS

Luo] *

So va)

“wslS

i=)

Eol Tt

. avy n

SE] *

21S

i=)

can

* x

Salam

compared with S. occidentalis, S. brownei, S. oreas and S. orientalis

TABLE 3. SIMOSTHENURUS SP. Il, TOOTH DIMENSIONS, (LENGTH x MAXIMUM WIDTH)

MACROPODID SKELETONS, INCLUDING SIMOSTHENURUS TEDFORD

#3 > i & 3

S Qa _ is] mn

ok | fa * “ “

5S 2 ce |o |a

De ~ tt w wn

Fe +t inal a

oe + toa) oe ae As :

“ t t S q

ao io 00. poral = — 4

os > tad bad * bd “

~ silt |e |e |e le [sz

“ ire) + S al t oO

< I = — 4 —

Oo an al aT So a

CQ bch S = = = °9 so wn = a }

Lol n Lon rc cael Sol ~ an co font fo.) Col

* “ “ » x * * x be x x *

a 2 6 st 09 ™ DQ t+ a Q > “o

vay —) 3} = 4 _ oO + a S = a

4 — = = -_ 4 _ — a) WH

~ Ne) - a a é~ s nn a

a 3S _ = a 4 S I a 4 — S

aol = ao oa al tan is] ool 00 loa] - -

* * * * * = = ba ~ * ~ ”

o |e [a Jo [= |e eo |> [oe |e [a Jo

~ an o - nN rm On so a i=) a4 —

kon “=~ = = a Sol foal Sal _

owlan [ast [am ac lan |ouw

Soo pea rast [| ae ~~ exvee, |S | Oe | Oe

TS D8 | ey +m

SQ (SS 14S )48 7) R | we aq |S /S8ea/e2

| | | we | we | ee | ww | we mies | | | | wwe] wee | ew | em

TO /er lag [of | ae hes mo 19S [GS] eH

ang laa [4H [Hw | ts a os [ttle lau

Se I of ee an ae ae ee re Se aed Sn al -—<—

to mo lot [om |e ee SO | am | om

aa a tt [tt [an So > fh mae faa | aa

aM helen! ad an aS w Aan noe) ac ae

we] | | | ww | ee | ee | ee a De ee

‘GAO Qe pom | com PANS L eo." So} Te | on

mo Ag lst [Hwy [ast ww ma | on +

mre ere me ae a<- Cael ae ae ae

00 COD | Ort | COD | Ors ey Dw pcm paw9

aa at lah [tt [os roa) i) ir) alam

ae nar ae oe 5 nl onl Ww re ce ce ae

wee] | | | we | oe | me me | oe Ql || jlexd | | | we] ee | ee

990 SM 1 SSD | OS | MK a 200 So laga tag

ee am [ow [tt | tt Vana an [sa | oo

ae cer Ke ree Ciao coer ae ae re

an oo lanm [to |g TF TO [MoS | oer

ac an | + tt [a n tT aa mn

ae ae — 5 aioe] <4 ao Calon aA 5 ni eel ae

xe | | | | em | we | me | Ke QS] |] tweed | | | we | ewe | ew

70 wr | me |ow fon ra ad |ome |an

00 aa fest ltt [om var aa les [ow

ne nae mre re te oe | ee ae tei el ee

wen to, jor / tam ‘S bt] > ;

am + Vala) ++ ow | Y S a at +

Se oe | ei ool as [ian 4 wn On Sol Se) Smal Smal

wm | | | | eu | eK | oe | oe be Pye l ddd del|el dx «|

>t com | aa | ao loa mle a |o fm [re

00 00 a ta [+ “ aa i a + Val

te ool ae Se ee Cian Set onl Cel Sal te rn rt

ae 7 € 2: Li ae S . 3 ia: * 2 2 3 :

ant ne a4 rar res v ¢ 2 : : : a ;

35 tee ; ~ zt 3) > et r xt "

F : : o] : : : :

; : : ol] o: co 8

t : i aay

pn 1 ry &¢ f€ g€

Be & EE Se EE

14-6 x 12-7

11-3 x 10-3

10:8 x 10-7

137

REC. S. AUST. MUS. 18(6): 131-141 May, 1980

138

‘saul] JO ssauajatduios

Aq payesrpur uorsisaid ‘sayojays pue sajou ,siaarp wo payidwios yied sajeasopun ‘usw “A “4 Aq Aaains ssedurod pue

ade} uo paseq dew aavjing ‘says Suroaj[oo Jo suoltsod ajeutxoidde Surmoys ‘dAvd [ISSO] ay JO UOTJ9as JeUIpNIsuO] puke ue]|g—|] “OA

7 >"

wee 4 \

ma /, et

a ga Se

iF} 149% : !

nad \ V 7

‘ / vY’

\ ‘ v

\ of

See

— os \\ | }

, — el ier we S 4

bart Tel —_—_— —— —_— >

ise

6a => _-

mS.

{Zl =, ,* byw

a a (Li (Ey 7 —

“7

itl “Ll #91

Wee

TH ds ot cst

a va

“Ss

~ —

— ————

= 6

“XN

(sasjeuwW)

uydep yods

' | i | jl

a ate ee a

—- Nw !

a, | jeunousoo

SNS eee EE ae

$01}OWW

MACROPODID SKELETONS, INCLUDING

FIG, 2—A. Simosthenurus gilli Merrilees), P17248 Skull and

jaws. Area 1.

B. Simosthenurus sp. I, P17247 Skull and jaws. Area 4.

SIMOSTHENURUS TEDFORD

140 REC, S. AUST. MUS. 18(6); 131-141 May, 1980.

FIG, 3—Simosthenurus sp. Il, P17245 Skull and jaws. Area 2,

MACROPODID SKELETONS, INCLUDING SIMOSTHENURUS TEDFORD 141

FIG, 4—Simosthenurus sp.1, associated bones (approx. X1/3)

Area 1,

A right scapula (s), humerus (h), ulna (u) and radius (7)

P17476

B. right pelvis P17494

C, left femur (f), tibia (t) and fibula (fl)—anterior view

P17475

D. ditto, lateral view

RECORDS oF THE

SOUTH AUSTRALIAN

MUSEUM

VOLUME 18 NUMBERS 7—9 _ 30 January 1981

No. 7 THE GENERA LACCOBIUS AND NOTHYDRUS (COLEOPTERA, HYDRO-

PHILIDAE) IN AUSTRALIA AND NEW ZEALAND

by ELIO GENTILI

No. 8 THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND

COLEOPTERA IN THE BANKS COLLECTION AT THE BRITISH

MUSEUM (NATURAL HISTORY)

by WINIFRED P. K. RADFORD

No. 9 SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

1. NOTATION. 2. BIFEMORATA and PTYCTIMA (CRYPTOSTIGMATA)

by DAVID C, LEE

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

THE GENERA LACCOBIUS AND NOTHYDRUS (COLEOPTERA,

HYDROPHILIDAE) IN AUSTRALIA AND NEW ZEALAND

BY ELIO GENTILI

Summary

A key to the Australian and New Zealand genera of the water beetle family Hydrophilidae,

subfamily Hydrobiinae, tribe Hydrobiini, subtribe Hydrobiae is given and the Australian and New

Zealand species of Laccobius Erichson, 1837 and Nothydrus Balfour-Browne, 1939 are described

or redescribed and figured. Laccobius is represented in Australia by the species L. (Notoberosus)

zietzi (Blackburn, 1895). L. (Platylaccobius) clarus n.sp., L. (Platylaccobius) decipiens n.sp., L.

(Macrolaccobius) marmoratus Macleay, L. (Macrolaccobius) matthewsi n.sp., L. (Macrolaccobius)

bicaudatus n.sp., L. (Macrolaccobius) collium n.sp., and L. (Macrolaccobius) brittoni n.sp., and in

New Zealand by L. (Platylaccobius) arrowi d’Orchymont, 1925. Nothydrus is found only in

Australia where it is represented by N. australis (Blackburn, 1891) and N. montanus (Blackburn,

1891). The differences between Nothydrus and Laccobius are discussed and other genera to which

Nothydrus may be closely related are noted.

THE GENERA LACCOBIUS AND NOTHYDRUS (COLEOPTERA, HYDROPHILIDAE) IN AUSTRALIA

AND NEW ZEALAND

by ELIO GENTILI

Museo del Seminario, Venegono Inf, 21040, Italy,

Abstract

GENTILI, Elio. 1980, The genera Laccohius and Nothydrus

(Coleoptera: Hydrophilidae) in Australia and New Zealand,

Ree. S. Aust, Mus. 18 (7): 143-154,

A key to the Australian and New Zealand genera

of the water beetle family Hydrophilidae, subfamily

Hydrobiinae, tribe Hydrobiini, subtribe Hydrobiae

is given and the Australian and New Zealand species

of Laecobius Erichson, 1837 and Nothydrus Balfour-

Browne, 1939 are described or redescribed and

figured, Laccobius is represented in Australia by the

species L, (Notoberosus) zietzi (Blackburn, 1895). L.

(Platylaccabius) clarus n.sp., L. (Platylaccobius)

decipiens n.sp,, L, (Macrolaccobius) marmoratus

Macleay, L. (Macralaccobius) matthewsi n.sp,, L,

(Macrolaccobius) bicaudatus n.sp., L. (Macrolac-

cobius) callium n.sp., and L. (Macrolaccobius)

brittoni np., and in New Zealand by L,

(Platylaccobius) arrowi d'Orchymont, 1925. Nathyd-

rus is found only in Australia where it is represented

by N. australis (Blackburn, 1891) and N. montanus

(Blackburn, 1891), The differences between Nothyd-

rus and Laccobius are discussed and other genera to

which Nothydrus may be closely related are noted.

INTRODUCTION

Members of the genera Laccobius and Nothydrus

lire aquatic Coleoptera found in both stagnant and

running freshwater. The first genus is widespread,

hemny absent only from South America, the second is

Australian. They belong to the family Hyd-

rophilidae, subfamily Hydrobiinae, tribe Hydro-

hint, subtribe Hydrobiae (see d'Orchymont 1942).

The genera of Hydrobiae d'Orchymont. 1919, in

Australia and New Zealand may be distinguished by

the following key.

fev te Australian and New Zealand Hydrobiae

1, Hind teachanters with an clongate apex and separaled from

the femoru at their tip, Elytra withoul purasutural furrows.

Six abdominal segments, the sixth somewhat retractile into

the ile _Laceobius Erichson, 1837

Hind frochanters not elongate. Elytra with parasutiural

furrows, Five dhdominil segments —. , . . 2

2 (1) Regolur longitudina| tines of punctures on elyrra,

Antenne 9 segmented, Meso- und meta-tursi with the first

article shorter than second, Larger body sive: length more

thha+ pn 3

Flyteal pinerires noi, or only on hind and lateral surface.

arninged in longitudinal tines but randomiy disposed

Antennue 7-9 segmented. Size normally less than 4 mm

3. (2) Mesa- snd metie-tars’ without long swimming setae.

Meso- and meta-lernora without hydrofugal hairs. Body

posteriorly enlarged , Hybogralius WOrchymont,, 1942

Upperside of meso- and meta-tarsi with long swimming

setae. Meso-and meta-femora with hydrofugal hairs... 4

4. (3) Pro-, meso-, and anteriorly meta-sternum keel-shaped in

the middle Limroxenus Motsehulsky, 1853

Only the mesosternum sometimes keel-shuped jn the

middle Hydrobius Leach. 1818

5S. (4) Prosternunt keel-shaped in the middle. Hind femora

without dense hydrofugal hairs Paracymus Thomson, 1867

Prosternum Without any longitudinal keel. Hind temora, ut

least anteriorly, with dense hydrofugalhairs,........ 6

6. (5) Labium deeply excavated. Fifth antennal article

cupuliform Elytral rows postertorly conspicuous

Nolhydrus Balfour-Browne, 1939

Labjum anteriorly nor excavated. Elytra without longitudinal

tows of punctures, Fifth antennal article normally

shuped Memmi: Anacaena Thomson, 1859

SYSTEMATICS

The venus Laccobius is recognizable chiefly by the

following features. A convex, oval bady, not capable

of rolling up. Head with a Y-shaped suture and

entire eyes; sometimes the male with two specula

under the labium; antenna 8-segmented (53), the

filth article asymmetrically cup-shaped; maxillary

palpi shorter than antennae, with penultimate article

shorter than ultimate, Pronotum without furrows or

keels; prosternum keel-shuped in the middle;

anterior tarsi expanded in the male. Scutellum a

subequilateral triangle; mesosternum usually with a

longitudinal keel and a small tooth in front of the

keel; middle femora without hydrofugal hairs

(sometimes these are represented by a small basal

tuft, but not in Australian specimens): elytra, as a

rule (and always in Australian species), without

parasutural furrows, Wings provided with can-

tharidiform neryation. Metasternum without any

keel, in the middle with a small glabrous area: hind

trochanters with an clongate apex and separated

from the femora at their tip; hind femora without

hydrofugal hairs; hind tibiae usually curved. Six

abdominal segments, without any keel; the sixth

somewhat retractile into the fifth, Pro, mid- and

hind tarsi with the first article shorter than the

second, and the Jast shorter than the others

combined.

The subgenera in our region are the following:

|. Notoberosus Blackburn, 1895 (syn. fh. |

Ortholaceobius Ganglbauer, 1904), It has

straight hind tibiae. This feature is alsa

i44

. Hind tibiae straight: subg. Notaberosus .. .

REC, 8. AUST. MUS. IS (7):

typical of the subgenus Ortholaccobius

Ganglbauer, 1904; the two subgenera, in my

opinion, are to be united. The differences are

chiefly two, both in the male of zietzi

(Notoberosus); the specula and the lack of a

mesoternal keel. These differences disappear

in the female, and are therefore inadequate

to form the basis for the separation of two

subgenera. Notoberosus are very numerous in

New Guinea and in surrounding islands. The

other three subgenera have curved hind

tibiae.

Microlaccobius Gentili, 1974 is characterized

by the pattern of the elytral series,

alternately more regular, more impressed,

more punctate, and less regular, less impres-

sed, less punctate, The subgenus appears to

be the most primitive, owing to lack of

specula, and to the small, weakly chitinized

and simply structured aedeagus.

Macrolaccobius Gentili, 1974 is, like the

preceding, a subgenus widespread in the

Palaearctic, and Oriental regions. It has

more complex features; the elytral series

more irregular, an evolved aedeagus, and

sometimes specula and tufts of hairs on the

middle femora.

Tit.

IV. Platylaccobius Gentili, 1974 is characterized

by the lack or reduction of the elytral

seriation, The body is normally flat, more so

than in Micro-and Macrolaccobius, and the

aedeagus more complex than in Microlac-

cobius,

Key lo species

zietzi Blackburn

Hind tibiae curved | .,,.-..,.)-- ene ff, Lae. 2

. (1) Elytral punctures randomly placed and not forming any

ong of taws or lines, rately in ee | and ai ak

ines, rows or whorls subg, Platylaccobius . 3

Elyral punctures in about 20 longitudinal curved lines, © 5

. (2) Pronotam without a dark spot, or the dark Spot is very

feeble and indistinct; aedeagus asin Fig 2a, clarusa. sp.

A distinet dark spot on the pronotum._.. iCeee vent 4

. (3) Body length twice (or more than twice) the breadth.

Australian species, Acdeagus asin Fig. 2c, d

decipiens n.sp

Body length less tham twice the pik or New Zealand

species. Aedcagus as in Fig. 3.4a,b |. arrowid'Orchymon(

(2) Elytra! punctures alternately (in primary series) more

regular, more impressed and more punctate. and in

secondary series, not régular, weakly imprestent and

scarcely punctate. Size smal], subg. Microlaccabius . 6

Elytral punctures are in series not so alternated, same of

them are irregular, with mixed small nig large punctures,

subg. Macrolaceobius , a

(43-154 January [YS]

6. (5) Labium extended posteriorly in the shape of a V reaching

the postlabium, hind margin of propygidium deeply

excised: aedeagus asin Fig. 4a,6., . marmoratus MacLeay

Hind margin of the labium slightly sinuous medially;

propygidium not excised; aedeagus as in Fig. 4 ed

matthewsi 1.sp.

7, (5) A large preocular patch, Hind apex of each elytron

separated from the other, leaving a large triangular space

between them .... . bicaudatus n.sp.

No prevcular patches, Hind apex of each alesis close lo the

other in resting position. Smaller body size - ee,

8. (7) Aedeagus wider, more complex, 46 1h Fig Sa, b

collin n-sp-

Aedcapus narrower and simple, asin Fig. 3 ¢.d briranin-sp

1. Laeccobius (Notoberosus) zietzt (Blackburn 1895)

(Figs | a, bz 7)

Blackburn, 1895S (Noroberosus); Zaitzev, [908

(Notoberosus); Knisch, 1924 (Notoberosus); d'Orchy-

mont, 1925; 1937; 1948.

Body length 3-0-3-9 mm; breadth 1-35-1-73 mm-

Elongated, more than twice as long as wide.

Labium dark or metallic, sometimes with pale

sides, or entirely pale except in middle; surfuce

ulutaceous; anterior edge sinuous in male, straight in

female, the posterior curved. Head dark or metallic,

with large preocular patches reaching eyes; the

patches sometimes united; surface very slightly

alutaceous, with fine and scatiered punctures, Eyes

taken together scarcely smaller than intervening

distance. Pronotum yellowish, sometimes reddish,

ot with some small evanescent dark spots in central

area; smooth in male, hardly shagreened in female.

Corners of pranotum rounded, the hind corners very

widely so (more than in minutus). Scutcium pale or

dark, with very thin punctures, Elytra yellowish,

with or without a number of dark spots, but always

pale on edges; the punctures are slightly darkened,

numerous. irregularly scattered (but tending to form

5-6 series, chiefly near the suture), of various size,

Elytral apex nearly vertical.

Cnderside dark, except head, pronetum~ palpi,

antennae and legs. Labium of male has two specula

nearly circular in shape. The postlabium (or

mentum) is rugose. its anterior margin sinuous.

Prosternum short, keel-shaped; mesosternum

weakly but distinctly keel-shaped in female, gibbous

in male, the keel not anteriorly abrupt. Hind margin

of fifth abdominal segment deeply sinuous in male,

Antennae 8-segmented (Blackburn (1895) enumera-

ted 6 segments); legs slender, the hind tibiae

straight. Aedeagus (Fig. 1 a, b) 1/3-5 of body length;

tegmen longer than parameres, which are rounded

at apices; median lobe sulcate and pointed at apex.

South Australian species (Fig. 7), with the

following distribution: Lake Callabonna (type

locality, 10 ex. leg, A. Zietz); River Wakefield,

THE GENERA LOCCOBIUS AND NOTHYDRUS IN AUST. AND NEW ZEALAND

1.0

O05

145

Fig. 1—Aedeagus of Laccobius zietzi, lectotype, Lake Cal-

labonna, South Australia: A, frontal view, B, lateral view.

Prepared in euparal, soluble in absolute alcohol, like the

following aedeagi. The same scale applies to all of the text.

Figures 1-6,

near Adelaide (2 ex.); Levi Creek, 6 km NW Big

Perry Spring 28° 19° 2S, 136° 16" 1E (17 ex., leg. J.

A. Herridge, 7.X11,1974); Coopers Creek, Ferry

Crossing (1 ex., leg. J. A. Herridge, 30.XI.1974).

The specimens are in the South Australian Museum,

Adelaide, and in the Museums of Verona and

Milano, Italy.

Lectotype: South Australian Museum, Adelaide:

3-4x 1-55 mm; labels: L. Mulligan/L. Callabonna A.

Zietz/9.6580 Notoberosus zietzi Blkb. L. Callabonna

type /l1. d Paratypus 2. Lectotypus; Laccobius

(Notoberosus) zietzi (Blackburn 1895); det. E.

Gentili 1976. Another 7 specimens in the S.A.

Museum labelled in the same manner as the

lectotype are to be considered paratypes.

2. Laccobius (Platylaccobius) clarus n.sp.

(Figs. 2 a, b: 8)

Body length 2-4-3-1 mm; breadth 1-2-1-7 mm.

Elongated, nearly twice as long as wide, very slightly

convex.

Fig. 2—A, B, Aedeagus of Laccobius clarus, holotype, Adelaide,

South Australia: A, frontal view, B, lateral view; C,D, Aedeagus

of Laccobius decipiens, holotype, Tamworth. New South Wales:

C, frontal yiew, D, lateral view.

146

Labium large, dark or metallic with pale angles,

alutaceous, with anterior edge straight, the posterior

curved. Head dark or metallic, with large preocular

patches reaching eyes, the patches taken together

are of the same width as. intervening space, the

whole surface shagreened and closely punctured.

Prothorax testaceous, with a small evanescent dark

spot in its central-anterior area; the surface

shagreened and punctured like the head; fore

margin sinuous, slightly prominent if centre (in

arrowi it is straight). Seurellum pale or scarcely

darkened, with a number of very thin punctures.

Elytra pale, testaceons (sometimes darkened owing

to ‘actio pest mortem’); the punctures are numerous,

larger than those of pronotum, but similar to each

other in size, irregularly scattered, darkened on

elytral disc, clear at the edges; the humeri are

conspicuous. Posterior apex ogival.

Underside dark, excepting head, pronotum,

abdominal sides. palpi, antennae and legs. Post-

labium flat, nor rugose, with very fine punctures; the

fore margin curved. Prasternum short (its length ts‘

that of the pula), keel-shaped. Mesosternum keel-

shaped on its anterior 4, with an anteriorly abrupt

tooth on its fore apex. Hind margin of propygidium

nearly straight in female, lightly curved in male.

Palpi. antennae and legs of the normal pattern.

Aedeagus (Fig. 2 a, b) |i of body length, with a

large median lobe, shorter than parameres, the

median lobe sometimes shows a small mobile strut

near the apex.

The species is known from South Australia, New

South Wales and Queensland (Fig. 8), and is among

the commonest Australian Laecabius. South Aus-

tralia: Adelaide (type locality, 11 ex., leg. T-

Blackburn et al.); Hindmarsh Falls (2 ex., leg. E. B.

Britton-N. Tindale, at light, 23-24.X11.1961); Well,

near Oraparinna (1 ex,, leg. G. F. Gross, at light.

12.11.1956); Karoanda to Peebinga (2 ex., leg. G. E.

H, Wright); Murray River, E Adelaide (1 ex,, leg,

H, S, Cope); Frome River crossing of Birdsville

Track, near Maree (4 ex., leg. G. FP. Gross, at light,

25.X. 1966); 10 km N Maynards Well. N Flinders

Range (1 ex., leg. J. A, Herridge, at light,

11.X1L.1974); Brachina Gorge, Flinders Range (6

ex., leg, J. A. Herridge, at light, 14,X11,1974);

Muddle Bore, 35 km NE Billa Kalina (3 ex., leg. J.

A. Herridge. at light, 4.X1I.1974): Margaret River,

10 km SE Coward Springs (2 ex., leg. J. A

Herridge, at light, 3.XI1.1974); Marree Racecourse

(13 ex., leg. J. A. Herridge, at light, 1.061974),

Levi Creek, 8 km NW Big Parry Spring 28° 19° 2S,

136° 16° JE (J ex, leg. J. A. Herridge, at light,

7.X1I. 1974); Coopers Creek. Ferry Crossing (38

ex., ley. J. A. Herridge, at light, 30.X1.1974); 5-5

km WNW Myrtle Springs (1 ex., leg. E. G.

Matthews, 2-3.111.1975); 37 km W of Anna Creek (1

REC, & AUST, MUS TS (7): 193-154

January, 198]

ex., leg. E. G. Matthews, 8-9.[t1.1975), Coward

Springs (1 ex., leg, G. F, Gross, at light, 9, X1, 1966).

New South Wales: Murray River, 50 mi. W of

Wentworth (14 ex.. leg. A. Neboiss 22.X1.1967)

Queensland: Cunnamulla (1 ex,. leg. AH. Hardeas-

tle); Dear River, 23° 13'S, 144° 04 E, 31 km NW by

N of Longreach (3 ex., leg. M. S. Upton,

22.X.1975); Biedsville (4 ex., leg, J. Blyth,

11-V.1975). Australia (2 ex. Call. Fairmuire; 2 ex.

Pascoe Call, ).

Holotype: British Museum; 2-7 * 1-35 mim:

labels: Australia; Adelaide /Sharp Call. 1905-313 /

Laccobius (Plarylaccohius) clarus n. sp. E. Gentili

det, 1976 / ¢ Halotypus.

Allotype: British Museum; 2-9 » 1-45 mmj similar

labels. the Paratypes ace those of the former list:

they are in the following Natural History Museums:

British Museum; Musée de Paris; Museo di Verona;

Museo di Milano; South Australian Museum,

CSIRQ: National Museum of Victoria.

3. Laccobius (Platylaccobius) decipiens n, sp,

(Fig. 2 c, d; 7)

d'Orchymoent, 1925 (L. marmoratus).

Body length 2-9-3-9 mm; breadth 1-5-2-0 mm.

Flongated, oval, nearly twice as long as wide.

Labium entirely dark or metallic, alutaceous, with

a straight anterior margin. Head dark or metallic,

with preocular patches reaching eyes, but somewhat

smaller than in clarus; surface shagreened and

closely punctured, the punctures more numerous

and more impressed than in clarus. Pronotum pale,

festaccous, with a dark spot longer than wide,

reaching the fore margin more widely than the hind,

and slightly expanded in the hind half; surface

similar to that of head, shagreened and punctured;

fore margin sinuous, slightly prominent in the

centre. Seutellum dark. Elytra pale, sometimes. with

nebulous spots or darker lines (post mortem

changes?); the punctures are seattered irregularly

and very densely, sometimes suggesting longitudinal

series. Apices of elytra agiva), but often the elytra

are separated.

Underside dark, except antennae, palpi and most

of legs, and sometimes borders of prothorax and

abdomen, Labium without specula; postlabium flat,

granulate, Prosternum keel-shaped; mesosternum

keel-shaped on the hind 2/3, with an anterior tooth

on the keel. Hind margin of propygidium straight in

both sexes. Palpi. antennae and legs of the normal

pattern, Aedeagus (Fig. 2 c,d) ca. 1/47 of the body

length, the median lobe somewhat narrower than in

clarus, but slighty longer than the parameres.

THE GENERA LOCCORIUS ASD NOTHYDRLS IN AUST. AND NEW ZEALAND

The species is known [rom Victoria, New South

Wales and Queensland (Fig. 7). and seems to be

rarer than clarus. Victoria; Thurra River, Cape

Everard (2 ex., leg. A. Neboiss, 22.11,1970);

Deddick River, |) km above Snowy R. junction (10

ex., leg, A. Neboiss, 13.XIL. 1976); Thomson River,

L km upstream Cowwarr Wier (2 ex., leg. A.

Neboiss, 26,X.1973); Snowy-Deddick River junc-

tion (1 ex., leg. A, Neboiss, 14.11.1976); Snowy

River, 5 km below Deddick River junction (1 ex.,

leg. A. Neboiss, 14,X11,1976); Rainbow Ck.,

Cowwarr—Seaton Rd,, 18.11.1977, N.M.V, Survey

Dept., GRES (1 ex.), New South Wales: Tamworth

(type locality, 1 ex., leg. A.M. Lea); Jenolan Caves

(14 ex., leg. J. C, Wiburd 1864); Volumla (1 ex., leg.

J. Sedlacek, 1.1954); Tumut River (2 ex., leg. J.

Sedlacek. 1955). Queensland: Cairns (1 ex., leg. J.

Sedilacek 1951); N Queensland (3 ex., leg. T.

Blackburn):

Holotype: Institut R. Sciences Naturelles de

Belgique, Brussels; 3-5 x 1-75 mm; labels: Coll. R.1.

Sc. N.B, Australia 1864 Tamworth N.S.W. Ex Coll.

d’Orchymont/Coll, et det. dA. Orchymont;

Notoberosus marmoratus M. 2 R.M.H.N.B. 15.962/

Laccobius (Platylaccobius) decipiens n.sp. E. Gentili

det, 1976 / ¢ holotypus.

The Paratypes are those of the former list; they

are jn the following Natural History Museums:

South Australian Museum; National Museum of

Fig. 3—Aedeagus of Laceobius urrowi, Rotorua Co,, Waimangu

Stream, N, Island, New Zealand: A, frontal view, B, lateral view.

147

Victoria; California Academy of Sciences; Bishop

Museum, Honolulu; Museo di Verona; Museo di

Milano.

4. Laccobius (Platylaccobius) arrowi

dOrechymont, 1925

(Figs, 3 a, b; 7)

d@Orchymont, 1925; 1937, Winterbourn,

(mnineralis); 1973 (mineralis).

1970

Body length 2:3-3-5 mm; breadth 2-2-1-7 mm-

Oval, slightly convex, not much elongated (breadth

less than half of length).

Labium dark or metallic, alutaceous, punctured,

with fore margin straight, the hind curved. Head

also dark or metallic, alutaccous, with fine scattered

punctures; preocular patches pale and wide, as in

decipiens. Pronotum dark on disc, pale on sides, the

border of the colours uncertain; the dark spot is

normally wider than in decipiens. Anterior margin of

pronotum, between the lateral corners, almost

straight (in contrast to clarus and decipiens); the

surface alutaceous and punctured as on head.

Scutellum dark. Elytra pale, sometimes darkened by

post-mortem colour; the punctures are scattered,

rarer than in decipiens, darkened on disc, pale on

edges of elytra.

Underside dark, except palpi, antennae and most

of legs (femora are for the most part darkened),

Labium without specula; postlabium flat, with

scarcely visible punctures. Prosternum keel-shaped;

mesosternal keel with a tooth on fore end. Palpi.

antennae and legs of normal pattern. Aedeagus (Fig.

3 a, b) of body length, the median lobe shorter

than parameres and very complex,

The species is known from New Zealand (Fig. 7):

Otago, S Island (type locality, 10 ex., leg. Lewis

1901); Copland River, Welcome Flat, South

Westland, § Island (Winterbourn, 1973); Rotorua

Co,, Waimangu Stream, N Island (4 ex., leg. K, A.

J. Wise, 10.V.1971); Taupo, Waipuwerawera

Stream, N Island (type locality of mineralis; 2 ex.,

leg. M. Winterbourn VI.1966; efr. Winterbourn,

1970): Taupo, Waipahihi Stream, N Island (2 ex..

efr.. Winterbourn, 1970); Waiotapu (Winterbourn,

1973); N. Swale (1 ex., of the British Museum).

These specimens are in the British Museum,

Brussels Museum, Auckland Museum and Verona

Museum.

Holotype: British Museum; 2:75 x 1-5 mm; labels:

Holotype/type/Otago Lewis 1901/A. d’Orchymont

det. Laccobius s.str. arrowi d'Orchym./Sharp coll.

1905-313 (the anterior tarsi are all lost, but the

description is of a male),

(48

Paratypes: I know of two paratypes from the type

locality in the British Museum and in Brussels

Museum.

Laccobius mineralis Winterbourn, 1970 is in my

opinion the same species as arrawi. This opinion is

based on the observation of both the typical

specimens of arrowi and of ane specimen fram the

type locality of mineralis, determined by Wirter-

bourn and accompanied by 4 *‘mineralis” taken trom

the same area by K. A.J. Wise, as well as on the

description of mineralis, These aré the differential

characters proposed by Winterbourn (1970): (1) The

size: 2-3-2-4 mm for a@rrawi; 3-2-3-5 mm_ for

mineralis. But I obtained, on 4 beetles from 4

different stations, these measurements: 2°65 x 1-35;

2-75 x 1-45; 2:95 x 1:55 3:15 x 1:55 mm. They

evidently are a continuous series. (2) The anterior

angles of the pranotum are rounded in arrowi , and

the pronatum is deeply curved anteriorly, while in

mineralis the anterior angles are acute giving the

anterior margin a sinuous appearance. This is really

the same thing: the anterior angles are really

rounded, but observed from above they appear

acute; the sinuate appearance is due to the articulate

corners that make the fore margin deeply curved, (3)

The colour of the pronotum of arrawiis pale on the

borders, of mineralis completely brown. Winter-

bourn observed possibly a melanie specimen. (4)

The antennae are 8-segmented, not 6-segmented as

observed by Winterbourn. The Holotype of

mineralis is in the Auckland Institute and Museum.

New Zealand.

5. Laccobius (Microlaccobius) marmoratus (Mac-

leay. 1873)

(Figs. 4 a, b: 7)

Macleay, 1873 (Philhydrus); Zaitzey, 1908 (Enoc-

hrus): Knisch, 1924 (Enochrus); ctr. d’Orchymont.

1925

Body length 2-1-2-4 mm; breadth 1-1-1-3 mm-

Elongated, oval, nearly twice as long as its width.

Anterior margin of labium arcuate. Labium and

head dark, with a greenish metallic reflection:

preocular pale patches before eyes are half the

breadth of each eye. Surface (observed at 60 x)

shagreened, without punctures, Ratio between

breadth of eyes and of interposed space 1s ‘h,

Prenotum testaceous, with a dark metallic spot

which reaches posterior but not anterior margin; this

spot is narrower than space between eyes, and its

fore border is divided inta two parts, Surface

shagreened more conspicuously than head. and with

some impressed punctures on discal area. Scutellum

alutaceous like the head, with very fine punctures.

Elytra testaceaus, with some dark nebulosity on disc

and paler edges: the series of punctures alternately

more and less impressed.

REC; §, AUST, MUS I (7). 143-154

January, }981

Labium without specula, but with a border

attached to the postlahiuym tracing a triangle.

Postlabium flat, slightly wrinkled when observed at

60 x, Prosternum keel-shaped; mesosternal keel is

pyramidal and present only on hind part of the

mesosternum; glabrous metasternal line somewhat

excavated. Hind margin of propygidiunt deeply

excavated. Palpi, antennae and legs tesiaceous; the

legs are normally shaped, elongared. Aedcagus |); of

body length; ratio tegmen/parameéres Ys; structure

simple (fig. 4 a, b). the parameres scarcely longer

than median lobe.

Lectotype: CSIRO, Canberra; 2:2 x 1-1 mm

(elytra 145 mm); labels: Gayndsh/On permanent

loan from Macleay Museum, University of Sydney/

Laccobius marmoratus (Macleay) d Lectotypus EB.

Gentili 1975/syntype/Philhydrus marmoratus, Macl.,

Gayndah. Paratypes: The Australian Museum,

Sydney; labels; red circular label/719509/Philhydrus

marmoratus Me, L, W. Gayndah the Holotype/

1, Allolectotypus 2 2. Paratypus @ Laccobius

marmoratis Macleay, E, Gentili 1976.

The species ts Known only from the type locality.

Gayndab is a mountain town about 320 km north of

Brisbane, Queensland (Fig. 7).

6. Laccobius (Microlaccobius) matthewsi

(Figs. 4 c, d; 8)

Body length 1-6-2-5 mm) breadth 0-9-1-35 mm,

oval, moderately elongated, the length less than

Twice: the width.

Head and labium dark metallic, with two

yellowish preoecular spots before the Y-shaped

suture Or, sometimes, before the eyes; these spots of

same width as eyes or sometimes less. Surface

clearly alutaceous and weakly punctured. Anterior

border of labiurn regularly curved; distance between

eyes more than width of eyes. Pronotum testaceous,

with a dark or metallic central patch; the patch has

uncertain borders and is normally larger anteriorly,

reaching level of eyes, Surface shagreened like that

of head, but with stronger punctures. Scutellum dark

or metallic, micro-reticulated. Elytra testaceaus,

sometimes with a dark spot near centre of suture.

The serics of punctures aré clearly alternating

strangly impressed (primary), or Weakly (secon-

dary). Elytral apex rounded.

Underside dark; palpi, antennae and legs.

testaceous-yellowish, Labium without specula, the

hind margin straight (in contrast with mnarmoratus).

Prosternum keel-shaped; mesosternum with a keel

on posterior part, the keel having an anterior tooth;

propygidium posteriorly straight (another contrast

ta marmoratus). Acdeagus '/-'/ of body length;

parameres narrower than maprmeratus in frontal

view, and nor narrowing out in apical part in lateral

view (hig, 4 c, d).

nesp.

THE GENERA LOCCOBIUS AND NOTHYDRUS IN AUST. AND NEW ZEALAND 149

Fig. 4—A, B, Aedeagus of Laccobius marmoratus, lectotype,

Gayndah, Queensland: A, frontal view, B, lateral view; C, D,

Aedeagus of Laccobius matthewsi, holotype, Parachilna, Flinders

Range, South Australia: C, frontal view, D, lateral view,

This is one of the more common Laccobius of Tumut River (1 ex., leg. J. Sedlacek, 1955);

Australia (Fig. 8). Western Australia: Mullewa (1

ex., leg. Harvard Exp. Darlington, 2.1X.1931);

Carson Escarpment 14° 49’ S, 126° 49" E (2 ex., leg.

I.F.B. Common-M.S. Upton, 9-15. VIII.1975);

Langi Crossing, 10 m (5 ex., leg. E.S. Ross-D.Q.

Cavagnaro, 13.X.1962); Milly Milly, 300 m (1 ex,

leg. E. S. Ross-D, Q. Cavagnaro, 6.X.1962);

Northern Territory: Bessie Spring, 16° 40’ S, 135°

51' E, 8 km ESE of Cape Crawford (7 ex., leg. M.S.

Upton 26.X.1975); Nabarlek Dam, 12° 20' S, 133°

19‘ E,15 kmS by W of Nimbuwah Rock (1 ex,, leg.

E.G. Matthews, at light, 2.X1.1973); South

Australia: Parachilna, Flinders Range (1 ex., type

locality); Queensland: Coen, C. York (4 ex., leg.

Harvard Exp. Darlington, V. 1932); Crystal Creek,

23 mi. SSE of Ingham, 18° 58' 5” S, 146° 16’ E (7 ex.,

leg. Britton-Misko, at light, 9.XII,1968); Cairns

Distr. (2 ex., leg. A.M. Lea); New South Wales:

Moruya River, Araluen (9 ex., Jeg. E.B. Britton,

27.X11.1965); Manar Creek, Canberra Coast Road

(1 ex., leg. E.B. Britton-S. Misko, 18.V.1967);

Vicinity of Jenolan Caves (4 ex., leg. J. C. Wiburd);

Victoria; Thurra River, Cape Everard (1 ex., leg. A.

Neboiss, 22.111.1970); Deddick River, '2 km above

Snowy River junction (4 ex., leg. A. Neboiss,

13.XI1. 1976); Meredith (1 ex., leg. A. Neboiss, 12-

13.11,1959); Eppalock Res. Redesdale (1 ex., leg. A.

Neboiss, 27.X1. 1967).

Holotype: South Australian Museum, Adelaide;

2-3 x 1-25 mm; labels: Parachilna, Flinders Range /

Laccobius matthewsi n.sp, E. Gentili 1976 / ¢

Holotypus.

Paratypes: those of the preceding list; they are in

the following Museums: South Australian Museum;

National Museum of Victoria; CSIRO; Museum of

Comparative Zoology of the Harvard University;

California Academy of Sciences; Bishop Museum,

Honolulu; Museums of Verona and Milano, Italy.

7. Laccobius (Macrolaccobius) bicaudatus n.sp.

Body length 3-2 mm; breadth 1-5 mm; elongated,

more than twice as long as wide.

150 REC. S. AUST. MUS 18 (7): 143-154

Labium and head dark with metallic reflections;

two pale patches as wide as eyes, situated before the

latter; the whole surface shagreened, punctured, the

punctures thicker and stronger behind lateral

branches of Y-shaped suture. Fore margin of labium

regularly arcuate. Pronotum pale, testaceous, the

dark metallic spot oval, filling only central area of

disc without reaching borders, the surface shag-

reened and punctured like vertex. Fore margin

sinuous, with the median convexity equal to the

lateral ones (these are scarcely developed, less than

in other Laccobius). Elytra testaceous, hardly paler

on apex and sides; the punctures without dark rings,

the series not always conspicuous, more irregular on

the apical zone. Elytral apices divergent.

Under side dark, except hind abdominal seg-

ments, palpi, antennae, legs, and to a certain extent

the prosternum,. Postlabium flat, shagreened and

punctured; prosternum keel-shaped. The mesoster-

nal keel ends anteriorly in an arrow-point, scarcely

protruding. Hind border of fifth abdominal sternite

scarcely arched, almost straight. Palpi short;

January, 1981]

antennae much longer, legs elongated, normally

shaped.

Holotype: Bishop Museum, Honolulu, Hawaii; it

comes from the Northern Territory and is the only

specimen known to me. Labels: Australia: N.T.,

Edith Ck., 2.XII.1963/J. Sedlacek Collector, Bishop

Museum / E. Gentili det. 1977, Laccobius bi-

caudatus n.sp. / ¢ holotypus.

8. Laccobius (Macrolaccobius) collium n.sp.

(Figs. 5 a, b; 8)

Body length 2-3-2-4 mm; breadth 1-2-1-25 mm:

oval, moderately elongated.

Dark, labium and head greenish-black, without

preocular patches, micropunctured at 60 x, particul-

arly in front of the Y-shaped suture; some larger

punctures behind this suture. Labium and head

large; fore margin of labium regularly arcuate; space

than eyes

between eyes broader as a whole.

THE GENERA LOCCOBIU'S AND NOTITYDRUS IN AUST

Pronotum pale on the sides. the central patch

greenish-black with uncertain borders, of the same

widih as space between eyes, hardly reaching fore

and hind borders of pronotum at middle. Fore

margin convex in middle; the hind also, but very

little, on the scutellum, The whole surface smooth at

60 x, scarcely punctured. Scutellum dark. with

metallic reflexes, sometimes finely punctured.

Elyira testaceous, with some uncertain dark patches

(one near middle of suture; another near base of

elytron; a third between the others, but slightly

external), Punctures darkened (except on sides and

upex); the series are formed by punctures different

in size, and alternately more and less regular,

Sutural angle of each elytron acute, approximate to

the other elytron,

Underside dark, the abdomen sometimes lighter;

palpi, antennae and legs testaceous. Labium withour

specula: postlabium flat or lightly undulated, with

some impressed punctures, near the borders lightly

wrinkled. Prosternum keel-shaped; mesosternum

with a tubercle on posterior half; abdomen and legs

normally shaped, the claws nearly straight. Acdea-

gus \) OF body length; parameres appear to merge

with euch other; median lobe very complex

(Pig. S a, b).

The typical seric¢s comes from the Northern

Territory (Fig. 8). Holotype: CSIRO, Canberra; 23

x [-2 mm; labels: Australia, 12° 31° S, 132° 54° E,9

km N by E of Mudginbarry Hs., N.T., 10.V1.1973,

Upton-Feehan/E. Gentili det. 1976. Laccabius

collum wsp.d Holotypus. Allotype and 4 Para-

types labelled in the same way; they are in the

CSIRO and Verona Museums’ Collections.

9 Laveobius (Macrolaccobius) britton) nesp.

(Figs. 5 c, dy 7)

Body length 2-2-2-4 mm); breadth I:}-1:3 mm.

Closely similar to collin: the chief difference is in

the wedeagus, which is smaller, narrower, and with a

simpler inedian lobe (Mig. 5 ¢, d): the ratio

aedeagus/body length is about |\\. Externally, the

sole difference appreciuble is # greater convexity and

a denser punctuation on the pronotum,

The species comes from Northern Australia

(Fig. 7); Northern Territory; Bessie Springs, 16° 40"

S$. 135° STE. 8 km BSE of Cape Crawford (type

locality, 4 ex.. leg. M. 8. Upton 26.X,1975);

Nabarlek Dam, 12° 20' 8, 133° 19 FE, 15 km S$ by

Nimbuwah Rock {1 ex., leg. B. G. Matthews;

2.V1,1973); Queensland: Mary Creek, 16 33° S,

145° 12° FE (2 ex., leg. Britlton-Misko. al light.

4.X17,1968),

Holotype: CSIRQ, Canberra; 2-2 x L-l mm;

labels: Australia, 16° 40° S, 135" 51° E. Bessie

AND NEW 7FALAND 15|

Spring, 8 km ESE of Cape Crawford. 26.%.1975, M,

S. Lipton, N.T/E. Gentili det, 1976. Laccohius

brittont o.sp./% Holotypus,

Paratypes: the above list; they are in the CSIRO

and Verona Museums Collections,

THE BLACKBURN “LACCOBIUS” (NOTHYD-

RUS BALF.-BR,, 1939, EMENDED)

Blackburn (1891) described two ‘“Laccobius”,

montanus and australis, affirming that they “may be

attributed to this genus... although they differ from

the European members of ttin having --. characters

... that may justify a new generic name”. Ln effect,

after the study of the typical females in the British

Museum collechons, Balfour-Browne (1939) pro-

posed for these insects the new generic name

“Norohydrus", emended by d'Orchymont (1942) to

Nothydrus. 1 agree with these coleopterists; the

differences [rom the nearer genera are in my opinion

great cnough to separate Norhydrus as an mdepen-

dent genus,

Nothydrus has chiefly the following characters.

Labiium shorter than in Laeceobius, widely excavated

anteriorly; prefrons flat, long, with a straight lore

margin{ upper surface of the head punctured.

Postlabium plate-sshaped. Antennae &-segmented

(5 | 3). the fifth article nearly cupuliform, like the

Cereyonini, maxillary palpi robust, nearly as long as

the antennae, the fourth article longer than the

third, The fore margin of the pronotum is widely

excavated for the reception of the head, the corners

widely rounded, the surface punctured, The

prosternum is lacking a keel. The fore femora are

pubescent on the whole under surface, glabrous only

near the apices; the fore tarsi do not show any sexual

dimorphism, Secutellum nearly equilateral, Elytra

with parasutural suleron /) of their length, these sulci

continuing anteriorly by meaus of some large

punetures; the elytral punctures are scattered, but

showing about ten series of larger punctures, more

conspicuous and engraved on the hind surface; as in

Laeccobius, the epipleura are anteriorly large, and

reduced on the posterior half of the eclytron. The

mMesosternum is tuberculate, not sharply curinute.

The mid femora ure pubescent like the fore. The

wWiitgs are present, provided with a cantharidiform

nervation. The metasternum is pubescent, slightly

elevated in the middle. The hind trochanters are

small, not separated at their upices from the femora;

these are almost entirely pubescent, save at the

apices and sometimes the hind margin; the hind

libiae wre straight; the fifth metatarsal segment is

shorter than the four basal segments taken together.

Five abdominal segments are visible, the sixth at rest

is internal. The male sexual organs are similar to

those of Ayaceaena in their general pattern,

152 REC, $8. AUST, MUS Ik (7)

The genus differs from Laccobius in having five

abdominal segments, a prosternum without a keel,

small hind trochanters not separated from the

femora, pubescent femora, sexual dimorphism

absent on fore tarsi, and a sutural stria on the elytra,

From Paracymus, it differs in having the prosternum

without any keel and the mesosternum without an

arrow-head shaped keel, It differs from Anacaena in

having an elongated and depressed body form, a

deeply excavated lablum, conspicuous elytral series,

and a cupuliform fifth antennal article. In contrast to

Balfour-Browne (1939), who suggested an affinity

with Laccohius, placing the genus Norhydrus at the

end of the subtribe Hydrobiae, d’'Orchymont (1942)

suggested an affinity with Anacaena. The pattern af

the aedeagus (basis of the tegmen pointed; aedeagus

flat, with 4 simple trilobate structure) confirms his

opinion.

Nothydrus australis (Blackburn, 1891)

(Fig. 6 a)

Blackburn, 1891 (Laccubius); Knisch, 1924

(Laccobius); Balfour-Browne, 1939 (Notuhydrus);

d'Orehymont, 1942.

Body length 2:9-3-5 mm, breadth |:5-1-8 mm;

oval elongated, less than twice as long as its width.

Fig, 6—A, Aedeagus of Nothydrus australis, Dondangadale,

Victoria: frontal view; B, Aédeagus of Nothydrus sientanus,

Erica, Victoria: frontal view.

The beetle has the characters of the genus. It is

enough to record here the differences from

montanus. The dorsal colour is pale, testaceous,

143-1454 Junuary, 198l

only the discs of the head and of the pronotum being

dark, The puncturation of the head and pronotum is

finer and sparser than that of the elytra, while on the

elytra the puncturation of the series is quite notably

distinct from that of the general surface, ‘The sides of

the pronotum are not transparent. The greatest

width is at the level of the middle of the elytra. The

plate of the postlabium (flat, smooth, without

punctures) and the sides of the abdomen are

yellowish, the remainder of the under surface is

dark, The hind femora are pubescent near the fore

margin. Aedeagus nearly '/s of the body length,

tegmen longer than parameres; these are narrow and

sharp al their apices like the median lobe (fig. 6 a).

Australia, Victoria; Ovens River (type locality,

Blackburn, 1891); Bogong Village (1 ex., leg.

A, Neboiss, 23,1.1960); Upper Nariel (6 ex. leg,

A. Neboiss, 29.1,1957); Dondangadale (2 ex., leg.

A. Neboiss, 11.1.1955); Marwall River (3 ex., leg

§. Trout, XII.1945); Sth. Neerim (1 ex., leg. PF. E.

Wilson 11.1,1941, det. A. d’Orchymont). These

specimens are jn the National Museum of Victoria,

Melbourne, and in the Museum of Verona, Italy,

Holotype: British Museum; 3:0 * 1:5 mm; labels:

T 3564 Al / Holotype / Australia, Blackburn Coll.,

B. M. 1910-236 / Laecobius australis Blackb. /

Laccobius australis Blkb., M. BE. Bacchus det, 1977,

Holotype / Nathydrus australis Blackb., E. Gentih

det, 1977,

Nothydrus montanus (Blackburn, 1891)

(Fig. 6 b)

Blackburn, 1891 (Laccobius); Knisch, 1924

(Laccobius); Balfour-Browne, 1939 (Notohydrus):

d’Orchymont, 1942.

Body length 3-5-4:5 mm; breadth 1-9-2-3. mm;

Jarger than australis but like tt in shape,

The chief differences from australis are the

following. Dorsal colour darker, blackish-brown,

more shining; apart from the preocular patches, only

the sides of the pronotum are clear and transparent,

The upper puncturation is the same on the head,

pronotum and elytra, consisting of finer and sharper

punctures than in australis, and it includes the

punctures of the series, confusing the latter with

those of the interseries. The greatest width js at the

level of the base of the clytra. The plate of the

postlabium (flat, without punctures, but slightly

alutaceous) is pale, the remaining underside is dark.

Hind femora pubescent at the bases, Aedeagus

neatly \ of body length; tegmen long like the

parameres, these larger than in australis, with

rounded apices; the median lobe very wide at the

base (Fig. 6 b)

Australia, Victoria! Victorian Mountains (type

THE GENERA LOCCOBIUS AND NOTHYDRUS IN AUST, AND NEW ZEALAND 153

locality, Blackburn, 1891); Bogong, High Plain (1

ex., leg. A. Neboiss, 26.1.1960); Erica (4 ex., leg. A.

Neboiss, 29.1.1960); Millgrove (1 ex., leg. A.

Neboiss, 26.11.1958); Cumberland Falls (1 ex.. leg.

A. Neboiss, 16.11.1958); Black’s Spur (2 ex., leg. C.

Oke, 1.1934); Fern Tree Gully (1 ex., leg. C. Oke);

New South Wales: Kosciusko (2 ex., leg. H. G.

Carter). The specimens of this list are in the

National Museum of Victoria, Melbourne, in the

South Australian Museum, Adelaide, and in the

Museum of Verona, Italy.

Holotype: British Museum; 3-6 x 1-85 mm;

labels: Holotype / T 3563 Al. / Australia, Blackburn

Coll., B. M. 1910-236 / Laccobius montanus Blackb.

/ Laccobius montanus Blkb., M. E. Bacchus det.

1977 Holotype / Nothydrus montanus Blackb. E.

Gentili det. 1977.

ACKNOWLEDGEMENTS

I am indebted to the following colleagues and

institutions for their generous assistance:

Dr. E. G. Matthews, South Australian

Museum, Adelaide (Loan of material and

revision of the manuscript).

Mr. M. E. Bacchus, and Mr. P. M. Hammond,

British Museum, London.

Dr. A. Bons, and Dr. H. Perrin, Muséum

National d’Histoire Naturelle, Paris,

Dr. E. B. Britton, CSIRO, Canberra.

Dr. A. A. Calder, National Museum of

Victoria, Melbourne.

Mr. E. C. Dahms, The Queensland Museum,

Fortitude Valley.

Dr. R. Damoiseau, Institut R. des Sciences

Naturelles de Belgique, Bruxelles.

* zietzi

e decipiens

= arrowi

a brittoni

| —

Na 120 130 140

—J

150 160 170 180

Fig. 7—Geographical distribution of Laccobius zietzi, decipiens,

arrowi, marmoratus, and brittoni.

REC. S. AUST. MUS 18 (7): 143-154

January, 1981

clarus

matthewsi

collium

100”t”~été“‘C;‘SCO.O!OWOWOWOWOWOOOOO#COWSO

140 150 140

Fig. 8—Geographical distribution of Laccobius clarus, matthewsi,

and callium,

Mr. G. A. Holloway, The Australian Museum,

Sydney.

Dr. D. H. Kavanaugh, California Academy of

Sciences, San Francisco.

Dr. J. Krikken, Rijksmuseum van Naturlijke

Historie, Leiden,

Dr. M. Pearce, Museum of Comparative

Zoology, Harvard University, Cambridge,

Mass.

Dr. G. A. Samuelson, Bernice P. Bishop

Museum, Honolulu.

Dr, M. Winterbourn, University of Canterbury,

Christchurch.

Dr. K. A. J. Wise, Auckland Institute and

Museum, Auckland.

Special thanks also to Mr. A. Bertocchi, who

prepared the drawings.

REFERENCES

BALFOUR-BROWNE, J. 1939. Contribution to the study of the

Palpicornia, If. Ent. Month. Mag. 75: 5-6.

BLACKBURN, T, 1891. Further notes on Australian Coleop-

tera, with descriptions of new Genera and Species. Trans. R.

Soc. S. Austr. 14, (1): 67-68,

1895. Further notes on Australian Coleoptera, with

descriptions of new Genera and Species. Trans. R. Soc. S,

Austr. 19 (1): 20-73,

BRITTON, E. B. 1970. Coleoptera (Beetles) In; Waterhouse D.

F. et al. ‘The Insects of Australia” (Melbourne University

Press, Melbourne).

GANGLBAUER, L. 1904, Die Kifer von Mitteleuropa’’. TV,

251, note. (Goecke & Evers. Krefeld).

GENTILI, E, 1974, Descrizione di nuove entité appartenenti al

genere Laccobius Erichson, 1837 e proposta per un nuovo

inquadramento sottogenerico. Mem. Mus. Civ, Sr, Nat.

Verona 20 (1972): 549-550, 1 fig.

GENTILI, B., and CHIESA, A. 1975, Revisione dei Laccohius

palearetici. Mem. Soc. Ent. It. 54: 6-22.

KNISCH, A, 1924. Hydrophilidae. In: Junk, W., Coleopterorum

Catalogus: 181-192 (W. Junk, Berlin).

MACLEAY, W. 1873. Notes on a collection of Insects from

Gayndah. Trans. Ent. Soc. N.S. Wales 2: 130-131.

ORCHYMONT, A. d’. 1925, Contribution a l'étude des

Hydrophilides, 1. Bull. Ann, Soc, Ent. Belg. 65: 66-70.

ORCHYMONT, A. d’. 1937, Check list of the Palpicornia of

Oceaniu, Occas. Papers Bern, P. Bishop Mus. Honolulu 13:

13, 153.

ORCHYMONT, A. d’. 1942. Contribution a l’étude de la tribu

Hydrohiini Bedel. Mem. Mus, R. Hist. Nat. Belg. I, 24: 3-

10, 39.

ORCHYMONT, A. d°. 1948. Report on the Palpicornia, Mr.

Omer-Cooper’s Investigation of the Abyssinian Fresh Waters

(Hugh Scott Expedition). Proc. Zool. Sec, Landon 117: 727-

728.

WINTERBOURN, M. J. 1970. The Hydrophilidae of New

Zealand's Thermal Waters. Trans. R, Sec. New Zealand 12:

21-28, 11 figs.

WINTERBOURN. M. J. 1973. The larva of Anacaena tepida

from a Rotorua Hot Spring. New “ealand Ent. 5: 171-174. |

fig.

WISE, K, A. 1973. A list and Bibliography of the aquatic and

water-associated insects of New Zealand. Rec. Auckland

Inst, Mus, 10: 156.

ZAITZEV, P. 1908. Catalogue des Coléoptéres aquatiques des

familles des Dryopidae, Georyssidae, Cyathoceridae,

Heteroceridae et Hydrophilidae. Horae Soc, Ent, Rassicae

38: 391-395,

THE FABRICAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND

COLEOPTERA IN THE BANKS COLLECTION AT THE BRITISH

MUSEUM (NATURAL HISTORY)

BY WINIFRED P. K. RADFORD

Summary

The Banks Collection is a collection of Fabrician types housed in the British Museum (Natural

History). It was presented to the British Museum in 1863 by the Linnean Society. In this paper the

Australian and New Zealand Coleoptera are classified as far as possible according to their present

status. This classification includes nine new synonyms and eighteen species omitted from recent

catalogues. In the discussion of the species they are recorded alphabetically. The original Fabrician

reference and the 1801 reference are given, together with the Junk Coleopterorum Catalogus

reference where applicable, and the Zimsen reference. As far as possible known synonyms of the

species are listed. The labels and condition of the types are given and in some cases comments are

made as to identification and locality. Sometimes it has been found that the Fabrician locality is

incorrect. There is rarely a locality label on the specimen — but is only given in the original

description. In the cramped ocnditions on the H.M.S. Endeavour there must have been some mixing

of material.

THE FARRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

IN THE BANKS COLLECTION AT THE BRITISH MUSEUM (NATURAL HISTORY)

by WINIFRED P. kK. RADFORD

la Cheviot Avenue, Lower Mitcham, South Australia 5062

ABSTRACT

RADFORD, WINIFRED P. K. 1980, The Fabrictan types of the

Australian and New Zealand Coleoptera in the Banks

Collection at the British Museum (Natural History), Rec. §

Aust. Mus. 18 (8); 155-197,

The Banks Collection is a collection of Fabrician

types housed in the British Museum (Natural

History), It was presented to the British Museum in

1863 by the Linnean Society. In this paper the

Australian and New Zealand Coleoptera are

classified as far as possible according to their present

status. This classification includes nine new

synonyms and eighteen species omitted trom recent

catalogues. In the discussion of the species they are

recorded alphabetically. The original Fabrician

reference and the 1801 reference are given, together

with the Junk Coleopterorum Catalogus reference

where applicable, and the Zimsen reference. As far

as possible known synonyms of the species are listed.

The labels and condition of the types are given and

in some cases comments are made as to identifica-

tion and locality. Sametimes it has been found that

the Fabrician locality is incorrect. There is rarely a

locality label on the specimen—but is only given in

the original description. In the cramped conditions

on the H.M.S. Endeavour there must have been

some mixing af material.

INTRODUCTION

In 1928 Dr. R. J. Tillyard, who was the chief of the

Division of Entomology in the then recently created

Australian Council of Seience and Indusirial

Research, suggested that while J] was studying in

London ] should examine the Australian and New

Zealand species of Coleoptera in the Banks

Collection located in the British Museum. This [ did

and received much help from G. J. Arrow, and K.

G. Blair of the Museum and G, E. Bryant of the

Entomological Bureau, On my return to Australia,

late in 1929, 1 was fully occupied in establishing the

Coleoptera Collection of the C.S.1.R. in Canberra.

My manuscript on the Banks Collection was laid

aside and only came to light again in 1976.

In the meantime specialists in various groups have

studied some of the species, tracing out their present

status and in some cases their synonymy. These are,

as far as IT have been able to ascertain, H. J. Carter

(1926, 1929, 1930) on Austrahan Tenebrionidae,

Buprestidae, and Alleculidae; P. Lepesme (1939) on

Dermestidac; HL E. Hinton (1945) on Dermestidae;

G. Ochs (1949) on Australian Gyrinidae; P. B.

Carne (1957) on Australian Dynastinae: G. Kuschel

(1969, 1970) on New Zealand Curculionoidea; E.G.

Matthews (1972, 1974) on Australian Scarabaeinac,

Y¥. Lobl (1976) on Australian species of Scaphidium.

The purpose of this paper is to present a list of the

Fabrician Types. of the Coleoptera in the Banks

Collection im the British Museum (Natural

Museum), for which Fabricius recorded the localities

as ‘'nova Hollandia”, “nova Zelandia’, and ‘terra

Diemenii”. A systematic list has been drawn up and

the species classified, as far as possible, into families,

sub-families, and their present generic status, It has

been found that some species have been omitted

from recent catalogues, This may be due to the very

brief and inadequate Original descriptions, which are

often no more than twa lines, together with the

“habitat” for the locality giving only the name of the

country, €.g., “nova Hollandia’ and not mentioning

any particular regian, Sometimes the name of a

collector or a collection is given, e.y,, “Mus.Dam.

Banks”, or just “Dom.Banks’~ Tn some instances a

camparison with a selated species is given or,

especially in his later publications, a fuller

description is added.

Fabricius at first only separated the insects he

studied into the large groups or orders as we now

know them. Then, on the characters of the mouth

parts, he classified them into genera only, not into

families and sub-families although he did put them

into groups according to the antennal structure. As

he studied more and more specimens he realised his

initial genera contained many species. which could be

better classified inta smaller graups and so added

more and more genera. Thus we find in his later

publications the original generic name of the species

was sometimes changed.

E. Zimsen (1964) published a book on ''The Type

Material of J, C, Fabricius". In this work she states

where the specimens on Which Fabricius based his

descriptions are to be found. Fabricius did not label

any particular specimen as a type and sometimes his

descriptions were made from more than one

specimen. S. L. Tuxén (1967: p. 8), who worked

closely with Ella Zimsen, explains that trom the

references given by Fabricius in his original

descriptions she “‘inferred, though it is nowhere

stated explicitly in his books" that where the name

of a collector or a collection is given that is where the

“type specimen or specimens” are to be found, and

where no such reference is given the type specimen

is in his own collection, now housed in the

Zoological Museum at Copenhagen, Tuxén adds,

“Her work. based on this axiom. showed this to be

the case ., . There may be reasons of labelling for

this difficulty (choosing the type), but there is also

156

another reason which Fabricius himself mentions in

1803 (Tuxén 1967; pp, 9-10), In later years Fabricius

himself got so confused in describing so many

species (Zimsen lists 9 776), that he sometimes put

one insect in two different genera or used the same

name for different species. Tuxén quotes Fab-

ricius:—""In my youth this did not easily happen, my

memory was beiter and my eye sharper, now both

are weaker...” It is understandable that in dealing

with so many Specimens mistakes occurred. Where

there is only one known specimen then it can be

taken as the holotype, but where there is more than

one specimen later workers have sometimes selected

a specimen as the lectotype and the others in the

senes are known as paralectotypes. Where no such

study has been made the series are known as

syntypes. In the late 19th century C, O, Waterhouse

studied the Fubricius material in the Banks

Collection and put type labels on those specimens

which he was reasonably able ta ascertain were the

specimens ta Which Fabricius’s description referred.

Where the species is represented in the collection by

only one specimen lt have called it the holotype as

Fabricius’s reference states in “Mus, Dom. Banks",

If there are two specimens | have called them

syntypes according to the present day nomenclature,

If one of these has a museum type-label an it, in the

comments | have referred to that specimen as “type”

as future workers may designate them as hoilo-,.

lecto- or syntypes. Where there was more than one

specimen of a species the owner of the collection

often gave Fabricius specimens for his own

collection, Thus some “type material” occurs in

more than one place. The references for the

synonyms of each species are not given in the

bibliography,

This paper brings to light 18 Fabricius species

which have been overlooked of omitted from recent

catalogues, records nine new synonyms, and gives a

modern classification for the Australian and New

Zealand species in the Banks Collection.

JOHANN CHRISTIAN FABRICIUS

Johann Christian Fabricius was born act Tender in

Denmark in 1745. He was fortunate that he was able

to travel throughout Europe and England, where he

had access to many museums and private collections,

London became his second home. At the British

Museum he met Dr. C. Solander who, like himself.

had studied under Linnaeus at Uppsala. When

Solander and Banks sailed on H.M.S, Endeavour,

Fabricius returned to Copenhagen where he had

been appointed a professor at the Charlotlenberg

Naturalickabenet. In 1771 TLM.S. Endeavour

returned to England. From 1772-1775 Fabricius

divided his time between Copenhagen and London,

continuing his studies of the insects in the collections

RECS ALIST. MUS, 18 (8), 455-107

January, [98]

of Hunter, Drury and Banks—the latter now

included the insects brought back tram the voyage

an the Endeavour. In 1775 his Systema

Entomologiae was published. which cansisted

mainly of descriptions of inseets he collected in

England, or of insects he had studied in the English

collections. He stated of this publication, 'Entamol-

opy was at that time in its infancy’. Up till then the

only classification known was that of Linnaeus, who

had based his classification on the wings of insects

Which was not satisfactory as Linnaeus himself

realised. Also Linnaeus did not have sufficient

specimens fo make an entirely satisfactory classifica-

tion, Fabricius used the mouth parts as a basis for

classification. He was aware of its limitations bur

considered his classes were “more natural and my

species were more numerous and more ably defined

and the number of described genera considerably

greater” Fabricius had now become the entomolog-

ical authority of the day, and in his classifications

introduced the coneise language of the Linnaean

“school” into this compartment of Natural History.

He continued ta travel in Europe and in England

where he spent the summers of 1780-81 and again in

1787 (Tuxén 1967: p. 3). During these travels he had

access tt) many museums and private collections. He

published many systematic warks on insects and in

the later part of his life published monographs of the

larger orders of insects, He died in 1808.

SIR JOSEPH BANKS

Joseph Banks was born in 1743 into a wealthy

family. He was educated at Harrow. Eton and

Oxford where Botany was one of his chief studies-

Although Botany was his life interest he had an

extensive knowledge of all the Natural Sciences, His

house at 32 Soho Square was open to all who wished

to study his colleetians aud work in his library. His

wealth enabled him to undertake and finance three

voyages. The first in 1766 to Newfoundland and

Labrador, the second an the Endeavour from 1768-

1771, and the third to Iceland and Staffa, It was due

to the Royal Society that he obtained permission for

himself and a staff of seven, including Dr. C.

Solander of the British Museum. to accompany

Lieutenant (jater Captain) Cook on the Endeavour,

He was fully aware of the task he was undertaking to

make collections. of Natural History specimens,

which would be mostly new to Science, on the long

voyage ahead of him. His preparations were

thorough, John Ellis F.R.S. in a letter to Linnaeus

gives us the only known account of these

preparatians:— “No people went to sea better fitted

out for the purpose of Natural History. They had got

a fine library of Natural History; they have all sorts

of machines for catching and preserving insects, all

kinds of nets, trawls, drags and hooks for coral

fishing; they even have a curious contrivance of a

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 157

telescope. by which, put into water, you can see the

hotiom at a preat depth where it is clear. They have

many cases of bottles with ground glass stoppers, of

several sizes to preserve animals in spiric. They have

several sorts of salts ta surround the seeds. and wax

both beewax and that of Myrica, Besides there are

many people whose sole business is to attend them

for this very purpose”, He was a true scientist in that

he was a keen observer and a careful recorder, All

specimens collected were recorded and preserved.

He was also a keen student of the ethnic races that

they encountered. Banks made friends easily and

throughout the three years voyage there was no

friction with Cook and the crew.

The voyage of the Endeavour enabled the

scientists ta collect at Madeira, Rio de Janeiro, in

the region of Cape Horn, Tahiti, round the coast of

New Zealand, where Cook discovered it to be twa

islands. not one; alang the east eoast of Australia

landing for short periods at Botany Bay, Bustard

Bay, Thirsty Bay, then for two months at Cooktown

while the ship had to be repaired after running on ta

a reef; then sailing round Cape York, proving New

Guinea to be a separate island, stopping briefly on

the west of New Guinea at Cook's Bay; then on ta

Suva and Batavia, and home round the Cape of

Good Hope with a stop at St. Helena.

The voyage was primarily undertaken to observe

the transit of Venus across the sun at Tahiti. This

was nol successful as the instruments af those days

did not make it possible to fake accurate

observanions. However, the work of Burks and his

staff, the collections and observations of Natural

History and of the ethnie races they encountered,

were of inestimable value, After this, scientists were

always part of the personnel of the British voyages of

discovery. Banks himself did nol publish uny papers,

He did keep and file all his correspondence which

showed how wide his interests were. In 1779 he was

asked to give evidence before a commiltee of the

House of Commons concerning the suitability of

Botany Bay as a convict settlement, Nothing was

done till 1786 when the over-crawding of the prisans

made the matter urgent. and on Banks’ advice plans

were made and the new colony was established

shortly afterwards. For some time the governors of

the colony wrote to Banks for help and advice and

he acted as a go-between for the Goyernment and

the calony. Banks was by then scientific adviser te

the Royal Gardens at Kew and his work there made

the Gardens a centre for botanical research into

plants of all the colonies of the Crown, which he

foresaw would be necessary for their development.

Today the Gardens are still a centre for botanical

research.

Unfortunately Sir Joseph Banks ended his life as

an invalid and a cripple, but to the end, his interest

in all around him never flagged. His papers were left

to his wife's nephew, and at his death were to go to

the British Museum, Through misunderstanding as

to the final destination and ownership, the papers

were sold at auction, many were lost or destroyed

and those remaining were distributed all aver the

warld. Foriunately for Australia many of the papers

relating to Australia, including the original copy af

the Journal of the Voyage of the Endeavour, were

procured and are now in the Mitchell Library, Apart

from his papers. his library and collections were left

to the botanist, Reahert Brown. Later via the

Linnaean Society they were removed to the British

Museum which ensured their preservation.

THE BANKS COLLECTION OF AUSTRALIA

AND NEW ZEALAND CGLEOPTERA

In a report by C. O. Waterhouse (1906: p. 580) he

states that the Banks Collection “was received in the

original Banksian cabinet , . . the specimens are,

however, kept in the arder of Fabricius’ Systema

Entomolagiae.” This is the 1775 publication. Many

species in the collection were described after this

date. Waterhouse adds, “Some of the species

mentioned by Fabricius as being in this collection

were not in it When it was received by the Museum,”

Recently (1979) some Banks material has been

found in the Hunterian Collection in Glasgow, Jn

correspondence with Dr. H. Brock of the University

of Glasgow she writes; “There are in all over 100

specimens in the Hunterian Collection that must

have come from the Cook voyages.’ I agree with her

that Fabricius when working on both the Banks and

Hunter Collections ‘enabled Hunter to acquire

Banks duplicates." Banks was a generous man both

with his money and possessions and shared them

with others. Fabricius certainly added specimens

fram the collections he studied to his awn collectian,

Now two of the six missing ‘types’ from the Banks

Collection, namely Dennestes navalis and Lampyris

australis are found to he in the Hunterian

Collection, As he worked with so many specimens it

is very probable that some specimens were put back

in the wrong collection. The Banks Collection

contains over 3.000 insects in all. The Systema

Eleutheratorum is Fabricius’s record of the Coleopt-

era he described.

The Banks Collecuon of Coleoptera is naw

arranged in the order given by Fabricius in the

Systema Eleutheratorum. 1801, except for one

specimen which was omitted from the

Eleutherstorum, and was found by EB, 4imsen

(1964) among the unnamed specimens. It was then

added at the end of the named specimens im the

collection,

[58 REC, & AUST

Where they exist, the references given for cach

species are;—

1. The original reference,

2, The Systema Eleutheratorum.

3. Junk’s Coleepterorum Catalogs

4. The reference in “The Type Material af J-C.

Fabricius” by E. Zimsen.

5. The reference of a recent reviser al the

group,

The species with the original ceference in Systema

Entomologiae, 1775, and four Curculionaidea from

New Zealand with the original reference in the

Species Insectorum, 1781, were collected during the

voyage of the Endeavonr. G, Kuschel (1920: p. 192)

states that “the more Spectacular weevils’ were

described first in 1775 and “the remamder™ were

described later, The species with the original

reference in the Manitissa Insectorum, L787. were

collected by D. Nelson, who was sent by Banks on

Captain Cook’s third voyage on H-M.S-. Discovery.

After the return of H.M.S. Endeavour many more

voyages of discovery were made. Banks himself anly

made one more voyage after his return and that was

to Iceland and the Shetland Islands, However, when

opportunity offered, he sent aut men at his own

expense or from the Royal Gardens ut Kew to

collect for him and for the Gardens (Maiden 1909: p.

124). David Nelson was one of these collectors.

In this paper 131 Fabrician specific names are

listed, ten of these are synonyms as Fabricius

renamed those species in his later publications for

one reason or another. Of the remaining 121;—

i. Seventy-nine species are recorded by Fab-

ricius from “‘nova Hollandia’. The localiry

of seven of these were erroneously

recorded, three are probably from the East

Indies, and four from New Zealand. As G.

Kuschel (1970: p. 191) points out, there

must have been some mixing of matenal on

H.M.S, Endeavour.

. Thitty-two species are listed fram “nova

Zelandia™. Of these, the locality of ane is

doubtful, and one erroneously recorded

from “Brasilia’’ has been included (Kus-

chel 1970: p, 199).

i, Hight species are recorded from “‘terra

Diemenit’ one of which is labelled

Kerguelen.

iv, Two are recorded as being from both ‘‘nova

Hollandia et Zelandia”’,

wot

In each case an exact copy of the Fabrician label,

which is in his own handwriting, has been given,

Sometimes the label is no more than a scrap of

paper, but according to Zimsen (1964: p, 10)

Fabricius in is own collection used “a small secap of

paper oo which was written the name of the species.

MUS 18 (Ry: 155.197

January, Fost

never mention of the genus’, In the Banks

Collection, however, the genus is usually given, but

very rarely the localily or name of a collectar. The

Fabrician labels in this Collection are usually writren

in two lines, the upper one giving the name and the

lower one the reference, In some cases the reference

1s the number only of the species in the genus being

described, with no page reference, This would

indicate that the label was attached when the species

was described and before publication; where (he

pageé also is given would denote that the label was

added after publication, In the introduction of both

his publications, Species Insectorum, 1781, and

Mantissa lnsectorum, 1787, it is stated that these

works contain dese¢riptians of species studied in

“Angha™ (England), These descriptions were not

only of wew species but also fuller descriptions of

some species alrgady described in the Systema

Entamoelogiae, 1775. When compiling his Species

Insectorum, 1781, he says ‘My system of Insects

gained ground considerably, as well by the more

exact definition of the species, as by the addition of a

considerable number of geners.’” In doing so

Fabricius must have changed the labels on some of

the species he redeseribed, so we find references toa

later publication instead of the original one, and

sometimes a change of the generic name. As nore

and more specimens were examined he realised his

classification required further divisions, Therefore,

sometimes the BMNH Jabel has a different genus to

the original one, in fact the genus for the species as

given in Systema Eleutheratorum, Itis probable that

the BMWNH label was written when the collection

was arranged acearding to the order in that

publication, A few specimens also haye a BMNH

Museum name-labels are not attached to the

specimen but are pinned below it, except for C. OQ.

Waterhouse's type-lahels, Kuschel (1970) did not

add any holotype or lectotype labels when he studied

the New Zealand Curculionoidea in the collection.

The specimens are usually pinned through the right

elytron and in most cases the pin has been glued on

the ventral surface, often obliterating the structures.

In the discussion of the species they are listed in

alphabetical order avcording to the specific name.

Known synonyms are listed, The numerous varieties

of the chrysomelids are not listed as [ beheve that

there is much work to be done on the separation of

these varieties, ‘The localities given are those

published by Fabricius, but corrections to these

localities have sometimes had to be made.

The following list of places visited by Cook around

the coasts of New Zealand and Australia will give

some indication Where the type specimens were

collected, A, Musgrave (1954; p. 232-7) points aut

that when the Endeavour was sailing up the coast of

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

Australia it was late in the season for collecting

insects at Botany Bay, so probably most of the

insects from Australia would have been collected at

Endeavour River (Cooktown), However, Banks’

journal does record that collecting was done at both

Bustard and Thirsty Bays,

Round the coast of New Zealand:—

1769 9-10 October at Poverty Bay

20 October at Anaura Bay

23-29 October at Tolaga Bay

4-15 November at Mercury Bay

20 November at Waihou (Thames) River

29 November- 5 December at Bay of

Islands

16 January- 6 February at Ship Cove,

Queen Charlotte Sound

27-30 March at D’Urville Tsland

31 March left New Zealand waters and

sailed to Australia

27 April- 6 May at Botany Bay

23-24 May at Bustard Bay

29-31 May at Thirsty Bay

16 July- 4 August at Endeavour River

(Cooktown)

3 September at Cooks” Bay, New Guinea

18-2] September at Savu (Sawu) Indonesia

10-26 October at Batavia (Jakarta) Java

4-14 January at Princes Island

13. May- 7 June at Ship Cove, Queen

Charlotte Sound, New Zealand

3-25. November at Ship Cove, Queen

Charlotte Sound, New Zealand

23 March- 11 May at Dusky Sound, New

Zealand

24 December- 7 January at Kerguelen’s

Land

26-30 January at

Tasmania

12-25 February at Ship Cove, Queen

Charlotte Sound, New Zealand

The letters BMNH are used to denote the British

Museum (Natural History)

1770

1771

1773

1776-77

Adventure Bay,

CLASSIFICATION OF THE AUSTRALIAN AND

NEW ZEALAND COLEOPTERA IN THE BANKS

COLLECTION

This classification follows that of E. Britton in

“Insects of Australia’ published by the C.S.1.R,O.

(1970: pp. 517-8),

New synonyms, and those species Fabricius

renamed in later publications, now regarded as

synonyms, are given in this classification. A full list

of synonyms is given under the heading of each

species,

The original generic names are in parentheses.

159

*Denotes that the species has been omitted from

recent catalogues.

*Denotes a new synonym.

FAMILY CARABIDAE

sub-family Scaritinae

Laccopterum cyaneum (F.) (Scarites)

sub-family Cicindelinae

Cicindela tuberculata F.

FAMILY DYTISCIDAE

Platynectes decempuncratus (F \

(Dytiscus)

FAMILY GYRINIDAE

Dineutes australis (F.) (Gyrinus)

FAMILY HISTERIDAE

Saprinus cyaneus (F.) (Hister)

* Saprinus detritus (F.) (Hister)

FAMILY SCAPHIDITIDAE

Scaphidium quadripustulaium (F,)

(Sphaeridium)

FAMILY STAPHYLINIDAE

Creophilus aculatus (F,) (Staphylinus)

Creophilus erythracephalus (F.) (Staphy-

linus)

FAMILY LUCANIDAE

Lissotes cancroides (F.) (Lucanus)

FAMILY SCARABAEIDAE

sub-family Scarabaeinac

Onthophagus quadripustulatus (F.) 2

(Scarabaeus)

= O, bipustulatus (F.) (Scarabaeus) ?

Tesserodon novaehollandiae (F.)

(Scarabaeus)

= T-. hollandiae (F.) (Scarabaeus)

sub-family Cetoniinae

Glycyphana stolata (F.) (Cetonia)

= G. fasciata (F.) (Cetonia)

sub-family Dynastinae

Haploscapanes barbarossa (F.)

(Scarabaeus)

Pericoptus truncatus (F.) (Scarabaeus)

sub-family Melolonthinae

Calonota festiva (F.) (Melolontha)

C. festiva var, laeta (F.) (Melolontha)

Chlorochiton suturalis (F.) (Melolontha)

Liparetrus sylvicolus (F.) (Melolontha)

Liparetus monticolus (F.) (Melolontha)

sub-family Rutelinae

Repsimus aeneus (F.) (Meloloniha)

FAMILY PSEPHENIDAE

* Sclerocyphon collaris (F.) (Tritoma)

+ = §S. hicolor Carter

FAMILY BUPRESTIDAE

sub-family Buprestinae

Cisseis s.g. Neospades eruciata (F.)

(Buprestis)

160)

sub-family Chalcophorinae

Cyphogastra farinosa (F.) (Buprestis)

FAMILY RHIPICERIDAE

Rhipicera mystacina (P.) (Hispa)

FAMILY LAMPYRIDAE

Luciola australis (F.) (Lampyris) in

Glasgow

FAMILY MELYRIDAE (DASYTIDAE)

Dasytes minutus (F.) (anobium)

= D. subcyaneus Broun

FAMILY DERMESTIDAE

Dermestes carnivorus F. (carniforus F.)

Dermesies felinus F. is a synonym of D.

ater Degeer

FAMILY BOSTRYCHIDAE

Bostrychopsis jesuita (F.) (Apate)

Dinoderus minutus (F.) (Apate) Type

lost

FAMILY LYCIDAE

Trichalus serraticornis (F.) (Lycus)

FAMILY CLERIDAE

Eunstalis percata (F.) (Notoxus)

Phymatophaea violacea (F.) (Noloxus)

= Balcus niger Sharp

FAMILY NITIDULIDAE

Macroura abbreviata (F.) (Nitidula)

FAMILY PHYCOSECIDAE

* Phycosecis limbatus (F.) (Dermestes)

FAMILY COCCINELLIDAE

Coccinella leonina F.

Coelophora inaequalis (F.) (Coccinella)

Coelophora novemmaculata (F,) (Coc-

cinella)

= Coelophora novempunctata (F.) (Coc-

cinella)

Epilachna guttato-pustulata (F,) (Coe-

cinella)

Veranis striola (F.) (Coccinella)

= V. lineola (F.) (Coccinella)

FAMILY COLYDIDAE

Ulonotus scaber (F.) (Dermestes)

= U. integer Sharp

FAMILY TENEBRIONIDAE

sub-family Ulominae

Adelium porculatum (F.) (Calosoma)

=A. porcatus (F.) (Carabus)

Alphitobius laevigatus (F.) (Opatrum)

Celibe laevicallis (F.) (Silpha)

Dermestes navalis (F,) is a synonym of

Triboloides (Blair 1913) ferrugineus (F.)

* Uloma sanguinipes (F.) (Tenebrio)

sub-family Amarygminae

Amarygmus bicolor (F.) (Eratylus)

* Amarygmus morio (F.) (Erotylus)

Chaleopterus amethystinus (F.)

(Erotylus)

REC. §. AUST. MUS 18 (8): 155-197 January, 1981

Chalcopterus cupreus (F.) (Erotylus)

= Ch. setoasus Blackburn

Chalcopterus smaragdulus (F,)

(Erotylus)

FAMILY LAGRITDAE

Lagria tomentosa F.

FAMILY ALLECULIDAE

*“ Homotrysis rufipes (F.) (Helops)

FAMILY OEDEMERIDAE

* Dohrnia tristis (F.) (Necydalis)

Selenopalpus cyaneus (F.) (Lagria)

+ = S. subviridus White

Sessinia lineata (F.) (Lagria)

FAMILY CERAMBYCIDAE

sub-family Prioninae

Toxeutus arcuatus (F.) (Prionus)

sub-family Lamiinae

Depsages solandri (F.) (Lamia)

Hybolasius cristus (F.) (Lamia)

Xylotoles griseus (F,) (Saperda)

Xylotoles lynceus (F.) (Saperda)

sub-family Cerambycinae

Coptomma variegata (F.) (Callidium)

Hesthesis variegata (F,) (Leptura) Type

lost

Navomorpha lineata (F,) (Callidium)

Navomorpha sulcata (F.) (Callidium)

Oemona hirta (F.) 3 (Saperda)

= O, villosa (F,) 2 (Saperda)

Pachydissus obscurus (F,) (Callidium)

Phoracantha semipunctata (F.)

(Stenocorus)

Zorion minuta (F.) (Callidium)

Lamia bidens F. Present status

unknown as type missing

FAMILY CHRYSOMELIDAE

sub-family Cassidinae

Aspidomorpha deusta (F,) (Cassida)

Aspidomorpha interrupta (F,) (Cassida)

sub-family Chrysomelinae

Calomela crassicornis (F.) (Chrysomela)

Chalcolampra octodecimguttata (F.)

(Chrysomela)

Paropsis s.g. Chrysophtharta detrita (F.)

(Coccinella)

= P. laesa Germ,

Paropsisterna moria (F,) (Chrysomela)

Phyllocharis cyanicornis (F.) (Chry-

somela)

Phyllocharis cyanipes (F.) (Chrysomela)

Phyllocharis nigricornis (F.) (Chryso-

mela)

sub-family Criocerinae

Crioceris nigripes F.

Lema bifasciata (F.) (Crioceris)

Lema aculata (F.) (Crioceris)

Lema unifasciata (F.) (Crioceris)

-b

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 161

sub-family Eumolpinae

Encolaspsis brunnea (F.) (Chrysomela)

Rhyparida didyma (F.) (Crypto-

cephalus)

sub-family Galerucinae

* Monolepta cyanocephala (F.) (Crioceris)

+ = M, antennalis Lea

Synodita melanocephala (F,) (Crioceris)

sub-family Halticinae

Lieyllus 4-punctata (F,) (Galleruca)

= Altica albicollis F.

ANTHRIBIDAE

Phloeobius griseus (F.) (Anthribus) Type

missing

> Ptinus gigas F.

BELIDAE

Agathinus tridens (F.) (Curculio)

Belus semipunctatus (F.) (Curculio)

BRENTHIDAE

Lasiorrhynchus barbicornis (F,) (Cur-

culio)

= Curculio assimilis F.

FAMILY CURCULIONIDAE

sub-family Aterpinae

Aterpus cultratus (F.) (Curculio)

Chrysolopus spectabilis (F.) (Curculio)

sub-family Brachydeérinae

Protomus scutellaris (F.) (Curculio)

sub-family Cossoninae

* Cossonideus lineolus (F,) (Curculio)

sub-family Cryptorrhynchinae

Euthyrhinus meditabundus (F.) (Cur-

culio)

Onmtydaus luridus (F.) (Curculio)

* Tentegia stupida (F.) (Curculio)

= Rhynchaeus strepidus F.

* = T. ingrata Faust

Curculio cruciatus F, Present

unknown

+ = Acalles doriae Pasc.

sub-family Curculioninae

Curculio amoenus F.

Orthorrhinus cylindrirostris (F.)

(Curculia)

sub-family Erirrhininae

Nyxtes bidens (F,) (Curculio)

Stephanorrhychus attelaboides (F.)

(Curculio)

Tysius bicornis (F,) (Curculie)

sub-farmly Haplonychinae

“ Haplonyx haemorrhoidalis (F.)

(Curcullo)

sub-family Leptopiinae

Catoptes interruptus (F.) (Curculio)

Leptopius clavus (F.) (Curculio)

Leptopius quadridens (F.) (Curculio)

Leptopius tribulus (F.) (Curculio)

Stenocarynus crenulatus (F.) (Cureulio)

FAMILY

status

sub-family Otiorrhynchinae

* Oribius adspersus (F.) (Curculio)

sub-family Rhadinosominae

Rhadinomus acuminatus (F.) (Curculio)

sub-family Rhynchophorinae

Cureulio bituberculatms F. is a synonym

of Sitophilus oryzae Linné

sub-family Rhyparosominae

* Cecyropa modesta (F.) (Curculio)

The present status of the following species has not

yet been determined:—

* Rhynchaenus curvirostris F, (Curculia)

* Rhynchaenus exclamationis F. (Curculio)

DISCUSSION OF SPECIES

ABBREVIATA, Nitidula

Fabricius 1781, p. 91, n, 3

Junk’s Catal. 1913, 15, 56, p. 155 (Aethina)

Zimsen 1964, p. 81, n. 1138

Holotype labels:

i. Fabricius ‘‘Nitidula abbreyiata Fabr, Spec.

Ins. n. 3”

ii. BMNH ‘“‘Nitidula abbreviata’’

Locality: “nova Zelandia”’. It is doubtful that this

locality is correct,

Current status: Macroura abbreviata (F.):

Nitidulidae.

Comment: Except for the mesothoracic legs, mast

of which are missing, the specimen is in fairly good

condition.

This species belongs to the genus Macroura and

not to the genus Aethina in which it has previously

been placed. The genus Macroura is represented by

species from India, East Indies, Australia and New

Guinea. It is, therefore, possible that if the locality

New Zealand is incorrect that it should be one of the

above localities where Banks collected. The locality

New Zealand has been queried by Broun (1880: p.

171) who stated “I have not seen any insect agreeing

with Fabricius’s description”, also by Sharp (1891: p.

350) and by Grouvelle (1912; p. 398), In Grouvelle’s

table of the genus (1912: p. 574), it is very close to

M, javanica Grouy. and M. orientalis Nictur.

ACUMINATUS, Curculio

Fabricius 1775, p. 152, n. 132; 1801, 2. p. 535, n. 168

Junk’s Catal. 1937, 28, 154 p. 1 (Rhadinosomus)

Zimsen 1964, p. 219, n. 3781

Kuschel 1970, p. 203

Lectotype (Kuschel) labels:

i. Fabricius ‘Cure, Acuminatus Fab. Entom-

p. 182 n. 132"

142 REC. $8. AUST, MUS [8 (&): 155-197

ii. BMNH “Curculio acuminatus”, ‘‘Type”’

Locality: “nova Zelandia”

Current status: Rhadinosamus acuminatus (F.):

Curculionidae, Rhadinosominae.

Comment: Lectotype is a male. It is very dirty and

in poor condition, pinned through left elytron part of

which is missing, right elytron and left meta-thoracie

leg are also missing, There is a second male

specimen, a paralectotype, which is very badly

damaged, head, prothorax and right metathoracic

leg are missing,

Broun (1880: p. 430), following Lacordaire (1866:

p. 63) attributed this species to Schdénherr, since

then the correction has been made. Kuschel

designated the lectotype.

ADSPERSUS, Ctirctilio

Fabricius 1775, p. 149, n. 118; 1801, 2, p. 329, a.

131.

Zimsen 1964, p. 217. n. 3752,

Holotype labels:

i, Fabricius ‘Curc. Adspersus Fab. Sp. Ins.

n. 165.

ii, BMNH “Curculio adspersus”.

Locality: ‘nova Hollandia”’. This species may

possibly be from New Guinea or the East Indies.

Current status: Oribius adspersus (F.) Cur-

culionidae, Otierrhynchinae-

Comment: The holotype is in poor condition,

head and prothorax have been broken off and glued

on to the rest of body: left antenna, tarsi of

prothoracic legs, both mesothoraci¢ legs and right

metathoracic leg are missing. C.O. Waterhouse did

not identify this specimen as a “type”. There has

been a great deal of contusion in the literature about

this species.

Zimsen records three specimens under the name

Curculio adspersus, namely the numbers 3473, 3638,

and 3752.

Number 3638:—The type is recarded as “in

Cajenne Dom.V.Rohr” with the reference Mant

Ins. 1, p. 106. It is listed in Junk’s catalogue as

lleomus mucoreus L. This species does not concern

the Banks Collection.

Number 3473:—This was named by Fabricius as

Rhychaenus pallux, which is now recorded by both

Zimsen, and Csiki in Junk’s catalogue 1934, 29, 137,

28 as a synonym of Hypera (Erirhinomorphus)

adspersus (F.). This is not the species in the Banks

Collection and should not be known by the specific

name of adspersus as it is a jumior homonym, but by

the name given hy Fabricius in 1801, 2, p. 457, 0. 94,

namely R. pollux,

Number 3752:—Curculia adspersus F. is recorded

by Zimsen and as “in nova Hollandia in Dom.

January, IYS1

Banks—London 1 specimen”, and gives the original

reference of 1775. This species ts not listed by Csiki

in Junk's catalogue, but he does give two references,

namely Systema Eleutheratorum 2, 1801, p, 529, n.

131, and Herbst, Nat, Ins. Kal. 6, 1795, p. 258 for

the species H. (Erirhinomorphus) adspersus (F.)

which is incorrect. These two references are for the

Fabrician species Oribius adspersus (F.), number

3752 in Zimsen, It was classified into the penus

Oribius by R. T, Thompson of the BMNH (pers.

comm.), O. adspersus (F.) is the senior homonym,

The other two species which Fabricius names

Curculio adspersus were renamed by him in Syst. El.

as Rhynchaenus pollux and Lixius roreus, and these

specific names should now be used for them. L

roreus F, is a synonym of Ileomus mucoreus L,

AENEA, Melolontha

Fabricius 1775, p. 34, n, 11; 1801, 2, p. 166, n 30

aeratus Linne 1790, p, 1570 (Scarabaeus)

purpureipes Macl. 1871, p. 197 (Repsimus).

Junk’s Catal. Suppl. 66 p. 285 (Repsimus).

Zimsen 1964, p, 144, n. 2360.

Carne 1958. p, 178-9.

Holatype labels:

ij. Fabricius “Melal. aenea Fabr. Sp. Ins. n-

iii BMNH ‘“‘Melolontha Aenea”, “Type”.

Locality: “nova Hollandia".

Current status; Repsimus aeneus (F.);

Scarabaeidae, Ruterlinae.

Comment: The condition of the holotype is good,

except that the mouth parts are covered with dried

mycelial hyphae. This species is the type of the genus

Repsimus. The reference on the label is not the

original reference. This probably accounts for the

fact that Fabricius would have re-examined this

species before publishing Spec. Ins. 1781 and when

compiling this list put this reference on the specimen

before actual publication as there is no page

reference given. Carne (idem) discusses the

synonymy of this species in full.

ALBICOLLIS F., Altica

See QUADRIPUNCTATA F., Galleruca

A. albicollis F, is the junior homonym,

AMETHYSTINUS, Erotylus

Fabricius 1775, p. 124, n. 7; 1801, 2. p- 13, n. 6

(Cnodulon)

Junk’s Catal, 1911, 18, 28, p. 579 (Chalcopterus)

Zimsen 1964, p. 116, n, 1806

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

Holotype labels:

i. Fabricius “‘Erot, Amethystinus

Entom. p. 124 n, 7”

ii; BMNH "Cnodulon amethystinus"’,

“Type”

Fab.

Locality: “nova Hollandia”’

Current status: Chalcopterus amethystinus (F.);

Tenebrionidae, Amarygminae

Comment: The condition of the holotype is good,

though the head is retracted somewhat into the

prothorax, parts of both antennae are missing.

The spelling in the 1775 reference is Erotylus

amethystines though on the label the specific name is

spelt amethystinus, So the ending “‘es" is probably a

misprint.

Blair (1914; p. 489) studied this species in relation

to the specimens identified by Blackburn and Carter

as Ch. amethystinus (F.). He points out the

differences between these specimens and the

Fabrician type.

“|. In the type, the legs are entirely black, in

the other specimens the femora is red and

sometimes there are traces of red coloura-

tion on the tibiae.

2. In the type there is a small but distinct

depression in the centre of each half of the

prothorax, this is absent in the other

specimens.

3. The anterior margin of the prothoran is

slightly sinuate in the type but not so in the

other specimens.

4. Punctures of the elytra are heavier in the

type than in the other specimens.”

Blair considered that the Fabrician species is

nearest to Ch. pulcher Blkb. but is “not identical

with it’. In his opinion the specimens identified by

Blackburn and Carter (1913: p. 10) as Ch.

amethystinus F, are not the Fabrician species.

‘This is another species Fabricius reclassified in his

later publications.

AMOENUS, Curculio

Fabricius 1775, p. 142, 0. 81: L801, 2, p. 489, n. 239

(Rhynchaenus)

Junk’s Catal. 1923, 29, 123,

Zimsen 1964, p. 208, n. 3582

Holotype labels:

i, Fabricius "Cure. Amoenus Fabr. §. Ent. p,

142, 0. 81"

i. BMNH “Rhynchaenus amoenus”, “Type”

Locality: “nova Hollandia”

Current status: Cureulid amoenus F,:

Curculionidae, Curculioninae.

Comment: Holotype is in good condition. all

structures are complete. At one time this species was

placed in the genus Balaninus but is now back in is

original genus Curculio,

ARCUATUS, Prionus

Fabricgius 1787.1, p. 129, 0.9; 1801, 2, p. 259, n. 10

eurvous Gmel. 1789, p. 1817. After McKeown

1947

Junk's Catal, 1913, 22, 52, p. 38

Zimsen 1964, p. 163, n. 2717

Holotype labels:

i. Fabricius “Prionus Arcuatus vy. Diem. land

Nelson Fabr. Mss”’.

ii. Fabricius “Prionus arcuatus Fabr. Mant.

Ins, n, 9”

iii, BMNH ‘Priomuas arcuatus”, “Type”

Locality: ‘“‘Terra Diemenii"'

Current status: Toxeufes arcuatus (F.): Ceramby-

cidae, Prioninae.

Comment: Holotype has been pinned twice, now

through the right elytron, previously through the

prothorax; terminal joints of left antenna and some

tarsi of mesothoracic legs ure missing.

Lameere (1904 p. 24) studied the holotype and

gives a detailed description of it.

The name Nelson on the label, is that of the

collector who sailed with Cook on his third voyage,

and collected at Adventure Bay, Tasmania.

ASSIMILIS, Curculio

Fabricius 1775, p. 134, n. 42; 1801, 2 p. 546, n, 3

(Brentus)

Junk’s Catal. 1927, 26, 89, p. 58 (Lasiorhynchus)

Zimsen 1964, p. 221, n, 3833

Kuschel 1970, p. 194

Lectotype (Kuschel) label;

i. Fabricius ‘Brent. Assimilis Fabr. M. Ins.

n. 9"

ii. BMNH *'Brentus assimilis”, ““Type”’

Locality: “nova Zelandia”’

Current status: synonym of Lasiorrhynchus

harbaricornis (F.)

Comments: “Type” is a female, pinned through

left elytron which has been glued on to specimen,

Otherwise specimen is in good condition, except for

terminal joints of both antennae which are missing,

The reference number 9 on the label is incorrect, it

should be number 2, Kuschel (idem,) designates this

specimen as the lectotype and the specimen in

Fabricius'’s own collection as a paralectotype,

ATTELABOIDES, Curculio

Fabricius 1775, p, 156, n. 162; 1801, 2, p, 545, n. 227

osculator Broun 1909, p. 134

Junk's Catal. 1934, 28, 140, p. 81 (Stephanorrhyn-

chus)

164

Zimsen 1964, p. 221, n. 3832

Kuschel 1970, p. 199

Holotype labels:

i. Fabricius “Curc. attelaboides Fab. Entom.

156 n. 152

ii. BMNH “Cureulio attelaboides”

Locality: ‘Brasilia’. This locality is incorrect. [tis

a New Zealand species.

Current status: Stephanorhynchus atielaboides (F.)

Curculionidae, Erirrhininae

Comment: Holotype is a female. This. species is

endemic to New Zealand and was identified in the

Collection by Kuschel (1970; p. 191) as being

erroneously recorded from Brazil, Kuschel (1970: p.

199) gives a key to the indentification of this species

and its closely related species.

AUSTRALIS, Gyrinus

Fabricius 1775, p. 235, n. 2; 1801, 1, p. 275, n. 3

rufipes F. 1801, 1, p. 276, n. 13

dentipennis Macl, 1825, p. 30

limbatus Mae}. 1825, p. 30

iridus Hope 1842, p. 428

leucopoda Montr, 1860, p. 245 (Dineutus)

janthinus Reg., 1882, p. 421 (Dineutes)

dentatus Sutfr, 1842, p. 256 (Dineutes)

Leucopus Montr. 1860, p. 245 (Dineutes)

Junk'’s Catal. 1910, 4, 21, p. 4 (Dineutes)

Zimsen 1964, p. 70, n. 927

Ochs 1949, p. 192

Holotype labels:

i. Fabricius ‘‘australis’’

ii. BMNH “Gyrinus australis’’ museum regis-

ter number 63-46.

Locality: ‘fin novae Hollandiae aquis”’

Current status: Dineutus australis (F.); Gyrinidae,

Comment: The holotype has been broken

between the meso and metathorax and then glued

together. The full list of synonyms and literature

concerning this species is given by Ochs (idem.).

Fabricius described the female of this species at a

later date under the name of G., rufipes F.. which is

now a synonym, the type of which is in ‘“‘Dom

Billardiae". D. dentatus Suffr and D, leucopus

Montr. are not listed by Ochs as synonyms as they

are not Australian species.

AUSTRALIS, Lampyris

Fabricius 1775, p. 201, n. 11; 1801, 2, p. 104, n, 23

Junk’s Catal, Suppl. 1966, 9, p. 99 (Luciola)

Zimsen 1964, p. 133, n, 2141

REC, 8. AUST. MUS 18 (8): 155-197

January, 198]

Holotype is missing from the Collection, “Type” has

recently been found in the Hunterian Collection

in Glasgow.

Locality: “nova Hollandia”

Current status: Luciola australis (F.) Lampyridae.

Comment: In Gemminger et Harold Catalogue

this species is recorded under the name Luciola

australis (F.) and the locality is erroneously given as

“Noy. Island” (New Ireland). Boisduval (1835; p.

125) gives the same locality. However, Masters (p.

312) and MeDermot in Junk's Suppl (idem.) records

the Fabrician locality, ‘Australia’. Masters records

two synonyms L, guerini Laporte and L. nigripennis

Latr. McDermot states that L. guerini Laporte is

from New Guinea and is not a synonym of L.

australis (F.). L. nigripennis Latr is not listed by

MeDermot and Lea (1909: p,108) regards L.

nigripennis Latr as merely a “catalogue name’.

Therefore there is no recognised synonym of L.

australis (F.).

BARBAROSSA, Scarabaeus

Fabricius 1775, p, 12, n, 35: 1801, 1, p. 47

(Geotrupes)

var similis Prell 1934, p. 57.

Junk’s Catal. 1937, 21, 156, p. 93 (Haploscapanes)

Zimsen 1964, p. 22, n. 28 (Geotrupes)

Carne 1957, p. 190

Syntypes labels;

i. Fabricius “Scarab. Barbarossa” Fabr, Sp

Ins, No. 41.

ii. BMNH “Geotrupes barbarossa’”

Locality; ‘Nova Hollandia”’

Current status: Haploscapanes barbarossa (F,),:

Scarabaeidae, Dynastinae.

Comment: There are two specimens, a male and

female. The condition of the male is fair; only two

basal joints of the antennae are present and the

terminal joints of the left labial palp and most of the

tarsi are missing. The female is an exceptionally

large specimen and is in good condition, The

reference on the label is the 1781 reference so

Fabricius must have re-examined this specimen in

1780-1.

BARBICORNIS, Curculio

Fabricius 1775, p. 134, n. 41; 1801, 2, p, 545, n. |

(Brentus)

assimilis F. 1775, p. 134

Junk’s Catal. 1927, 26, 89, p, S& (Lasiormhynchus)

Zimsen 1964, p, 221, n, 3833

Kuschel 1970, p. 193-5

Lectotype (Kuschel) labels:

i. Fabricius “‘Brentus Barbicornis Fabr.

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

Mant, Ins, n. L"

ij BMNH ‘Brentus barbicornis’’, “’Type"’

Locality: “nova Zelandia™

Current status: Lasiorrhynchus barhicornis (F.):

Brenthidae

Comments: There are two specimens both males.

The “type” specimen designated by Kuschel as the

lectotype is very dirty and terminal joints of lett

antenna are missing. In the 2nd specimen, a male, a

paralectotype (Kuschel) head is missing. Broun

(1880: p. 543) notes that there is great variation in

length of this species. The reference on the Jabel

indicates that Fabricius must have re-examined this

species after the original 1775 publication and before

the Mantissa Insectorum was published as there is no

page number given, the page number is 95.

BICOLOR, Erotylus

Fabricius 1775, p, 124, n, 8; 1801, 2, p. 14, n. 7

(Cebrio)

tardus Blk. 1889 p. 1271

var torridus Pasc. 1869 p. 351

Junk’s Catal, 1911, 18, 28, p, 578 (Amarygmus)

Zimsen 1964, p, 116, n, 1807

Carter 1926, p. 158

(Amarygmus)

Holotype labels;

i, Fabricius “Erotylus Bicolor Fab. Entom.

p. 124 n. 8”

ii, BMNH “Cnodulon bicolor’, “Type”

Locality; “nova Hollandia”

Current status: Amarygmus bicolor (F.): Teneb-

rionidae, Amarygminae

Comment; The holotype is in a poor state of

preservation. The ventral surface is covered with dirt

and mycelium, except for the terminal joints of the

right antenna all structures appear to be present but

are difficult to discern, Carter (1915: p. 33) discusses

the synonym of this species fully, This reference

includes a copy of the correspondence between

Carter and Blair of the BMNH who compared the

Fabrician type With the type of A. tardus Blkb.

which is in the BMNH, Fabricius changed the genus

of this species in his later publication. This species is

recorded in Junk’s Catalogue (idem.) as a queried

synonym of Chalcopterus affinis Bless. This is not

accepted by Carter (idem.).

BICORNIS, Curculio

Fabricius 1781, 1, p. 180, n. 111; 1801, 2, p. 489, n.

241 (Rhynchaenus)

amplipennis Pasc. 1875, p. 218

purus Broun 1893, p. 1224 (Tysiius)

Zimsen 1964, p. 208, 108, n. 3584

Kuschel 1970, p. 202

165

Lectotype (Kuschel 1970) labels:

i. Fabricius “Cure, Bicornis Fabr. Sp. Ins. n.

ya"

ii. BMNH “Rhynchaenus bicornis’, “Type”

Locality: “nova Zelandia”

Current status: Tysius bicornis (F,): Cur-

culionidae, Erirrhininue

Comment; The lectotype is badly damaged; head,

tarsi of right prothoracic leg, Jeft prothoracic leg

except for coxa and trochanter, and right

metathoracic leg are missing. This species was for

sometime omitted from the catalogues. G. Kuschel

places this species in the genus Tysius, He regards

this “type”, a male, as the lectotype although the

Fabrician reference says “Mus. Dom. Banks’’ and

there is only one specimen in the collection. There is

a female in Fabricius’s own collection which he

names as a paralectotype.

BIDENS, Lamia

Fabricius 1775, p. 177, n, 30; 1801, 2, p. 304, n. 124,

Junk’s Catal. 1923, 23, 74, p. 604.

Zimsen 1964, p. 172, n. 2890.

Labels of specimen in collection;

i. Fabricius “Lamia Bidens Fab. Entom.

p. 177 n. 30",

ii. BMNH “Lamia bidens’’. This label and

the ‘Type label” have been crossed out

and the word “Bidens” queried.

iii, A mutilated label—'‘an bidens fabricii?"’

Loeality: “nova Hollandia”.

Comment: This specimen cannot be the **Fabri-

cian type of L. bidens. It is in good condition but has

been badly pinned. I have been unable to identify it.

Aurivillus (Junk's Catal. idem.) and McKeown

(1947; p. 174) both list this species among those

‘doubtful or unknown” genera. Oliver (1789: p. 67

Fab. 17 fig. 125) figures L. bidens F. as a small

narrow insect with the apices of the reddish-brown

elytra distinctly bidentate.

BIDENS, Curculio

Fabricius 1775, p. 136, n. 51; 1801, 2, p. 457, n. 96

(Rhynchaenus)

rufipes Broun 1881, p. 718 (Nyxetes)

Junk’s Catal. 1934, 28, 140, p. 84 (Nyxetes),

Zimsen 1964, p. 203, n. 3474.

Kuschel 1970, p. 201,

Lectotype labels:

i. Fabricius ‘Cure,

p. 136 n, 51".

ii. BMNH “‘Rhynchaenus”, ''Type”

Bidens Fab. Entom.

Locality: “nova Zelandia"’.

166

Current status: Nyxeles bidens (F.): Cur-

culionidae, Erirrhininae.

Comment: Glue is present on both dorsal and

ventral surfaces; spine-like clusters of hairs on right

elytron are broken, abdomen twisted and lying at

right angles to the rest of body; antennae, most tarsi

and right metathoracic leg are missing: The colour of

this species varies from reddish brown to black,

“type” is black. Fabricius changed the genus in his

later publications. Kuschel designated this specimen

as a lectotype.

BIFASCIATA, Crioceris (Lema)

Fabricius 1775, p. 120, n. 12; 1801, 1, p. 476, n_ 29

(Lema).

Junk's Catal, 1924, 24, 51, p. 55 (Lema).

Zimsen 1964, p. 107, n. 1645,

Syntype labels:

i. Fabricius “Crioc, bifasciata Fab. Entom,

p. 120, n. 12”.

ii. BMNH “Lema bifasciata’’, ‘’Type"’.

Locality: “nova Hallandia”,

Current status: Lema hifasciata

melidae, Criocerinae.

F.: Chryso-

Comment: The condition of the “type” is fair,

parts of the antennae and tarsi of right metathoracic

legs are missing. The second specimen is not in as

good a condition as the “type". and it is pinned

through the left elytron, Fabricius re-classified this

species in 1801 into the genus Lema.

BIPUSTULATUS, Scarabaeus

Fabricius 1775, p, 30, n. 121; 1801, 1, p. 62. n 37

(Ateuchus)

Junk's catal. 1927, 19, 90, p, 209 (Onthephagus)

Zimsen 1964, p. 31, n. 188

Matthews 1972, p, 220 (Onthophagus)

Holotype labels:

i, Fabricius “Scarab. 2-pustulatus’’ Fab. Sp.

Ins. No. 152

ii. BMNH *‘Ateuchus bipustulatus”’

Locality: “‘nova Hollandia”

Current status; synonym of Onthophagus quad-

ripustulatus (F.),

Comment: Holotype is badly pinned through the

left elytron. Several legs are without tarsi and the

club of the left antenna jis missing. O, bipustulatus

(F.) is a female and the synonym has been

established by Matthews (idem).

BITUBERCULATLUS, Curculio

Fabricius 1781, 1 p. 171, n. 58; 1801, 2, p. 438, n, 42

Junk’s Catal. 1936, 30, 149, p. 111

REC §. AUST, MUS 18 (8): 155-197

January. [98]

Zimsen 1964, p. 199, n. 301

Kuschel 1970, p. 197

Holotype labels:

i, Fabricius “Curc. 2-taberculatus Fabr. Sp-

Ins. n. 58"

ii. BMNH “Calandra 2-tuberculata’’, “type”

Locality: “nova Zelandia”

Current status: C. bituberculatus F. is a synonym

of Sitophilus oryzae Linné: Curculionidae, Rhyn-

chophorinac.

Comment: The holotype is badly pinned, head

and most tarsi are missing. Oliver (1807: p. 954. [67)

describes this species, stating that the body ts a

ferrugineus uniform colour and without marks, and

that the specimen in the Bank's Collection is without

any raised tubercles on the prothorax as described

by Fabricius. Herbst (1795: p. 29) and Broun (1880:

p, 504) both note the same absence of tubercles.

Kuschel (1970; p, 197) makes no reference to this.

Kuschel determined the synonymy. He also explains

the confusion caused in identifying this species as it

had been erroneously associated with the weevils

Dryophthorus ecrenatus Boisd. and Mitrastethus

baridioides Redtenbacher.

BRUNNEA, Chrysomela

Fabricius 1781, 1 p. 123, n, 44; 1801, 1, p. 439, n, 104

Junk’s Catal, 1914, 24, 59, p. 22

Zimsen 1964, p, 100, n. 1504

Holotype labels:

i. Fabricius ‘Chr, Brunnea Fabr, Sp, Ins, n,

44"

ii, BMNH “Chrysomela brunnea”, Type”

Locality: “nova Zelandia''—Zimsen (1964) and

Musgrave (1952) record the locality as ‘tnova

Hollandia’” as given by Fabricius in his 178]

publication, In his later publications of 1792 & 1801

Fabricius corrects this and gives the correct locality

as “nova Zelandia”.

Current status: Eucolapsis brunnea (F.): Chry-

somelidae, Eumalpinae,

Comment: The condition of the holotype is poor,

and the structures on the ventral surface are covered

with glue and dirt. Parts of both antennae and parts

of all legs are missing. There are several colour

varieties of this species. The holotype is reddish and

there is no trace of any green colour, as on some

specimens, this may be due to fading. This species is

described by Broun (1880; p. 622) and his account

corresponds very closely to it, and the three

specimens in his own collection are identical with the

“type”, Broun records the habitat as ‘‘New Zealand

on the flowers of Leptospermum"’. Musgrave (1932:

p, 87) incorrectly records the Fabrician genus as

being Coccinella.

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

CANCROIDES, Lucanus

Fabricius 1787, l, p. 2, n. 9; 1801, 2, p. 252, n. 18

Junk's Catal, 1960, 8, p. 39

Zimsen 1964, p. 162, n, 2071

Holotype labels:

i, Fabricius ‘Lucan,

Mant, Ins, n, 9"

li, BMNH “‘Lucanus Crancroides”, ''Type™

Crancroides Fabr.

Locality; “Terra Diemenii”

Current status: Lissetes crancroides (F.);

Lucanidae.,

Comment: Holotype has been pinned twice, now

through the right elytron but previously through the

prothorax: mouth parts and tarsi covered in dirt;

tarsi of right prothoracie and left metathoracic leg,

and right mesothoracic leg are missing,

Westwood (1871: p, 371) gives a full description of

this species. and compares it with closely allied

species.

CARNIVORUS, Dermestes

Fabricius 1775, p. 55, n. 2, (D. carniforus); 1801, 1,

p. 31,22

Junk’s Catal. 1911, 14, 33, p. 42

Zimsen 1964, p, 64, n. 1048

Hinton 1945, p. 287

Locality: ‘tin nova Hollandia et Zelandia™ in 1775.

“in nova Hollandia et in Germania" in

Ta01,

Current status: Dermestes carnivorus F.: Dermes-

tidae

Comment: Fabricius changed the ‘‘habitat” in

1801 and added that the “type’’ was in ‘Dom

Smidt”, having previously in 1775 stated it was in

“Mus. Dom. Banks”. The spelling in 1775 of the

species as “carniforus’’ is possibly a misprint as in all

later publications it is spelt carnivorus. Hinton

records the distribution of this species as ““N. & S.

America, Europe and India’. According to Fauvel

(1889) it is indigenous to America, It seems probable

that this insect may have been found on the

Endeavour, having been brought aboard with stores,

thus giving rise to the confusion of its place of origin,

This then is not an Australian or New Zealand

species,

CLAVUS, Curculio

Fabricius 1775, p. 154, mn. 140, 1801, 2, p,

elegans Lea 1916 p. 332

Junk’s Catal, 1931, 28, 114, p. 18 (Leptops)

Zimsen 1964, p. 219, n, 3789

536,n. 177

Holotype Jabels:

i. Fabricius “Curcul.

p, 154 n, 140'

Clavus Fab. Entom.

167

ii. BMNH “Curculio clavus’’, ‘“Type”,

Locality: “nova Hollandia”

Current status: Lepropius (Oke 1951) clavus (F.):

Curculionidae, Leptopinae.

Comment: Holotype is pinned through the left

élytron, tarsi of left prothoracic leg and right

mesothoracie ley are missing. This species varies in

colour and intensity of red markings and in the

number of scales present.

COLLARIS, Tritoma

Fabricius 1775, p. 69, n, 2

bicolor Carter (Sclerocyphan) new synonym,

Zimsen 1964, p. 237, n. 4109

Holotype labels: Fabricius ‘'Tritoma Collaris Fab

Entom, 69 n, 2”

Locality: “nova Hollandia"

Current status: Sclerocyphon collaris (F.);

Psephenidae,

Comments: There is no BMNH label. This

speciment was added at the end of the named

specimens in the collection after it was found in 1964

by E. Zinsen “in the last box’ of the collection,

which consists of unnamed specimens.

The condition of the type is very poor, The

antennae and the abdomen are missing. The right

side of the prothorax 1s broken, but the broken piece

is lying on top of the prothorax. [t is badly pinned

and the right elytron is cracked, There is a large

lump of glue on the ventral surface and the insect is

covered with dried mycelial hyphae and debris.

This species has not only been omitted from the

catalogues but Fabricitis omitted it from his 1801

publication, the Systema Eleutheratorum, It is listed

in bis Species Insectorum and in Mantissa

Insectorum., However, the omission from the

Eleutheratorum may account for the fact that it was

not found in the arrangement of the Banks

Collection which was based on the order given in

that publication, While I was studying the collection

R. D. Pope of the BMNH classified this specimen as

a member of the family Psephenidae and as

belonging to the genus Sclereeyphon, (pers. com.)

While at the BMNH [ compared it with H. J,

Carter's (1935: p. 191) description of S. bicolor. This

described the Fabrician species, T. collaris F, There

was no Speciment of S. bicolor Carter in the BMNH,

so no comparison could be made, On returning to

Australia. ] was eventually able to track down a

paratype of S. bicolor Carter at the Zoology

Department at the University of Tasmania, where

Mrs, J, Smith is making a study of the genus

Sclerocyphon, The specimen was sent to R, D, Pope

at the BMNH and he made a comparison of the two

168

species, and agrees that S. bicolor Carter is a

synonym T. collaris F. In a letter received from Mr.

Pope he states: “I have compared the Carter

paratype with the specimen in the Banks Collection

and am quite sausfied that they are the same species.

The overall size, pronotal colour pattern and shape,

elytra punctation and colour all agree well. The

teeth on the hind margin of the pronotum from

margin of the scutellum and basal borders of the

elytra all correspond in size and distribution. The

colour pattern and general morphology of the head

also agree”.

CRASSICORNIS, Chrysomela

Fabricius 1775, p. 99, n. 27; 1801, 1, p. 437, n. 94.

Junk’s Catal. 1916, 24, 68, p. 193 (Calomela),

Zimsen 1964, p. 99, n, 1496,

Holotype labels;

i. Fabricius “Chrys. crassicornis Fab. Entom

p. 99, 27,

ii. BMNH “Chrysomela

“Type”.

crassicornis”’

Locality: “nova Hollandia’”’.

(F.);

Current status: Calomela crassicornis

Chrysomelidae, Chrysomelinae.

Comment; The holotype is not in good condition.

Glue on the ventral surface has covered the

metathorax and abdomen; lelt mesothoracic and

right metathoracie legs are missing. Baly (1855;

p. 249) gives a detailed description of this species.

However, the black markings on the “type” differ

somewhat from what Baly describes as typical of the

species. The general colour is fulvous, the “type”’

being paler than most specimens in the BMNH, but

this may be due to fading. The “type” differs from

Baly’s description in the following respects:—

1. no dark frontal patch on head

2. no black marking on prothorax

3. markings on each elytron are a_ single

rounded black basal spot, and an elongated

black streak posterior to it. Baly describes

the elytral markings as a “‘subtriangular

‘spot near the scutellum and a sinous vilta

.. extends nearly to the apex of the

elytron”.

CRENULATUS, Curculio

Fabricius 1775, p, 147, n. 105; 1801, 2, p. 518, n. 64,

australis Boisd. (Gastrodus),

Junk’s Catal, 1913, 28, 114, p. 16.

Zimsen 1964, p. 214, n. 3696.

Holotype labels:

i. Fabricius “Cure, Crenulatus Fab, Entom.

p. 147, n. 105”.

ii. BMNH ‘Curculio crenulatus"’, *“Type’’.

REC, S, AUST. MUS t& (8); [55-197

January. S98T

Locality: “nova Hollandia™,

Current status: Stenocorynus crenulatus (F.):

Curculionidae, Leptopiinae.

Comment: Holotype is pinned through left

elytron; terminal joints of both antennae and tarsi of

left metathoracic leg are missing, it is also very

abraded only a few scales are present.

CRISTA, Lamia

Fabricius 1775, p. 170, n. 3; 1801, 2, p. 282, n. 6.

Junk’s Catal. 1922-23, 23, 74, p. 323.

Zimsen 1964, p. 167, n. 2787-

Holotype labels:

i, Fabricius ‘Lamia

p. 170 n. 3".

ii. BMNH “Lamia crista"’, *‘Type"’.

Crista Fab, Entom.

Locality: “nova Zelandia”™.

Current status: Flybolastus cristus (F.): Ceramy-

cidae, Lamiinae.

Comment: The holotype is in fairly good

condition; pinned through the left elytron which is

cracked; terminal joints of both antennae and tarsi

of right metathoracie leg are missing. Broun (1880:

p. 609-10) states that he examined the Fabrician type

and noted that Fabricius made an error when he

described the basal tubercles of the elytra as

tridentate, The mistake is due to the fact that the

tubercle is surmounted by “a compressed pencil of

hairs”.

CRUCIATA, Buprestis

Fabricius 1775, p. 222, n. 36; 1801, 2, p. 210, n. 134.

Junk’s Catal. 1935, 12, 143, p. 844.

Zimsen 1964, p, 155, n, 2568.

Carter 1923, p. 172 and 1929 p. 277.

Holotype labels:

i. Fabricius Buprestis cruciata Fab. Entom.

p. 222 n, 36".

i. BMNH “Buprestis cruciata’’, ‘Type’.

Locality; “nova Hollandia”’.

Current status; Cisseis s.g. Neospades cruciata

(F.): Buprestidae, Buprestinae.

Comment: The holotype is in good condition

except for a large amount of glue on the ventral

surface. Carter (1929: p. 277) classified this species

in the genus Neospades which is now a sub-genus of

Cisseis.

CRUCIATUS, Curculio

Fabricius 1775, p. 129, n. 8; 180], 2, p. 434, no. 23

(Calandra)

doriae Pasc, 1885 p. 257, new synonym

Junk’s Catal. 1936, 29, 151, p. 151

Zimsen 1964, p. 198, n. 3386

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 169

Holotype labels:

i, Fabricius "Cure, cruciatus Fabr. Entom.

p. 129 n, 8”

ii, BMNH “Calandra cruciatus”

li. “Crypto-rhynchus”’

Loeahty: “nova Hallandia”

Current status: unknown: Curculionidae, Cryp-

torrhynchinae.

Comment: All structures appear to be complete

though the legs are all tightly contracted and folded

inwards, there is a great deal of glue on the ventral

surface. There is no museum ‘type’ label on this

speciment although the label has the original

reference on it. In correspondence with R. J.

Thompson of the BMNH he sent the following

communication to me “‘I have compared the types of

cruciatus and Acalles doriae Pasc, and confirm that

they belong to the same species, Although doriae

stands in Ophrythyreoces in Junk’s Catalogue

(Hustache 1936), it certainly does not belong there.

In his description of Ophrythyreocis, Lea draws

special attention to the ‘‘conspicuously elevated

scutellum” (1913: p. 245) but in doriae the scutellum

is absent. Lea (idem, p. 247 note) placed doriae in

Pseudoporopterus, along with P, irrasus (idem,

p. 246). Although doriae and irrasus are probably

congeneric, [ doubt if they really belong in

Pseudoporepterus”.

CULTRATUS, Curculio

Fabricius 1775, p. 153, n. 136; 1801, 2, p. 536, n, 173

bicristatus F, 1801, 2 p, 517

Junk’s Catal. 1936, 28, 150, p. 3 (Aterpus)

Zimsen 1964, p. 219, n, 3785

Holotype labels:

i. Fabricius “'Cure. cultratus Fab. Entom,

p, 153 n. 136"

ii. BMNH "“Curculio cultratus”, “Type”

Locality; “nova Hollandia™

Current status; Aferpus cultratus (F.); Cur-

eulionidae, Aterpinae.

Comment: All structures are complete, but the

ventral surface is covered with dirt, This is a well

known species. Fabricius (1775: p. 153) describes the

elytra with ‘‘tuberculis sex v. septem elevatis”’ (six to

seven elevated tubercles). The holotype has only

five distinct tubercles.

CUPREUS, Erotylus

Fabricius 1775, p. 123, n. 5; 1801, 2 p. 12, n. 1

(Cnodalon)

venereus Gmel p. 1728 (Chalcopterus)

rusticus Blkb. Mon. p. 63, 76 (Chalcopterus)

setosus Blkb, by Blair (1914: p. 489) (Chalcop-

terus)

var cupriventris Carter (Chalcopterus)

Junk's Catal. 1911, 18, 28, p. 580 (Chalcopterus)

Zimsen 1964, p. 115, n. 1801

Carter 1926, p. 160

Holotype labels:

i, Fabricius ‘‘Cnodalon cupreum Fab. Ent.

p. 123 n, 8”

The original name of Frotylus cupreus

on the label has been changed to

Cnodalon cupreum in the same handwrit-

ing. After re-examination of mare

species Fabricius must have done this

himself, As his work progressed he often

re-classified species in new genera.

ii. BMNH ‘Cnodalon cupreus’’, “type”

Locality: “nova Hollandia’’

Current status: Chaleopterus cupreus (F.); Tene-

brionidae, Amarygminae.

Comment: The condition of the holotype is good

except for the head which is retracted within the

prothorax. Only the terminal joints of the left

antenna and terminal tarsi of left metathoracic leg

are missing.

The synonyms of this species are given by Carter

(idem,) as Ch, venereus Gmel and Ch, rusticus Blk.

H. J. Carter (1913: p. 14) had some correspondence

with K. G. Blair of the BMNH about this Fabrician

species. In this reference Blair’s description of the

Fabrician type is given, Blackburn’s type of Ch.

rusticus and Ch. setasus are both in the BMNH, Blair

(1914; p. 489) published that Ch. setosus Blk. is also

a synonym. Carter (1926: p, 160), who had not seen

the types still published them as separate species,

omitting Blair’s work on this species. The full

synonymy should be as above.

This species shows great variation in colour from

deep purple, almost black with a metallic tinge

through various shades of purple with a green

metallic tinge, to reddish copper, to an emerald

green. The holotype is black with thorax and elytra

coppery,

CURVIROSTRIS, Curculio

Fabricius 1781, 1, p. 166, n, 29; 1801, 2, p, 446, n. 40

(Rhynehaenus).

Zimsen 1964, p. 201, n. 3436,

Holotype labels:

i. Fabricius: *‘Curc. Curvirostris F, Sp, Ins.

N. 23". This number 23 is incorrect: it

should be 29,

ii, BMNH "Rhynchaenus— curvirostris™,

“Type”.

Locality: “nova Hollandia",

Current status; unknown. Family Curculionidae-

17u

Comment: The holotype is in poor condition;

head twisted through an angle of 90° to the left. Both

antennae and some of the legs are missing, and the

rest twisted and coyered with dirt,

R. curvirostris F. does not appear in any recent

catalogue. Olivier (1807: p. 152) described and

figures this species. As Fabricius studied more and

more species he realised that the large genus of

Curculio had to be divided up into separate genera.

This species was put in the genus Rhynchaenus in his

later publications.

CYANEUS, Hister

Fabricius 1775, p. 52, n, 3; 1801, 1, p, 86, n, 13

speciosus Boisd. 1835, p. 148

laetus Er. 1834, p. 179.

Junk’s Catal. 1910. 8, 24, p. 93 (Saprinus),

Zimsen 1964, p. 35. n. 253.

Labels; There are 3 specimens over the BMNH label

“Hister cyaneus”’.

1. i. Fabricius ‘“‘Hister cyaneus Nova Hol-

landia’”’.

ii. BMNH “Type”,

2. No labels.

3. Fabricius “Hister eyaneus C.B.S. Fab. Ent,

p- 52 n, 3”.

Locality of “Type™: “nova Hollandia”.

Locality of 3rd specimen: “C.B.S." (Cape of

Good Hope).

Current status; Saprinus cyaneus (F.): Histeridae,

Comment: Both labelled specimens are pinned

through the left elytron with a large amount of glue

on the ventral surface. The “type is complete

except for the tarsi of the right meta-thoracic leg.

The specimen labelled as from “C.B.S."" is in poor

condition with right antenna and most legs missing.

The unnamed specimen is the same species as the

“type”, so both specimens may be taken as syntypes.

The second specimen is in much the same condition

as the “type’’. These specimens were compared with

the large series in the BMNH collection of S,

cyaneus (F.). It is a species that varies a good deal in

size and colour from green to blue, also in degree

and form of striation on the elytra, In none of the

species examined was the sutural and second striae

united anteriorly, as they are in the specimen with

the “habitat” given as “C.B.8." (Cape of Good

Hope). This specimen was compared with the

specimens of S, bicolor (F.), the type of which jis in

the BMNH collection and the “C.B.S.". specimen

proved to be that species, The type of S. bicolor (F.)

of South Africa is given in Syst. El. 1801, 1 p, 86, n,

14 as being in Mus. D. de Schestedt, and the locality

is given as “C.B.S.” Jt appears then that this

specimen has been erroneously labelled Hister

cyaneus with the reference “Fab. Ent. p. 52 n. 3”

REC § AUST. MUS 18 (8): 155-197

January, L981

which is far H, cyaneus. The locality C.B.S. on the

label is correct for H. bicolor. There has abviously

been some mixing of material here. This specimen of

H. bicolor may be a syntype as the label is in

Fabricius’s handwriting-

CYANEUS, Lagria

Fabricius 1775, p. 125. n. 4: 1801, 2 p. 68, n. 5

(Dryops)

chalybeus White & (Selenepalpus)

subviridus White 2 (Selenopalpus) new synonym

Junk’s Catal. 1915, 17, 65, p. 24 (Selenopalpus)

Zimsen 1964, p. 127, n. 2026

Holotype labels;

i. Fabricius ‘‘Lagria Cyanea Fab, Entom, p.

125 n, 4"

ii, BMNH “Dryops cyanea”, “Type”

Locality; ‘nova Hollandia” is the locality given by

Fabricius and recorded as such by Zimsen. It is,

however, a New Zealand species (Broun 1880; p,

420).

Current status: (F,):

Oedemeridae

Selenopalpus cyaneus

Comment: Pin has forced the elytra apart; glue on

the ventral surface has entangled meso and

metathoracie legs; abdomen has been broken off and

glued on at a slight angle; joints of both antennae

and left maxillary palp are missing.

White's types of S. chalybeus and S, subviridus are

in the BMNH callection. 8. chalybeus is a male and

this synonymy was published by Broun (1880: p.

420), On examination S, subviridus white is found to

be the female of S. chalybeus White and is,

therefore, another synonym of 8. cyaneus (P.)

Schenkling in Junk's Catalogue published all three

as separate species, ignoring Broun’s publication,

Sclenopselophus is another generic synonym.

CYANEUS, Scarites

Fabrieius 1775, p. 249, n, 2; 1801, 1, p. 125, n. |S.

Jabriciti Westw 1842, 1, p. 8&5

var denescens Sloane 1923, p, 19.

Junk’s Catal. 1927, 1, 91, p. 451 (Lacecaprerurn)

Zimsen 1964, p. 41, n, 376

Holotype labels:

i, Fabricius “‘cyaneus”

ii. BMNH Scarites cyaneus

Locality: “nova Hollandia"

Current status: Laccepterum (F.):

Carabidae, Searitinae

cyaneum

Comments: The specimen is badly pinned through

the mid-dorsal line forcing the elytra apart. The

ventral surface is covered with glue. The left

THRE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 71

mesothoracic leg is missing, the metathoracic leg is

detached and lying in the glue.

CYANICORNIS, Chrysomelta

Fabricius 1775, p. 99. n. 24; 1801, 1 p, 436, n. &5-

Junk’s Catal. 1916, 24, 48%, p. 200 (Phyllocharis)

Zimsen 1964, p, 99, n. 1489

Holotype labels:

i, Fabricius “Chrys. Cyanicornis Fab.

Entom. p. 99”.

ii. “Chrysomela cyanicornis”, “Type”

Locality: ‘“‘nova Hollandia”’

Current status; Phyllocharis cyanicarnis (F.)

Chrysomelidae, Chrysomelinae

Comment: The holotype is pinned through the left

élytron, the posterior half of which is missing, only

the coxa and trochanter of left mesothorae leg is

present. There are several varieties of this species.

As in the other species of this genus there is

tremendous variation in colour patterns, This

holotype has a single central prothoracic spot and

eight spots distributed over the elytra, one basal spot

and two marginal ones on each elytron, and one

central and one triangular apical spot each with half

on each elytron. Lea (1902 p. 402) suggests that this

species is a variety ot P. cyanipes (F.). It needs a

specialist in this group to decide which are valid

species and which are varieties.

CYANIPES, Chrysomela

Fabricius 1775, p. 98, n. 23: 1801, 1 p. 436 n. 84,

Junk's Catal. 1916, 24, 68, p. 201 (Phyllacharis)

Zimsen 1964, p. 99 n. 1488

Holatype label:

i. Fabricius ““Chrys. Cyanipes Fab. Entom.

Pp. 98 n. 23".

ii, BMNH “Chrysomela cyanipes” “Type”

Locality: “nova Hollandia"™

Current status: Phyllocharis

somelidae, Chrysomelinae

cyanipes. Chry-

Comment: The condition of the holotype is fair.

Left antenna and sore tarsi are missing, and there is

a large crack between the metathorax and abdomen,

also there is a great deal of glue on the ventral

surface

There are a number of colour pattern varieties

of this species listed in Junk'’s Catal (idem). Also

Baly (1855: p. 172) describes and discusses these

varieties fully. He states that the Fabrician type was

founded on a single specimen which he, Baly,

regards as an extreme variety. There are two blue

spots on each elytron in the holotype specimen, one

basal spot near the sutural angle and one slightly

posterior to it nearer the external margin, The

posterior half of the elytra is dark blue with a ramus

running up each side of the suture. There is a small

ferrugineous apical spol. This species is recorded

from New Guinea and Northern Australia.

CYANOCEPHALA,. Criaceris

Fabricius 1775, p. 121, n. 18; 1801, 1, p. 461, n. 58

antennalis Lea 1923: p. 52] and p. 543, new

synonym,

Zimsen 1964, p. 104, n. 1591,

Syntype label:

i. Fabricius “Cr. Cyanocephala Fab. Entom

p. 121 p. 18”.

ii. BMNH “Crioceris cyanucephala”’

Locality: “nova Hollandia™

Current status: Monolepra

Chrysomelidae. Galerucinae.

cyanocephala (F.)

Comment: There are two specimens on the

Fabrician Jabel, neither of which has been identified

by Waterhouse as the type, therefore both must be

regarded as syntypes. Zimsen records only one.

Both are in a very bad State of preservation, One

specimen, being badly pinned is very distorted and

the structures so twisted and superimposed on one

another, it is difficult to distinguish the characters;

both antennae are damaged. In the other specimen

the membraneous wings are unfolded and protrude

beyond the elytra, the antennae are damaged and

parts of the right one are carded. The ventral surface

is in a better condition than the previous specimen

and only the tarsi of the metathoracic legs are

missing. This isa male; the sex of the other specimen

is indiscernable.

This species has been omitted from recent

catalogues. Hope (1840: p.. 145) mentions it in his list

of Fabrician species under the original genus of

Crioceris. J, W. Wilcox does not mention it in Junk’s

Supplement 1971-74. In 1929, G. E. Bryant of the

Entomological Bureau, London, identified this

species as belongong to the genus Monolepta. T.

Blackburn (1896; p. 99) states “I have not seen any

Galerucid agreeing with the description of this

species, but it does not seem at all likely to be a true

Monolepta as its author states that the third joint of

its antenna is as long as the fourth”. On examination

of this specimens this is not the case, The second and

third joints are very small, the fourth and fifth joints

are longer and equal in size, and joints six to eleven

are longer than the second and third joints, but

shorter than four and five,

in the BMNH there are four cotypes, two males

and two females, of M. antennalis Lea, On

comparing these with M, cyanocephala (F.) they are

found to be identical. M, antennalis Lea then

172

becomes a synonym of M. cyanocephala (F.). Mrs.

8. L. Shute of the BMNH verified this synonymy.

CYLINDRIROSTRIS, Curculio

Fabricius 1775, p. 137, n. 55; 1801, 2, p, 463, n, 125

(Rhynchaenus)

innubus Herbst 1792-5, p 172-

longimanus Boisd. 1835, p. 408 (Orthorrhinus)

var. simulans Boh. 1836, p_ 245

var. laetms Saund. and Jekel 1855, p. 297

var, aspreda Pasc, L882, p, 380

var, carbonarius Pase. 1882, p. 381

var. lateralis Pasc, 1882, p, 381

vac. euchramus Fairm. 1883, p. 36

var. patreulis Pasc, 1885, p. 225

var, vagus Olliff. 1889, p, 89

var. pomicola Lea 1897, p, 624

var. albiceps Lea 1897, p. 624

var, howensis Lea unpublished,

Junk’s Catalogue 1932, 28, 122, p, 46 (Orthorrhinus)

Zimsen 1964, p. 204, n, 3491

Holotype labels:

i. Fabricius “Cure. eylindrirostris

Entom. p. 137 n, 55",

iii BMNH “Rhynchaenus

“Type”

Fabr.

cylindriostris”’,

Locality: “nova Hollandia’'

Current status: Orthorrhinus cylindrirastris (F.):

Curculionidae, Hylobjinae.

Comment; Holotype is black with scattered

greyish and brownish scales, This is a cammon

species which varies considerably in size and number

of scales present. In Junk’s Catalogue O. laetus

Saund. and Jekel and O, lateralis Pasc. are listed as

separate species. Before A.M. Lea died in 1932 he

sent me a copy of his MS notes on Lord Howe Island

Coleoptera, These have not been published, In these

notes he states that both O, lateralis Pase and O,

laetus Saund. are varicties of O. cylindrirostris (F.). 1

quote from Lea's MS:—

“O. cylindrirostris Fab. var. lateralis Pasc.

Six specimens were obtained; four with white

markings conspicuous, tWo With them rather vaguely

indicated,

O. cylindrirostris, var. vagus Oll. This var. was very

abundant on freshly felled trees at night time, and

was a common victim of Cormodes darwini.

O. cylindrirostryis, var howensis Lea. a mew variety.

A large var. 14-18 mm. Scales nearly all of a

creamy whiteness except in the vicinity of the

fascicles; elytral fascicles more or less ochreous, only

the median pair very canspicuous, apical fascicles of

prothorax feeble or absent. Rostrum long and thin;

front legs long in both sexes, but in male

considerably longer than in female and the front

tarsi wide, flat, and with conspicuous fringes.

REC 5. AUST MUS 18 (8): 155-197

January, [981

Specimens of this yar, in good condition have a

curious speckled appearance owing to the abundant

granules showing through the scales; but greasy or

dirty specimens look very different. Twelve males

and seven females were obtained tn company with

specimens of the preceding var. i.e. var. vagus Oll.”

These MS notes were sent to me in December

1931. Ina covering letter to me A.M, Lea stated that

var. howensis Lea was close ta var_ laetus Saund and

Jekel fram the New Hebrides.

In the BMNH Collection under the label O, laetus

Saund and Jekel which is in Jekel’s handwriting is a

specimen with the label O. cylindrirastris (F.)

identified by A.M. Lea as var. howensis Lea. There

are 2 locality labels on this specimen, (1) Lord Howe

Is. A.M. Lea and a BMNH label, Australia. Variety

howensis Lea is therefore a variety of O.

cylindrirostris and according to Lea QO. laetus Saund

and Jekel and O. lateralis Pasc, should be cegarded

as varietics also.

DECEMPUNCTATUS, Dytiscus

Fabricius 1775. p, 232, n. 11; 1801, 1, p, 263, n. 26

ab. flavescriptus Zimmerm, 1917, p, 213

(Platynectes)

ab. lugubris Blanch. 1853, p. 49 (Platynectes)

var, masteri Macl, 1871, p. 126 (Platynectes)

var. octodecimmaculatus Macl. 1825, p. 31

(Platynectes)

var. semperi Reg. 1899, p. 285 (Platynectes)

var, spilopterus Germ, 1848, p, 172 (Platynectes)

var, variegatus Reg. 1899, p. 286 (Platynectes)

Junk's Catal, 1920, 4, 71, p. 149 (Platynectes)

Zimsen 1964, p. 68, n. 879

Zimmermann 1917-19, p. 213

Syntype labels:

i. Fabricius “‘Dytise. 10—punctatus Fab.

Entom, p. 232 n, 11”

i. BMNH “Dytiscus 10—punctatus’’,

“Type”

Locality: “nova Hollandia", This species is very

widespread and has also been recorded from the

islands of Indonesia, Philippines and New Guinea.

Current status: Platynectes decempunctatus (F.):

Dytiscidae

Comment; The condition of the “'type’’ is good,

only the left mesothoraci¢ leg and tarsal joints of the

right and left metathoracic legs are missing. This

species has a great many varieties which are

recarded ty Zimmermann in Junk’s Catalogue

(1920). Both Sharp (1882: p. 540) and Zimmermann

(1917, 1919; p, 213) give full accounts of this species.

The varieties differ mainly in the markings of the

yellow spots. The “type” has a large distinct anterior

spot on the head, one spot on each anterior angle of

the prothorax, five yellow spots on each elytron

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 173

arranged in two longitudinal rows—three in the

inner row near the suture, and two outer lateral

spots situated slightly posterior to the inner anterior

and medial spot. The dorsal surface of the type is

dull, this may be due tw dirt and age. All the

specimens in the BMNH collection are shiny,

DETRITA, Coccinella

Fabricius 1775, p. 85, 9. 36: PSO, 1. p. 373, 0. 98

Yamica Newman 1842, p. 415 (Paropsis)

laesa Germ. 1848, p. 235 (Paropsis), new

synonym,

conferta Chap, 1877, p. 81 (Parcpsis)

Zimsen 1964. p. 76, 0. 1237

Holotype labels:

i, Fabricius “‘Coccin. Detrita Fab. Entom, p.

85 n, 36"

ii. BMNH “Coccinella deirita”

Locality: ‘nova Hollandia”

Current status: Paropsis

somelidae, Chrysomelinae

detriia (F.): Chry-

Comment: Condition of holotype is poor. The

insect has been broken and roughly glued together.

Parts of the antennae, the labial palps and parts of

all legs except for those of the right meso and

metathoracic legs are missing.

This species was classified as a coccinellid by

Fabricius. Hope (1840; p, 87) refers it to the genus

Paropsis but makes no further comment on it. lt does

not appear in either Gemminger et Harold's

catalogue nor in Junk’s catalogue. In 1929 G. E.

Byrant of the Entomological Bureau of London

checked this synonymy and agreed that it was the

same as.the species. P, (Chrysophtharta) laesa Germ.

(1948: p, 235). P. detrira (F.) is therefore the valid

name and P. laesa Germ is a new synonym,

Blackburn's (1899: p. 493) description of P. laesa

Germ. describes the “type” specimen of P, detrita

(F.). The above synonyms are given in Junk’s

Catalogue 1916, 24, 68, p. 164 for P. laesa Germ.

DETRITUS, Hister

Fabricius 1775, p. 53, n. 10; 1801, 1, p. 89, mn, 28

Junk’s Catal. 1910, 8, 24, p, 93 (Saprinus)

Zimsen 1964, p. 35, 0. 262

Holotype labels:

i. Fabricius ‘‘Hister detritus Fab. Entom. p.

53 n, 10"

ii. BMNH “Hister detritus’, “TYPE”

Locality: “nova Hollandia”’. This is probably a

New Zealand species.

Current status: Saprinus detritus (F.): Histeridae,

Comment; The condition of the type is fair except

for the legs, one of which is entirely missing and the

others are in a bad state of preservation. There is

also a good deal of dried fungal hyphae present.

H. Bickhardt in Junk's Catalogue (idem,) lists S.

detersus \lliger, (1807: p. 365) and queries S. detritus

Rossi (1790: p. 29) as asynonym, Fabricius (1927: p

76) refers to Rossi's and Olivier’s descriptions. Rossi

was not the author of the species. Both Fabricius and

Olivicr (1789: p. 12) give “nova Hollandia’ as the

locality. The locality of S, detersus Ill. is a

Mediterranean and Western European species. R.

Staig (1931: p. 102) lists S. detritus (F.) as the species

and S. detersus (ll. as 4 synonym. He notes that in

the Fabrician reference 1787 p. 33 the specific name

is spelt as detricus, which is probably a misprint, 3.

detritus (F.) has been omitred from tecent

catalogues. On comparison of the type of S. derrints

(F,) with specimens of S. detersus El. in the BMNH

collection, it is evident that these two species are not

identical and should be regarded as two distinc?

species.

Characters of S. detritus (F.):—Head and thorax

black, elytra deep red. Head:—punctation slight and

scattered. Prothorax:—anterior margin fringed with

hairs, transverse, slightly bisinuate at base, anterior

arigles rounded but projecting, lateral and posterior

margins punctate with punctures fading out towards

centre on posterior margin, disc smooth and shiny

and under magnification very fine punctures are

visible, Elytra;—oblong, almost rectangular. slighty

swollen at humeral angle, lateral edges bent sharply

at right angles, posterior half of elytra coarsely and

densely punctate, on lateral margins punctures

extend up to two-thirds of length of elytra and reach

anterior margin between third and fourth striae,

Five striae are distinguishable in anterior half, and

first and second striae are united anteriorly. Ne

striae are distinguishable in posterior half,

Legs:—External margin of tibiae af pro- and

mesothoracic legs have four strong denticles and two

small ones at apex, The metatibiae have spines on

external margin,

Abdomen:—Ventral surface punctated, propy-

gidium and pygidium also punctated.

In the BMNH there are several unidentified

specimens from New Zealand. On comparing them

with the type of S. detritus (F,) they appear to be the

same species. It is probable therefore that the

Fabrician locality “nova Hollandia” is incorrect and

should be ‘nova Zelandia”™.

DEUSTA, Cassida

Fabricius 1775, p. 89, n. 8; 1801, 1, p. 396, n. 44

corallina Boisd 1835, p, 541

angulifera Blanch. 1853, p. 324

Junk’s Catal. 1914. 28, 62, p. 70 (Aspidomorpha)

Zimsen 1964, p 90. n- 1313

174 REC. 8. AUST. MUS [8 (8): 155-147

Syntype labels:

i. Fabricius ‘‘Cassida deusta Fab. Eniom. p.

89 yn, R

li. BMNH "Cassida deusta’ “Type”

Locality; ‘nova Hollandia™

Current status! Aspidomorpha deusta (F.): Chry-

somelidae, Cassidinae.

Comment; The “‘type’’ is in fair condition, The

terminal joints of the left antenna, the terminal larsi

of the mesothoracic legs and the left metathoracic

leg are all missing. There is a second specimen which

is cleaner than the “type” but both right meso- and

metathoracic legs and the tarsi of the left

metathoracic leg are missing.

The number of spots arranged in three longitu-

dinal rows on the elytra varies considerably in this

species, They may be absent or two or three may

unite, If present, the position of the spots remain

constant. The holotype has tive spots in the sutural

and median lines and four in the marginal line with

three of them fusing,

A, nigrodorsaia Boh., given In Junk's Catal,

(idem.) as a subspecies, has two black median spots

missing on the elytra but otherwise is in close

agreement with A. deusta (F.).

DIDYMUS Cryptocephalus

Fabricius 1775, p. 107, n. 9; T8OT, 2. p. 43, n. 11.

ab. fulvoplagiata Jac. 1884, p. 210)

Junk's Catal, 1914, 24, 59, p. 92 (Rhyparida)

Zimsen 1964, p, 122, n. 1923

ees

Syntype labels;

i, Fabricius “Crypt, didymus Fab, Entom, p,

107 n. 9”

iii BMNH “Cryptocephalus didymus"’,

“Type”

Locality: “nova Hollandia”

Current status: Rhyparida didyma (F.) Chryso-

melidae, Eumolpinae.

Comment: The “type’’ is badly damaged. Both

antennae and all legs are incomplete, postenor hulf

of right elytron is missing. There is a second

specimen but it also is in a poor condition with the

glue on the ventral surface obscuring most of the

structures, both elytra are complete.

The three spots on the eltyra may vary in size and

shape in this species, may also be united or obsolete,

Jn the “type” they are distinct, the basal spot which

Fabricius describes as ‘‘postice didyma”’ (posteriorly

divided into two) is semicircular in shape with one

branch extended towards the sutural margin and a

slight indication of a branch given off medianly

towards the outer margin, the central spot is roughly

triangular and the apical one is elongale.

January. 1Ys8T

ERYTHROCEPHALUS, Staphylinus

Fabricius 1775, p. 265, n. 6; 1801, 2, p. 593, nm. 19,

punctatus Hope

Junk’s Cutal. 1914, 5, 57, p. 398

Steel 1949, p, 57-61

Zimsen 1964, p, 231, n. 3986

Holotype labels:

i, Fabricius “‘Staph, erythraceph,

Entom, p, 265 n. 6"

ii. BMNH “Staphylinus erythrocephalus”,

“Type”

Fah,

Locality: ‘‘nova Hollandia”

Current status: Creophilus erythrocephalus (F.)

Staphylinidae, Staphyliniae

Comments; Holotype, a female, head is very

much deflexed, membraneous wings are unfolded,

terminal joints of left antenna missing and left

prothoracie leg is broken at base of tibia, The second

specimen, a female, had been identified by W. O,

Steel (idem,) as C. lania Er, It is in fair condition,

only right metathoracic leg is missing and membran-

ous wings are unfolded.

C. erythrocephalus (F.) is widely distributed

throughout Australia and is recorded from Tonga,

Tahiti and Chile, In the BMNH there is the type of

C. punctatus Hope which is marked as a synonym,

W.Q, Steel (idem,) records this and points out the

differences between C, erythrocephalus (F,) and

C, lania Er,

EXCLAMATIONIS, Curculio

Fabricius 1775, p, 133, n. 37; 1801, 2, p. 456, n. 89

(Rhynchaenus)

Zimsen 1964, p. 203, n. 3471

Holotype labels:

i, Fabricius ‘Cure, Exclamationis Fab,

Entam, p. 133 n, 37”

ii. BMNH ‘‘Rhynchaenus exclamationis’’,

“Type

Locality: ‘nova Hollandia’

Current status: unkown, Fam. Curculionidae

Comment: The holotype is badly pinned through

position of Jeft elytron which is missing, pin emerges

on ventral surface between metathorax and

abdomen so that abdomen is nearly detached from

the thorax, antennae hidden beneath rostrum; left

prothoracie leg is missing,

There is no record of this species in recent

catalogues, Olivier (1801 p. 155, £165) describes this

species but the figure ts poor and does not shaw any

detailed structure. Fabricius changed the genus in

his later publications to Rhynchaenus.

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

FARINOSA, Buprestis

Fabricius 1775. p. 219, n, 16; 1801, 2, p. 195. n. 50

var. auroimpressa Cast. and Gory. 1835, p. 20

(Chrysodema)

vat, impressa Kerr, 1898, p. 118 (Cyphogasira)

var. borneensis kerr. idem, p. 251 (Cyphogastra)

Junk's Catal, 1926, 12, 84, p. 117 (Cyphogastra)

Zimsen 1964, p. 152, n, 2509

Holotype labels:

i, Fabricus ““Buprestis farinosa Fab, Entom.

p. 219 n, 16”

i, BMNE “Buprestis farinosa’, ‘Type’

Locality; “nova Tlollandia’, This Fabrician

locality is probably incorrect, This species has only

been recorded in the East Indies.

Current status: Cyphogastra farinosa(F.): Buprest-

idae, Chalcophorinae,

Comments: Holotype is pinned through the left

elytron; terminal joints of both antennae and tarsi of

left mesothoracie and both metathoracic legs are

missing: ventral surface is covered with a fine golden

powder adhering to the pubescence, There are

several specimens of this species in BMNH

collection and they are all from Java and Borneo.

Banks may have collected this specimen when the

Endeavour stopped at Batavia for repairs.

FASCIATA F., Cetonia

see STOLATA F. Cetonia

C, faseiata F. is a synonym of C. stolata F,

FELINUS Dermestes

Fabricius 1781, 1, p. 34, n. 11; 1801.1, p. 314.0. 13

Junk's Catal, 1911, 14, 33, p, 43

Zimsen 1964, p. 77. 1, LOS7

Hinton 1945, p, 296

Syntype labels:

i. Fabricius “Felinus Fabr. Mant. Ins. n, 11”

Kergulandel”

i. BMNH- “Dermestes

number ‘'63-46"

felinus’ Register

Locality; “‘Terra Diemenii” in 1781 reference,

“Kergulandel” on the label, Kergulen Island or

Land is an island in the South Indian Ocean. Cook

charted jts coast on his Third voyage before calling

in at Adventure Bay, Tasmania. Nelson, the

collector, was on hoard, It appears that Fabricius

must have thought this island was off the coast of

Tasmania.

Current status: D. felinus F. is a synonym of D.

ater Degeer (1774: p . 223): Dermestidae, Lepesme

(1939: p, 192) and Hinton (idem.) discuss this

SVnONYMy-

175

Comment: There are two specimens in the Banks

Collection, one of which has the BMNH type label.

This is in the better condition, Head is sharply

deflexed and some of the tarsi are missing. Zimsen

only records one specimen in the Banks Collection.

FESTIVA, Melolontha

Fabricius 1775, p. 36, n, 23; 1801, 2, p. 171, n. 63

festa Gmelin 1788, p. 1561 (Calonota)

munda Sharp 1876, p. 73 (Pyronota)

var laeta F. 1775, p. 36, n. 24

Junk’s Catal, 1912, 20, 47, p. 90 (Calonota)

Zimsen 1964, p. 146, n, 2390

Syntype label:

1. Fabricius “Melo. Festiva Fabr. Sp, Ins

No. 31”

i. BMNH "Melolontha festiva’, “‘Type”

Locality: “nova Zelandia”

Current status: Calonota festiva (F.):

Scarabaeidae, Melolonthinae

Comment: The “type” is in good condition, It is

pinned through the left elytron; right prothoracic leg

except for coxa, and tarsi of left metathoracic leg are

missing, There is a second specimen which is not

recorded by Zimsen, the condition of which is only

fair, it has been broken at the base of the prothorax

and then glued together, This species must have

been re-examined by Fabricius who put the later

reference 1781 on the label when compiling the list

for this publication.

There are a number of colour varieties of this

species of which Melolontha laeta F. is one.

GIGAS, Ptinus

See GRISEUS F. Anthribus

P. gigas F. is the junior homonym.

GRISEA, Saperda

Fabricius 1775, p. 186, n, 9; 1801, 2, p. 324, n. 37

heteromorpha Boisd. 1835 (Lamia)

lentus Newm. 1840, p. 12 (Xylotoles)

westwoodi Guer. 1847, p. 27 (Xylotoles)

Junk’s Catal. 1922, 23, 73, p. 3 (Xylotoles)

Zimsen 1964, p. 175, n. 2966

Syntype labels;

i. Fabricius ““Saperda Grisea Fab. Entom. p.

186 n. 9"

ii. BMNH ‘‘Saperda grisea’, “Type”

Locality: “nova Zelandia”

Current status; Xylotoles griseus (F.): Ceramby-

cidae, Lamiinae

Comment; This is a small insect, The “type”

specimen is pinned through the right elytron which it

has displaced, There ts a great deal of glue present

on both dorsal and ventral surfaces and entangling

the legs; both antennae, except for the basal joint of

the left one, and the left elytron are missing.

There is a second specimen which, is cleaner than

the “type’’ and left antenna and left elytron are

complete, but right elytron is broken and abdomen

is displaced.

Broun (1880; p, 593) attributes this species

erroneously to Westwood (1843; p, 27) and gives S,

grisea F. as a synonym. Westwood's description has

a much later date, i.e., 1843,

GRISEUS, Anthribus

Fabricius 1775, p. 63, n. | (Ptinus gigas); L801, 2.

p, 410, n, 22

gigas F. 1775 p. 63 (Ptinus)

longicornis F. 1798 Suppl., p. 160

subsp. cervinus Klug. 1833, p. 188

nigroungulatus Fahrs. 1839, p. 241

Junk’s Catal, 1929, 26, 102, p. 86 (Phloeobits)

Zimsen 1964, p. 193, n. 3297

Locality: ‘nova Hollandia”. Its known distribu-

tion is naw from New Guinea to Australia.

Current status: Phloeebius (F.)

Anthribidae,

griseus

Comment: The holotype is missing from the

collection. In the original description (1775)

Fabricius adds ‘tan proprii genesis’’, a genus of its

own, In the 1792 and 1801 publications, Fabricius

changed not only the generic but also the specific

name to Anthribus griseus. This species is now in the

genus Phloeobius (Schk. 1826: p. 36) and the valid

specific name is griseus according to the Interna-

tional Code.

GUTTATOPUSTULATA, Coccinella

Fabricius 1775, p. 87, n. 51; 1801, 1, p. 385, n. 153

Junk’s Catal. 1931-32, 16, 118, p. 32 (Epilachna)

Zimsen 1964, p. 88, n. 1273

Holotype labels:

1, Fabricius “Cocc. guttato-pustulata Fab,

Entom, p. 87 n. 51”

ii. BMNH “Coccinella guttato-pustulata”

Locality: “nova Hollandia”’

Current status: Epilachna guttatopustulata (F.):

Coccinellidae

Comment: The holotype is very dirty and there is a

great deal of glue on the ventral surface,

The species is a well known leaf-eater

176 REC, 8, AUST. MUS 18 (A): 155-197

January, [98]

HAEMORRHOIDALIS, Curculio

Fabricius 1775, p. 140, n. 71, 1801, 2, p. 484, n. 215

(Rhynchaenus)

Zimsen 1964, p. 208, n, 3567

Syntype labels:

i, Fabricius “Cure. haemorrhoidalis Fab.

Entom. p. 140 n, 71"

ii, BMNH ‘'Rhynchaenus haemor-

rohoidalis”, “Type”

Locality: “nova Hollandia™

Current status: Haplonyx haemorrhoidalis (F.)

Curculionidae, Haplonychinae.

Comment; They are two specimens, male and

female, The male has the “type” label, it is badly

pinned through left elytron, the abdomen has been

pushed out of place and lies at right angles to the

body plane; the specimen is badly abraded,

Female specimen is in good condition but is also

badly abraded,

This species has been omitted from recent

catalogues. In the BMNH there are specimens of

this species labelled Melanterius haemorrhoidalis

(F.).

Both E, C, Zimmermann of C.§.1,R.O. Canberra

and R. T. Thompson of BMNH (pers, comm,) place

this species in the genus Haplonyx.

HIRTA, Saperda

Fabricius, 1775, p. 184, n. 4; 1801, 2, p. 320, n, 13

(S. villosa)

villosa F. 1801

Junk’s Catal, 1912, 22, 39, p. 115 (Oemona)

Zimsen 1964, p. 174, n, 2945

Syntype labels:

i, Fabricius “Saperda hirta Fab, Entom p.

184 n. 4”

ii. BMNH “‘Saperda hirta’, “type"’

Label of 2nd specimen; BMNH “‘Saperda villosa

F.. Syst. El.”

Locality; ‘nova Zelandia”

Current status: Oemona hirta (F,); Cerambycidae,

Cerambycinae.

Comment: The ‘‘type” is a male, its condition 1s

fair, terminal joints of both antennae, right

prothoracic Jeg except for coxa, tarsi of left

prothoracic leg and both metathoracic legs are

missing, The second specimen is a female, its

condition is poor and it is badly rubbed, It has the

BMNH label ''Saperda villosa F, Syst. El.” This

labelling indicates that this was labelled when the

collection was arranged in the order given in the

Systema Eleutheratorum, as in that reference

Fabricius gives the specific name as villosa and hirta

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 177

as the synonym as nov, nom., as he considered the

name was preoccupied. He had used the specific

name of hirta for a second species of Saperda,

described first in 1792, p, 317, the locality of which is

given as Italy. The New Zealand species however, is

the senior homonym and therefore is the valid name

for this specimen and villosa is the synonym. There

is a large series in the BMNH of this species, and it

shows great variation in size. Broun (1880; p, 570

and p. 1275) gives a detailed description of it.

However, the type differs from this description as it

has a smooth vertex, a dense patch of bright yellow

hairs in the emarginate portion of the eye and

another posterjor to the eye, There 1s no indication

of a smooth dorsal line on the prothorax. Also there

is no pubescence along the dorsal line, but this may

be due to rubbing. The elytra are also devoid of

hairs,

HOLLANDIAE F., Scarabaeus

See NOVABHOLLANDIAE, F., Scarabaeus

INAEQUALIS, Coccinella

Fabricius 1775, p, 80, n. 8: 1801, 1, p, 362, n. 40

Junk’s Catal. 1931-32, 16, 120, p. 292 (Coelophora)

Zimsen 1964, p. 84, n. 1197

Holotype labels:

i. Fabricius “Cocc, inaequalis Fab. Entom,

p. 80 n, 8"

i. BMNH “Coccinella inaequalis’, “Type”

Locality: “nova Hollandia”

Current status: inaequalis (F.):

Coccinellidae

Coelophora

Comment: Condition of “type”’ is fair. It has been

pinned twice through the right elytron. The ventral

surface is covered with dirt and dried hyphae, rhe

antennae are missing. There are a number of

varieties recorded for this Species in Junk’s

Catalogue (idem.). The Fabrician type is taken as

the typical structure and is described fully by

Mulsant (1850; p.404), The large series in the

BMNH collection show considerable variation of

markings on thorax and elytra, This species has a

wide distribution from Japan and China, through the

Malay Archipelago to Tasmania.

INTERRUPTA, Cassida

Fabricius 1775, p. 89, n. 7, 1801, 1, p. 395, n. 40

plasoni Spaeth, 1898, p. 539 (Aspidomorpha)

var badeni Wgenr. (Aspidomorpha)

var planipennis Blkb. (Aspidomorpha)

Junk’s Catal, 1914, 25, 62, p. 71 (Aspidomorpha)

Zimsen 1964, p. 90, n. 1309

Holotype labels:

i. Fabricius “Cassida interrupta Fab. Entom.

p. 89 n. 7”

ii. BMNH “Cassida interrupta”’

Locality; “nova Hollandia™

Current status; Aspidomorpha interrupta (F.)

Chrysomelidae, Cassidinae

Comment: The holotype is complete except for

the terminal joints of the right antenna. The legs are

covered in dried mycelial hyphae.

The black markings on the elytra of this species

varies as to degree and arrangement.

INTERRUPTUS, Curculio

Fabricius 1781, p. 188, n, 148; 1801, 2, p, 519, n, 70

obliquesignatus Boh. (842, p. 244

griseus White 1846, p. 14 (Otiarhynchus)

apicalis Broun 1881, p. 701 (Empaeotes); loc.

cit. 1893, p. 118 (Platyomida)

Zimsen 1964, p. 214, n. 3701

Kuschel 1969, p. 789 and pp, 792-5 (Catopres)

Lectotype (Kuschel) labels:

i. Fabricius “Curcul

Ins. n. 148

ii, BMNH “Curculio Interruptus’, “type”

Interruptus Fabr. Sp-

Locality: Fabricius recorded this specimen from

“nova Hollandia’’.. This has proved incorrect as it 1s

from New Zealand (Kuschel idem,).

Current status: Catoptes interruptus (F.): Cur-

culionidae, Leptopiinae

Comment: There aré two specimens in the

Collection. The one that originally had Water-

house's “type” label is in very poor condition having

metathoracic legs and abdomen missing, The

“second specimen” is in better condition and has

only right metathoracic leg missing, Glue obscures

the ventral surface. This specimen is a male and has

been designated by Kuschel (idem, ) as the lectotype

and a Jabel as such has been added to the specimen

by R. T. Thompson of the BMNH, The former

“type” specimen Kuschel now designates as a

paralectotype (1970; p, 204),

There is a female specimen in Fabricius’s own

collection which Kuschel designates as a paralec-

torype (Kuschel 1970: p. 204).

G. A, K. Marshall (1944; p, 73) transferred all

New Zealand Catoptes species from the sub-family

Eremninae to the sub-family Leptopiinae.

JESUITA, Apate

Fabricius 1775, p. 54, n, 3; 1801, 2, p. 380, n. §

canarii Nordl. 1880 App., p. 66

Junk’s Catal, 1938, 10, 161, p, 39 (Bostrychopsis]

Zimsen 1964, p. 186, n. 3175

178

Holotype labels:

i. Fabricius ‘‘Apate Jesuita Fab. Entom. p.

54 n. 3"

ii. BMNH “‘Apate jesuita”

Locality; “nova Hollandia™

Current status: Bostrychopsis jesuita (F.): Bos-

trychidae

Comment: Only the metathorax and abdomen of

the holotype is left and neither metathoracic leg is

complete.

There is another specimen in Fabricius’s own

collection, now at the Zoological Museum in

Copenhagen, This is a well known species and is

widely distributed throughout North and South

Australia, New South Wales and Victoria, The

colour is usually uniformly black, but some

specimens may be reddish,

LAETA, Melolontha

Fabricius 1775, p. 36, n. 24; 1801, 2, p. 171, n, 64

Junk’s Catal, 1912, 20, 47, p. 90 (Calonata)

Zimsen 1964, p, 146, n, 2391

Holotype labels:

i, Fabricius ‘‘Melol, laeta Fabr. Sp. Ins. n.

32”

ii. BMNH ‘‘meloiontha laeta’’, ‘‘type”’

Locality: “nova Zelandia”

Current status: Calonota laeta (F.) is a variety of

C. festiva (F.); Scarabaeidae, Melolonthinae.

Comment: The condition of the holotype is fair,

both antennae except for two basal joints, tarsi of

left prothoracic and right metathoracic legs are

missing.

This is a colour variety of C. festiva F, It is a pink

opalescent colour with red thoracic stripe and red

scutellum. Fabricius must have re-examined this

species before publishing in 1781 and put this later

reference on the label,

LAEVICOLLIS, Silpha

Fabricius 1775, p, 73, n. 7; 1801, 1, p. 338, n. 8

reticulatus Haag 1878, p. 659 (Saragus)

Junk’s Catal. 1910-11 , 18, 28, p. 424 (Saragus)

Zimsen 1964, p. 80, n. 1117

Carter 1926, p. 142

Holotype labels:

i, Fabricius ‘'Silpha laevicollis Fab. Entom.

p. 73 n. 7”

ii. BMNH “Silpha laevicollis”

Locality; ‘‘nova Hollandia”™

Current status: Celibe laevicollis (F.): Tene-

brionidae, Ulominae

REC. & AUST. MUS 18 (8): 155-197

January, LIST

Comment: The holotype is in good condition, only

a few tarsi and the right metathoracic leg are

missing.

Schenkling in Junk's Catalogue (idem.) incor-

rectly attributes this species to Olliff, but Carter

(idem.) corrects this. This species was formally

classified in the genus Saragus; Watt (1968; p, 36)

made Saragus a synonym of the genus Cellbe.

LAEVIGATUM, Opatrum

Fabricius 1781, 1, p. 90, m. 4; 1801, 1, p. 117, n. 8.

?mauritanicus F, 1792, p. 113 (Tenebrio)

piceus Ol, 1792, p. 50

Joryzae Herbst 1799, p, 18

picipes Steph, 1832, p. 11 (Alphitobius)

granivorus Muls. et God, 1868, p.

(Alphitobius)

striatulus Fairm, 1869, p, 231 (Cartaphronetis)

rufipes Mad. 1872, p. 286 (Microphyes)

Zimsen 1964, p. 39, n. 334

Carter 1926, p. 137

288

Syntype label;

i, Fabricius “Opatrum Jaevigatum Fab. Sp.

Ins, n. 4 Hab, in Nova Zel”

iii BMNH ‘‘Opatrum laevigatum"

There are two specimens under this label, the

“type™ specimen has the Fabrician label

Locality: ‘nova Zelandia”’

Current status: Alphitobius laevigatus

Tenebrionidae, Ulominae.

(F.);

Comment: The condition of the “type” js fair, the

legs are bunched together and tarsi are missing. This

species is not listed by H. Gebien in Junk’s

Catalogue (1910, 18, 15, p. 458) but the synonym

Microphyes rufipes Macl. is. This synonymy was

identified by Blair (1914: p. 486), The synonyms of

this species are listed in Junk’s Catalogue as

synonyms of A. piceus Ol. The distribution of this

species is now known to be cosmopolitan.

LEONINA, Coccinella

Fabricius 1775, p. 87, n. 54; 1801, 1, p. 384, n. 160.

tasmanii White 1846, 11. p. 23 (tasmaniae)

Junk’s Catal. 1931-32, 16, 120, p. 508.

Zimsen 1964, p. 88, n,. 1279

Syntype labels:

(i) Fabricius “Cocc. leonina Fab. Entom.

p. 87 n. 54”

(Gi) BMNH “Coccinella leonina"™, “Type”

Locality: "nova Hollandia’’. It is doubtful if this

Fabrician locality is correct. Both Mulsant (1850: p.

128) and Schenkling in Junk’s Catalogue (idem,)

give the locality as New Zealand. Out of twenty

specimens in the BMNH collection seventeen have

IHE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

locality labels as New Zealand. Gemminger et

Harold and Masters Catalogues both give the

Fabrician locality,

Current status: Coccinella leonina F.: Cocci-

nellidae.

Comment: The “type” is in a very poor condition.

It has been badly pinned and the specimen is

covered with dried fungal hyphae. There is a second

specimen which is in much the same condition,

The specimen in the BMNH Collection of the

synonym C. tasmanii White is labelled “Coccinella

Tasmanii White, Zool. Ereb. and Terror'’ (1846:

p. 23). The habitat of this specimen is labelled New

Zealand, This is probably White’s type specimen

although it is not labelled as such, In Junk's

Catalogue this synonym is incorrectly spelt as

‘Tasmaniae’,

LIMBATUS Dermestes

Fabricius 1781, 1, p. 66, n. 23; 1801, 1, p, 318, n. 36

atomaria Pase, 1875, p. 210-223

discoidea Pase. 1875, p, 210-223.

Zimsen 1969, p. 78, n, 1076

Crowson 1964, p, 313 (Phycosecis)

Syntype labels:

(i) Fabricius “‘Dermestes limbatus Fabr. Sp.

Ins. n, 23"

(ii) BMNH ‘*Phycosecis limbatus” “Type”

Locality: "nova Zelandia”

Current status: Phycosecis limbata (F.): Phyco-

secidac.

Comment: The “type” is in a very dirty condition,

so thar it is difficult to determine its structures,

especially those of the legs. It is a very small insect

and has been pinned through the suture. The head is

deflexed sharply. Head is black, main body dark

brown and lateral margins of elytra pale testaceous,

Prothorax as broad as long, rounded, posterior

angles obsolete and anterior angles very acute.

Elytra twice length of prothorax, pubescence scanty,

punctate and striate. This species is not recorded in

Junk’s Catalogue, in Broun’s manual of New

Zealand Coleoptera, nor in Huttons Index Faunae

Novae Zealandiae. There is a second specimen, It is

in a worse state of preservation than the “type”.

LINEATUM, Callidium

Fabricius 1775, p. 189, n, 10; 1801, 2, p, 340, n. 40

australis Gmel, 1790, p. 1849

Junk’s Catal. 1912, 22, 39, p. 488 (Navomorpha)

Zimsen 1964, p. 179, n. 3036

Holotype labels;

i. “Callid. lineatum Fab, Entom. p. 189 n.

10"

ii, “Callidiam lineatum’”’, “Type”

179

Locality; “nova Zelandia’. Fabricius in 1801

erroneously published the locality as “nova Hol-

landia’’. In 1775 the locality is correctly given as

“nova Zelandia”’.

Current status: Navomorpha

Cerambycidae, Cerambycinae.

lineata (F.):

Comment: The holotype is in good condition, only

most of the right antenna, and left metathoracic leg

are missing. See Broun (1880: p. 590) for detailed

desenption of the species. In Gemminger et Harald

Catalogue (1873; p. 2984) the genus is spelt

Naomorpha,

LINEATA, Lagria

Fabricius 1775, p. 124, n. 3; 1801, 2, p. 68, n, 4

(Dryops)

Junk’s Catal, 1915, 17, 65, p. 33 (Sessinia)

Zimsen 1964, p. 127. n, 2025

Syntype labels:

i. Fabricius “‘Lagria lineata Fab. Entom p

124 n. 3",

ii, BMNH “Dryops lineata’, '"Type”™

Locality: “nova Zelandia”’

Current status: Sessinia lineata (F.);

Ocedemeridae,

Comment: The “type” is a female and is in a good

state of preservation. the antennae except for basal

joint of left antenna and right methatoracic leg are

missing. There is a second specimen, it is not in a

good condition; it is pinned through the left elytron

4s of which is missing, together with both prothoracic

légs; not one leg is complete. This species has been

recorded under a number of different genera.

Fabricius put it in the genus Dryops in his later

publications. It has also been recorded in the

following genera Oedemera (Ol. 1794: p, 6),

Sclenopselaphus (Gemm, et Har, Cat, 1870: p. 2168)

and Thelyphassa (Blair 1920: p. 153)

LINEOLA F.,, Coccinella

See STRIOLA F,, Coccinella

C. lineola F. is a junior homonym

LINEOLA, Curculio

Fabricius 1775, p. 136, n. 48; 1801, p. 503, n. 28

(Lixus)

Zimsen 1964, p, 211, n, 3631

Syntype labels:

i, Fabricius ‘‘Curc. lineala Fab, Entom, p,

136 n, 48”

ii. BMNH “Lixus lineola”, "Type"

Locality: ‘nova Hollandia’’

180) REC, S. AUST, MUS 18 (8); 155-197

Current status: Cossonideus lineolus (F.): Cur-

culionidae, Cossoninae,

Comment: There are two specimens in the

Collection, Zimsen records only one. The specimen

with the type” label is in very poor condition; head

and prothorax are intact, but posteriorly it is very

badly damaged, only a few remnants of abdomen

and elytra remain, The second specimen is in a

better condition, only elytra are missing. This

species is not listed in Junk’s Catalogue. In

discussion with EF. C, Zimmermann of C.S.1,R.0.,

Canberra, he placed this species in the genus

Cossonideus,

LURIDUS, Curculio

Fabricius 1775, p, 138, n. 63; 1801, p. 470, n. 155

(Rhynchaenus)

fulginosus Boh, 1835, p, 431 (Crypterrhynchus)

immansuetus Boh. 1837, p. 328 (Acalles)

plinthoides Pasc, 1871. p. 199 (Omydaus)

Junk’s Catalogue 1936, 29, 151, p. 163 (Omydaus)

Zimsen. 1964, p. 205, n. 3514

Lea 1918, p, 272

Holotype labels:

j. Fabricius ‘Cure. luridus Fab, Entom., p.

138 n. 63”,

li, BMNH “Rhynchaenus luridus”, ‘type’,

Locality; “nova ‘‘Hollandia™

Current status; Omydaus luridus (F.): Cur-

culionidae, Cryptorrhynchinae

Comment; The holotype is badly abraded, there

being no scales present: tarsi of right prothoracic leg

and left mesothoracic leg, and the right metathoracic

leg are missing.

This species has at times been placed in various

genera (Lea idem.), It is now listed in the genus

Omydaus.

LYNCEA, Saperda

Fabricius 1775, p. 185, n. 8; 1801, 2, p. 323, n, 35

Junk’s Catal, 1922, 23, 73, n. 3 (Xyotales) ,

Zimsen 1964, p. 175, n, 2964

Holotype labels:

i, Fabricius “Saperda lyncea Fab, Entom.

p 185 n. 8”

ii. BMNH “Saperda lyncea’’, *‘Type”’

Locality: “nova Zelandia”’

Current status: Xyotoles lynceus (F.): Ceramby-

cidae, Laminae.

Comment: The condition of the holotype is poor;

head is covered in dirt; neither anterinae is

complete, and every leg has parts missing; ventral

surface is covered in glue. In the copy of Broun's

Manual in the BMNH on p. 593 Dr. Gahan has

Jartidry, 198]

made a note that White's description (1846; p, 22)

which Broun followed “refers ta X. humeratus

Bates, the Fabrician species being wrongly identified

by him". Broun does quote a description given by

Bates after he, Bates, had examined the Fabrician

type, and this is the correct description of X. lynceus

(F.). X. lynceus (F.) is larger than either X. griseus

(F.) or X. humeratus Bates, It is elongate, narrow,

sides are nearly parallel along most of entire length

of insect, tapering at the apices of elytra;

ferruginous, head and prothorax are slightly darker

than the rest; prothorax is longer than wide,

margined anteriorly, and postetiorly reticulose with

an indication of an horizontal anterior and posterior

depression; base of elytra is not wider than the

prothorax, elytra striate with large scattered

punctures on the anterior half, more concentrated

towards the base, apices produced and divaricate.

MEDITABUNDUS, Curculio

Fabricius 1775, p, 139, n. 64; 1801, 2, p. 473, n. 169

(Rhynchaenus)

monachus Boisd. 1835, p. 430 (Cryptorrhyn-

chus)

Junk's Catalogue 1936, 29, 151, p. 241 (Euthyrhinus)

Zimsen 1964, p. 206, n, 3526

Holotype labels:

i, Fabricius. “Cure. meditabundus Fab.

Entom, p, 139 n, 64"

ii BMNH “Rhynchaenus meditabundus”,

“Type”

Locality: “nova Hollandia’”’

Current status: Euthyrhinus meditabundus (F.):

Curculionidae, Cryptorrhynchinae

Comment: Holotype is in good condition; it has

been pinned twice, now through the right elytron but

previously through mid-basal line of prothorax.

Some authors have spelt the genus Euthyrrhinus.

Schénherr who named and described the genus

(1837; p, 371) spelt it with only one “r’’, This species

is widely distributed throughout the Pacific Islands,

MELANOCEPHALA, Crioceris

Fabricius 1775, p. 119, n, 8; 1801, 1, p. 450, n, 7

Junk's Catal, Suppl. 1973 78, 3, p. 445 (Synodita)

Zimsen 1964, p, 102, n. 1544

Holotype labels:

i. Fabricius “Crioc, meélanocephala Fab,

Entom. p. 119 n. 8"

ii, “Crioceris melanocephala"', “Type”

Locality: “nova Hollandia”’

Current status: Synodita melanocephala (F,):

Chrysomelidae, Galerucinae.

Comment: The condition of the holotype is poor:

glue obscures much of the ventral surface, The right

THE -FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 18]

antenna, parts of the left antenna and metathoracic

legs, half of the elytron and the abdomen are

missing, J, A, Wilcox gives a full list of references in

Junk’s Catal. Suppl, (idem,). The colour of the

elytra of this species varies from metallic green-blue

to violet-blue. The elytra of the holotype is violet-

blue, but rather faded with age.

MINUTA, Apate

Fabricius 1775, p. 54, n. 4; 1801, 2, p. 383, nm. 25

substriatus Staph 1839, p. 352

vertens Walk. 1859, p, 260

siculus Baudi 1873, p. 336

bifoveolatus Zoufal 1894, p. 42

aponicus Matsum. 1915, p. 184

Junk’s Catal, 1938, 10, 161, p. 24

Zimsen 1964, p, 187, n. 3192

Locality: “nova Zealand”

Current status: Dinoderus minutus (F.): Bos-

trychidae

Comment: Specimen under this label in the

Bank's Collection is not Apale minuta F. but is a

species of Sinoxylon, C, Waterhouse recognised this

was not the type of A, minutus F, and has added a

label “This cannot be the type.”” There is no

specimen of this species in Broun’s collection but

there is a space left for it, P, Lesne (1897: p. 329) in

his revision of this group records this species as being

cosmopolitan but does not record New Zealand as

one of the countries where it has been found. There

is a large series in the BMNH showing its wide

distribution.

MINUTUM, Anobium

Fabricius 1781, 1, p. 72, 9. 8: 1781 Spec. Ins. App.,

pr. 499. n, 13-14 (Lagria), 1792, 2, p. 81, n. 17

(Lagria), 1801, 2, p. 74, n. 12 (Dasytes)

subcyanens Broun L880. p. 328 (Dasyles) new

synonym.

Junk’s Catal. 1937, 10. p 75 (Dasytes)

Zimsen 1964, p, 129, n, 2051

155

w~)s

Syntype labels:

i, Fabricuis ~“Lagria Minuta Fabr. Sp. Ins.

App. p. 499 n, 13-14”

ii. BMNH “Dasytes minula’, “Type”

Locality: **nova “#elandia”

Current status: Dasytes minutus F. Melyridae

(Dasytidae)

Comment: The “type” is in very poor condition,

pinned through the left elytron, structures appear to

be complete bur the specimen is very dirty and

covered with mycelium, The second specimen is also

pinned through the left elytren and is badly damaged

with head and prothorax missing. The Fabricius

reference on the label is not recorded by Pic in

Junk’s Catalogue nor by Zimsen. It appears that

When compiling the Species Insectorum Fabricius

put this species into the genus Anobium, then,

presumably after re-examination changed the genus

to Lagria and published this in the Appendix. In

1792 he still published it under the genus Lagria but

in 1801 put it in the genus Dasytes, the genus in

which it is how stands, Pie (idem.) records D, minuta

F. with the earliest reference but gives the incorrect

page number of 79 which should be 72. Page 79 is the

reference for the species Sphaeridium minuia F, D,

minuta F, does pot appear in Broun's manual

(1880), norin Hutton's Index Fauna Novae Zelandia

(1904). Broun (1880 p. 328) describes D, subcyaneus

as anew species, This account decribes the Fabrician

type of D. minuta F, and on camparison D,

subcyaneus Broun proves to be a synonym of DB.

minuta F, Pic. (idem.) lists them as separate spectes,

MINUTUM, Callidium

Fabricius 1775 p. 192 n. 23; 1801, 2 p. 346, n. 2

(Clytus)

exiguum Gmel, 1788 p. 1852, (Zorion)

fabricianum Westw. (Obrium) 1845, p, 28,

Junk’s Catal. 1912, 22, 39, p. 156 (Zorion)

Zimsen 1964, p. 180, n. 3061,

Holotype labels:

i, Fabricius ‘‘Callid Minutum Fab. Entom p

192, n, 23"

ii, BMNH “Clytus minutus’, “Type

Locality: “nova Zelandia™

Current status: Zorion minutum (F.): Ceram-

bycidae, Cerambycinae

Comment; The holotype is badly damaged, right

elytron is almost detached, except for 3 basal joints

of left antenna and abdomen are missing, and right

metathoracic leg is detached. See Broun (1880:

p. 584) for detailed description, Fabricius himself re-

classified this specimen in 1801 into the genus

Clytus.

MODESTA, Curculio

Fabricius 1781, p. 186, n, 138; 1801, 2.p. 512. n 30

brevipennis Brotin 1880, 1, p. 438 (Cecyropa)

maritima Broun 1880, 1, p. 438 (Cecyropa)

alba Broun 1881, 2, p. 698 (Cecyrepa)

varia Broun L881, 2, p, 698 (Ceeyropa)

albicans Sharp 1886, 2, 3 p. 416 (Cecyropa)

macularia Broun 1886, 4, p. 961 (Cecyropa)

lineifera Broun 1903, 7, 12 p. 72 (Cecyropu)

alternata Broun 1904, 7, 14, p, 10S (Cecyropa)

dicors Broun 1904, 7, 14, p. 106 (Cecyropa)

striatella Broun 1917, 1, 5, p. 399 (Cecyropa)

jucunda Broun 1917, 1, 5, p. 400 (Cecyropa)

laticollis Broun 1917, 1, 5, p. 400 (Cecyrapa)

182

Zimsen 1964, p, 213. n, 3670

Kuschel 1969, p, 789 and p. 807; 1970, p. 205

(Ceeyropa)

Holotype labels:

i, Fabricius “Cure. Modestus Fabr. Sp. Ins

no. 138”

ii, BMNH “Curculio modestus”, “Type"

Locality; ‘nova Zelandia”’

Current status: Ceeyropa modesta (F,); Cur-

culionidae, Rhyparosominae

Comment; Holotype a male, is very dirty; parts of

all legs are missing. This species was omitted from

recent catalogues, G, Kuschel describes and

discusses this species and its synonyms fully. Zimsen

records that there are 2 specimens in BMNEI. There

is only one, therefore it is the holotype,

MONTICOLA, Melolontha

Fabricius 1775, p. 39, n. 38; 1801, 2, p, 184, n. 138

Junk’s Catal. 1912, 20, 47. p. 96 (Liparetrus)

Zimsen 1964, p. 150, n. 2460

Holotype labels:

i. Fabricius ‘‘Melol. monticola Fabr. Sp. Ins.

n. 67"

ii, BMNH ‘Melolontha monticola’, Type”

Locality: “nova Hollandia”

Current status: Liparetrus monticolus (F.);

Scarabaeidae, Melolonthinae,

Comment: The condition of the holotype is fair;

but for the legs which are missing except for a few

remains lying in the glue. In Junk's Catalogue Dalla

Torres records this Fabrician species, but all earlier

references until Macleay’s publication in 1886, query

the author of this species, though there is a labelled

“type” specimen in the Banks’ Collection. Blackburn

(1905: p, 315) states that the two badly broken

specimens in the Macleay Museum labelled as

“monticola Fab.” are incorrectly labelled. The

specimens are two distinct species and one is labelled

elsewhere in the same collection as L. atriceps Macl.

and are not L. monticolus (F.), There are two

specimens labelled L. atriceps Macl, in the BMNH

but these are distinct from the Fabrician type L.

monticolus (F.), L. monticolus (F.) is smaller than L.

sylvicolus (Fab.), but they are closely related,

punetation on prothorax and striation and punctua-

tion on elytra is very similar. This species must have

been re-examined by Fabricius as the label has the

later reference 1781.

MORIO, Chrysamela

Fabricius 1787, 1, p. 66, n. 2; 1801, 1, p. 424, n, 5

Junk’s Catal. 1924, 24, 68, p, 161 (Paropsisterna)

REC, §.. AUST, MUS 18 (8): 155-197

January, LYS]

Zimsen 1964, p. 96. n. 1425

Syntype labels:

i. Fabricius “Chr, Morio Fabr, Mant. Ins.

n, 2”

i, BMNET “Chrysomela mario”, “Type”

Loeality: “terra Diemenii”’

Current status: Parapsisterna mario (F.), Chry-

somelidae Chrysomelinae

Comment: The condition of the “type’’ is fair.

parts of the antennae and tarsi of the right

metathoracie leg, which is detached and lying in the

glue, are missing. There is a second specimen which

is in very good condition, all parts are complete, In

this species there is a good deal of colour variation

from pitch black to dark lerruginous or reddish, The

type is black with brownish antennae, while the

second specimen has a reddish colouration,

Specimens from New South Wales, Tasmania and

South Australia in the large series in the BMNH

collection show the same variation in colour, They

also show variation in the punctation of the elytra. Tt

is possible that there is more than one species or

variety under this label. Blackburn (1898; p, 225)

describes this species in detail but states that the

punctures on the elytra ‘‘are fine, finer than those of

P. rubrosignata Boh. and P. sexpustulata Marsh. P.

rubrosignata Boh. is now regarded as a synonym of

P. beta Newm. Tn the Fabrician type, however, the

punetures are stronger and coarser than in either of

those species.

MORIO, Erotylus

Fabricius 1775, p. 123, n. 4; 1801, 2, p, 19. n. 14

(Cistela)

fovea-seriaius Fairm. 1877 p, 187 (Amarygmus)

curvipes Geb, (alienus BIK, nom, praeaece, 1893

p. 89 and 93) A’narygmus)

var, uniformis Blk. 1889, p. 1272 and 1893, p, 93

(Amarygmus)

var, fasmanicus Blk. 1893, p. 105 (Amarygnius)

Zimsen 1964. p. 117, n, 1825

Carter 1913, p. 37 and 1926, p. 158

Holotype labels:

i. Fabricius “Erotylus morio Fab. Entom. p,

123 n, 4”

ii, BMNH “Cistela morio”, Type”

Locality: “nova Hollandia”

Current status: Amarygmus morio (F ); Tene-

brionidae, Amarygminae.

Comment; The holotype is in a poor state of

preservation, It has been broken between the pro

and mesothorax and glued together again: glue

obscures most of the ventral surface; all legs, except

for the left metathoracic leg, are damaged and parts

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

missing. Carter (1913, p. 6) discusses the synonymy

of this species and on page 35 he quotes a letter from

Blair of the BMNH describing how the Fabrician

type was so dirty the colouration was completely

obscured but after cleaning the true colours were

revealed, Fabricius (1775; p. 123) describes the

colour as ‘ater’ (dull black), this must have been

due to the dirty state of the specimen which he must

have described without cleaning it, This species as

Carter notes was omitted from the catalogues,

MYSTACINA, Hispa

Fabricius 1775, p- 70, mn. 1; 1801, 1, p. 328, nm. 1

(Prilinus)

Junk’s Catal, 1925, 11, 81, p. 6 (Rhipicera)

Zimseo 1964, p. 79, n. 1102

Holotype labels;

i. Fabricius “Ptil. Mystacinus” written on the

underside af the label. “Derm. navalis

Fab. Entom. p. 56 9" was originally

written on the label but has been crossed

out, The writing on both sides of the

label is the same and appears to be that

of Fabricius,

ii, BMNH “Ptilinus mystacinus’” and the

Locality: “nova Hollandia”

Current status!

Rhipiceridae,

Rhipicera mystacina (F.):

Comment: Holotype is in good condition, most of

the left antenna is missing and also tarsi of the left

metathoracic leg, This was one of the specimens

Fabricius must have re-examined during his Jater

visit to London, By then, he had studied many more

species and had created new egencra as his

knowledge of insects increased, Therefore we find

that this species is no longer in the genus Hispa but is

put in the genus Ptilinus and is published as such in

1801. How the label of Dermestes nabalis F. came to

be on this specimen we do not know, As the "type"

of D. navalis F. was evidently missing when

Fabricius made his last study of the Banks

Collection, he meant only to use this label

temporarily When he changed the generic name of

H. mystacinus F.

NAVALIS, Dermestes

Fabricius 1775, p, 546, n. 9; 1801, 1 p. 155, n, 23

(Lyctus)

Junk’s Catal, 1910-11, 18 28, p, 395 (Tribolium)

Zimsen 1964, p. 48. n. 497

Locality: “nova Zelandia”

Comment: Zimsen records “'D, navalis in Nova

Hollandia, Mus D, Banks, type lost’, The 1775

reference is given in which the habitat is “nova

Ib3

Zelandia”. Zimsen records that D. navalis F. is a

synonym of Tenebrio ferrugineus F. the locality of

which fs given as ‘African aequinoctali” and the

type as being in Mus. Dom. Banks.”

In the Fabricius reference 1801, 1, p. 155 n. 23 for

Trogosita ferruginea there are fwo synonyms

listed—~Ips festacea Ent, Syst, Suppl, p. 179 n. 14 and

Lyetus navalis Ent. Syst. 1, 2 p. 504 n. 10 and the

“habitat ts piven as “India utrague’, In Junk's

Catalogue (idem) Tenebrio ferrugineus F. is listed

under the generic name of Tribolium with the

reference Mant. Ins. 1787, 1 p. 212. Zimsen gives an

earlier reference still for this species, Spec, Ins.

1781, Lp. 324. In Junk’s Catalogue T. navale Herbst

(1792: p, 138) 1s listed as a synonym of T, ferrugineus

F., but in the index of the catalogue T. navalis is

correctly attributed to Fabricius. K, G, Blair (1913;

p. 222) proposed the generic name of Triboloides for

T. ferrugineus F,

The “Type"' of D. navalis F. has recently been

found jn the Hunterian Collection in Glasgow. It is

in # poor state of preservation and is carded in a

larged amount of glue. See Introduction (p. 157),

NIGRICORNIS, Chrysomela

Fabricius 1775, p. 98, n. 20; 1801, lL. p. 435, n. 79

Junk’s Catal, 1924, 24, 68, p, 201 (Phyllocharis)

Zimsen 1964, p. 99, n. 1485

Holotype labels:

i. Fabricius “Chrys. Nigricornis Fab, Entom.

p. 98, n. 20".

ii, BMNH “Chrysomela nigricornis”, “Type”

Locality: “‘nova Hollandia”’

Current status: Phyllecharis

Chrysomelidae, Chrysomelinae,

nigricornis (F.):

Comment; The condition of the “type ” specimen

is fair, It has been pinned twice through the right

elytron which is cracked. The antennae are missing

except for the basal joint of the right one, the left

metathoracic leg is also missing. The condition of the

second specimen is much the same as that of the

“type” only the left antenna is complete, and the left

not the right metathoracic leg is missing, There is 4

marked Variation inh eolour and size of the

ferrugineous spots in this species, The ‘type’ is

brassy-black; head ferrugineous with a black trontal

patch. antennue black: thorax brassy-black with

ferrugineous lateral margins; elytra finely punctu-

ated, brassy-black with two ferrugineous humeral

spots placed obliquely; abdomen blue-black, base

and apex ferrugineons, Baly (1855; p, 175) in his

discussion of this genus described a new species P.

cyannipennis, but realised that it was “very closely

allied’ to P. nigricornis (F.). Baly's type in the

BMNH Collection is labelled P. nigricornis (F.)

thd REC. S AUST. MUS I8 (8): 155-197

Weise in Junk’s Catal, (idem) lists P, cyannipennis

Baly as a variety of P. cyanipes (F.),

A study of the large series of this genus in the

BMNH Collection showed that there are a great

many Variations in colour and colour patterns, often

one grading into another, It seems likely that there

are a number of geographical subspecies or varieties

of the different species which have yet to be worked

out,

In the second specimen in the Banks Collection

there is a ferrugineous apical spot as well us the two

humeral spots on the elytra, Pabricius in his original

(1775) description does not mention this so the

specimen labelled “Type” by C, O, Waterhouse

must be the holotype. Fabricius’ description is "C.

ovata, nigro-aenea, capite, thoracis lateribus elyt-

rorumque macula duplici, baseos ferrugineis’—

an oval Chrysomela, brassy-black, head, sides of

thorax, two spots on elytra, base ferrugineous.

NIGRIPES, Crioeeris (Lema)

Fabricius 1775, p. 120, n, 14; 1801, 1, p. 476. n. 30

(Lema)

australis Jac. 1876, p. 807 and 878, p. 152

Junk’s Catal, 1924, 24, S51, p. 48

Zimsen 1964, p. 107, n. 1646

Syntype labels:

i, Fabricius “Croceris nigripes Fab, Entom,

p. 120, n. 14".

i. BMNH “Lema nigripes’, “Type”

Locality: “nova Hollandia’”’

Current Status: Crioveris

somelidae, Criocerinac,

nigripes F.2 Chry-

Comment: The condition of the “type” is good

and all structures are complete but meso and

metathoracic legs are entangled in gluc. The

condition of the second specimen is much the same,

but the ventral surface is not so clean, Fabricius

reclassified this species into the genus Lema in the

1801 publication but since then it has been restored

to original genus. Zimsen only records one specimen

as being in the collection.

NOVAEHOLLANDIAE, Scarabaeus

Fabricius 1775, p. 29, n, 113; L801. 1, p. 57, n. 15

(Ateuchus hollandiae)

hollandiae F, 1781, 1, p. 32;

piceum Hope 184], p. 44 (Tesserodon)

Junk’s Catal. 1911, 19, 38, p. 40 (Tesserodon)

Zimsen 1964, p. 30. n. 173

Matthews 1974, p. &7 (Tesserodan)

Holotype labels:

i, Fabricius “Scarab. N. Hollandiae” FPabr,

Sp. Ins. No. 144.

iw BMNH “Ateuehus N. Hollandiae”’

January. L98L

Locality: “nova Hollandia”

Current status; Tesserodon novachollandiae (F,):

Scarabaeidae. Scarabaeinae.

Comment: The specimen is very dirty. C. O.

Westwood found this type specimen among the

unarranged specimens in the last drawer of the

Bank's Collection. The reference on the label is the

1781 reference indicating that Fabricius re-examined

this Specimen in 1780-1. In this reference Fabricius

omitted the N. for “nova”, although it is on his

label, and gave the specific name as Scarab.

hollandiae. This was copied in his later publications

and in later catalogues. The valid name now is T.

novaehollandiae (P.).

NOVEMMACULATA, Coccinella

Fabricius 1775, p. 81, n. 15; 1781, 1, p. 97, no. 26:

1801, 1, p, 366 and 61,

novempunctate F. 1775, p. 81;

picta Muls. Voet. 1796, p. 61 (Coelophora)

Junk’s Catal. 1931-32, 16, 120, p. 294 (Coelophora)

Zimsen 1964, p. 85, n, 1212

Holotype labels;

i. Fabricius ‘Coccinella 9-maculata Fabr.

Spec. 97 n, 26"

nu) BMNH "Coccinella 9-maculata”’

Locality: “nova Hollandia”’. This species has been

recorded from the Philippines and Indonesia, not

from Australia. It is probable that the “type’’ was

collected in Java, as Banks collected at Batavia when

the Endeavour stopped there for some weeks for

repairs,

Current status; Coelophora novemmaculata (F.)

synonym C. novempunctata (F.). Coccinellidae,

Coccinellinae

Comment: Holotype has been badly pinned

through left elytron, and there is a great deal of glue

on ventral surface obscuring most of the abdomen

and entangling the metathoracie legs, The right

membranceous Wing is unfolded and lying out

beyond the elytra. The apical segments of both

antennae, the right prothoracic leg and the terminal

tarsi of the right mesothoracic leg are missing.

Musgrave (1932; p, 87) in his listings of Fabrician

types from “nova Hollandia’ recorded in the

Species Insectorum (1781), states, C. 9-maculata

(1781: 1, p. 97) nom. nov. for Cece. 9-punctata Fab,

preoccupied by Cocc: 9-punctata Linne 1758".

Fabricius, however, although changing the specific

name, continued to publish both names in the

publication 1781, and in 1787 and 1801 as separate

species, Although the Linnean and Fabrician species

are now in different genera, the valid name tor the

Fabrician species is C. novemmaculata F. according

to the International Code, There is no specimen in

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZRALAND COLEOPTERA TKS

the Bank's Collection labelled 9-punctata F. The

type is marked with two median dark brown or black

spots on the thorax. There is no basal black line nor

is the sutural margin black, There are four black

spots on each elytron and one posterior spot which is

divided so that one half lies on each elytron.

NOVEMPUNCTATA, Coccinella

See NOVEMMACULATA, Coccinella

C, novempunctata is the junior homonym.

OBSCURUM, Callidium

Fabricius 1787, 1, p. 131. n..1, 1801, 2, p. 333, n. 1.

lentiginosus Newm, 1841, p. 7 (Phacodes)

Junk’s Catal. 1912, 22, 39, n, 70 (Phacodes)

Zinsen 1964, p, 178, n, 3008

McKeown 1947, p. 22

Syntype labels;

i. Fabricius “Call.

Ins, n, 1".

iii BMNH “Callidjum obscurum’, “Type”

obscurum Fabr. Mant.

Locality: ‘terra Diemeni”™

Current status; Pachydissus obseurus (F.):

Cerambycidae, Cerambycinae

Comment: The condition of the “type” is good.

There is a second specimen, which is very dirty;

terminal joints of antennae, left metathoracic leg

and all tarsi are missing. This large well known insect

has a wide distribution throughout Tasmania, South

and Eastern Australia,

OCTODECIMGUTTATA, Chrysomela

Fabricius 1775, p. 100, n. 31; 1801, 1, p, 439, n. 101,

keyserlingi Ws. (Phola)

sedecimpustulata Stal. (Phyllophila)

Junk’s Catal. 1924, 24, 68, p. 199 (Chalcolampra)

Zimsen 1964, p, 99, n, 1502

Syntype labels:

i. Fabricius “Chrys. 18-Guttata Fab. Entom.

p. 100 n, 31”

ii, BMNH “Chrysomela 18-guttata’’, “Type”

Locality: ‘“‘nova Hollandia”

Current status: Chalcolampra octodecimguttata

(F.): Chrysomelidae, Chrysomelinae.

Comment; The specimen labelled "Type" is not in

good condition; ventral surface is in a bad state of

preservation; right antenna and all but five joints of

left antenna, tarsi of left mesothoracic and right

metathoracic legs are missing, The condition of the

second specimen is not as good as the ‘type’. Baly

(1855: p. 186) described this species and put it into

its present genus, He notes that there are eight not

nine spots on the elytra, The depth of colour of this

species varies considerably, and the elytral spots

may or may not be distinct. These spots are arranged

in two longitudinal lines. Ou examination of the

large series in the BMNH it is seen that these spots

may or may not coalesce, In the “type” the spots in

the outer or marginal line have coalesced and

therefore definition of the actual number of spots is

obscure, The prothorax in this species varies in

colour from dark ferrugineous to pale, with or

without three black spots arranged medianly in the

form of an inverted triangle, as it is in the “type’’

OCULATA, Criaceris

Fabricius 1775, p. 121, n, 1S (C, oculata); 1801, 1,

p. 458, n, 43

sculpta Bikb 1890, p. 363 (Candezea)

Junk’s Catal. Suppl. 1973, 78, 3, p. 580 (Monolepre)

Zinsen 1964, p, 103, n, 1578

Syntype labels;

i. Fabricius ‘Crioc.

p. 121 n, 15"

ii, BMNH “Crioceris oculata”, “Type”

oculata Fab, Entom,

Locality; “nova Hollandia”

Current status: Lema oculata (F.):

somelidaé, Criocerinac.

Chiry-

Comment: The condition of the ‘‘type"' is poor,

mainly due to the presence of a large amount of glue

on the ventral surface. Right antenna and terminal

joints of the left antenna, the right prothoracie leg

except for coxa, tarsi of the left metathoracic leg and

the right metathoracic leg are missing. The condition

of the 2nd specimen is also jn a very poor state, both

antenna and all legs are missing.

In Junk's Catal, (1924, 24, 51, p. 72) this species

Was incorrectly attributed to Olivier (1791: p, 200),

Fabricius's later references are given but not the

original (1775) one. Olivier attributes tt 1o Fabricius

and gives the original reference in which the name of

the species is spelt octulataia, Fabricius himself

changed the spelling to oculata, “a justified

emendation”, first, in his Species Insectorum (1781)

and then in his later publications.

N.B. Wilcox in Junk's Catalogue Suppl, (idem,)

gives full references and synonyms,

OCULATUS, Siaphylinus

Fabricius 1775, p. 265, n, 4; 1801. 2, p, 592, n, 2

Junk's Catal, 1914, 5, 57. p, 399 (Creaphilus)

Zimsen 1964, p. 230, n, 3982

Steel 1949, p. 59

Syntype labels;

i. Fabricius “Staphyl. oculatus Fab. Entom.

p. 265 n. 4”

186

ii, BMNH *Staphylinus oculatus’’, “Type"

Locahty: ‘nova Hollandia et Zelandia™

Current status: Creophilus oculatus (F.): Staphy-

linidae, Staphylininae,

Comments: “‘Type", a male, has been pinned

twice, previously through the base of the prothorax

and now through the right elytron; terminal joints of

tight antenna are missing, membranous wings are

unfolded and extend out beyond the abdomen.

Second specimen, a female, is pinned through the

left elytron, terminal joints of both antennae and

tarsi of right metathoracic leg are missing,

prothoracic legs are immersed in glue and

membranous wings are unfolded.

See W. ©. Steel (idem.) for description of this

species and table of Australian species of this genus,

PEDICORNIS, Lamia

Fabricius 1775, p. 170, n. 1; 1801, 2, p, 282. n, 4

tuberculata Hope 1841, p. 49 (Rhytophora)

obscura Breuning 1938. p. 363 (Saperdopsis)

bispinosa Breuning 1938, p, 364 (Saperdopsis)

Junk’s Catal, 1922-23, 23, 73, p. 267 (Platyomopsis)

Zimsen 1964, p. 167, n. 2784

McKeown 1947, p, 153

Holotype labels:

i. Fabricius ‘‘Lama pedicornis Fab. Entom.

p. 170 n. 2”

ii. BMNH "Lamia pedicornis’’, ‘“Type”

Locality: “nova Hollandia”

Current status; Platyomopsis pedicornis (F-):

Cerambycidae, Lamiinae.

Comment: The holotype condition is fair:

terminal joints of both antennae and tarsi of left

metathoracic leg are missing, right metathoracic leg

is detached and lying in the glue.

PORCATUS, Carabus

See PORCULATUM, F,, Calosoma

C. poreatus. F. is the junior homonym,

PORCATUS, Notoxus

Fabricius 1787, 1, p. 127, n. 1; 1801, 1, 287, n. 1

cribricollis Spin, 1844, p. 203 (Natalis)

var inconspicus Blk, 1890, p. 124 (Natalis)

Junk's Catal. 1910, 10, 23, p. 37 (Eunatalis)

Zimsen 1964, p. 72, n. 965

Holotype labels;

i, Fabricius ‘‘Not. Porcatus I, Mant, n, 1"

ii, BMNH ‘“Notoxus porcatus”

Locality: “terra Diemenii”

REC. 8, AUST, MUS 18 (&)r 155-197

January, [981

Current status: Eunatalis porcata (F.); Cleridac

Comment: The type has been badly pinned once

through the prothorax and onee through the right

elytron. The specimen has been broken between the

pro and mesothorax and then glued together, the

glue spreading over parts of the elytra and ventral

structures. Both head and legs are covered with dirt.

This specimen would have been collected by David

Nelson who travelled with Cook on his third voyage

when the Discovery called in at Adventure Bay,

Tasmania.

PORCULATUM, Calosoma

Fabricius 1775, p. 239, n, 16 (Carabus porcatus);

1801, 1, p. 211, n. 3

poreatus F. 1775, p. 239,

caraboides Kirby 1818, p, 466. (Adelium)

Junk’s Catal. 1911. 18, 28, p, 510 (Adelium)

Zimsen 1964, p. 60, n. 727

Carter 1926, p. 154

Holotype labels;

i. Fabricius “Carabus porcatus Entom, p

239 n, 16"

ii. BMNH “‘Carabus porcatus’’ A later label

has been added “Tenebrionid”.

Locality; “nova Hollandia”

Current status! Adelium porculatum (F.): (Tene-

brionidae, Ulominae,

Comment: Thé holotype is pinned through the left

elytron which is cracked, Glue conceals most of the

right metathoracic leg. Neither antenna is complete,

3 joints are missing of the left antenna, and 5 from

the right one. Most tarsi are missing and remnants of

the left meso and metathoracic legs are lying in the

glue. E. Zimsen lists Calesoma poreulatum F. with

the 1801 reference as a synonym, In this reference

Fabricius lists the name C. porecatus as the synonym

with the reference of Ent, Supt, 1792, 1, p, 147.

Fabricius himself changed the original name porcatus

to porculatumt in 1801 and this now stands as the

valid name according to the International Code,

QUADRIDENS, Curculio

Fabricius 1775, p. 153, n. 139; 1801, 2, p. 536, n. 175

Junk’s Catal, 1931, 28, 114, p, 21 (Leptops)

Zimsen 1964, p. 219, n, 3787

Holotype labels:

i. Fabricius "Cure, 4-dens Fab, Entom, p.

153 n. 139

ii. BMNH “Curculio 4-dens”

Locality; **nova Hollandia"’

Current status: Leptopius (Oke 1951) quadridens

(F,): Curculionidae, Leptopiinae.

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 187

Comment: The condition of the type’ is good,

There is a second specimen minus its head and

prothorax so I have designated the better specimen

as a holotype rather than a syntype. There is a large

series in the BMNH under the label L. quadridens

(F.), However, these specimens were considered by

G. J, Arrow in 1929 to have been incorrectly

identified, The posterior region of the elytra in the

Fabrician type is vertically declined, while in all the

above-mentioned specimens it is not nearly so

abruptly declined,

QUADRIPUNCTATA, Galleruca

Fabricius 1775, p. 112, n. 4 (Altica albicollis); 1801,

1, p. 499, n. 110

albicollis F. 1775, p. 112 (Altica)

splendidus Jac, 1885, p, 928 (Licyllus),

Synonymy after Bryant 1923, p. 143

Junk's Catal. 1940, 25, 166, p, 7 (Licyllus)

Zimsen 1964, p, 113, n, 1748

Holotype labels:

i. Fabricius “Altica Albicollis Fab. Entom.

p. 112, n. 4"

ii. BMNH “Crioceris albicollis’, ‘'Type”

Locality: “nova Hollandia”’

Current status: Licyllus quadripunctata (F,)-

Chrysomelidae, Halticinae.

Comment; The “type” is in a poor state of

preservation. It is a very small insect and has been

pinned twice, formerly through the right elytron and

now through the suture; left elytron, right and left

antennae except for basal joints, tibiae and tarsi of

metathoracic legs are missing. Fabricius re-classified

this species into the genus Galleruca (1792: p. 29)

and in 1801 he changed the specific name to 4-

punctata due fo the pre-occupation of the specific

name albicollis which he had given to a Brazilian

species. Though both species are now in different

genera, according to the International Code

guadripunctata is the valid name.

QUADRIPUSTULATUM, Sphaeridium

Fabricius 1775, p. 67, n, 6; 1801, 2, p, 575, n, 2

(Seaphidium)

hbimaculatum Mael. 1864, p. 119 (Scaphidium)

Junk’s Catal. 1910, 8, 13, p & (Scaphidiwm)

Zimsen 1964, p, 229, n, 3943

Lébl 1976, p, 287-8

Holotype labels:

i. Fabricius “Sphaerid 4-pustulatum Fab,

Entom. p. 67-6"

ii, BMNH “Type”

Locality: ‘nova Hollandia”

Current status: Scaphidium

(F.): Scaphidiidae

quadripustulatum

Comments: Holotype is a male, synonym

himaculatum Macl, is a female, The condition of the

holotype is fair, both antennae and mesothoracic

legs is are missing. It seems that when the Bank's

Collection was arranged in the order of that in

Systema Eleutheratorum this specimen was not

included, Later it must have been discovered,

perhaps in the last drawer among the unnamed

specimens by C, O, Waterhouse when he was adding

type-labels to the collection and as the label is

Sphaeridium 4-pustulatum he put it in that genus and

added a type-label to it but no other BMNH label. In

1929 when I was studying the collection K, G. Blair

of the BMNH placed it in its correct position

according to the order in the Systema

Eleutheratorum.

E, Csiki in Junk’s catalogue (idem) and in some

literature it has been attributed to Olivier, This is

incorrect and has been corrected by Y. LGbl] (idem)

in his revision of the Australian species of

Scaphidium,

QUADRIPUSTULATUS, Scarabaeus

Fabricius 1775. p. 27. n, 107; 1801, 1, p. 53, n. 105

(Copris)

bipustulatus F, 1775, p, 30

albertisi Harold 1877, p, 71 (Onthaphagus)

minusculus Macleay 1888, p. 903

(Onthophagus)

Junk's Catal. 1927, 19, 90, pp. 215 (Onthophagus)

Zimsen 1964, p, 29, n, 155

Matthews 1927, p. 220 (Onthophagus)

Holotype labels:

i, Fabricius “Scarab. 4-pustulatus Fabr. Sp.

Ins, No, 137"

ii. BMNH “Copris 4-pustulatus’’, “‘Type’’

Locality: “nova Hollandia”

Current status; Onthophagus quadripustulatus

(F.): Scarabaeidae, Scarabaeinae.

Comment! The holotype is badly pinned; bead is

sharply deflexed, right elytron and meso and

metathoracic legs are detached and lying in glue.

The club of the left antenna and some tarsi are

missing. The reference on the label is the 1781

reference indicating that Fabricius re-examined this

specimen in 1780-1. O. bipustulatus (F-) is the

female of O, quadripustulatus (F,) (Matthews idem),

RUFIPES, Gyrinus

see AUSTRALIS, Gyrinus

G, rufipes F. is a synonym of G, australis F,

188

RUFIPES, Helops

Fabricius, 1775, p. 258, n. 5; 1801, 1, p. 161, n. 34,

australis Boisd 1835, p, 282 (Allecula)

angusticollis Boh. 1858, p. 100 (Allecula)

Carter 1915, p. 79, and 1930, p. 269

Zimsen 1964, p. 50, n. 531

Holotype labels:

i. Fabricius “Helops rufipes Fab, Entom, p,

258 n, 5"

i, BMNH “Helops rufipes’’ and the BMNH

Locality: “nova Hollandia”

Current status: Hometrysis rufipes (F.):

Alleculidae

Comment: Holotype has been broken between

meso and metathorax and badly glued together

again so that the anterior part is lying at an angle to

the posterior region, The terminal joints of the

antennae are missing and the right antenna is

detached and lying in the glue. H. J. Carter (idem.)

established that Allecula angusticollis Boh. is a

synonym, Blair (1920: p. 154) referring to this

synonym notes that “this is another name (H. rufipes

F.) that seems to have disappeared from recent

catalogues’’.

SANGUINIPES, Tenebrio

Fabricius 1775, p. 256, n. 7; 1801, 1, p. 148, n, 20

consentanea Perroud 1864, p. 119

depressa Pasc, 1886, p. 454

laticornis Pasc, (Achthosus) 1869, p, 294

Carter 1926, p. 137

Zimsen 1964, p. 46, n. 474

Holotype labels:

i. “Fabricius Tenebrio

Entom. p. 256, n. 7”

ii. BMNH “Tenebrio sanguinipes" and the

iii, Achthosus laticornis Pasc,

sanguinipes Fab,

Locality: “nova Hollandia"

Current status: Uloma sanguinipes (F.); Tene-

brionidae, Ulominae

Comment: The condition of the holotype is poor,

Tt has been broken and glued together at the

junction of the meso and metathorax, All legs on left

side are missing. The holotype is a female and differs

from the male described by Pascoe (idem,) as

Achthosus laticornis in that the prothorax has no

éxcavation, the anterior tibiae are without internal

emargination and it is smaller in size, LU. sanguinipes

(F,) is not listed by H, Gebien in Junk’s Catalogue

bul if is in Carter’s check list (idem.) together with

its synonyms,

REC S. AUST MUS 18 (8); 155-197

January, 181

SCABER, Dermestes

Fabricius 1775, p. 37, n. 16; 1801, 1, p, 318, n, 32

integer Sharp (Ulonotus), new synonym

Junk's Catal. 1930, 15, 107, p, 39 (Ulonatus)

Zimsen 1964, p, 77, n, 1073

Holotype labels:

i. Fabricius “Derm. Seaber Fab, Entom, p,

57, n. Léa”

ii, BMNH ‘Dermestes scaber”

Locality; “nova Zelandia”’

Current status; Ulonotus scaber (F.): Colydiidae

Comment: Specimen is very dirty and pinned

badly through left elytron. Hope (1840: p, 145)

created a new genus Pristoderus for this species.

However, both Lacordaire (1854, p. 359 nota) and

Sharp (1876, p. 17) state that this name should be

dropped as the description given by Hope was

insufficient for indentification, The species is now

classified in the genus Ulonotus Er, see Junk’s

Catalogue (idem.). A. Hetschko in Junk’s Catalogue

lists U, integer Sharp (1877, p. 268) as a separate

species. Sharp’s type is in the BMNH on comparing

this type with D. scaber F. it is found to be

synonymous with it. This is a new synonym,

SCUTELLARIS, Curculio

Fabricius 1781, 1, p, 115, n. 196; 1801, 2, p. 519, n.

exsertus F. 1801, 2, p. 534, n, 163

var, murinus Lea 1911, p. 63 (Prypnus)

Junk’s Catal, 1939, 27, 164, p. 259 (Prostomus)

Zimsen 1964, p. 215, n, 3702

Holotype labels:

i, Fabricius “Cure. Scutellaris Fabr, M, Ins,

n. 196"

i. BMNH “Curculio seutertlaris", Type"

Locality: ‘'Terra Diemenii"’

Current status: Prostomus scutellaris (F.): Cur-

culionidae, Brachyderinae

Comment: Holotype has been pinned twice,

previously through prothorax, now through right

elytron. [ft is in fair condition; both antennae, left

prothoracic leg except for coxa and trochanter, tarsi

and remaining legs except for right mesothoracic leg,

are missing. Al one time this species was placed in

the genus Prypnus. In Junk’s Catalogue C. exsertus

F. is given as a synonym. Fabricius records this

species from ‘nova Cambria’’, and the type as being

in “D. Billardiere”’.

SEMIPUNCTATUS, Curculio

Fabricius 1775, p. 135, n, 43; 1801, 2, p. 200, n. 11

(Lixus)

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 189

cyaneipennis Boh, 1859, p, 118 (Belus)

lineatus Donoy. 1805 (Brentus)

var. poverus Lea 1917, p, 597 (Belus)

Junk’s Catal. 1935, 28, 144, p. 11

Zimsen 1964, p, 210, n, 3615

Holotype labels;

i. Fabricius “Cure. semipunctatus

Entom, p, 135, n, 43”

it, BMNH “Lixus semipunctatus’, **fype”

Fab.

Locality; “nova Hollandia”

Current status: Belus semipunctatus (F,); Belidae

Comment! Holotype is in fairly good condition,

only right antenna and tarsi of right prothoracic leg

are missing, This is 4 well known species. The

number of small whitish tufts of hairs varies

tremendously.

SEMIPUNCTATUS, Stenocorus

Fabricius 1775, p. 180, n. 8; 1801, 2, p, 306, n. 8.

inscripta Germar. 1848, p. 226 (Phoracantha)

Junk’s Catal. 1912, 22, 39, p, 90

Zimsen 1964, p, 172, n, 2901

McKeown. 1947, p. 26

Syntype labels:

i. Fabricius “Stenoc, Semipunctatus Fab.

Entom, p. 180 n. &”

ii, BMNH ~*‘Stenocorus

“Type”

semipunctatus”’,

Locality: ‘“‘nova Hollandia”

Current status: Phoracantha semipunctata (F.):

Cerambycidae, Cerambycinae,

Comment: The “type” is in fairly good condition;

some joints of both antennae and the left

metathoracic leg are missing, There is a second

specimen, this is in much the same condition.

The series in the BMNH Collection show that this

Species varies in size and in the yellow markings on

the anterior half of elytra. It has a wide distribution

throughout Australia and also been recorded from

New Guinea.

SERRATICORNIS, Pyrachroa (Lycus)

Fabricius 1775, p. 203, n. 8; 1801, 2, p, 111 a. 6

(Lyeus)

Junk’s Catal, 1910, 9, 128, p. 73 (Trichalus)

Zimsen 1964, p. 134, n. 2155

Holotype labels:

i. Fabricius *‘Lyc. Serraticornis Fab. Mant.

Ins. n, 5”

i, BMNH “Lycus serraticornis’, “Type”

Locality: “nova Hollandia”

Current status: Trichalus serraticornis (F,):

Lycidae.

Comment: Very little remains of the type and the

pieces have been carded, The left antenna is

complete, there are seven segments of the right one.

The elytra are broken, The legs are missing except

for parts of the left pro and mesothoracic legs, The

abdomen is detached but still has remains of the

wings attached to it. K. G. Blair in 1929 compared

this type with the only specimen then in the BMNH,

which was from Queensland, The reference given on

the label is not the original reference but must have

been put on by Fabricius after he had changed its

genus but before the publication of Mantissa

Insectorum as there is no page number which is p,

164. In this reference he also gives the original genus

Pyrochroa with the 1781 reference.

SMARAGDULUS, Erotylus

Fabricius 1775, p. 123, n. 6; 1801, 2, p, 13, n, 5

(Cnodolon)

cupricollis Hope 1803, omitted from Carter’s list

semtiticus Pasc, 1869, p. 349 (Chalcopterus)

triangularis Haag 1878, p. 104 (Chaleopterus)

vigilans Blkb. 1892, p. 432 (Chalcoprerus)

cairnst Blkb. 1893, p. 60 (Chalcopterus)

Junk'’s Catal, 1911, 18, 28. p, 578 (Amarygmus)

Carter's Check list 1926, p. 162

Zimsen 1964, p. 116, n. 1805

Holotype labels:

i, Pabricius “Erot. smargdulus Fab, Entom,

p. [23 n, 6"

i. BMNH “Cnodulon smaragdulus”, “Type”

Loeality! “nova Hollandia”

Current status: Chalcopterus smaragdulus (F.):

Tenebrionidae, Amarygminae,

Comment; The condition of the holotype is fair;

the mouth parts are covered in dirt and parts of both

antennae, left mesothoracic and both metuthoracic

legs are missing. As in most species of this genus

there is a good deal of colour variation, In this

species it varies from a golden coppery metallic

colour through various shades of purple to greenish

copper, which is the colouring of the holotype.

In Carter’s Revision of the Australian Amary-

ginae (1913: p. 10-11) he noted that Ch, cupricollis

Hope was a synonym of Ch. smaragdulus (F-), the

synonym being determined by K. G. Blair (see

Carter’s reference 1913: p. 11) byt omitted it in his

check list (1926: p. 162). This is another species

which Fabricius re-classified into another genus.

SOLANDRI, Lamia

Fabricius 1775. p. 177, n. 31: 1801, 2, p. 304, n- 126

Junk’s Catal, 1922, 23. 73, p. 269 (Depsages)

1) REC, Ss. AUST

McKeown 1947, p. 160

Zimsen 1964, p. 172, n. 2892

Holotype labels:

i, Fabricius “Lamia Solandri Fabr. Eatom.

p. 177, n, 3t"

ii. BMNH "Lamia Solandri’, “Type”.

Locality: “nova Hollandia”

Current status: Depsages solandri (F.): Ceramby-

cidae, Laminae

Comment: The specimen is badly rubbed, most of

the pubescence on dorsal surface is absent; terminal

joints of right antenna and tarsi of both metathoracic

legs are missing. This specimen Was named after Dr,

C. Solander of the British Museum who went with

Banks on the Endeavour as another naturalist,

Froggatt 1907 p. 199 describes this Cerambyeid “as a

large beetle clothed with a dense coat of buff hairs”.

Due to abrasion of the “type” the punctation is

distinct. Disc of prothorax is rough and rugose with

no distinct tubercles, humeral angle of elytra well

marked, striae indistinct but there are large

scattered punctures diminishing in size towards the

apex.

SPECTABILIS, Curculio

Fabricius 1775, p, 155, n, 144; 1801, 2, p. 537, n, 184

Junk’s Catal. 1936, 28, 150, p, 4 (Chrysolopus)

Zimsen 1964, p. 219, n. 3796

Syntype labels:

i, Fabricius “Cure, spectabilis Fah. Entom,

p. 155, 144".

ii, BMNH “Curculio spectabilis’, “Type”

Locality: “nova Hollandia™

Current status: Chrysolopus spectahilis (F.):

Curculionidae, Aterpinae.

Comments: The condition of the "'type’’ is fairly

good though most of the tarsi are missing. In a

second specimen all structures are complete . It is

probable that the type locality may be Botany Bay as

it is a common species in that region and it is where

Cook made his first landing in Australia. The genus

Chrysolopus was put in the sub-family Cleoninae by

E.. Csiki in Junk’s Catalogue 1931, 28, 134, p, 3, In

1936 S. S. Schenkling and G, A. K, Marshall

transferred it to the sub-family Aterpinae, see Junk's

Catalogue (idem.) with a foot note that belore, il

had been “incorrectly placed in the sub-family

Cleoninae and is now put in its correct position’.

STOLATA, Cetonia

Fabricius 1775, p. 50. n. 33 (Cetonia fasciata); T781-.

p. 58, 0, 45; 1801. 2, p. [53, n. &9

fasciata F. 1775, p. 50

perversa Schaum 1844, p. 371 (Glycyphana)

MLIS 18 (8): (SS-197

January, 1YsT

hrunnipes Kirby 1818, p, 454-478 (Glycyphana),

Tunk’s Catal. 1921, 21, 72, p. 271 (Glycyphana)

Zimsen 1964, p. 142, n. 2312

Bacchus 1974, pp. 113 and 122, (Glycyphana)

Holotype labels:

i. Fabricius “Stolata Ins. n. 45"

ii, BMNH “Cetonia stolata’, Type”

Locality: “nova Hollandia”

Current status: Glycyphana - stolata

Scarabaeidae, Cetoninae.

(Fi);

Comment: The Fabrician label has been mutilated

and only part of it is left. The condition of the

holotype is good and all structures are complete, but

its colour has faded. Fabricius changed the name of

this species from fasciata in the 1775 publication to

stolata in 1781 because the name fasciata was

preoccupied (Master’s 1932: p. &7), The full

reference on the Jabel should be Spec. Ins. p. 58

n, 45. In this reference Fabricius also gives the

original 1775 reference with the name C. fasciata,

M. E. Bacchus (idem,) gives a full discussion and

explains the confusion that has arisen in recognising

this species and ifs synonyms.

STREPIDUS F., Rhynchaenus

see STUPIDUS F., Curculio

R, strepidus F. is a synonym of C. stupidus F.

STRIOLA, Coccinella

Fabricius 1775, p. 79, n. 5; 1801. 1, p, 359, no. 17;

1803, Index p. 32

lineola F, 1775, p, 79

Junk’s Catal. 1931-32, 16, 120, p. 309 (Verania)

Zimsen, 1964, p. 83, n. 1179

Holotype labels:

i. Fabricius “Coce, Lineola Fab. Entom, p.

79, mn, 5”

it. BMNH “Coccinella lineola”

Loeality: “nova Hollandia™

Current status: Verania striola (F,): Coccinellidae

Comment: The condition of the holotype is only

fair. There is a large amount of glue present on both

dorsal and ventral surfaces, both antennae and left

elytron are tnissing. The pro and mesothoracic legs

are covered in dirt and the methathoracic legs are

immersed in glue. There has been some confusion in

the past over the name of this species, In

Gemminger et Harold catalogue it appears as Alesia

lineola (F.). Mulsant (1866: p. 17) deseribed it under

the specific name siriola, given by Fabricius in the

Index L803 und used by Schonherr (1808: p. 156),

Whe put it in his new penus Verania, Korschetsky

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 19}

lists it in Junk’s Catalogue under this genus. Mulsant

recognised the Fabrician name Cocc. lineola F. as

the type name but used the name Y. striola, In his

book (1850; p. 367) he gives no reason for this but

just followed Schanherr’s nomenclature. Both

Zimsen and Ikorschefsky give Cocc. striola F. as a

synonyin of Coce. lineala F. Zimsen records a

second type for Cace. lineola F. The two references

are:—

I. No, 1179 “Coee. lineola F. in nova

Hollandia, Mus. Dom. Banks., synonym

Coccinella striola Index Alphabeticus 1803,

p. 32".

2. No, 1235 “Coec. lineola Ent. Syst, 1, p, 283,

77” in Americac meridionalis [nsulis dam.

Smidt, Syst. El. p. 373.96". The type is in

Copenhagen where there are seven speci-

ments, This species is now placed in the

genus Psyllobora.

Somenmes Fabricius named two species in the

same venus with the same name, then realising his

error he changed the specific name of one. In this

case he changed the specific name lineola to striola

for his later publications, Now even though the two

species named fineola are placed in different genera,

auceording to the International Code the valid name

for this species is striola.

STUPIDUS, Curcullo

Fabricius 1775, p, 139, 0. 65; 1801, 2, p. 473, n. 172

(Rhyachaenus strepidus)

strepidus F. 1801, p. 473

ingrata Faust 1892, p.

synonym

Zimsen 1964, p, 206, 9, 3529

182 (Tentegia) new

Holatype labels:

i, Fabricius “‘Curc. stupidus Fab, Entom p.

139 n. 65"

li, BMNH “Rhynchaenus strepidus”

Locality: “nova Hollandia™

Current status; Tentegia Cur-

culionidae, Cryrtorrhynchinae.

stupida (F.):

Comment: Holotype is in good condition, all

structures complete though terminal joints of right

antenna are hiddén beneath rostrum. It is pinned

through left elytron.

The name of this species has bee’ omitted from

recent catalogues, The type of T, ingrata Faust, is at

Dresden, There are two specimens in the BMNH

identified by Lea as T, ingrata Faust. G. J. Arrow in

1929 verified for me that 7. ingrata Faust. is a

synonym of T. stupida F. This is another species for

which Fabncius changed the name in his later

publications, using the name R, strepidus in Syst. El.

1801. It is possible that this was merely a spelling

error in copying down fram one list to another.

Therefore the synonyms of T. stupida (F.) are

Rhynchaenus strepidus F, and T. ingrata Faust, The

BMNH label R. strepidus would have been added

when the collection was arranged in the order given

in Systema Eleutheratorum,

SULCATUM, Callidium

Fabricius 1775, p, 189, n. 11; 1801, 2, p. 340 n. 41

aculipennis White 1846, p. 20 (Coptomma)

Junk's Catal, 1912, 22, 39, p. 489 (Navomorpha)

Zimsen 1964, p, 179, n, 3037

Syntype labels:

i, Fabricius “Callid. sulcatum Fab, Entom,

p. 189 n, 11"

ii. BMNH “Callidium suleatum”, “Type”,

Locality: “nova Zelandia”’

Current status: Navomorpha

Cerambycidae, Cerambycinae.

sulcaia (F_):

Comment: There are two specimens, a male and

female. The ‘‘type”’ label is on the male, which is

dirty, pinned through left elytron and left

metathoracic leg is missing. The female has several

parts missing—Both antennae, left prothoracic,

right meso and metathoracic legs; left mesothoracic

leg is complete but is detached. The male specimen

is just over half the length of the female, Broun

(1880; p, 590) describes this species fully,

SUTURALIS, Melotontha

Fabricius 1775, p. 34, n- 2; 1801, 2, p, 166, 1, 31

chlorophylla Boisd, 1835, p. 188 (Micronyx)

prasina Cast, 1840, p. 143 (Paranonca)

Junk’s Catal. 1912, 20, 47, p. 89 (Chlorochitan)

Zimsen 1964, p, 145, n. 2361

Holotype labels:

i. Fabricius ‘‘Melolontha Suturalis Fabr. Sp.

Ins, no. 15”

ii, BMNH ‘Melolontha suturalis”’, “‘Type™

Locality: “nova Zelandia”™’

Current status: Chlorochitan

Scarabaeidae, Melolonthinae.

suturalis (F.):

Comment: The condition of the holotype is fair;

except for the Jeft antenna (only 2 basal joints

present), and rhe tarsi of all legs are missing, one set

of tarsi are lying in the dried glue.

Broun (1880: p. 261) describes this species and

Arrow (1903; p, 305) gives the characters of the

genus Chlorochiton and its relation to nearby related

veneta.

SYLVICOLA, Melolontha

Fabricius 1775, p. 39, n. 34; 1801, 2, p. 181, , 123

convexus Boisd, 1835, pl 209 (Lipareirus)

salebrocus Macl. @ 1886, p. 27 (Liparetrus)

192

macleayi Blkb, 3 1888, p. 27 (Liparerrus)

Junk’s Catal. 1912, 20, 47, p. 98 (Liparetrus)

Zimsen 1964, p. 148, n. 2445

Syntype labels;

i. Fabricius “Melolontha sylvicala Fab. Sp.

Ins. n. 57”

i. BMNH “Melolontha sylvicola”, “type”

Locality: ‘nova Hollandia™

Current status: Liparetrus sylvicolus (F.);

Scurabaeidae, Melolonthinae.

Comment; The condition of the holatype is poor:

it is pinned through the left elytron; ventral surface

is covered with a thick felt of dried mycelial hyphae

and glue; Jeft mesothoracic and beth metathoracic

legs are missing, Fabricius must have re-examined

this Species before publishing in 1781 and put the

later reference on the label.

Blackburn (1905: p. 311) in his revision of the

Australian specics of Liparetrus gives the following

synonyms for L. sylvicolus (F):—L. salebrocus Macl.

2.1L. basalis Blanch. 3d and L. macleayi Blkb. 2.

Burmeister (1855: p. 198) describes both the male

and female L, sylvicolus (F,.)— He says, ‘‘nigro

pilasus, pronoto canaliculato, pygidio rugosa variol-

oso carinato’—black, hairy, pronotum grooved,

pygidium rough and variously ridged, Fabricius’s

description states, “Caput et thorax glabra, nigra,

immaculata’—head and thorax glabrous, black and

unmarked, From Blanchard’s descriphon (1850: p.

105) of L. basilis and that of Burmeister’s of L.

sylvicolus (F.) they could be the same species, In the

BMNH there ace eleven specimens labelled L,

hasalis Blanch., the habitat labels are from

Tasmania, South Australia and Nova Hollandia. In

1929 G. J. Arrow compared these specimens with

Blanchard’s and Burmeister’s descriptions and

agreed with me that they were all L. basalis Blanch,

Burmeister’s descriptions were made trom speci-

mens sent to him by Hope. He had both sexes and

they are not L. sylvicolus (F.), but are probably L.

basalis Blanch., which is now regarded as a distinct

species. In Junk’s Catalogue 1912 it is listed as a

synonym of L, sylvicolus (F.). Blackburn never saw

the Fabrician type and was evidently misled by

Burmeister’s description of the specimen he

obtained from Hope, as Burmeister stated that he

had examined the Fabrician types of the Melolon-

thinae in London. Blackburn assumed he would

have studied and described the type of L. sylvicolus

(F.), although in his descriptian Burmeister states

that he obtained the specimens from Hope,

Blackburn did note that Burmeister's description did

not coincide with Fabricius’s description as regards

sculpture and pubescence, but was so sure that

Burmeister's description was of the Fabrician type

that he concluded that Burmeister’s specimens were

REC. § AUST. MUS 18 (8): 155-197

January, 1981

of the same of species. Fabricius had more than one

specimen. there is ane in his own collection in

Copenhagen, as he adds to his description, “variat

interdum elytris fuscus’-—at times there is a

variation in the elytra which may be dark ta

tawny.—The type, be describes as “elytra abdomine

breviora, laevia mgra’’—elytra shorter than abdo-

men, smooth, black—He also states that the type

was in the Banks Collection, but it is possible that he

kept the “type” specimen in his own collection and

returned another specimen, a variety, to Banks. The

specimen, in the Banks Collection has reddish or

fuscus elytra, therefore these two specimens should

bath he regarded as syntypes.

L. basalis Blanch is not a synonym of L. sylvicalus

(F.) but is a separate species. N. B. In Junk’s

Catalogue the species is spelt silvicola,

TOMENTOSA, Lagria

Fabricius 1775, p. 125, n. 9; 1801, 2, p. 70, n, 9

pulchrivaria Lea. 21917, p. 175

Junk’s Catal, 1910, 17, 2, p. 13

Zimsen 1964, p, 128, n, 2037

Holotype labels;

i. Fabricius ‘‘Lagria tomentosa Fab. Entom,

p, 125, n. 9”

ii. BMNH “‘Lagria tomentosa”, ‘“Type”

Locality: ‘nova Hollandia”’

Current status: Lagria s.g. Ecnolagria (Borch,

1936 p. 142) tomentosa F.: Lagriidac.

Comment; The condition of the halotype is poor,

pinned at the sutural margin of left elytron and

cracking it; ventral surface distorted; mouth parts

covered in dirt; both antennae, terminal tarsus of

left prothoracie leg, right prothoracic leg, both

mesothoracic legs and tarsi of both metathoracic legs

are missing. Lea (1917: p. 175) described the male

under the name L, pulcharivaria, Blair (1920: p,

155) determined this synonymy. Champion. (1895; p.

229) pointed out that the species commonly known

as L, tomentosa F. from Western Australia does not

agree with the Fabricius type, it should be known as

EF. aeneoviolacea Champ. In Masters Catalogue

1885, p. 387 the locality is erroneously given for the

Fabrician species as Western Australia

TRIBULUS, Curculio

Fabricius 1775, p, 193, n. 138; 1801, 2. p. 536. n. 174

echidna Macleay 1863-66, p. 280

Junk’s Catal, 1931, 28, 114, p. 23 (Leptops)

Zimsen 1964, p. 219, n. 3786

Holotype labels:

i. Fabricius “Curc. Tribulus Fab. Entom. p.

153, n. 138"

ii, BMNH ‘Curculio tribulus’’, "Type"!

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA

Locality: “nova Hoallandia”

Current status: Leptopius (Qke 1951) tribulus (F.):

Curculionidae, Leptopiinae.

Comment: Holotype is in good condition, only

tarsi of prothoracic legs are missing. There has been

confusion about the identity of this species, G. A. K.

Marshall (1952; p. 265) compared the types of L.

tribulus (F.) and L. ferus (Pasc.), both of which are

in the BMNH and stated that they are separate

species, What A, M, Lea (1906 p, 326) describes as

L. ferus (Pasc.), agrees with the Fabrician type of L.

tribulus. Lea in his table, states that L, ferus (Pasc.)

has “a rostrum carinate in the middle” which applies

to L. iribulus (F.) not to L. ferus (Pasc.) which is

grooved. The description given by Lea of L. tribulus

(F.) describes that of L, duponti(Boisd,), L. duponti

Boisd. is the common species often erroneously

called L. tribulus (F.), and mentioned as such in R.

J. Tillyard’s book of Insects (1926: p, 242), There is

a large series in the BMNH labelled L. duponti

(Boisd,) from Austraha, Two specimens in this

series have been identified by Lea as L. rribulus (F.)

but are definitely L, dupontt (Boisd,). L. duponti

(Boisd.) (1835: p. 333) is given as a female synonym

of L, tribulus (F.) in the Gemminger et Harold

catalogue. [t is not a matter of sexual differences, it

is a distinct species.

Therefore, L. duponti (Boisd.), L. tribulus (F,),

and L, ferus (Pasc,) are distinct species. L. echidna

(Boisd.) is given as a synonym in Dejean’s

Catalogue and Junk’s Catalogue and ts incorrectly

attributed to Macleay. The specimen in the BMNH

labelled L. echidna (Macleay) (Boisd.) is the same

as L, dupont (Boisd.) not L. ribulus (F,).

TRIDENS, Curculio

Fabricius 1787, 1, p. 122, n, 271; 1801, 2, p. 537, n.

186

sextuberculats White 1846, p. 13 (Rhinaria)

Junk’s. Catal, 1935, 28, 144, p. 2 (Agathinus)

Zimsen 1964, p. 219, n. 2798

Kuschel 1970, p. 197

Holotype labels;

i. Fabricius ‘Cure, Tridens Fabr. Mant, Ins,

n. 271”

ii, BMNH “Curculio tridens”, ““Type’’ There

is another label but it is unintelligible. [t

may read “‘variety sexual of C. expor-

ti—Mus Dom Labrielle”’

Locality: “nova Zelandia”’

Current status: Agathinus tridens (F.): Belidae.

Comments: Holotype is a female, Synonym

sextuberculatus White is a male (Kuschel idem,)

Holotype has been pinned twice, now through left

elytron, previously through prothrorax; most of the

193

antennae, and all tarsi except those of the left

mesothoracic and right metathoracic legs are

missing.

TRISTIS, Necydalis

Fabricius 1787, 1, p. 170, n. 5; 1801, 2, p. 370, n. 14

miranda Newm. 1851 App., p. 133 (Dohrnia)

anthina Olliff 1887, p. 154 (Ithaca)

Zimsen 1964, p. 185, n. 3148

Holotype labels:

i. Fabricius “‘Necy, Tristis Fabr. Mant. Ins.

n 5"

ii. BMNH “Necydalis tristis”, Type”

Loeality: “terra Diemenii"’

Current status: Dohrnia tristis (PF): Qedemeridae.

Comment: Only the abdomen of the holatype

remains. This species has been omitted by

Schenkling in Junk’s Catalogue 1915, 17, 65, p. 72

and the two synonyms are listed as separate species.

Blair (1929; p. 158) determined the above synonymy

but it has been overlooked by subsequent workers of

the species, Blair states, ‘“‘Unfortunately all that

remains of the type is the abdomen attached to the

pin. The description in conjunction with Olivier's

figure, leaves no doubt that the insect was the female

of the species better known us Dohrnia miranda

Newm., and an examination of the abdomen makes

this identity certain. Olliff evidently did not know

Newman’s insect, but his description is. so full and

detailed as to leave the synonymy beyond question.”

This reference is omitted from Musgrave's Bib-

liography of Australian Entomology, Also R. J.

Tillyard used the name D. miranda Newm. in his

book ‘Insects of Australia and New Zealand” an

p. 225.

TRUNCATUS, Scarahaeus

Fabricius 1775, p. 4, nm. 12; 1801, 1, p. 7, a. 18

(Geotrupes)

Junk’s Catal. 1937, 21, 156, p. 58 (Pericoptus)

Zimsen 1964, p. 21, n- 12.

Holotype labels:

i. Fabricius “Scarabaeus Fabr, Sp. Ins, No.

16”

i. BMNH ‘Geotrupes truncatus”

Locality: “nova Zelandia’ as recorded in 1775

publication.

Current status: Pericoptus truncalus (F.);

Scarabaeidae, Dynastinae.

Comment: The condition of the holatype ts good.

only the club of the left antenna and the tarsi of the

metathoracic legs are missing. The reference on the

label indicates that Fabricius re-examined this

specimen in 1780 and put the 1781 reference on the

label. At the same time he must have recorded the

locality erroneously as ‘nova Hollandia’ which was

copied in his later publications.

TUBERCULATA, Cicindela F-

Fabricius 1775, p. 225, n. 7: 1801, 1, p. 238, n. 32.

tuberculosa Ol. 1790, p. 732

latecincta White 1846, p. 1

Junk’s Catal. 1926, 1, 86, p. 200.

Zimsen 1964, p. 64, n. 800.

Holotype labels:

i. Fabricius ‘‘Cicindela

Entom. p. 225 n. 7”

ii. BMNH “Cicindela tuberculata”’

tuberculata Fab.

Locality: “nova Zelandia”

Current status: Cicindela tuberculata F.; Cicin-

delidae.

Comment: The condition of the holotype is fair. It

is pinned through the left elytron. Only the three

basal joints of the antennae are present, the right

mesothoracic leg is entirely missing and none of the

others are complete.

There is a second specimen labelled “nova

Zelandia, Nel.”. This specimen is without head, pro

and mesothorax. ‘‘Nel!."’ would stand for Nelson, the

collector who sailed with Cook on his third voyage.

This specimen would, therefore, have been collected

at a later date than the holotype.

E. Zimsen records that 2 specimens are in the

Banks Collection and one at Kiel but incorrectly

records the locality as ‘nova Hollandia’’. In both the

1775 and 1801 references Fabricius records the

locality as ‘nova Zelandia™. Tt is a New Zealand

species.

UNIFASCIATA, Crioceris (Lema)

Fabricius 1775, p. 120, n. 13; L801, 1, p. 474, n. 28

(Lema)

Junk’s Catal. 1924, 24, 51, p. 82 (Lema)

Zimsen 1964, p. 107, n. 1644

Holotype labels:

i. Fabricius “Crioc. Unifasciata Fab, Entom,

p, 120 n. 13,

ii. BMNH “Lema unifasciata’, “Type"’.

Locality: “nova Hollandia”

Current status: Lema' unifasciata F. Chry-

somelidae, Criocerinae.

Comment: Holotype is pinned through left

elytron; left antenna, and terminal segment of right

antenna are missing. Femora and tibia of left

mesothoracic leg and both metathoracic legs are

entangled in the glue. Fabricius re-classified this

species into the genus Lema in 1801.

REC, S& AWST MUS 18 (4); 155-197

January, 1987

VARIEGATA, Lepiura

Fabricius 1775, p. 199, n. 19; 1801, 2, p. 375, n. 2

(Molorchus)

Junk’s Catal. 1912, 22, 39, p. 286 (Hesthesis)

Zimsen 1964, p, 186, n, 3163

Type specimen has been lost from the Collection

Locality: ‘nova Hollandia’”’

Current status: Hesrhesis variegata (F.): Ceramby-

cidae, Cerambycinae.

Comment: There is a large series in the BMNH

General Collection, McKeown (1947: p. 89) gives a

complete list of references but he erroneously

records the type as being in the BMNH. This

specimen was re-classified by Fabricius in 1801 into

the genus Molorchus.

VARIEGATUM, Callidium

Fabricius 1775, p. 189, n. 9; 1801, 2, p, 340, n, 39

viratum Newm. 1841, p, 18 (Coptomma)

Junk’s Catal, 1912, 22, 39, p. 488 (Coptomma)

Zimsen 1964, p. 179, n, 3035

Syntype labels:

i, Fabricius “Callid variegatum Fab, Entom.

p. 189, n, 9”

ii, BMNH “Callidium variegatum"”, ‘‘Type"

Locality; “nova Zelandia”’

Current status: Coptomma variegata (F,):

Cerambycidae, Cerambycinae,

Comment: There are two specimens a male and

female, The male specimen has the Museum “type”

label. It is pinned through the left elytron; neither

antenna is complete and the legs are entangled in

glue on the ventral surface. The condition of the

female is in an excellent state of preservation and all

structures are complete. Broun (1880; p. 589) gives a

detailed description of this species,

VILLOSA F,, Saperda

see HIRTA F. Saperda

§, villosa F. is a synonym of S, hirta F.

VIOLACEUS, Notoxus

Fabricius 1787, 1, p. 127, n, 1; 1801, 1, p. 287, n. 2

niger Sharp 1877, p. 7 (Balcus) new synonym

Junk’s Catal. Suppl, 1950, p. 297 (Phymatophaea)

Zimsen 1964, p. 72, n, 966

Holotype labels:

i. Fabricius ““Notoxus violaceus Fabr, Mant.

Ins. n. 2”

ii. BMNH ‘“Notoxus voilaceus™

THE FABRICIAN TYPES OF THE AUSTRALIAN AND NEW ZEALAND COLEOPTERA 185

Locality; “nova Zelandia™

Current status: Phymatophora violaceus (F.):

Cleridae

Comment: Ventral surface is covered with dirt

and all legs ure closely intertwined. The type of

Balcus niger Sharp is in the BMNH, A comparison

of this type with N. violaceus F. shows them to be

synonymous. This is a Tew synonym and was

confirmed by K. G. Blair when | was working in the

BMNH in 1929, A study of the series of this species

in the BMNH shows that this species varies

considerably in size. colour and markings on the

elytra, which may be three pale spots, two, one of

none at all. The Fabrician type has a metallic violet

tinge and three pale sports om the elytra, tibiae, tarsi

and hase of the femorae are ferruginous. The type af

B. niger Sharp is black and there are no colour spots

on elytra; tibiae and tarsi are black. The Fabrician

type is devoid of hairs, probably due to abrasion,

while the Sharp type is scantily covered with hairs.

ACKNOWLEDGEMENTS

Many thanks go ta Dr G, F, Gross and Dr E, G

Matthews al the South Australian Museum and Mr

M. S. Upton of the Division of Entomology,

C.S.1.R.0., in Canberra, who read the manuseript

and have given me valuable help and advice, and

also to the authorities of the South Australian

Museum, especially to Dr G. F. Gross, who made it

possible for me to use the facilities in the

Entomology Section, | also thank Dr E, C,

Zimmerman, and Dr E. G. Britton of the Division

of Entomology, C.S.1.R.0., for their help and

advice. My thanks also go to my husband for his help

in checking references, and translating many of the

original Latin descriptions. In the final stages, on my

return to London to complete the study, I thank the

authorities of the British Museum (Natural History)

for allowing me to work there once more, and in

particular 1 thank Mr R. D. Pope and Mr R, T,

‘Thompson.

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SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

1. NOTATION 2. BIFEMORATA AND PTYCTIMA (CRYPTOSTIGMATA)

BY DAVID C. LEE

Summary

A study of sarcoptiform mites from surface soil (usually greatest depth = 4cm) at 9 florally diverse

sites in South Australia is introduced. A modified notation for mite morphology is presented. The

following 6 species of Bifemorata or Ptyctima were collected and are either newly described or

annoatations are given: Stomacarus abresi n.sp., Loftacarus siefi n.gn., n.sp., Ctenacarus araneola

Grandjean, Protoplophora palpalis Berlese, Hoplophthiracarus shealsi n.sp. and Rhysotritia

wallworki n.sp. Some relevant higher taxa are redefined. New synonyms are Aphelacaridae under

Adelphacaridae, Ctenacaridae under Palaeacaridae and Andacarus under Stomacarus. New

combinations are campbellensis, ligamentifer and watsoni (ex Andacarus) with Stomacarus,

longicaudatus (ex Stomacarus) with Loftacarus and africanus (ex Ctenacarus) with Beklemishevia.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS.

1, NOTATION.

2, BIFEMORATA and PTYCTIMA (CRYPTOSTIGMATA)

By DAVID C, LEE

South Australian Museum, Adelaide, South Australia 5000

ABSTRACT

LEE. D, C. 1980), Sarcoptiformes (Acari) of South Australian

soils |. Notatian. 2, Bifemorata and Ptyciima (Crypto-

stigmuta). Ree. 8. Aust. Mus, 18(9)) 199-222,

A study of sareoptiform mites from surface soil

(usually greatest depth = 4 cm) at 9 florally diverse

sites in South Australia is introduced. A modified

notation for mite morphology is presented. The

following 6 species of Bifemorata or Ptyctima were

collected and are either newly described or

annotations are given! Stomacarus abresi n.sp,,

Loftacarus siefi n.gn., n.sp., Ctenacarus araneola

Grandjean, Proteplophora palpalis Berlese.

Hoplophthiracarus shealsi n.sp. and Rhysotritia

wallworki n.sp.. Sofne relevant higher taxa are rede-

fined. New synonyms are Aphelacaridae under

Adelphacaridae, Ctenacaridae under Palaeacaridae

and Andacarus under Stomacarus. New combin-

ations are campbellensis, ligamentifer and watsoni

(ex Andacarus) with Stomacarus. longicaudatus (ex

Stomacarus) with Loftacarus and africanus (ex

Ctenacarus) with Beklemishevia

INTRODUCTION

The aim of this study is to give a broad (yet,

because of resources, necessarily superficial) indica-

tion of the sarcoptiform mite (Cryptostigmata and

Astigmata) fauna of South Australian soils. This

group was selected because it is startlingly poorly

known in Australia and because its members, with

two small groups of mites (Notostigmata and

Endeostigmata), are the only arachnids capable of

ingesting solid particles.

The limited extent of knowledge about Australian

Cryptostigmata 3s illustrated by comparing the

known mite fauna of Australia and that listed from

New Zealand by Spain and Luxton (1971). Of the

832 species of New Zealand mites, 380 species are

Cryptostigmata. Whilst, of approximately 1 300

species of Australian mites, 19 species are

Cryptostigmata. This situation will be partially

rectified by a study of Cryptostigmata of the Pacific

area by Professor Janos Balogh and Dr. Sandor

Mahunka of Hungary. It is to be pubhshed in the

near future, and refers to 200 or more species from

Australia, mainly from east coast forest regions

(personal communication). As elsewhere, the

Astigmata of Australian sails are neither as

abundant nor as diverse as the Cryptostigmata, In

fact the only abundant species found in this study,

and then anly at the pasture site, was the

casmopolitan Tyrophagus similis Volgin.

The Cryptostigmata and Astigmata in soils feed

on decomposing plant material, fungi, algae,

bacteria and frass, Solid particles are fragmented

and ingested, and are evident as relatively opaque

boli in cleared specimens. Unlike purely Jiquid-

ingesting mites they deposit faecal pellets similar to

the frass of some insects and containing living micro-

organisms. In other wards, this group mechanically

breaks down plant material, metabolize some of it,

and feeds on and disperses organisms that also

metabolize it. Furthermore, it is likely that the food

preferences of particular species are limited, some,

for example, feeding on only fungi, or even only

certain fungal species. Therefore a study of

variations in their fauna between different sites is

pertinent to assessing variations in the decomposing

pracesses that Jead to nutrients being made available

to growing plants,

During the sorting of samples neacly 30 000 adult

Cryptostigmata and Asligmata were separated oul

and grouped in about 120 species. Relatively few

immatures were collected and these are referred to

only when they can both be identified and belong to

4 species that would not otherwise be represented

from a particular site,

The author intends to classify the species in a

number of publications and then comment on their

distribution.

The higher classification used will be detended

only as it deviates from that in “The oribatid genera

of the world’ by Balogh (1972). Since Balogh’'s

classification, although comprehensive, does nat

answer the plea by Woolley (1971) for improvement

in the ill-defined and complex system of classifica-

tion for the Cryptostigmata, definitions of higher

taxa will be presented and there will be a bias

towards simplification by reducing the number of

supecfamilies and families.

All material is deposited in the South Australian

Museum, Adelaide.

This particular publication presents the notation

used for mite morphology and collection data. Then

the six species of mites collected that belong to the

Bifemorata and Ptyetima are classified and

described when necessary,

200

NOTATION FOR MORPHOLOGY

Unless specified, attributes belong to adults.

Much of the description of the soma is dealt with

under four headings related to the areas illustrated

(Fig. 1). The term external mala (see Hughes 1959:

139) is used rather than ‘rutellum’, and spermapositor

rather than ‘penis’ since the relevant structure is not

an intromittant organ. The term cowl is used for a

proteronotal rostral hood which encompasses the

retracted gnathosomal appendages, a condition

previously referred to as ‘stegasimy’. A partly new

notation for hairs, chaetotaxy and pores is presented

below, plus that for mensuration.

REC. S. AUST. MUS, 18 (9): 199-122

January, 1981

Hairs

Hairs are regarded either as setae if they have

actinochitin and are hollow with no protoplasmic

core, solenidia if they have no actinochitin and a

protoplasmic core, plasmic setae (‘eupathidia’,

‘famuli’ and ‘sensilli’) if they have both actinochitin.

and a protoplasmic core. All hairs examined in this

study exhibited only peripheral birefringence under

polarized light, so they apparently always lack an

actinochitin core. Solenidia usually occur dorsally on

the distal segments of appendages, and are all

referred to by the symbol so, and numbered, the

more proximal and anterior solenidia first.

HYSTERONOTUM Oe eae tole

OPISTHOSTERNUM ~— \

~ jaF

PROTERONOTUM

_- GNATHOSTERNUM

4 a: ™

IDIOSTERNUM

Chaetotaxy of Soma

Up to four systems of notation for dorsal setae of

Cryptostigmata have been used in a_ single

publication, and a fifth system is usually used for the

Astigmata. This has sprung from a desire to link the

systems of notation with only confident proposals of

homology. On the other hand, confusion results

from multiple systems, and this has led me to follow

a single system with less stringent proposals of

homology. This system was initially applied to the

opisthonotal shield of species of Zercon by Sellnick

(1944) and then modified and extended to the notum

of other members of the Gamasina (Mesostigmata)

by Hirschmann (1957) and Lindquist and Evans

(1965). Amongst sarcoptiform mites it has been used

only for the Pelopoidea (Cryptostigmata) by

Schweizer (1956) and Hammer (1972). It is based on

a grid in which the ranks are represented by numbers

and the files by letters derived from German words

(J, inner = central; Z, zwischen = between; S,

seiten = lateral; R, rand = edge). Capitals are used

for hysteronotal files and lower case letters for

proteronotal files. The complete dorsal chaetotaxy

(holotrichous condition) is regarded as three pairs of

~~. PODOSTERNUM ~~

Fig. 1—Zones of soma surface.

aft

PROSTERNUM

files with two proteronotal ranks and six hys-

teronotal ranks. Further setae may be present: if

lateral to file S, and possibly representing a further

file, they are referred to file R; if they clearly reflect

a hypertrichous condition they are represented by

the notation for the file lateral to them combined

with ‘x’ as in Jx. When only a few setae are missing

compared with the holotrichous condition, the

absence of a setal pair is illustrated by a circle and

superimposed cross in order to indicate the

numbering used in the text. For example, in Fig. 2,

the third seta in file Z is Z4 not Z3, Z3 being

regarded as missing and represented by a circle and

cross. Also on illustrations, a number as a prefix to

the file signature, for example as in 5Z, indicates

how many setae are present in the file.

The ventral setae are either strictly somal setae or

they are born on coxal regions which merge in with

the soma. The opisthosternal setae belong to the

former group and have notations similar to those of

the hysteronotal setae except that they are regarded

as either aggenital or adanal, the g or arepresenting

one or other group being prefixed by either J, Z or

S. The genital setae, those actually on the ovipositor

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS elt]

or spermapositor, are either proximal (pg), median

(mg) or distal (dg) and are numbered from the

anterior. The podosternal setae are labelled by a

roman numeral representing the leg articulating with

their coxosternal zone, sometimes followed by a

number representing ranks of which the first is at the

adaxial end. and then by either @ (anterior), mm

(median), p (posterior) or d (dorsal). The

gnathosternal setae are regarded as adoral (ao),

postoral (po), palp coxosternal (¢) and supracoyal

(cd), the numbers and letters applied to palp

coxasternal sefae are as for the podosternal setae,

Chaetotaxy of the Appendages

The widely used notation for the leg setae of

sarcoptiform mites is based on french words such as

‘unguinale’ and ‘fastigiale’. It was. devised by

Grandjean in 1935 and later modified ta make the

notation for the lateral surfaces of legs I and II

match the homologous surfaces of legs TIT and IV

(Grandjean 1940a). Because the system is esoteric

and linked to debatable proposals of homology, T

have used a system which is a manifest reference to

setal position. Although in some instances I have not

been able to rigidly keep to this, For example, the

proximal dorsal seta, dl, on tarsus 1 (‘famulus’) has

been regarded as occurring in different posidons

(compare Figs. 7, 27 and 42).

The widespread convention athongst Specialists

deseribing characters on arthropod appendages is 10

refer to their position as if on a cylinder-like segment

and to use co-ordinates of two signature systems

based on their location bath an a hypothetical

circular cross-section of the cylinder and on the

straight proximal-distal axis. Two such notations

have been ignored by acarologists, One was

proposed by Newell (1956 and 1957) for Oppia

(Cryptostigmata) and the other by Southcott (1961)

for the Erythraeoidea (Prostigmata), Qn the other

hand, a notation that dramatically improved

methods of distinguishing families of Gamasina,

proposed by Evans (1963), is now used generally for

the Anactinochaeta, The Anactinochaeta is the

other of the two major groups of mites, Sarcop-

tiform mites belong ta the Actinachaeta which have

optically active chitin in their setae. Amongst the

Anactinochaeta a single whorl of setae al a

particular position on the proximal-distal axis is

considered (Evans 1963) ta be basically made up of 6

setae (2 dorsal, | anterior, 2 ventral, 1 posterior),

Amongst the Actinochaeta a single complete whorl

is Considered as including 7 setae (1 dorsal, 2

anterior, 2 ventral, 2 posterior) as an tibia | of

Palaeacarus (see Grandjean 1940b; Fig. 1). Qn the

other hand the ‘fastigiale’, ‘tectale’ and ‘itérale’

setae on tarsus T (see Grandjean. 1954c; Fig. SA) are

apparently 3 pairs of dorsal setae. Therefore, 1 am

repardinp a single whorl as including setae that cari

be referred to one of 8 positions (2 dorsal, 2

anterior, 2 ventral, ? posterior), although a complete

whorl of § setae has not be deseribed, and no claim is

made here that such a whorl is a lost primitive

condition, The notation used for the eight positions

is ad, pd; dp. vp; pu, av; va, da_as illustrated (Fig.

19). In each pair of letters the last one represents the

major area, Tf there is only one seta present in a

particular quarter of the circumference then only

this last letter is piven. Since paired dorsal setae only

aceur in the absence of paired anterior and paired

posterior setae, itis possible that one or both of the

paired dorsal setae fad_ pd) are homologous with a

dorso-anterior (da) or 4 dorso-posterior (dp) seta or

both on other segments. The apparently dorsal setae

on the tarstis, for example. having migrated

dorsalwards from a primitive lateral location. The

whorls are numbered fram the proximal to the distal

end, tarsus I is regarded as having 4 whorls, or a

maximum potential or 32 setae. Whilst there are

usually considerably fewer than 32 setae, members

of the Bifemorata may have more, a condition Which

is regarded here as hypertrichous.

Pores

Balogh (1972) classifies pores into the following

groups (notation in parentheses); slit-like pores (ia.

ip, ips, th); reduced sacculi pores (Pa, P;, P;, P;):

sacculi (Sa, S;, §,, S;); areae porosae (Aa, Ay. As,

Ay, Ad, Al, App). Because some of these letters are

already in use, | am using ‘f' (foramen) as a signature

for all pore types. Slit or punctiform pores are

prefixed by ht (hysteranotal), Za (lateral adanal) or

Ja (central adanal). Sacculi, which are always

hysteronotal are prefixed by s and similarly area

porosae by m (multiporose area), A number

following the signature equals the rank of a

particular pore and a number before indicates. how

many pores are jn the file, The pore of the lateral

hysteronotal gland duct is referred to as hGf.

Mensuration

Measurements are in microns (4m), The

idiosomal length is an average of the number of

specimens which are jndicated afterwards in

parentheses with their range of lengths. For

inembers Of the Ptyctima, bysterosamal and

proterosomal lengths are given separately. The

idiosomal length of the holotype, or about average

size specimen for established species, from Which the

appendage measurements were taken, is given in

parentheses befare these measurements. Lengths for

the chelicera are the longest axis of the movable digit

and for the palp and legs are the distances from the

base of the trochanter to the tip of the tarsus. The

breadths are the greatest width of the femur or, in

202 REC. S, AUST. MUS 18 [9); 199-122

the case of the Bifemorata, the telofemur. In the

case of the mites longer than 250 jam measurements

are to the nearest 5 pm for most lengths and nearest

2.5 wm for the cheliceral length and all breadths, For

mites under 250 jum in Jength the measurements are

to the nearest 2-5 wm for most lengths and the

nearest 1-0) wm for the cheliceral length and all

breadths.

NOTATION FOR DISTRIBUTION DATA

Under the heading of ‘Material examined’,

callectian data and registration numbers are given

for all mites examined in detail. Under ‘Distribution’

a wider indication is given of where 4 species is

known to occur using a notation for geographical

location, South Australian sites that are part of this

study, and numbers of mites and their distributian

amongst samples. This notation is explained under

three headimgs: ‘samples’, ‘sites’, ‘geographical

location’. The sampling process and sites will be

considered in more detail in the final paper of this

series that will be concerned with the distribution of

the relevant mites.

Samples

Samples were collected in plastic bags in a

manner similar ta a method | previously used (Lec

1973), They were, therefore, disturbed and would

not have yielded such a high proportion of their mite

fauna as undisturbed core samples. The top 4 cms of

soil, plant litter, moss and other plants with a similar

growth form were collected from 25 ® 25 cm

squares. An exception to this was the Knoit Hill

cultivated pine forest site from which, because of the

unusual depth of the litter, four'samples of different

layers to. a depth of 16 cms were taken from each

25 * 25 cm square, Usually samples of eight 25

25 cm Squares were taken from each site, so that the

number of adults of a particular species of mite ata

site represents an extraction from half af a square

metre of soil surface area, but collections from only

two 25 = 25 cm Squaces were taken from the Knott

Hill cultivated pine forest site.

The numbers given after the site, for example 10

(5/8), equal the number of adults collected and, in

parentheses, the number of 25 * 25 cm squares of

soil surface that the adults were taken fram as a

fraction of the total number sampled (usually &, but

2 in the case of the pine forest site).

Sites

Sites were selected in order to represent floral

diversity using the description of South Australian

vegetation by Specht (1972). They also present a

wide range of geographical locations. All nine sites

are given a brief description below, The mites

January. 19871

referred to in this publication were only callected

from five of the nine sites, but the value of the

negative results is emphasized. No attempt has been

made to register seasonal variations in populations.

in all instances sampling was done when the ground

was relatively damp and therefore the sarcaptiform

mite population was presumably relatively large.

The sites with rhe lowest rainfall are listed first for

those with native flora, whilst the two final

cultivated sites have the one with the highest rainfall

listed first. Annual rainfall figures given below are

ranges of average annual rainfalls that include such a

value for a particular site. Prior to cultivation the

flora of the cultivated sites would haye been similar

to the savannah woodland in the case of pasture and

sclerophyll forest in the case of pine forest.

. Arid tussock graysland: Vietoria Desert, 28° 41'S, 132°. 08' E.

Bases of love grass (Bragrosfis eriopda) tussocks amongst

patches of tall mulga (Acacia aneura) shrubland, on deep

red silicequs sand of swale between ridges. Annus rainfall:

[25-150 mm, sporadic 11.10.1976.

2. Semi-arid Jow shrubland: Koonamore, 32° 07’ §, 139° 21' B.-

Litter, muss and other law growth plants under bladder

saltbush (Avriplex vesicaria) amongst false sandalwood

(Myaporum platycarpum) low open-woodland, on rolling

plain, with sandy clay loam on limestone pan, Annual

rainfall! 750-200 mm, sporadic. 27.4.1974,

3. Mallee-broombush open-scrubland Ferries-MeDonald, 35°

15' S. 139° 09' E. Litter and sparse moss, under ridge-

fruited mallee (Eucalyprus incrassala) clamps amongst

broombush shrubs (Melaleuca uncinata) on low hill with

shallow caleareous sandy soil and sparse clayey subsoil on

limestone pan. Annual rainfall: 350-4) mm. mainly

winter, 20.6,1974.

4, Mallee-heath tall open-shrubland> Tamboore, 35° 57'S. 140°

29’ E. Liner under banksia shrubs (Banksia ornata)

amonest sclerophyllous shrubs and sparse brown stringy

back mallee (Bucalyptus baxter’) on tidge in area of deep

caleareous sand. Annual rainfall: 450-500 mm, mainly

winter. 4.71974,

5. Coastal closed-scrubland: Piccaninnig Ponds, 38" 03" S, 14uU°

57’ E Litter and sparse grass under coastal wattle (Acacia

sophorae) with little light beneath canopy on black

calcareous sand at landward edge of coastal dunes,

sometimes inundated from permanent pond fed by fast

flowing underground strearns trom limestone hills. Annual

valotall: 700-750 mm, mainly winter. 3,7.1974 and

20,8 1975,

6, Savannah woudland: Chambers Gully, 34° 58’ 5, 138° 41' E.

Fither litter under manna gum trees (Eucalyptus viminalis)

or grass and moxs in between these trees on shallow red

brown loam of gully slopes in Mt. Lofty foothills. Annual

rainfall) 900-950 mm, mainly winter. 12.6,.1974.

7. Sclerophyll forest! Mt. Lofty. 34" 59° 8, 138° 45’ B. Litter and

sparse oioss under Sclerophyllous shrubs amongst

messmate Stringybark (Eucalyptus ebligua) in furest near

summit of Mt. Lofty on shallow, grey siliceous sand,

Annual rainfalf; 1150-1200 mm, mainly winter

9.51974,

&. Pine foresr Knott Hill, 357 12’ S, 138° 41° E. Litter under

cultivated Pinus pinea with little light beneath canopy on

siliceous sandy soil on rolling hills in Mt. Lofty Ranges,

Annual rainfall: 950-L000 mm, mainly winter.

22.5.1974.

9. Pasture: Glenthorne, 35° 02' $, 138° 32’ E. Bases of cultivated

Brass and plantain (Plantago lanceolata) on deep brown

loam on rolling hill in foothills of Mt. Lofty Ranges.

Annual rainfall: 500-550 mm, mainly winter.

12.6. 1974.

Geographical Lacation

Geographical regions are represented by 2 or 3

letters, the first af which, in capitals, equal the major

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

region, They are as follows: Nearctic (Nn—north-

ern. Ne—calilornian, Nr—rocky mountain,

Na—alleghanian); Neotropical (NTm—mexican,

NTa—antillean, NTb—brazilian, NTe—chilian);

Ethiopian (Ew—west, Ee—east. Es—south,

Em—malagasian); Palaearetic (Pe—european.

Pm—medilerranean, Ps—siberian, Pe—manchu-

rian); Orjental (Oi—indian, Oe—ceylonese, Os—

indochinese, Om—indomalayan); Australian

(Am—malayan, Aa—australian, Ap—polynesian.

An—newzealandian); subantarctic (Sm—magella-

nian, Sk—kerguelenian. Sa—antipodean); Antaretic

(ACr—rossian, ACs—scotian). For details of

divisions between these regions see the map thar I

have previously published (Lee 1970: p. 207).

SYSTEMATICS

Cohort BIFEMORATA

Diagnosis; Macropylina (Balogh 1972: 31). Pale

adult with weakly delineated shields not completely

encasing soma. Rostrum never longer than distance

between setae jl-/] and does not form cowl capable

of encompassing retracted gnathosomal appendages.

Genital and anal shields together almost as long as

opisthostermum, and never separated by more than

length of seta Jal. No conspicuous idiosternal

division. Coxae distally free from podosternum-

Femur divided into basifemur and telofemur. Either

tarsus I or tibia | or both with 4 solenidia, Between

20 and 40 setae on tarsus I. Immatures similar to

adult.

Remarks; The classihcation by Balogh (1972)

follows that of Grandjean (1969) to include six

families grouped in three superfamilies, but the

listing of Beklemishevia (under Ctenacarinac below)

and Tragardhacarus (under Palaeacarinae below)

under Aphelacaridae is original and probably

constitutes a typing error, [ have reverted to an

earlier classification (Grandjean 1954c) in regarding

the three superfamilies as families, But in

introducing new characters to delineate these

families, 1 have grouped Clenaearinae under

Palaeacaridae rather than with the Adelphacarinae.

There are also less important changes in the

classification. Whilst members af the Acaronychidae

and Palaeacaridae were collected in the present

study and are considered below, no adelphacarids

were collected so they are briefly considered here.

Adelphacaridae (Adelphacarinae Grandjean, 1954c;

198) includes two penetra: Adelphacarus Grandjean,

1952b (not 1932 as given by Balogh, 1972) and

Aphelacarus Grandjean, 1932b, Itis known from the

Nearetic and Palaearetic Regions. Aphelacaridae

Grandjean. 19S4c is newly synonymous with

Adelphacaridae and is regarded as junior since it is

listed after it on the same page, despite the fact that

213

its type genus is much betrer known and far a longer

time, Adelphacarus is based on a4 single specimen

from Sweden with the region of the first rank of

hysteronoral setae destroyed. Grandjean (1954c)

distinguished Aphelacaridae by its having only two

pairs of genital papillae and deep furrows rather

than fine striations behind the proteranotal shield,

which ts not considered adequate here to distinguish

a suprageneric taxon,

Key to Families of Bifemorata (Adult)

1. Aggenital file Ig includes 6 setae. If proteronotal shield

present, itis nol extensive enough anteriorly lo carry either

selae jl or 21. Some hysteronotal setae conspicuously

stouter than athers Sets Z2 sharter than J3 (1-75 = oF

less) and approximately level with J2 (21-22; 1-J2 = 0-9

or more:]) - Acaronychidae

Aggenital file Tg includes 7 setae. Proteronotal shield present

and carrying setae jl and zl. If some hysteronotal setae

conspicuously stouter then seta 22 is longer and

conspicuously displaced forward , . -

2. Aggenital seta Jg] hammer-like or claw-like. Some fytiehes

nota] setac conspicuously stouter than others. Seta Z2 is

subeqgval to or longer than J3 and conspicuously displaced

forward (Z1-22:N-J2 = 0:25-0:51) . Palaeacaridae

Aggenital seta Jel setose. Hysteronotal ‘setae all telatively.

slim. Seta 22 shorter than J3 (0-6 = or less) and

moderately Sieplarnd forward (21-22:N2 =

0-6-0-75:L) _.... Adelphacaridae

Family ACARONYCHIDAE Grandjean

Acaronychidae Grandjean, 1932b; 426,

Archeonothridae Grandjean, 1954a: 428.

Type genus; Acaronychus Grandjean, 1932b,

Diagnosis: Bifemorata. Gnathosternal seta aol

simple or bifid, Proteronotal shield absent or limited

ta posterior half of proteronotum. Rostrum has

veniral recess with bilabed refractile lining (equals

“oeil”’ of Grandjean, 1958b). Some hysteronotal

setae conspicuously stouter than others, with seta 22

shorter (0-73 »« of less) than J3 and approximately

level with J2 (distance ratio 21-272:J)-J2 = 0-9 or

more:!), Aggenital file Jg with six setose setae. On

tarsus I seta dl proximad to solenidium sol,

subequal ia length or longer than sel, and ciliate.

Tibia Ef with 2 solenidia, Pretarsi with 3 claws,

central claw less than quarter the length of lateral

claws,

Remarks: Acaronychidae is used here-as by Sheals

(1965) and 1s equivalent ta Archeanothroidea

Grandjean, 1969 which is used by Balogh (1972). Tt

is not known why Grandjean treais Archeono-

thridae as having priority over Acaronychidae ar if

there js any basis far Balogh (1972) giving it the

authority with the apparently incorrect date of 1932,

Sheals (1968) compared the genera of this family and

considered that it might be preterable ta consider

them as a single relatively homogeneous group. lam

currently retaining two subfamilies bur including a

new grouping of genera. A study of the four

unnamed species from South Africa referred to by

Grandjean (1958a; 76) might resolve which grouping

204

is best, One of these unnamed species was grouped

in Stomacarus and is commented on belaw in the

remarks on that genus, although it would be grouped

in the Archeonothrinae as defined here.

Subfamily ACARONYCHINAE

Acaronychidae Grandjean. 1932b; 426.

Type-genus: Acaronychus Grandjean, 1932b,

Diagnosis; Acaronychidae. Female genital setae

thickened to stout, curved spines, conspicuously

shorter than the aggenital setae. The longest

hysteronotal seta (J3) shorter than distance between

seta! bases J 3 and <I. Seta J2 marginally posterior to

£2, Opisthosternal setal file Sa includes 2 setae On

tarsus |, setae add and pd4 not plasmic, but similar

to setae d3.

Distribution: Southern temperate and subantaretic

regions (see Stemacarus). In narthern temperate

regions represented by Acaronvchus between

latitudes 30° and 50° North, mainly in hilly or

mountainous country (Oregon—Ne; North

Carolina—Na; France—Pe; Tangiers, Algeria,

Caucasus Mountains—Pm; Tadzhik Soviet Socialist

Republic—Ps),

Remarks: The modification of the female genital

setae is heavily weighted so that Stomacarus

(assurned synonymous with Andacarus) is grouped

in this subfamily for the first ime. Members of the

subfamily also have correlated attributes such as

relatively short hysteronotal setae. The following 2

nominal genera are included in the Acaronychinae:

Acaronychus Grandjean, 1932: Stomacarus Grand-

jean, 1942a.

STOMACARUS Grandjean

Stomacarus Grandjean, 1952a: 360.

Type-species; Stomacarus tristani Grandjean, 1952a,

by monorypy-

Andacarus Grandjean, 1958a: 81. nsyn.

Type-species: Stomacarus macfarlani Grandjean,

1957, by monotypy,

Diagnosis; Acaronychinae. Ciliated seta dl on

tarsus 1 with only slightly spatulate tip (no wider

than distance between tips of nearhy cilia),

Proteronotal shield conspicuous, with reticulate

markings and carrying at least seta j2. Setal pairs J1

and #1 all on single hysteronotal shield. Delineated

tip of mentum broad-based, wider than distance

between setal bases clp and ¢2. Lines from setal base

cla ta clp and c2 would enclose ai acute angle,

Opisthosternal setal file Sg mcludes 2 setae,

Distribution: Southern temperate and subantaretic

regians between latitudes 25° and 55° South. All

records are with the original descriptions of nominal

REC. 8. AUST. MUS 18 (9); 199-122

January, 1981

species except for S. macfarlani Grandjean. 1957. El

Bolsén, Argentina—NTc (Balogh and Csiszar_ 1963)

and S, ligamentifer (Hammer 1967), TahitimAp

(Hammer 1972).

Found on ground, with dead leaves and branches,

or On mosses, liverworts, low plants and lichens on

bark.

Remarks; Only male attributes are known for the

type species of Stomacarus based on a single

specimen from Tristan da Cunha. Grandjean

(1958a) examined two females of one amongst the

four new species of South African “Archeono-

thrinae’ before hitn. They were small, their genital

setae on the ovipositor were somewhat enlarged

proximally but slim medially and distally as on the

male spermapositor of similar species and, finally,

their hysteronotal setae Were similar to those of S,

tristant, From these few attributes, and because the

fauna of Tristan da Cunha is likely to be similar to

that of South Africa, he grouped this unnamed

species in Stomacarus. This led him to exclude S.

macfarlani fram Stomacarus because of its stout

curved, genital setae and establish it as the type of

Andacarus. If Grandjean (1958a) was correct, then

according to the classification presented here,

Stomacarus and Andacarus would be in separate

subfamilies. Although three of the five characters

used in the diagnosis of Acaronychinae are not

known for S, tristani, the other iwo da fit, and at

least one other mite, Parasiriphis aurora Lee, 1970,

in a genus with a similar far southern distribution to

Andacarus, is found in the Tristan da Cunha Island

Group. Therefore, ] have excluded the unnamed

South Ajirican “Stamacarus’ females (Grandjean

1958a) from Stomutcarts and regarded Andacarus as

synonymous with Stomacarus. If a female of 8S.

rristani is found and none of its genital setae are

thickened curved spines then the subfamily group-

ings as presented here will have to be disregarded.

The follawing 6 nominal species are included in

Stomacarus: 8. abresin.sp., South Australia—Aa; 8.

campbellensis (Wallwork, 1966) n.comb., Campbell

Island—Sa: 8S. ligamentifer (Hammer 1967) na.

comb,, New Zealand—An; S. macfarlani Grand-

jean, 1957, Argentina—NTe; S. tristrani Grandjean,

1952a, Tristan da Cunha—Sk; S. watsoni (Travé,

1964) n, comb., Macquarie Island—Sa,

Stomacarus abresi n.sp.

Figs. 2-9

Female

General appearance and measurements: Dull

ivory-white, with apricot-coloured shields and legs,

Extremities of chelicerae and external malae,

hysteronotal setae and female genital setae blackish

brown. Other setae and claws are light brawn.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

5.07 8,9

Figs. 2-9—Stomacarus abresi n.sp. 2-7 and 9, female: 2, notum; 3,

gnathosternum; 4, idiosternum; 5, left chelicera, anterior

surface; 6, part left palp, anterior surface; 7, legs, darsal

setae on genua, tibiae and tarsi, 9, oyipositor, ventral view,

partially extruded, 8. male; spermapositor, ventral view.

206 REC. & AUST. MUS IK (9): 199-122

Idiosomal length 350 (20, 305-370. amongst which

those with only one or no eggs are 355 or less):

appendage lengths (for 320)—eh 40. pa 105, 1290, [1

210, 17190, TV 275: telofemur breadths—pa 22,5, 7

40, 17 32,5, TT 30, IV 27.5.

Prosternum: Gnathosoma drawn (Fig. 3) is

dorsoventrally squashed, Palp coxite seta cd shorter

but stouter than seta aol and lying just dorsal ta

palp. Seta aol is simple and setose, Whilst

extremities of external mala are strongly refractile,

those of internal mala are hyaline. Coxosternal seta

Id shorter but stouter than seta aol and lying just

dorsal to leg [. Coxosternal seta I/| set in a poteh in

the edge of coxite Il. On coxite TV there is a median

seta (JV3m) in the distal whorl (unlike Acaranychus

tragardhi).

Proteronotum: Protéronotum drawn (Fig, 2) fore-

shortened because it was tipped ventralward,

Plasmic seta, 22, finer than illustrated, Apodemes

extend anteriorad from setal bases j/2 and 92,

Proteronotal shield between setae [2 has a

reticulated pattern of punetations, and extends

laterally to encompass base of seta z2 and carry setae

sl and s2,

Opisthasternum: Shields around genital orifice

only vaguely delineated, Pore-like structure on

ovipositor proximad to seta dgl (possibly equals

reduced seta on Stomacarus macfarlani Grandjean,

1957).

Hysterononim: Shields not so clearly delineated as

drawn and adaxial setae are foreshortened because

of the angle of their axis (Fig, 2). Larger setae with

two rows of small cilia,

Appendages: Chelicera with grooved notch an

proximal edge of anterolateral surface, Setae: ch (2),

pa (Q-2-1-3-18), I (0-4-6-5-6-37), IT (1-5-6-5-7-27).

IT (2-2-3-3-6-26), PV (3-3-3-4-5-25]. Solenidia: pa

(0-0-1) [ (2-4-4), TT (1-2-4), HT (1-1-0), TV (1-2-0).

On tarsus [1 solenidium so2 is very small (regarded

as “famulus’’ by Grandjean, 1957; 217—..e. sela

dl).

Eggs: Amongst 20 type females. four have 0 eggs,

five have 1 epg, six have 2 eggs, four have 3 epgs,

one has 4 eggs. Eggs about 130m long. ellipsoid

with uniform smooth surface.

Male

Measurements and spermapositor (otherwise as

female): Idiosomal length 340 (3, 345-360). All

genital setae are setose, shorter but of similar

diameter to aggenital setae, No equivalent of

anterior pore noted,

Material examined; Vfolotype female (N197613), 19

paratype females (N197614-N197632), allotype male

January, T98l

(N197633) and 2 paratype males (N197634 and

N197635), moss and litter under Eucalyptus obliqua,

sclerophyll forest, Mt. Lofty 9.5.1974, D. C. Lee.

Undesignated female (N197636) and male

(N197645) litter under Eucalyptus inerassata,

mallee, Ferries-MeDonald Reserve, 20.6,1974, D,

C. Lee. Undesignated protonymph (N197784), litter

under Pinus pinea, Knott Hill Forest, 22,5.1974, D,

C, Lee,

Distribution: South Australia—Aa:!

Ferries-McDonald, mallee-broombush open-scrub-

land, 10 (5/8); Mt. Lofty, sclerophyll forest, 23 (5/8),

Knott Hill, cultivaied pine forest, 1 protonymph

(—/2).

Remarks: In order to distinguish S. abresi. species of

Stomacarus can be regarded as including two groups:

those with a slim, tapering proteronotal seta z2, and

those with a tape-like or lanceolate sera 22, The

latter group (S. camphellensis, 8. ligamentifer and S.

walsoni) trom New Zealand and its lar southern

islands have a narrow, square-shaped proteronotal

shield only carrying seta j2. also the small

hysteronotal seta J4 is cither on the same shield as

Z4 or its base is.as close to setal base 24 as its length.

The former group includes S. abresi, S. maefarlani

and S. tristani. S. abresi can be distinguished from S.

macfarlani, which has a narrow proteronotal shield

and seta J4 on the same shield as 24 as in the latter

eroup. S. abresi can be easily distinguished from S.

tristant, Which differs from all other known species of

Sromacarus in having longer, sinuous, smooth

hysteronotal setae. On the other hand S. abresi and

S. fristani are similar in having a broader

proteronotal shield and seta J4 further from seta 74

and not on the same shield. S, abresi alsa differs

from both S, tristani and §. macfarlaniin having five

setac on basifemur I] rather than four setae. S.

dbrest also differs trom S, watsoni, as well as

Acaronychus tragardhi, in having a setose rather

than a bifid seta aol.

Subfamily ARCHEONOTHRINAE Grandjean

Archeonothridae Grandjean, 1954a; 428.

Type-genus: Archeonothrus Trigardh, 1906: §71

Diagnosis: Acaronychidac, Female genital setae

may be slighty thickened to spines, but, if so, then

straight and longer than aggenital setae. The longest

hysternonotal seta (J3) longer than distance between

setal bases J3 and 21. Seta J2 at least marginally

anteriar to 22, Opisthasternal setal row Sa includes

3 selae. On tarsus [, setae ad4 and pd differ from

setae d3 in being plasmic.

Distriburion: Southern temperate regions; South

America and South Australia (see Loftacarus) and

South Africa (Archeonothrus, see Grandjean,

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 207

shea SPs

J445+6.

11, 13,15, 16

Figs. 10-16—Loftacarus siefi n.sp. 10-14 and 16, female: 10,

notum; 11, left chelicera, anterior surface; 12, idiosternum; 13,

gnathosternum; 14, part left palp, dorsal surface; 16.

ovipositor, ventral view, partially extruded, 15, male:

spermapositor, ventral view.

208 REC. 8. AUST MOS 18 (Y); 199-122

1958a), In Northern temperate regions represented

by Zachvatkiniella and Amuracarus between 25°

and 50° Worth, and 40° and 150° East, in

mountainous country (Caucasus Mountains—Pm;

Nepal—Os; far eastern Siberia, east of Lake Baikal

and Sichote Alinja—Ps; Japan—Pc).

Remarks: I regard the relative lack of modifica-

tion of the female genital setae as the most

important attribute in diagnosing this subfamily.

When the four unnamed species from South Africa

referred to by Grandjean (1958a; 76) and probably

belonging to this subfamily, are better known they

may require changes in both the generic and

subfamilial groupings within the Acaronychidae.

The following 4 nominal genera are included in the

Archeonothrinae; Amuracarus Lange, 1975,

Archeonothrus Trigardh, 1906; Loftacarus n.gn.;

Zachvatkinelia Lange, 1954 (=Himalacarus Sheals,

1965).

LOFTACARUS n.gen.

Type-species: Loftacarus siefi n.sp.

Diagnosis: Archeonothrinae, On coxite ITV, 5§

setae including seta 7V3m. Opisthosternum with 2

setae in row Sg und 3 setae in row Sa. Proteronotal

shield absent, Hysteronotal setal pair J3 not on

single shield and bases not connected by thickened

strip of cuticle. Setae J4 and 44 om same shield. Seta

26 tapering from base and ciliate, Cheliceral fixed

digit With 6 or fewer teeth. Scta v present on

telofemur IL and 111, and genu [Il, so that these

segments carry 6, + and 4 setae respectively. Seta au

absent from basifemur TY, leaving only 2 setae,

Genu f with 2 solérnidia. Tarsus II with 2 solenidia of

which so 2. is very small.

Distribution; Southern temperate regions between

35° and 40° South (probably more extensive than

this), All records with original descriptions of

nominal species.

Remarks: Loftacarus appears most similar to

Amuracarus although it can be distinguished from it

by the two attributes given below. | have only seen

one diagnosis of Amuracarus (Lange 1975) which

quotes the authority of this name as ‘Lange, 1975”

but does not give a reference in the bibliography or

describe the single species, A, voskresenskii Lange,

1975. The description of L. longicaudatus is very

brief. It is grouped in Loftacarus rather than

Amuracarus because of the apparent absence of

both a proteronotal shield and a thickened cuticular

strip between the bases of setal pair J3.

The following 2 nominal species ate ineluded in

Loftacarus: L. longicaudatus (Balogh and Csiszar,

1963) n. comb., Argentina—NTe: L. stefi nsp.,

South Australia—Aa,

January. 198]

Loftacarus siefi n.sp.

Figs, 10-18

Female

General appearance and measurements: Dull

ivory-white, with shields and extremities of legs

tinged with pale brown, Extremities of chelicerac

and external malae, and most hysteronotal setae ate

pitch-black.. Most other setae are deep brown, A few

setae and claws pale brown, Idiosomal length 700 (2,

695 and 700); appendage lengths (for 700)—ch 47,5.

po 245, f 635, I 470, fH 465, TV 640: telofemur

breadths—pa 37.5. 185.0. 17 80.0, 7757.5, [TW 70.0.

Prosternum: Gnathosoma drawn [Fig. 13) ts

dorsoventrally squashed. Palp coxite seta cd shorter

but stouter than seta aol and lying just dorsal to

palp. Seta aol is simple and setose. Coxosternal seta

Id shorter but stouter than seta aol and lying just

dersal to leg 1, On coxite TY there is 4 median seta

(1V3m) in the distal whorl.

Proteronorum: Apodemes extend anteriorad from

setal bases j2 and s2, Proteronotal shield is absent.

Opisthosternum: Shields around genital orifice

only vaguely delineated. Describing genital setae ts

difficult because ovipositor only partially extruded.

All genital setae are slightly stouler at base than the

aggenital setae of file Jz. Setae del. mgl and mp2

are longer than the aggenital setae and appear to be

rigidly straight,

Hysteronotum: Shields not so clearly delineated as

drawn, Adaxial setae hardly fereshortened at all

because of the angle of their axes (Fig. 10),

Conspicuous apodeme at base of seta J3, Lurger

setae are ciliate.

Appendages: Movable cheliceral digit carries two

parallel rows of teeth, Setae: ch (2), pa (0-2-1-3-18),

T (0-4-G-5-6-39), IT (1-5-6-5-7-27), LIT (2-2-4-4-6-31),

TV (3-2-3-4-5-28), On tarsus [some ventral setae and

the d4 pair are plasmic setae, On tibia [, seta 1]

unusually slim and short, Solenidia: pa (0-0-1), 7 (2-

4-4), IT (1-2-2), TT (1-1-0), TV (1-2-0). Solenidium

so2 on tarsus VW is very small,

Eegs: None seen,

Male

Measurements and spermaposiror (otherwise as

female): [diosomal length 765(1). Spermapositor

hears 10 pairs of setose setae. All genital setae are

shorter than aggenital setae. Some genital setae in

file pe appear longer than other genital setae: it has

not been established whether this is true or if itis an

impression related to the angle of their axes.

Material examined: Holotype fernmale (N197646),

paratype female (N197647) and allotype male

(N197648), moss and litter under Eucalyptus

obliqua, sclerophyll forest, Mr. Lofty, 9.5.1974, D.

C. Lee.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 209

Figs. 17-20—Appendages, 17 and 18, Loftacarus siefi n.sp.,

female: 17, legs, except for tibia I, dorsal setae on genua,

tibiae and tarsi; 18, telofemur IIT and genu III, all setae. 19:

zones of appendage surface. 20: Protoplophora palpalis

Berlese, tarsus IT,

Distribution: South Australia—Aa: Mt. Lofty,

sclerophyll forest, 3(2/8),

Remarks: The other member of the genus,

Loftacarus longicaudatus, is known only by attri-

butes of the notum. There are a few apparent

differences in the relative sizes of its notal setae and

those of L. siefi, on which they appear somewhat

shorter and stouter. The best diagnostic attribute is

that seta 21 is longer on L. siefi, being about

0-75 x the distance between setal bases Z1 and J1,

while on L. longicaudatus Z1 is about 0-25 x this

distance.

Family PALAEACARIDAE Grandjean

Palaeacaridae Grandjean, 1932b: 426.

Ctenacaridae Grandjean, 1954a: 428, n.syn.

Type-genus: Palaeacarus Tragardh, 1932, 2.

210

Diagnosis; Bifemorata, Gnathosternal seta aol

simple, ciliate or pectinate. Proteronotal shield

present and extensive cnough to carry all pro-

teronotal setae, No bilobed refractile lining to a

recess under rostrum, Some hysteranotal setae

conspicuously stouter than others, with seta #2

subequal to or longer than J3 and conspicuously

displaced forward (#1-22(JI-J2 = 0:25-0-5:1),

Ageenital file Ig with 7 setae of which Jgl is

hammer-like or claw-like. On tarsus [, seta dl distad

to solenidium sol or even so4, simple or lanceolate,

may be considerably shorter than sol, Tibia IT with 1

solenidium. Pretarsus with 2 or 3 claws, central claw

may or may not be less than quarter the length of

Jateral claws.

Remarks: Palaeacaridae, as presented here, newly

includes members of the Ctenacarinae; Grandjean,

1954e: 198, thereby further separating the latter

from the Adelphacarinae and Aphelacarinae, which

are here regarded as a single separate family (see

remarks on Bifemorata) But, as pointed out below

in the remarks on the Ctenacarinae, there are

similarities between it and the Adelphacaridae,

Possibly the Adelphacaridae should in the future

alsa be included under the Palaeacaridae, Of the twa

palaeacarid subfamilies only ctenacarines were

collected in the present study, so the Palaeacarinae

is briefly considered here. Palaeacarinae ineludes

two genera: Palaeacarus Tragardh, 1932 (= Tra-

gardhacarus Zachvatkin, 1945a) and Palaeacaroides

Lange, 1972. It ts known from the Holaarctic

Region.

Subfamily CTENACARINAE Grandjean

Ctenacaridae Grandjean, 1954a: 428.

Type-genus: Ctenacarus Grandjean, 1939.

Diagnosis: Palacacaridae, Internal mala consists

of a broad proximal half and a4 slim distal half, the

former carries all three adoral setae, with setal bases

ad2 and a3 posteriorad ta base of external mala.

Seta aol sparsely ciliate, cihate or pectinate. Seta d1

on tarsus T never Janceloate, is either tapering or

blunt ended and distad to solenidium so4-

Distribution; In the Northern temperate regions it

is known from an area bounded by Tangiers (Pm).

East Germany (Pe), Ukrainian S.S.R. (Pe),

Turkmen S.S.R. (Ps) and Japan (Pc), Also it

includes the only Bifemorata known from the tropics

(Brazil and Venezuala—NTb; Rhodesia—Ee) as

well as the below record from South Australia (Aa).

Remarks: Because of the long thick hysteronotal

setae 22 and J3 with 22 conspicuously displaced

forward. Clemacarus looks superficially like

Palaeacarus. This similarity is even stranger in other

ctenacarine genera with less extensive hysteronotal

shields and ciliate setae 22 and J3, But because of

similarities in the setae of tarsus I and in the internal

malae and oral setae, Ctenacanmae has in the past

REC, S. AUST. MUS L8 (9): 199-122

January, 198]

been grouped with the Adeliphacaridae, Here the

bysteranotal similanties and those of aggenital setae

Jgl are regarded as important enough to regard

Clenacarits as not only superficially similar to, but

infact more closely allied to Palaeacarus than

Adelphacarus or Aphelacarus. The genera in this

subfamily, especially Gilarovella and Neoctenacarus,

are quite similar ta each other, The following 4

nominal genera are included in the Ctenacarinae:

Beklemishevia Zachvatkin, 1945b, Crenacarus

Grandjean, 1939; Gilarovella Lange, 1974; Neac-

tenacarus Moritz, 1974.

CTENACARUS Grandjean

Crenacarus Grandjean, 1939: 543.

Type-species: Palaeacarus araneola Grandjean,

1932, by original designation.

Grandjeanacarus Zachvatkin, 1945b: 70.

Type-species: Palaeacarus araneala Grandjean,

1932. by original designation.

Diagnosis: Ctenacarinae. No ciliate hysteronotal

setae, Setae J3, Z2 and Z6 simple or have

inconspicuous paired hyaline flaps running parallel

to main setal axis. Seta J6 is leaf-like, having

conspicuous paired hyaline flaps. Setae J4, J5 and

Z5 subequal in length to Zl, Opisthosternal

chaetotaxy hypertrichous—e.g. 8Sg.

Distribution: Possibly cosmopolitan, only one

nominal species (so see ©, araneola distribution) but

may be another species in Oregon—Nc, Found on

ground, in dry or swampy environments, in sail

mosses and other low plants and in a termite nest,

Remarks: Crenacarus is easily distinguished from

other genera in the subfamily by the form of its

hysteronotal setae. Beklemishevia africanus

(Mahunka, 1974: 206) n.comb, from Rhodesia was

included in Ctenacarus. Tt is known only by

attributes of the notum, although the venter is

recorded as being as "“Grundtypus". It is not

confidently distinguishable from the type, and only

other known species, B. galeodula Zachyatkin,

19456. There is apparently a second, unnamed

species of Crenacarus from Oregon (Krantz, 1978:

475) on which the internal mala has an anterolateral

flap ventrally obscuring the base of a pilose rather

than a pectinate adoral seta aol,

The following 1 nominal species is included in

Ctenacarus: C. araneola (Grandjean, 1932b).

Ctenacarus araneola (Grandjean)

Figs, none

Palaeacarus araneala Grandjean, 1932b; 417,

Ctenacarus araneola (Grandjean); Grandjean.

1954c: 248.

Ctenacarus araneola (Grandjean): Mahunka, 1977;

464,

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 2

Male

Idiosomal length 315 (9, 290-320); appendage

lengths (for 320}—ch 17.5, pa 5. T 160, 17 110, I

115, FV 190; telofemur breadths—pa 12,5, £25,

20, TT 17,5, TV 20. Although movable cheliceral

digit is unusually short, the cheliceral digits are

robust and distally pitch-black. as are the tips of the

external malae, Hysteronotal setae J3, 22 and 76

have a narrow hyaline flap on each side as wide as

the diameter of the seta. In five specimens the

positor was recognised and it was as decribed for the

male (Grandjean 1954c: fig, 22E).

Material examined: Nine undesignated males

(N19761-N19769),, litter or mass and litter, under

Eucalyptus incrassata, mallee, Ferries-McDanald

Reserve, 20.6.1974, D.C. Lee.

Distribution: Venezuela (NTb), Brazil (NTb).

Morocco (Pm), Algeria (Pm), Japan (Pc). Kenya

(Ee). South Australia—Aa; Ferries -McDonald,

mallee-broombush open-scrubland, 9(2/8).

Remarks; Specimens referred to fit Grandjean’s

(1954c) description of this species except that three

other pairs of hysteronotal setae, besides. J6. have

hyaline flaps, This observation is considered here to

either reflect the improved microscopy of interfer-

enee contrast or an intraspecific variation. All nine

specimens are regarded as male because none

contained eggs and five apparently had sperma-

positors and were similar to the other four on which

the positars were not recognised.

Cohort PTYCTIMA

Remarks: The grouping of mites within this taxon

as by Balogh (1972) reflects a long standing

classification that is still widely held to, This is

despite apparently well supported conclusions by

Grandjean (1967, 1949) that the Piyctima includes

three groups more closely allied to other taxa than to

each other, Using Balogh's (1972) taxa these

conclusions can be summarised as follows: the

Protoplophoroidea are allied to the Sphaeroch-

thoniidae (Arthronota), the Mesoplophoroidea are

allied ta the Eniochthoniidae (Arthronota) whilst

the Phthiracaroidea and Euphthiracaroidea are

allied to the Collohmanniidae (Holonota). As stated

jn the introduction, | intend to try and reduce the

number of superfamilies and families in the

Cryptostigmata, If this is done, in combination with

partly following Grandjean’s (1969) later conclu-

sions, the classification becomes very unfamiliar

because the superfamily names in the Ptyctima tend

to have older authorities. Therefore, for the time

being, I am only following Grandjean’s (1967) first

change in the classification; the split of the Ptyctima

into two subcohorts, the Arthroptyctima and the

Euptyctima. In the next stage of this study, when

members of the Arthronota and Holonota are

described, J will try and reach a decision about

Grandjean’s (1969) later conclusions. It is passible

that the Ptyctima should be disbanded and its

members grouped in both the Arthronota and

Holoneta,

Subcohort ARTHROPTYCTIMA

Diagnosis; Ptyctima, Hysteronotal shield divided

into separate anterior and posterior parts, but

sometimes. (Mesoplophoridae) posterior hys-

teronotal shield reduced and difficult to distinguish

from ventral shields. Anterior hysteronotal shield

carries only four to eight pairs of setae. When

posterior hysteronoral shield inconspicuous there 1s

an unbroken transverse shield between the genital

and anal smeld, On palp coxite, seta cd spine-like

and less than X 0:25 length of 2. Genua of all

appendages with either no or 1 solenidium. Nymphs

ptychoid with tigid shields, although not always

distributed as on adult-

Remarks; The aboye diagnosis includes the

Protoplophoroidea Grandjean, 1965 and Meso-

plophoroidea van der Hammen, 1959 as used by

Balogh (1972). These superfamilies are maintained

here because of Grandjean’s (1967, 1969) conclu-

sions, mentioned above, that they are each allied to

one of two quite distinct groups within the

Arthronota. The Protoplophoraidea includes one

family which is considered belaw because one of its

species was collected in the present study. The

Mescplophorocidea is considered briefly here.

Grandjean (1965) regards this superfamily as allied

to the Hypochthonioidea amongst the Arthranota,

partly because of the striking resemblance between

the unusual gnathosternum af Mesoplophora Bet-

lese, 1904. and that of Hypochthoniella Berlese, 1910

(= Enioechthonius Grandjean), Grandjean (1965)

also grouped a poorly known genus, Archoplophora

van der Hammen, in the Mesoplophoroidea, but in a

distinct family. Although the gnathosternum of

Archoplophora is not described and the genital and

anal shields only resemble those of the nymphs of

Mesoplophora, the hysteronotal structure and

setation with a lack of pleural shields provisionally

groups it clase to Mesoplophora. Members of

Mesoplophora look superficially like small Eup-

tyctima, but with anal and genital plates similar to

the Brachypylina, rather than members of the

Macropylina. The similarity to the Euptyctima is

because they appear to have a single, undivided

hysteronotal shield, with no ability to fold up

posteriorly, and a proteronotal shield which folds

downward to cover only a distinct anterior forward-

facing section of the genital shield, without any

pleural shields to provide lateral protective wings to

the join. On the other hand, the reduced

212 REC. $, AUST. MUS 18 (9). 199-122

hysteronotal chaetotaxy (8 setal pairs) indicates that

the hysteronotal shield is homologous with only the

anterior part of the hysteronotal shield of the

Euptyctima. This supports the concept of an

Arthronota-like ancestor, able to fold up the

posterior part of the hyst€érosoma, which has since

lost this ability because of the posterior hysteronotal

shield being reduced and merged with the ventral

shields. | am postponing an evaluation of the degree

of alliance between the Protoplophoroidea and

Mesoplophioroidea until a later stage in this study

when the Arthonota have been considered.

Family PROTOPLOPHORIDAE Ewing

Prataplapherinae Ewing, 1917: 199,

Type-genus, Protoplophora Berlese, 1910,

Diagnosis: Arthroptyctima. Hysteronotum clearly

divided into at Jeast 2 articulated shields, the

posterior shield capable of moving up under the

anterior shield, Anterior hysteronatal shield carries

4 pairs (J1, Z1, $1, $2) or 6 pairs (plus inconspicuous

J2 and 22) of setae. Large pleural shields cover

space between proteronotal and genital shields when

mite folded up. Genital shield flat but inclined

dorsalwards from plane of anal shield and

completely covered by proteronotal shield when

mite folded up. If adanal shield separates genital and

anal shields it is divided mid-ventrally into 2 parts.

Pretarsus with 2 or 3 claws which may be unusually

long, Palp genu carrying seta d. All genua Without a

solenidium, External mala is spatulate, broadening

out distally to hyaline flap.

Remarks: The protoplophorids are unusual in

being able to fold up their bodies posteriorly as well

as anteriorly, Van der Hammen (1959) indicates that

Berlese’s. protoplophorids, a major part of the

known family, are insufficiently described. The most

informative work is the redescription of Aedo-

plophora glomerata and Cryptoplophora abscondiia

by Grandjean (1954b). The classification is further

confused because of the similarity between nymphs

and adults and the possiblity that some attributes

used to diagnose taxa, such as nurmber of

hysteronotal divisions or sutures, vary between these

stages. With the exception of Proteplophora, all

genera are recorded in tropical latitudes, only the

tecord of Crypreplophora abscondita from Algeria

(Pm) is elsewhere, The extraordinary long claws of

some species suggest that they may sometimes live

on other animals, The following 6 nominal genera

are included in the Protoplopharidae: Aedaplophora

Grandjean, 1932a; Arthroplophora Berlese, 1910;

Crypioplaphora Grandjean, 19324; Hauseroplophora

Mahunka, 1977; Protoplophora Berlese, 1910;

Prototritia Berlese, 1916.

January, L98]

PROTOPLOPHORA Berlese

Protoplophora Berlese, 1910: 217.

Type-species; Pratoplophora palpalis Berlese, 1910.

by original designation,

Diagnosis: Protoplopharidae, Anterior hys-

teranatal shield carries 4 pairs of simple setae.

Posterior hysteronotal shields carry 8 pairs of simple

setae. Posterior hysteronotal shield either with twa

hinged divisions just posteriorad to setae J3, 23 and

J4, Z4 so that there are three articulating parts, or

these two divisions reduced to a suture not extending

to lateral margins. Lateral margins of adanal shields

approximately parallel so that they meet a wide

anterior margin to enclose almost a right angle. Gna

chelicera, seta ch2 simple and setose, On palp, distal

setae shorter than tarsus. Pretarsal claws on legs less

than x 0-5 tarsal! length, Palp with 5 segments and

relatively long (X 0-66 length of leg I),

Distribution: Previously known mainly from

around western mediterranean between latitudes 30°

and 40 North. The record below from South

Australia js within the equivalent southern latitudes.

Also recorded fram Central Asia.

Found on ground in humus, litter and moss.

Remarks; Van der Hammen (1959) was of the

opinion thal since Berlese (1910) did not mention

characters on the ventral surface of the type of

Protoplophora, it is impossible to decide whether or

not the specimen described by Grandjean (1932a)

belongs to this genus. In contrast, [ consider that

enough attributes of the type are known (more than

one hysteronotal division, simple hysteranotal

setae—noat hypertrichous, long palp without long

distal setae, short claws) for Grandjean’s grouping

of his specimen in Proteplophora to be accepted.

The following 2 nominal species are included in

Protoplophora; P, bivaginata Grandjean, 1932a, P.

palpalis Berlese, 1910,

Protoplaphora palpalis Berlese

Fig. 20

Protoplophora palpalis Berlese, 1910: 217.

Protoplophora palpalis Berlese: Grandjean,

L932a; 24.

Adult

Idiosomal length 185 (3, L85-190); appendage

lerigths (for 190)—ch 16, pa 40, 160, IT 57.8, WT S0,

IVS; femur breadths—pa 5, 713. 1710, 179, TV 9,

Proteranotal seta 22 broadly lanceolate and in its

normal drooping position the surface away from the

proteronotum carries a raw of inconspicuous. cilia,

One specimen (N197610) has a recognisable positor

with at least four pairs of setae and there are only

1wa pairs of genital papillae. It also has three clearly

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOTLS

delineated parts to the posterior hysteronotal shield

which appear to be capable of telescoping up to

overlap each other. Chelicerae have three teeth on

fixed digit and one tooth on movable digit, and setae

chl and chZ are inconspicuous and setose. The

external mala is spatulate with a long stalk (X 0.6

total length) anda pyriform hyaline flap distally. All

the legs are tridactyl (Fig, 20) with a refractile,

curved central claw and hyaline, slim and almost

straight lateral claws,

Material examined: Three undesignated adults

(N197610-N197612), litter. under Eucalyptus incras-

sata, mallee, Ferries-McDonald Reserve, 20.6.1974,

D.C Lee.

Distribution; Sicily (Pm). Spain (Pm), Central Asia

(Ps). South Australia—Aa: Ferries-McDonald,

mallee-broombrush open-scrubland, 3(1/8),

Remarks: The specimens referred to here fit the

desctiption of the specimen from Spain by

Grandjean (1932a) rather than the type. On the

basis of the standard of the descriptions this may

only indicate that Berlese (1910) was not so

accurate. The fact that both authors represent their

specimens as monodactyl is also regarded here as as

inaccurate, resulting from the smallness of the mite

and the use of only bright field rather than the more

revealing interference contrast illumination, But it is

possible that three distinct species are represented.

Subcohort EUPTYCTIMA

Diagnosis; Ptyctima. Hysteronotal shield un-

divided and carrying at least 12 pairs of setae,

Genital and anal shields either fused together or

abut onto each other. On palp coxite, seta ed setose

and more than X 0.25 length af ¢2, Palp with 3 ta 5

segments, if genu present it does notearry any setae,

Genu I carries 2 solenidia. Nymphs are not ptychoid

and. lack shields,

Remarks: Balogh (1972) follows Grandjean

(1967) and also Walker (1965) in regarding this

group as made up of four families. three of which are

grouped jnto one superfamily (Euphthiracaroidea)

leaving the other family in the Phthiracaroidea, This

grouping has been tollawed here except that the

superfamilies have been reranked as families and the

families as subfamilies.

Family PHTHIRACARIDAE Perty

Phthiracarea Perty, 1841: 574.

‘Type-genus; Phihiracarus Perty, 1841.

Diagnosis: Buptyctima, Clearly delineated pairs

of genital and anal shields, with the grestest width of

each of these 4 shields more than X 0-4 their length.

Lateral hysteronotal gland absent, On tarsus 1,

213

inconspicuous seta dF coupled to distal face of

solenidium so4. Bothridium has associated internal

tubes. Proteronotal setal file s with one seta. Palp

with 3 segments. Lay epgs containing strongly

sculptured, pigmented prelarvae.

Remarks: Following a computer-assisted Gowers

Principal Co-ordinates Analysis to study phenetic

affinity amongst 39 phthiracarid species, Sheals

(1969) concluded that this family included a

comparatively homogeneous group. The species

used represented five of the eight phthiracarid

genera listed by Balogh (1972). Sheals (1969)

indicated that there might be two clusters, one

representing Phihiracarus Perty and one

Steganacarus Ewing or similar genera, Untartu-

nately, the species collected in this study is

diagnosable as Haplophthiracarus, members of

which are placed between the two clusters, nearer to

one or the other,

HOPLOPHTHIRACARUS Jacot

Hoplophthiracarus Jacot, 1933: 239,

Type-species; Hoplederma — histricinum

L908, by original designation.

Berlese,

Diagnosis: Proteronatal seta j2 erect and subequal

in length or Jonger than seta JJ, Hysteronotum

usually dimpled, carrying 15-18 pairs of setae, with

only 3 pairs of pores and without conspicuous dorsal

apophyses. Notal setae either tapering ot parallel

sided except for a ciliate tip, Anal shields carrying 2

pairs of setae in file Ja and 3 pairs of setae in file Za,

At most, only 3 pairs of setae (file Ja and seta 4a3)

on median margin of anal shield, Seta Za2 is the

longest Seta on anal shield. Femur | carries 4 setae,

and genu IV curries 1 seta. On tibia [V. seta d may

be either longer than seta 1 and uncoupled, or much

smaller and coupled to a solenidium.

Distribution: Cosmopolitan. Na; NTb, NTo; Em;

Pe, Pm, Ps, Pe; Os; Aa, Ap; Sk, The Antarctic is the

only major region from which Hoplophthiracarys has

nat been recorded,

Found in soil, litter, moss and grass. Members of

the genus have been recorded in substantial numbers

from widely different environments such as Sphae-

nuni-fens, deciduous and coniterous forests and high

altitude Rhododendron-thickets,

Remarks: Hoplophthiracarus is a ditficult genus ta

diagnose. Three of the earliest named species,

including the type, are poorly known so it is not

certain that the diagnosis presented above includes

the entire range of attributes amongst species

grouped here in the genus, Van der Hammen (1959)

considered that Hoplophthiracarus is reminiscent of

Steganacarus and that possibly it is only an artificial

unit. On the other hand, Macfarlane and Sheals

214

(1965) indicate that, at that time, Hoplophthiracarus

was only distiguished from Phihiracarus by having

nearly erect interlamellar (/2) setae. Sheals (1969)

included four species of Hoplophthiracarus in his

study of phenetic affinity amongst phthiracarids. He

found a close affinity between H. nepalensis and

species of Atroapacarus Ewing (in the Streganacarus-

complex), The remaining three species (H, costai, an

unnamed species from France and another from

Argentina) were fairly widely scattered between the

Hoplophorella Berlese cluster (in the Steganacaris-

complex) and the Phthiracarus cluster, Sheals (1969)

concluded that ‘tas defined at present, the genus

Hoplaphthiracarus is not a natural group” and any

resajution of the problem by making a noamenc-

latural division would require a study of the type of

the genus.

One attribute often used to distinguish

phthiracarid genera, flattness as opposed ta the

protrusion of the anal shields, has not been included

in the abave diagnosis. This is because the species

described below has a mainly flat anal shield but

with a protruding median margin (Fig, 28), which in

some preserved specimens does not protrude

downwards beyond the ventral margins of the

hysteronotal shields, whilst in other specimens it

conspicuously protrudes,

Two species, Phthiracarus hamatus Hammer,

1973 from the Tonga Islands—Ap and P. tubulus

Hammer, 1972 from Tahiti—Ap, not included in

Hoplophthiracarus, should be considered as possibly

congeneric with species in this genus, The following

17 nominal species are included in Hoploph-

thiracarus; H. cavernosus Wallwork, 1977; H.

cazanicus Feider and Calugar, 1969; H. castai

Maefarlane and Sheals, 1965; H, dactyloscopicus

Mahunka, 1978b: A. grossmani Jacoi, 1933, HA.

histricinus (Berlese, 1908); H. kugohi Aaki, 1959; H.

minus (Krivolutsky, 1965); H. nepalensis Sheals,

1965; H. paludis Jacot. 1938: H. pavidus (Berlese,

1913)—tredescribed by van der Hammen, 1963; H.

regalis Mahunka, 1978a; H- rebustiar Jacot, 1933; H-

shealsi n.sp.; H. siamensis Aoki, 1965; H. variolosa

(Berlese, 1888); H, zebra Balogh, 1962. There is 1

nominal subspecies: H. histricinus nitidior (Berlese

1923),

Hoplophthiracarus shealsi n,sp,

Fig, 21-34

Female

General appearance and measurements; Dull,

straw-coloured or orange; darker specimens usually

partially covered in a thin adhering layer of debris.

Refractile parts (external malac, cheliceral

extremities, setae and claws) paler than general

integument. Much of soma covered in shallow

REC. 8. AUST. MUS 18 (9): 199-122

January, L981

dimples, whilst legs with very much smaller

(approximately X 0,05) dimples. As well as the

withdrawal of the legs and folding down of the

proteronotal shield, the anal and genital shields vary

in their degree of protrusion below the lateral

margins of the hysteronotal shield. Hysteronotal

length 600 (25, 550-665) and proteronotal length 301

(25, 270-345); appendage lengths (for 660 and 345

containing one prelarya)—ch 90, pa 120, J 260,

240, 117230, IV 230: femur breadth—pa 17.5, 140, 1

32,5, 10 27.5, IV 275.

Prosternum: External malae lie in a vertical plane

when in a natural position (as Fig, 31) rather than

tending towards lying in a horizontal plane as when

squashed (as Fig. 22). Gnathosternal setae slightly

ciliate, those in file ao stouter and more ciliate,

especially setae dol and ao? (Fig, 32). Small

sclerotized samal shield dorsal to where coxa J]

abuts onto coxa 11), Coxal seta I1pis inconspicuous.

Proteromotum; Seta z2 has rows of cilia distally on

ventral surface (Fig. 33) which is exposed outwards

and upwards when curled urcund in its natural

position (Fig. 21), Three lang hyaline bothridial

tubes curled over distally, possibly some similar but

short tubes also attached to bothridial chafnber (Fig.

33).

Opisthostermnum: Exposed anterior anal apophysis

extends from left shield in all type specimens.

Anterior forward facing diaxial flap of genital shield

carries four setae in file Jg, while Jg5 is an exposed

ventral surface (Fig. 29). In the drawing (Fig. 25) the

diaxial flap is squashed so that setal file Jgno longer

appears vertical. Furthermore, by comparing these

figures it can be seen that setae in file Jg are much

slimmer and more tapered in appearance under a

light microscope as compared with an electron

microscope, The two sides of both the genital and

anal shield also has a thickening which fits into a

notched tuberosity on the mid-venter of the

posterior margin of the hysteronotum. The anterior

pair of genital papillae are less than half the size of

the posterior two pairs. The oyipositor carries 16

setac. amongst which two pairs are regarded as

belonging to a file pg which has migrated distally, so

that seta pe2 is level with Migl (Fig. 25).

Hysteronotum: A zone ventralwards of seta R is

free of dimples (Fig. 28). The 15 pairs of setae do not

taper and are ciliate distally. No vestiges of setae J4

and Z4 were observed, Only three pairs of pores. in

file hf, hft apparently absent,

Appendages: Fixed cheliceral digit distally carries

two parallel rows of teeth, Setae: ch (2), pa (--1-6), 1

(3-1-4-2-5-17), I (O-1-3-2-3-12), IIT (1-2-2-1-2-L0),

TV (1-2-1-1-2-10). Solenidia; pa (--0-1), I (2-1-3), IT

(1-1-2), HI (1-1-0), TV (0-1-0). Seta d1 on tarsus 1 is

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 215

100um

23,26

22, 24, 25

Figs. 21-26—Hoplophthiracarus shealsi n.sp. 21-25, female; 21,

soma excluding prosternum, 22, gnathosternum; 23, part left

palp, dorsal surface; 25, left chelicera, anterior surface. 25,

right genital shield and ovipositor, 26, male: spermapositor.

regarded as having migrated distally to just Eggs: Amongst 25 type females, 12 contain 0

proximad of solenidium so 3. Legs monodactyl, all prelarvae in eggs (although some with clear patch

claws have two ventral spines on proximal half, possibly representing developing eggs), 10 contain 1

REC, §. AUST, MUS 18 (9): 199-122

January, 1981

Fig. 27—Hoplophthiracarus shealsi n.sp., legs, dorsal setae on

genua, tibiae and tarsi,

prelarva, three contain 2 prelarvae. Prelarvae about

220m long and similar to that described for

Hoplophthiracarus pavidus by van der Hammen

(1963).

Male

Measurements and spermapositor (otherwise as

female): Hysteronotal length 485 (25, 425-545) and

proteronotal length 250 (25, 210-280), Sperma-

positor carries 10 setae which are relatively even in

length compared to those on the ovipositor.

Material examined: Holotype female (N197649), 24

paratype females (N197650-N197673), allotype male

(N197674) and 24 paratype males (N197675-

N197698), litter under Pinus pinea, Knott Hill

Forest, 22,5.1974, D. C. Lee. Sixteen undesignated

goldcoated specimens on stub ARAS2, as above

except collected on 24.11.1976,

Knott

Mt,

Hill,

Lofty,

Distribution: South Australia—Aa:

cultivated pine forest, 319 (2/2);

sclerophyll forest, 2 (2/8).

Remarks: The diagnosis of this species is difficult

because of the inadequate description of some

previously named species. If the descriptions of the

latter are accurate as far as they go, H. shealsican be

recognised by possessing the following five groups of

attributes: 1—hysteronotal shield dimpled, without

numerous long furrows, also lacks a conspicuous

pair of posterior concavities each side of a knob-like

protruberance opposing posterior edges of anal

shields; 2—15 pairs of blunt, distally ciliate

hysteronotal setae; 3—solenidium on tibia IV

coupled with a small dorsal seta; 4—aggenital setae

of file Jg in a straight vertical line; 5—of the two

shorter adanal setae in file Za, Zal subequal in

length to aggenital setae (X 0.9-1.1)and Za3

subequal in length to setae in file Ja (X 0.9-1.1). The

setal alignment of file Jg in a straight vertical line is

only otherwise described on H, cavernosus amongst

species of Hoplophthiracarus, although it does occur

on Notophthiracarus australis Ramsay, 1966, from

New Zealand—An. If further studies support the

uniqueness of this species, then it may be that it is

endemic to Australia although occurring in large

numbers only in the litter of an introduced conifer. It

should also be noted that based on the two

characters used by Sheals (1969) to distinguish the

two groups into which he divides his four species of

Hoplophthiracarus, H. shealsi, by having both 15

pairs of hysteronotal setae and a coupled solenidium

on tibia IV, has one diagnostic attribute of each

group.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

Figs,

28-32—Hoplophthiracarus

shealsi female: 28,

n.sp,,

hysteronotal, genital and anal shields, «140; 29, anterior

genital shields as indicated in Fig. 28, x2100; 30, mid-venter

where anal and genital shields abut, «1400; 31, anterior

proteronotum and gnathasternum, x 700; 32, parts of right

internal and external malae, x3500.

Family EUPHTHIRACARIDAE Jacot

Euphthiracarini Jacot, 1930: 214.

Type-genus: Euphthiracarus Ewing, 1917.

Diagnosis: Euptyctima. Genital and anal shields

may or may not be separated from ventral shield by

a suture, but in both cases lateral margin of ventral

shields unbroken for entire length of opisthoster-

num, Lateral hysteronotal gland present or absent.

On tarsus I, seta d3 conspicuous and not coupled to

a solenidium. Bothridium may have associated

internal filaments or, rarely, internal tubes.

Proteronotal setal file s with 1 or 2 setae. Palp with

3, 4 or 5 segments. Either lay eggs with or without

ornate chorion, or containing a strongly sculptured,

pigmented prelarvae.

Remarks: The results of a computer-assisted

Gower’s Principal Co-ordinates Analysis of phenetic

affinity amongst 53 Euptyctima species indicated

that the Euphthiracaridae is not so homogeneous as

the Phthiracaridae, since the first generic separation

occurs at the 74 per cent rather than the 90 per cent

phenon line (Sheals, 1969). In that study the 14

species of euphthiracarid, representing 7 genera,

separate into three groups at the 74 per cent phenon

line: two within Oribotritinae, one within Euph-

thiracarinae. Therefore, it appears preferable to

follow Markel (1964) and group ‘non-phthiracarid’

Euptyctima in one family, but maintain subfamilies

equivalent to the families listed in Walker (1965).

The Oribotritiinae Grandjean, 1954a and Synichot-

ritiinae Walker, 1965 have not been collected in the

present study, while the other subfamily is

considered below.

218 REC, S. AUST.

33-36—Ventral

Figs.

Subfamily EUPHTHIRACARINAE Jacot

Euphthiracarini Jacot, 1930: 214

Type-genus: Euphthiracarus Ewing, 1917.

Diagnosis: Euphthiracaridae. Anal shield with

anterior triangular corrugated area. Anogenital

suture present but often limited to breadth of

corrugated triangle. Anal and genital shields fused

with ventral shield so that no line of demarcation

separates them. Lateral hysteronotal gland present.

Palp with 3 segments, Lay eggs with ornate chorion

or a smooth chorion and containing a larva.

RHYSOTRITIA Markel and Meyer

Rhysotritia Markel and Meyer, 1959: 329.

MUS 18 (9): 199-122

view of setae

January, 1981

a %

z2 and podosterna.

Hoplophthiracarus shealsi n.sp., female: 33, seta z2 plus

associated bothridial structures; 34, coxae and trochanters I-

IV. Rhysotritia wallworki n.sp., female: 35, seta z2 plus

associated bothridial structures; 36, coxae and trochanters I-

IV,

Type-species: Hoplophora ardua C. L. Koch, 1841,

by original designation.

Diagnosis: Bothridial flap dorsal to seta 22,

Distance between setal bases j2-j2 less than twice

distance between setal bases j2-z2, Seta j2

approximately twice as long as jl. On genital shields,

between 7 and 11 pairs of setae, Posterior

corrugations on anal shields. Pore Zaf subcircular

and never nearer triangular corrugated area than it is

to seta Zal. Seta Jal recognisable but small. Setae

Ja2 and Ja3 shorter than Zal and Za2. On genu IV,

no solenidium.

Distribution: Widespread. NTc; Pe, Ps; Aa, Ap.

Species from North America, previously included in

Rhysotritia, are now grouped in Microtritia Markel,

1964.

SARCOPTIFORMES (ACARI) DF SOUTH AUSTRALIAN SOILS

Found in decaying wood, humus and moss. A

number of ecological studies indicate some members

of this genus occur in the deeper more compact

layers of the lawer fermentation zone and the upper

part of the humus zone, whilst some species of

Phthiracaridae occur in zanes above them.

Remarks; One attribute used by Balogh (1972) to

distinguish Rhysotritia is not comprehensive enough

because R, clavata sextiana has only one pair of

setae on both trochanters III and ['V. The diagnosis

used here is based on that of Markel (1964),

The following 4 nominal species are included in

Rhysotritia: R. ardua (Koch, 1841), R. calvata

Markel, 1964; R, duplicata (Grandjean, 1953); R,

wallworki n.sp. There are 4 nominal sub-species! R,

ardua otaheitensis Hammer, 1972; R. ardua

penicillata Pérez-Inigo, 1969; R. calvata sextiana

Lions, 1966; R, duplicata limbara (Markel and

Meyer, 1959).

Rhysotritia wallworki n.sp,

Fig. 35-42

Female

General appearance and measurements; Dull,

ivary-white with pinkish edges to shields. Refractile

parts (external malae. cheljcera] extremities, setae

and claws) paler than general integument. Most of

soma covered by fine pores (Fig, 41), Hysteronotal

length 390 (5, 360-415) and proteronotal length 240

(5, 225-255); appendage lengths (for 410 and 245),

ch 60, pa 97.5, F185, 1 170, TN 167.5, IV 167.5;

femur breadth—pa 15.5, T 37.5, 1 27.5, IT 25, IV

22.5

Prosternum: External malae are curved, tending to

form a cylinder around the chelicerae. Only seta cd

on gnathosoma appears to be even slightly ciliate.

Long sclerotized somal. shield dorsal] to and between

coxae IJ and Ul. Coxa [ as drawn (Fig, 36) is

swivelled round so that anterior surface is facing

ventralwards.

Proteronotum; Seta 22 has rows of cilia distally on

ventral surface (Fig, 35) which is exposed forwards

and outwards when curled around in its natural

position (Fig, 37), A number of granular filaments

extend inwards from bothridial chamber (Fig. 35)

and disappeared when cleared, Presumably these

are ‘'tracheoles’. From their appearance in this

species it is possible that they are the outer layers of

a series of tubes. Only two ridges radiate forward

fram bothridium.

Opisthosternum: Small anterior forward facing

diaxial flap on genital shield. All setae in file Jg are

in a straight horizontal line inside grooyes on

opposing faces of midventral division between

219

genital shields. There are small interlocking

corrugations at posterior end of anal shield.

Qvipositor carries 12 setae; apparently file pg is

unrepresented. The positor is assumed female

because of the large size of the setae in file dg (Fig.

40).

Hysteronotum: The 14 pairs of setae are slighty

ciliate and tapered, The fine pores represented by

spots (Fig, 41) show up as lines in profile, passibly

being narrow ducts running as deep as the Jong outer

chambers at the setal bases. Five pairs of pores in file

hf, with a conspicuous duct to hysteranotal gland

opening near hf3.

Appendages; Chelicerae with relatively small

teeth. Setae: ch (2), pa (-2-8), J (5-1-3-3-5-17), IT (0-

1-3-3-4-13), U1 (2-2-2-23-11), TV (3-2-1-1-2-9).

Solenidia: pa (--0-1), F (2-1-3), (1-1-2), 17G-1-0),

TV (0-1-0), All solenidia on genua and tibiae are

coupled with a dorsal seta, but the bases of the

couples appear to be discrete rather than merged as

in Hoplophthiracarus shealsi described above, Seta

dl on tarsus I is stout with annular wrinkles and is

regarded as having migrated distally to be level with

solenidjium so4, The setal pair avl and pvl on tarsus

II are absent, Each pretarsus with a single claw

having 2 inconspicuous ventral spines on proximal

half.

Male

Unknown.

Material examined: Holotype female (N197699), 2

Paratype females (N1976100 and N1976101), litter

under Acacia sophorae, Piccaninnie Ponds Reserve,

3/7/1974, D. C. Lee. Two paratype females

(1976102 and N1976103) litter and grass under

Eucalyptus viminalis, Chambers Gully, 12/6/1974,

D. C, Lee.

Distribution: South Australia-Aa: Piceaninnie

Ponds, coastal shrubland, 3{2/8); Chambers Gully,

savannah woodland, 2(1/8).

Remarks: Considering the similarity between the

species of this genus and the relatively substantial

differences between some subordinate subspecies

and their type subspecies, itis difficult to decide how

to classify new material. R. wallworki can be clearly

distinguished from R. duplicata which is tridactyl,

has an extra ridge runming forward from dorsal

margin of bothridial flap, and setae aul and pvl

present on tarsus Il. R. ardua differs in being either

bidactyl or tridactyl and having a slim seta 22, On

the other hand R, clavaia is similar, hut R,

wallworki is distinguished by four attributes: longer

cilia on notal setae including z2, 11 pairs of setae on

the genital shields, only a very indistinct notch in the

bothridial flap and 17 setae on tarsus I, But it should

be noted. that two of the characters involved, the

220

REC, S. AUST. MUS 18 (9): 199-122

Figs, 37-42—Rhysotritia wallworki n.sp., female: 37, soma,

excluding prosternum; 38, left chelicera, anterior surface; 39,

gnathosternum; 40, right anogenital shield plus partially

extruded ovipositor; 41, hysteronotal seta J3; 42, dorsal setae

on genua, tibiae and tarsi.

January, 1981

SARCOPTIFORMES (ACARI) GF SOUTH ALISTRALIAN SOLLS 721

setation of the genital shield and tarsus I, vary within

the species R, clavaia (including R.c.sextiana),

although nor to the extent of overlapping with R,

wallworki.

ACKNOWLEDGEMENTS

1 am indebted to the Interim Council of the

Australian Biological Resources Study tor funds for

equipment, and to the Mark Mitchell Research

Foundation and to the Science and Industry

Endowment Fund for funds for travel in connection

with this project. Two species, Sromacarus abresi

and Loftacarus siefi, have names based’ on actonyms

for two of these funding bodies.

Special thanks are due to Dr J. G. Sheals for

arranging bench space for me at the British Museum

(Natural History), London, and to Dr J, A,

Wallwork, Westfield College, London, for advice in

the initial stage of this project. Thanks are also due

to Dr B, G. M, Jamieson, University of Queensland,

Brisbane lor commenting on the manuscript.

My greatest debt of gratitude is to Ms Linda

Blesing for preparing some drawings (Figs. 1-9) and

to Ms Jenni Thurmer for preparing most drawings

(Figs. 10-42), Thanks ure also due to Ms Jan Forrest

for preparing the photographic prints and ta Ms

Debbie Meltoy for typing this manuseript-

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RECORDS oF THE | O

SOUTH AUSTRALIAN Zan

MUSEUM oF

VOLUME 18 NUMBERS 10—13 SEPTEMBER 1981

No.

EVOLUTIONARY SYSTEMATICS OF XEWYLLA XII. REDESCRIPTION

OF X OCCIDENTALIS WOMERSLEY AND A COMPARISON OF THE

CHAETOTAXY OF X ARENOSA UCHIDA AND TAMURA AND X

LITTORALIS WWOMERSLEY (INSECTA: COLLEMBOLA)

by MARIA MANUELA DA GAMA

A NEW SPECIES OF THE GENUS CATACANTHUS SPINOLA

(HETEROPTERA: PENTATOMIDAE: PENTATOMINAE) FROM THE

NEW HEBRIDES WITH MORPHOLOGICAL NOTES ON TWO OTHER

AUSTRALASIAN SPECIES AND THEIR RELATIONSHIPS

by IMTIAZ AHMAD and SYED KAMALUDDIN

THE FOSSIL PELICANS OF AUSTRALASIA

by PAT VICKERS RICH and G. F. VAN TETS

ON SOME OVIPAROUS FILARIAL NEMATODES MAINLY FROM

AUSTRALIAN BIRDS

by ODILE BAIN and PATRICIA M. MAWSON

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

EVOLUTIONARY SYSTEMATICS OF ZENYLLA. XU. REDESCRIPTION

OF X. OCCIDENTALIS WOMERSLEY AND A COMPARISON OF THE

CHAETOTAXY OF X. ARENOSA UCHIDA AND TAMURA AND X.

LITTORALIS WOMERSLEY (INSECTA: COLLEMBOLA)

BY MARIA MANUELA DE GAMA

Summary

X. occidentalis is redescribed from Womersley material; the chaetotaxy of X. arenosa is given

showing it to be distinct from X. littoralis. The phylogenetic position of these species is also

considered.

EVOLUTIONARY SYSTEMATICS OF XENYLLA. XII. REDESCRIPTION OF X. OCCIDENTALIS

WOMERSLEY AND A COMPARISON OF THE CHAETOTAXY OF X. ARENOSA UCHIDA AND

TAMURA AND X. LITTORALIS WOMERSLEY (INSECTA: COLLEMBOLA)

MARIA MANUELA DA GAMA

Museu e Laboratério Zoolégico, Departmento de Zoologia, Universidade de Coimbra, Coimbra, Portugal

ABSTRACT Redescription

GAMA, M.M. da. 1981. Evolutionary systematics of Xenylla. Body length: 0.56-0.74 mm Blue. Cutaneous

XII. Redescription of X. occidentalis Womersley and a granulation relatively coarse.

comparison of the chaetotaxy of X. arenosa Uchida and

Tamura and X. littoralis Womersley (Insecta: Collembola).

Rec. S. Aust. Mus. 18(10): 223-226. Dorsal chaetotaxy showing the following charac-

X. occidentalis is redescribed from Womersley _ ters.

material; the chaetotaxy of X. arenosa is given Head: all setae present; L, equal in length to L;.

showing it to be distinct from X. littoralis. The Th.II-III: all setae present and central ones

phylogenetic position of these species is also arranged in five rows (characters h,, h)).

considered. There are 2 S.s. on each side, one in

position P,.

OBSERVATIONS AND DISCUSSION Abd. I-III: S.s. = P,; ps absent.

Abd.IV: S.s. = Ps.

Xenylla occidentalis Womersley, 1934 Abd SE. SPs: 9. present

Fig. 1

Pare Ventral chaetotaxy presenting the following features

Material examined Head: all setae present.

Western Australia: Red Hill, Kalamunda, on Th.II-III: without setae (character t).

fungus, D.C.S., 30.V.1931, 6 specimens identified Abd.II: p; and p, absent (character v).

by Womersley. Abd. III: without medial setae nor median seta.

Aa wwywwvnu

ed ww vun

weve

e uvuvy Mouse

aoe

MuUUC LUNN

—

rnaAA

RnRaARD NAKA

6 b

Fig. 1—Xenylla occidentalis. a—Mucrodens in profile; b—Mucro-

dens, postero-dorsal view.

September, 1981

224 REC. 8. AUST. MUS. 18(10): 223-226

Abd.IV: the following setae absent—m,

(character a,), m. (character a,) and m,

(character a,).

Antenna IV with four cylindrical sensillae, of

which three are in outer dorsal position and

one is in the inner dorsal position,

§ + 5 eyes.

Unguis without inner tooth, Tibiotarsus with

two dorsal and two ventral long, clavate

ténent hairs, Tenaculum with 3 + 3 barbs.

Mucro spoon-shaped (Fig. 1), about 1/4-1/5

of length of dens, separated from it, which

has two setae. The mucro is about half as

long as Unguis HT. Anal spines not able to

be observed,

Systematics and evolution

The chactotaxic characters (hy, ho, t, V, ay, as, a4)

mentioned in the redescription place this species in

the position indicated in the phylogenetic tree

(Guma and Greenslade 1981).

Xenylla arenosa Uchida and Tamura, 1967

(Fig. 2)

Material examined of X, littoralis (see Gama 1980a)

South Australia: Yorke Peninsula, Jolly’s Beach,

7 specimens (identified by Gama), Yorke Peninsula,

Royston Head, 8 specimens (identified by Gama);

Christies Beach, 4 specimens (identified by Womers-

ley); Marino Rocks, 2 specimens (identified by

Womersley).

Western Australia: Rottnest Island, |

(identified by Womersley),

cotype

Japan, Cape Tachimachi, 9 specimens (identified

by Gama).

Material examined of X. arenosa: Japan, Akkeshi, 3

paratypes, leg, H. Tamura, 28.VI.1965.

Systematics and evolution

XX. arenosa belongs to the most primitive group of

species (welchi, malayana, portoricensis, bellingeri,

subbellingeri, thibaudi and littoralis) in which the

central setae on thoracic tergites II-III] are arranged

in three rows (absence of characters h,hz)—see

phylogenetic trees in Gama 1980b and Gama and

Greenslade 1981, As well, these species have no

setae on thoracic sternites II-III (character t).

September, 198]

Dorsal chaetotaxy (Fig. 2): there are supernumer-

ary setae, not represented in Figure 2, on the

thoracic and abdominal tergites, principally on th.[,

lateral parts of th.TI-ITT and abd. [-TV, and one can

observe a differentiation into macrochactae and

microchaetac.

Head; p, absent (character b); L; equal in

length to L;; Ly as long as L; and L,

(unusual in the genus XNenylla).

Th. H-UT: central setae arranged in three rows,

m, absent (character k); there are 2 §,s, on

cach side, one of which is in position Py.

Abd, lll: S.s. = P.3 ps present,

Abd. IV: S.s. = Ps; m, absent (character 0); a,

present,

Abd. V: S.s. P,; a, present.

Ventral chaetotaxy:

Head: p, and m, absent (character rc, s).

Th. T-Ill: without setae (character t).

Abd, [f; all setae present.

Abd. [[l: without medial setae nor median seta.

Abd. IV: all setae appear to be present though

difficult to distinguish,

Comparison of the chaetotaxy of X. arenosa with that

of X. littoralis

A study of the descriptions of these two

halophilous species, the former known from Japan

and the latter from Australia and Japan (Gama

1980a, p. 123) might lead one to think that they are

one und the same, For a brief comparison of the

adaptive characters between X, arenosa and X,

littoralis (see Gama 1980a, p. 123-125),

However, an examination of their chaetotaxy

indicates that they are distinct species separated by

the following nonadaptative characters (Fig. 2, this

paper, and Fig. 1, in Gama 1980a).

Dorsal chaetotaxy: on the head p, is absent in X.

arenosa (character b) and present in X. littoralis; in

this latter species L, is longer than L, (character f),

whereas in the former these two setae are equal in

length. On the th. U-fIf m, is absent in X. arenosa

(character k), and present in X, littoralis. On the

abd. | V m, is absent in X. arenosa (character o) and

present in X. littoralis in which p, is absent

(character n), whereas in X. arenosa p, is present.

Ventral chactotaxy: on the head p,; and m, are

absent in X. arenosa (characters r, s) and present in

X. littoralis, On the abd, TJ all the setae are present

in X, arenosa, while in X. littoralis p, and p; are

absent (character v) and p, also (character w).

EVOLUTIONARY SYSTEMATICS OF XENYLLA

vee

Muu

PAOARR ANA Rm ARAnA

a . \, Se va ee

— “

~~ ro tial

Veuve eer ina

= naan

BR RAR AAAI NAA ARs nnn, We

accct AAR ARaA

aan

Ly ~

ererererereren ee eee ~

‘gee ewer Views

wes|eer er eeeewe eee renelereererererererenererenes enemas porenereeReerereneserenerenereteses >

a ?

u 2

)

pacer -

i y)

Ye / ‘\ ‘

. -

RL oe

Fig. 2—Xenylla arenosa. Dorsal chaetotaxy of head, th. I-II and

abd. I, IV-VI.

226 REC. S. AUST. MUS. 18(10): 223-226

REFERENCES

GAMA, M.M. da, 1969. Notes taxonomiques et lignées

généologiques de quarante deux espéces et sous-espéces du

genre Xenylla. WI. Mems Estud. Mus. zool. Univ. Coimbra

308: 1-61.

GAMA, M.M. da, 1980a. Evolutionary systematics of Xenylla.

XI. Species from the Australian region. Rec. S. Aust. Mus.

18 (5): 123-129.

GAMA. M.M. da, 1980b. Apergu évolutif d’une septantaine

d’espéces et sous-espéces de Xenylla provenant de tous les

continents. Proc. 1° Intern. Semin. Apteryg. Siena 1978: 53-58

September, 1981

GAMA, M.M. da, and GREENSLADE, P., 1981. Relationships

between the distribution and phylogeny of Xenylla

(Collembola, Hypogastruridae) species in Australia and New

Zealand. Revue Ecol. Biol. Sol (in press).

UCHIDA, H., and Tamura, H.,

Collembolen-Art von Hokkaido. J. Fac. Sci.

Univ. S. VI., Zoology, 16 (2): 234-237.

WOMERSLEY, H., 1933. A preliminary account of the

Collembola-Arthropleona of Australia, I Poduroidea. Trans.

R.Soc. S. Aust. 57: 48-71.

WOMERSLEY, H., 1939. Primitive Insects of South Australia.

Silverfish, springtails and their allies. 322 pp. Adelaide,

Govt. Printer.

1967. Eine neue litorale

Hokkaido

A NEW SPECIES OF THE GENUS CATACANTHUS SPINOLA

(HETEROPTERA: PENTATOMIDAE: PENTATOMINAE)

FROM THE NEW HEBRIDES WITH MORPHOLOGICAL NOTES ON

TWO OTHER AUSTRALASIAN SPECIES AND THEIR RELATIONSHIPS

BY IMTIAZ AHMAD AND SYED KAMALUDDIN

Summary

Catacanthus grossi Ahmad & Kamaluddin, n.sp. of the tribe Catacanthini Stal from the New

Hebrides is described and its metathoracic scent gland ostioles and genitalia examined and figured.

Morphological notes on two other Australasian species, C. carrenoi (Le Guillou) and C. punctus

(Fabr.), are given and the three species above are compared with C. incarnatus (Drury) from the

Oriental region and the relationships of the four species are briefly discussed.

A NEW SPECIES OF THE GENUS CATACANTHUS SPINOLA (HETEROPTERA: PENTATOMIDAE;

PENTATOMINAE) FROM THE NEW HEBRIDES WITH MORPHOLOGICAL NOTES ON TWO OTHER

AUSTRALASIAN SPECIES AND THEIR RELATIONSHIPS’

IMTIAZ AHMAD and SYED KAMALUDDIN?

Department of Zoology, University of Karachi, Karachi, Pakistan

ABSTRACT

AHMAD, L., and KAMALUDDIN, S, 1981, A new species of

the genus Catacanthus Spinola (Heteroptera: Pentatomidae,

Pentalominae) from the New Hebrides with morphological notes

on two other Australasian species and (heir relationships, Rec. S.

Aust, Mus, 18 (11); 227-233.

Catacanthus grosst Ahmad & Kamaluddin, nsp,

of the tribe Catacanthini Stal from the New

Hebrides is described and its metathoracic scent

gland ostioles and genitalia examined and figured.

Morphological notes on two other Australasian

species, C. carrenoi (Le Guillow) and C, punctus

(Fabr.), are given and the three species above are

compared with C. incarnarus (Drury) from the

Oriental region and the relationships of the four

species are briefly discussed.

INTRODUCTION

Gross (1976) placed the genus Catacanthus

Spinola in his general section of Pentatominue allied

to his Menida, Piezodorus, Pentatoma & Rhyn-

chocoris groups for want of sufficient male

specimens. Ahmad and Afzal (1978) studied the

external morphology, including the metathoracic

scent gland ostioles, and the male and female

genitalia and also the internal anatomy including the

digestive, male and female reproductive organs and

the scent and salivary apparati of C. incarnatus

(Drury) and proposed the resurrection of Catacan-

thint Stal within the subfamily Pentatominae Amyot

et Serville,

Recently by the courtesy of Dr G. F. Gross,

Principal Curator of South Australian Museum, the

authors were given the opportunity to examine a

unique male specimen of Catacanthus from the New

Hebrides which differs [rom all other species in

being a uniformly dark purplish-brown with the

venter and all femora yellow in contrast to the

generally patterned body of brilliant, shining colours

in other species. It is described here as C. grossi

n.sp., and the structure of its merathoracic scent

gland ostioles and genitalia have been investigated.

‘Financially supported by PARC-LISDA Research Project

A17-ENT-37 (EG-Pa-181),

“Research Officer ‘STUDIES ON THE RICE {[NSECTS OF

PAKISTAN WITH REFERENCE TO THEIR SYSTEMATICS

AND PHEROMONE GLANDS’ FG-Pa-310 (PK-ARS-139).

Male and female specimens of two other species,

C. punctus (Fabricius)—also by the courtesy of Dr

Gross—and C, carrenoi (Le Guillow) by the courtesy

of Dr W. J. Knight and Mr W. R. Dolling of the

British Museum (Natural History), London, were

examined with special reference to the above

characters, for there has been a general belief that

the two species may be synonymous (Grass,

personal communication), Diagrams of the male and

female genitalia of C. incarnatus (Drury) are also

given for comparison and for further clarification of

the structures and the relationships of the included

taxa are briefly discussed in this hght. For

dissections, drawings and measurements the conven-

tional procedures, especially those used by the

present authors (1976), were generally followed.

Catacanthus carrenoi (Le Guill.)

(Figs. 1-4, 12-18, 29, 31)

Raphigaster carrenot Le Guillow, 1841, p, 262,

Pentatoma tricalor Montrouziet, 1855, p. 96.

Catacanthus carrenoi, Stal. 1876, p, 89.

Coloration: Body pale, reddish except head

(excluding brownish-black eyes), pinkish ocelli,

pronotum with anterior and anteriolateal margins

including callosities and a wide posteriomedial

portion, scutellum with basal wide portion, each

corium with a Jarge transverse spot on middle

portion, clavus, legs, antennae and labium black,

connexiva reddish, membrane of hemelytra light

brown.

Head: Slightly broader than long; anteocular

region slightly shorter than postenor portion of head

including eyes; antennae with 3rd segments longer

than 2nd and equal ta 2/1 x basal, length of

segments, 1 1.2 mm (1.2-1.5 mm), 2 3.0 mm (3,0-

3.2mm), 3 3.3mm (3.1-3.3mm), 4 4.5mm, 5

4.) mm, antennal formula 1<2<3<4; labium

passing mesocoxae, basal segments shorter than

bucculae, 3rd segment longest and distinctly more

than L'/: x basal, length of segments 1 1,3 mm (1,3-

1.5 mm), 2 2.0 mm (t.9-2.2 mm), 3 2.35 mm (2.0-

2.4 mm), 4 1.75 mm (1.5-1.8 mm}, labial formula

1<4<2<3; length anteocular region 1-5 mm (1,5-

1.6 mm); length posterior of head including eyes

228

1.6mm (1.6-1.75 mm), width 3.15 mm (3.15-

3.9mm); interocellar distance 1.7mm _ (1.7-

1.8 mm); interocellar distance 1.0mm _ (0.9-

1,1 mm).

FIG, 1, Catacanthus carrenoi Le Guillou in dorsal view.

Thorax and Abdomen: Pronotum with humeral

angles rounded, length of pronotum 4.4 mm (3.8-

5.2 mm), width 10.4 mm (9.4-12.0 mm); scutellum

with apical lobe narrowed, apex acute, length of

scutellum 8.5 mm (8,2-10.0 mm), width 6.5 mm

(6.0-7.4 mm); mesosternum slightly raised and

rugulose; metathoracic scent gland ostioles (Fig. 2)

small, somewhat ovate, peritremes elongated

tapering anteriolateral, apically somewhat acute,

evaporating area distinct; distance base scutellum-

apex clavus 6.6 mm (6.2-7.6 mm); apex clavus-apex

corium 4.3 mm (3.5-4.8 mm); apex corium-apex

membrane 5.8 mm (5.2-6.1 mm); apex scutellum-

apex membrane including abdomen 8.1 mm (7.2-

9.0 mm); abdomen with basal abdominal spine only

reaching hind coxae; connexiva distinctly exposed in

repose; in 2 posterior margin of 7th abdominal

sternum medially shallowly folded in, laterally

almost straight. Total length ¢ 24.1 mm (22.3-

24.1 mm), 27.55 mm.

Male genitalia: Pygophore (Figs. 3 and 4) with

dorsomedian surface deeply concave, three spinelike

processes at inner side of posterior lateral margins,

ventroposterior margin folded in medially, laterally

distinctly sinuated, ventrolateral lobe with two

spinelike fused processes, directed inward; para-

meres (Fig. 12) F-shaped, inner process short, outer

REC. S. AUST. MUS. 18(11): 227-233

September, 1981

and inner margins entire, sinuated, apex much

narrowed, acute; theca (Figs. 13-15) with an

unpaired dorsal blunt thumb-like large median

thecal appendage, a pair of dorsal thecal appendages

with lateral, horn-like, semisclerotized lobes, a pair

of ventral thecal appendages proximally semi-

sclerotized and distally sclerotized and denticulated,

and a bilobed dorsal membranous conjunctival

appendages with distal portion sclerotized and

outwardly sinuated, vesica short, penial lobes ovate

medially and outwardly fused with conjunctiva.

Female genitalia: (Fig. 29) 8th paratergites fused,

medially almost straight, sub-equal to 1st

gonocoxae, latter with posterior margin sinuated,

9th paratergites lobelike, elongated, outer margin

medially concave, slightly passing beyond posterior

margins of fused 8th paratergites, proctiger post-

eriorly substraight, 2nd gonocoxae broad; sper-

matheca (Fig. 31) with pump region medially faintly

constricted, bulb with two equal elongated,

fingerlike processes.

Material examined: 2 36, 1 9%, N. Borneo,

Philippines and New Guinea: 1911; June 1936 leg

Distant, W. L. Cheesman, L. E., in British Museum

(Natural History).

Comparative Note: This species is most closely

related to punctus (Fabr.) in general appearance but

it can easily be separated in having the distal portion

of the anterior tibiae dilated, the distal scutellar

black patch distally rounded, the ostiolar peritreme

much narrower and other characters as noted in the

morphological description.

Catacanthus grossi n.sp.

(Figs. 5-7, 11, 16-19)

Coloration: Body coffee-coloured tinged with

purple, green and black, except eyes brownish;

punctate; ocelli reddish; legs excluding femora,

antennae and labium black with greenish tinge,

venter and connexiva pale except small portions at

connexival joints black, membrane of hemelytra

with distal half coffee coloured, proximal half

transparent.

Head: Distinctly more than 25% broader than

long; anteocular region distinctly shorter than

posterior portion of head including eyes; antennae

with 3rd segment distinctly longer than 2nd and

slightly longer than 2x the basal, length of segments,

1 1.15 mm, 2 2.8mm, 3 3.25 mm, 4-5 mutilated;

labium reaching hind coxae with basal segments

shorter than bucculae, 3rd segment distinctly more

than 1'hx basal, length of segments 1 1.4 mm, 2

19mm, 3 2.3mm, 4 1.7mm, labial formula

1<4<2<3; length anteocular region 1.3 mm; length

A NEW SPECIES OF THE GENUS CATACANTHUS SPINGLA pat!)

FIGS. 2-10

2-4—Catacanthus

carrenor Le Guillou,

2—melathoracie scent gland ostiole. evp. (evaporaturia).

(ostiole), per, (peritreme ): 3—pygopliore in dorsil view. pre

(prochiger), 1th abd. seg. (eleventh ahdominal segment),

ms. (dorsamedian surtace), ll, (uteral lobe); +— pvgophore

in Venrral view. 5-7—Catacanthus gross} nosp.

S—meltathoracie scent gland ostiole. 6 pygophore in. dorsal

view, 7—pyacphore in ventral view. 8-1-—Caracanthus

punctis (Pabr), Bemetithoracic scent gland ostoele;

9—pygophore in dorsal view; 10—pyguphore in ventral view.

posterior portion of head including eyes 1.6 mm,

width 3.8mm; interocular distance 1.7 mm;

jnteracellar distance 0.8 mm.

Thorax and Abdomen: Pronotum with humeral

angles subacute, length of pronotum 4.4 mm, width

10.7 mm; scutellum with apical lobe broad, apex

subacute, length of scutellum 9.0 mm, width

6.8 mm; mesosternum slightly carinated and rugul-

ase, metathoracic scent gland ostioles (Fig, 5) small,

ovate, peritremes much elongated, medially curved,

tapering anteriad, apically acute, evaporating area

distinct; distance base scutellum-apex clavus

7.4mm; apex clavus-apex corium 4.5 mm; apex

corium-apex membrane $5.5 mm; apex scutellum-

apex membrane including abdomen & 8 mm; abdo-

men with basal abdominal spine only reaching

mesocoxae; connexiva slightly exposed at repose;

fernale not available. Total length ¢ 25.10 mm,

Male genitalia: Pygophore (Figs. 6 and 7) with

dorsomedian surface shallowly concave, one bifur-

cated, short, spinelike process at inner sides of

posterolateral margins, ventroposterior miurgin

pushed in medially, latter raised into bilobed

processes following laterally into beaklike struc-

tures; parameres (Fig. 16) L-shaped, blade with one

short inner process and one dorsolateral out growth,

outer margin sinuated, inner margin of anterior half

of blade irregular; theca (Figs. 17-19) with an

unpaired, dorsal, blunt, thumblike, small median

theeal appendage, a pair of dorsal thecal appendages

with lateral hornlike semisclerotized lobes, a pair of

ventral thecal, proximally semisclerouzed and

distally sclerotized with larger teeth, a bilobed dorsal

membranous conjunctival appendages, distally

sclerotized and sinuated, vesica short, penial lobes

platelike inner margin straight outer margin

sinuated, medially and outwardly fused with

conjunctiva,

23u REC. §. AUST. MUS. 14(11)- 227-233

40mm.

FIG. Il. Cotacanthus: grossi rsp. in dorsal view.

Material examined: Holotype 3, at light, vicinity

of Anelgahaut, Aneityum L., New Hebrides, July 21,

1971 leg. G. F. Gross, Royal Society-Percy Sladen

Trust Expedition (in the South Austrahan Museum),

Comparative Notes: This new species appears

isolated in the entire genus and can be easily

separated from the other species in having the

chocolate body colour dorsally, elongated medially

curved swordlike peritremes and the other charac-

ters as listed in the description.

Catacanthus. punctus (Fabr,}

(Figs. 8-10, 24-28, 30, 32)

Cimex nigripes Sulzer, 1776, PI. 10 £9 (nee

Fubricius)

C. punctum Fabricius, 1787, p. 291.

Edessa punctum Fabricius, 1803, p, 149,

Coloration; Body orange, tinged red except head

(excluding brownish hlaek eyes), ocelli reddish,

pronotum with anterior and anteriolateral margins

including callosities and a wide posteriomedial

pertion, scutellum with basal wide portions, each

corium with a large transverse spot on posterior 2/3,

clavus with basal portions, legs, antennae and

September, 1981

labium black with greenish tinge, connexiva reddish,

membrane of hemelytra ight brown.

Head; Less than 1/2 again broader than long:

anteocular region distinctly shorter than posteriar

portion of head including cyes; antennae with 3rd

segments slightly longer than 2nd and distinctly

more than 2'4x basal, Jength of segments 1 1.3 mm

(1.3-1.4 mm), 2 3.0 mm (3.0-3.1 mm), 33.25 mm, 4

and 5 mutilated; Jabium and bueculae as in carreneai,

3rd segment about 1'® basal, length of segments |

1.5mm (1.5-1,6 mm), 2 1.9 mm (1.9-2.2 mm), 3

2.2 mm (2.2-2.5 mm), 4 1-7 mm, labial formula

1<4<2<3; length anteocular region t.4 mm (1.4-

1.4 mm); length posterior portion of head including

eyes 1.7mm (1.7-1.8 mm); width 3.6mm (3.6-

3.9 mm), interacular distance 1.7 mm (1.7-1.9 mm);

interocellar distance 1.0 mm (1.0-1.1 mim),

Thorax and Abdomen: Pronotum with humeral

angles rounded length of Pronotum 4.6 mm (4.6-

5.2 mm), width 11.0 mm (11.0-12,.8 mm), scutellam

with apical lobe narrowed, length of scutellum

9.1 mim (9.1-10.9 mm}, width 6.8 mm (6,8-9.0 mm);

mesosternum slightly sulcated and rugulose;

metathoracic scent gland ostioles (Fig. 8) compara-

tively large, slitlike, peritremes elongated tapering

lateral, apically sharp, acute, evaporative area

distinct; distance base scutellum-apex clavus 6.9 mm

(6.9-7.8 mm): apex eclavus-apex coriuny 4.3 mm (4,3-

5.2 mm); apex corium-apex membrane 6.0 mm (6.0-

6,5 mm); apex scutellum-apex membrane including

abdomen 8.0 mm (8.0-9.0 mm); abdomen with basal

abdominal spine passing mesocoxde; connexiva

distinctly exposed at repose; in 2 posterior margin

of 7th abdaminal sternum medially deeply folded in,

Jaterally concave. Toral Jength dd 24.8 mm,

2 28.5 mm,

Male genitalia: Pygophore: (Figs. 9 and 10) with

dorsomedian surface ecancave, three spinelike

processes at inner sidé of posteriolateral margins,

ventroposterior margin inpushed medially, latter

slightly convex, ventrolateral lobe with two spine-

Jike fused processes, directed inward; parameres

(Fig. 24) F-shaped, stem with inner process large,

outer and innet margins entire, sinuated, apex

narrowed, suhacute; theca (Figs. 25-27) with an

unpaired, dorsal, blunt, thumblike, compacatively

large median thecal appendage, a pair of darsal

thecal appendages with lateral comparatively small

horntike semisclerotized lobes, a pair of ventral

theeal proximally semisclerotized and distally

sclerotized and camparatively less denticulated and

a bilobed dorsal membranous conjunctival appen-

dages, distal portion sclerotized and concave; yesica

Short, penial lobes ovate medially and outwardly

fused with conjunctiva.

A NEW SPECIES OF THE GENUS CATACANTHUS SPINOLA 231

yosildth.app

Noxtoee — UME app.

AAD

15—

FIGS. 12-19, 12-15—Catacanthus carrenoi Le Guillou, 12—para-

mere in inner view, bl. (blade), st. (stem); 12—uninflated

theca in dorsal view; 14—inflated theca in dorsal view, ap.

(apodeme), bp. (basal plate), dmc, app, (dorsal membranous

conjunctival appendage), inf. p. (inflatory pump), slldth.

app. (semisclerotized lateral lobe of dorsal thecal appen-

dage), piv. (pivot), stt. (sclerotized tip), th. (theca);

1S—inflated theca in ventral view, gp. (gonopore), pl.

(penial lobe), splvth. app, (semisclerotized proximal lobe of

ventral thecal appendage); 16-19—Catacanthus grossi n.sp.,

16—paramere in inner view; 17—uninflated theca in dorsal

view; 18—inflated theca in dorsal view; 19—inflated theca in

ventral view.

Female genitalia: (Fig. 31) 8th paratergites fused,

medially slightly concave, longer than Ist

gonocoxae, latter with posterior margin convex, 9th

paratergites lobelike, elongated outer margin

sinuated, shorter than posterior margin of fused 8th

paratergites, proctiger posteriorly concave, 2nd

gonocoxae comparatively narrowed; spermatheca

(Fig. 32) with pump region medially swollen, bulb

with two unequal elongated fingerlike processes.

Material examined: 23, 12, Australian but

unlocalised (in the South Australian Museum).

Comparative Note: This species is most closely

related to C. carrenoi as noted above but it can

readily be separated in having a smaller body size,

the black patch on posterior of pronotum more

elongated and a narrowed distal half of the

peritremes beside other characters as noted under

‘Relationships’.

RELATIONSHIPS OF THE INCLUDED TAXA

Distant (1902, p. 218), although treating Catacan-

thus in his division Nezaria, indicated that further

studies might necessitate its separate treatment.

FIGS, 20-27, 20-23—Catacanthus incarnatus (Drury), 20—para-

mere in inner view; 21—uninflated theca in dorsal view;

22—uninflated theca in dorsal view; 23—inflated theca in

ventral view. 24-27—Catacanthus punctus (Fabr.); 24—para-

mere in inner view; 25—uninflated theca in dorsal view:

26—inflated theca in dorsal view; 27—inflated theca in

ventral view.

FIG. 28. Catacanthus punctus (Fabr.) in dorsal view.

pL fl REC. S AUST. MUS. 18(11): 227-233

——~—are

oe —tst.gox-

\ -— — Send gon

—— — fC.

September, 1941

—.—— =Pr« spd.

FIGS. 20.32. 29—Curacarthus carretoi Le Guillou—femiale

terminalia ip veutral view, Ist. gox (first gonocoxa), 2nd

gox, (secund gonocoxa), 8th pt, Ceigh| paratergite), 91h pr.

(ninth pardtergite). ape, (arcus), pre, (proctiger);

30—Culacanthus puncius (Fabr-)—temale terminalia in

ventral View, 31—Catacanthus carrenci Le Guillou—sper-

mathees, dis f (distal flange). dis spd. (distal spermathecal

duct), md, (median dilation), prsp.d. (process of sper

mathecal bulb), prox. f. (proximal (ange), prox, sp.d.

(proximal spermethecal duct), sel.nyd. (sclerotized median

duet), sp.b. (Spermiatheval bulb). sp.p. (spermathecal pump);

32—Catacanthus purens (Fabr.j--spermuatheci.

Ahmad

and Afzal (1978) in) their external

morphological and internal anatomical studies on

C. incarnatus (Drury) have clearly demonstrated the

distinctness of this genus within the subfamily

Pentatominae Amyot et Serville and on that basis

have resurrected the tribe Catacanthini Stdl to

accommodate the genus.

The present Australasian species C. carrenoi,

C. grossi and C. punctus are clearly related to the

type-Species C. incarnatus from the Oriental region

in their large size (24.8-28,.5 mm), reflexed Jateral

margins of head and pronotum, more prominent

clypeus than paraclypei, clongated daggershaped

ostidlar peritreme, in male pygophore having

dorsolateral inner processes, parameres with more

or less prominent inner knobs at the base of the

blade (not clearly shown by Ahmad and Afzal 1978),

theca with a single dorsomedian, a pair of dorsal

thecal appendages (with dorsolateral semisclerotized

lobes) and a pair of ventral thecal appendages with

ventrolateral sermisclerotized lobes, a bilobed dorsal

membranous conjunctival appendage with

sclerotized tips and penial lobes fused medially and

with conjunctiva and in the females with widely

exposed arcus and second gonocoxae and sper-

matheca with a small spherical bulb having a pair of

uniformly thin, elongated and curved fingerlike

processes, The above characters alsa isolare

Catacanthus in the entire subfamily Pentatominae

and suppor! Ahmad and Afzal’s (1978) resurrection

of Stal’s tribe Catacanthini as separate from the

Pentatomini.

However it shares with other pentatomines the

characters of rounded unprojected humeral lobes,

moderately narrowed head with sinuated lateral

margins, scutellum with a distinct apical lobe,

ostiolar peritreme highly developed and base of

abdomen always with a spinous projection, Some of

the vharacters of male and female genitalia including

the inflated aedeagus with thecal appendages and

spherical spermathecal bulb usually bearing finger-

like processes also show the two groups to be closely

allied,

C, punctus and C, carrenoi appear more closely

felated to the type species C. incarnatus in having a

brilliantly coloured and patterned body, 4 more or

less straight ostiolar peritreme, in’ males the

pygophore has the dorsolateral inner processes more

or less trilobed, (second and third lobes ill developed

in C, incarnatus), parameres somewhat F-shaped

with a conical inner knob at the base of distally

tapering blade, and in other characters as listed

under the comparative and morphological notes of

C. carrenot and C, punetus.

C. carrenoi and ©. punctus however, appear more

closely related to each other in having a shining

A NEW SPECIES OF THE GENUS CATACANTHUS SPINOLA 233

black posteriomedian patch on pronotum and a

poximal patch on scutellum (Figs. 1 and 28), in male

more prominent inner knob of parameres and more

or less spinulose comparatively large ventral thecal

appendages in the inflated aedeagus (Figs. 15 and

27) and broadly rounded apices of 9th paratergites,

in the females (Figs. 29 and 30) in contrast to no

patch on pronotum and two shining black lateral

patches on proximal portion of scutellum, in males

less prominent and very short inner knob of

parameres, and smooth comparatively smaller

ventral thecal appendages in the inflated aedeagus

(Fig. 4, labelled dorsal by Ahmad and Afzal, 1978)

and narrowed tapering apices of 9th paratergites in

females (Fig. 13, Ahmad and Afzal, 1978) in

C. incarnatus, The two species have been suggested

to be possibly synonymous (Gross, personal

communication), However the comparatively much

larger spinules on the ventral thecal appendages and

the more or less equal processes of spermathecal

bulb in C. carrenoi together with the other characters

listed under the morphological and comparative

notes clearly separate it from C. punctus,

C. grossi appears on the other hand isolated in the

entire genus in having uniformly dark brown body

(Fig. 11) with venter and all femora yellow, anterior

margin of ostiolar peritreme medially curved (Fig.

5), in males dorsolateral inner processes of

pygophore short, bilobed, more or less L-shaped

parameres with a large irregular inner knob at the

base of more or less uniformly wide blade of

parameres and the theca with a less prominent and

short dorsomedian thecal appendage (Figs. 17 and

18).

REFERENCES

AHMAD, I. and AFZAL, M. 1978. Some aspects of the

morphology of Catacanthus incarnatus (Drury) (Pen-

tatomidae: Pentatominae) with phylogenetic considerations.

Pakistan J. Sci, Ind. Res. 21 (3-4); 129-135.

AHMAD, I. and KAMALUDDIN, S. 1976. A new genus and

three new species of Phyllocephelinae (Pentatomomorpha;

Pentatomidae) from Pakistan with notes on their zoogeo-

graphy and phylogeny. Ent. Mitr. Zool. Mus. Hamburg 5

(93): 81-95.

DISTANT, W. L. 1902, The fauna of British India including

Ceylon and Burma, Rhynchota 1 Taylor and Francis.

London.

FABRICIUS, J. C. 1787. ‘Mantissa Insectorum sistens eorum

species nuper Detectas adiectis characteribus, Genericis,

Differentiis specificis, Emendationibus, Observersationibus’.

Proft, Hafnia.

FABRICIUS, J. C, 1803. “Systema Rhyngotorum’, Reichard,

Brunswick,

GROSS, G. F. 1976. ‘Plant-feeding and other bugs (Hemiptera)

of South Australia, Heteroptera.’ Part Il. Government

Printer, South Australia.

LE GUILLOU, E, J. F. 1841, Description des Hemipteres

nouvedux, receuillis pendant son voyage de circumnavigation

sur la Corvette Zelée. Rev, Zool,; 260-263.

MONTROUZIER, P, 1855. Essai sur Ja faune de Vile de

Woodlark ou Moiou. Ann. Soc. Agr. Lyon 7 (2): 1-114.

STAL, C. 1876. Enumeratio Hemipterorum 5. K.

Vetensk. Akad. Handl. 14 (4): 1-162.

SULZER, J. H. 1776. ‘Abgekirzte Geschichte der Insecten nach

dem Linneischen System’. H. Steiner and Co,, Winterthur,

svenska

THE FOSSIL PELICANS OF AUSTRALASIA

BY PAT VICKERS RICH AND G. F.. VAN TETS

Summary

A review of all known Australasian fossil pelecanid material suggests that four species have

occurred in the southwest Pacific region during the past 20 million years. The oldest, Pelecanus

tirarensis, a small robust pelican from Miocene sediments of central Australia, has a distinctive

trochlear arrangement of the tarsometatarsus. The younger material is all Pleistocene in age, and

most of it can be referred to P. conspicillatus, the living Australian Pelican. An exception is P.

novaezealandiae, originally described as a large New Zealand subspecies of P. conspicillatus; it is

as different, however, from that species as is P. onocrotalus. P. cadimurka, n. sp., is distinctly

smaller, being within the size range of P. tirarensis, but with a differently shaped tarsometatarsus,

more closely resembling that of P. conspicillatus.

THE FOSSIL PELICANS OF AUSTRALASIA

PAT VICKERS RICH

Earth Sciences Department, Monash University, Clayton, Victoria 3168 and National Museum of Victoria,

285-321 Russell Street, Melbourne, Victoria 3000

and

G. F. VAN TETS

CSIRO, Division of Wildlife Research, P.O. Box 84, Lyneham, A.C.T. 2602

ABSTRACT

RICH, P. V., and van TETS, G. F. 1981. The fossil pelicans of

Australasia. Ree. S. Aust. Mus, 38 (12): 235-264.

A review of all known Australasian fossil

pelecanid material suggests that four species have

aceurred in the southwest Pacific region during the

past 20 million years. The oldest, Pelecanus

firarensis, a small robust pelican from Miocene

sediments of central Australia, has a distinctive

trochlear arrangement of the tarsometatarsus. The

younger material is all Pleistocene in age, and most

of it can be referred to P. conspicillatus, the living

Australian Pelican, An exception is P. novaezealan-

diae, originally described as a large New Zealand

subspecies of P. conspicillatus; it is as different,

hawever, [rom that species as is P. onocrotalus. P.

cadimurka, n, sp., is distinctly smaller, being within

the size cange of P. tirarensis, but with a differently

shaped tarsometatarsus, more closely resembling

that of P. conspicillarus.

INTRODUCTION

Pelicans, all in the genus Pelecanus, have a long

history in Australia, occurring as early as the

Miocene in terrestrial deposits of the Lake Eyre

Basin, South Australia. Their first occurrence

elsewhere in the world is contemporaneous with this

antipodean record.

Pelican fossils were first reported from Australia

by De Vis (1892) when he recognized Pelecanus

proavus from Quaternary sediments of south-eastern

Queensland. Later he set up an addifional two

species, P. validipes (in Brown, 1894) and

P, grandiceps (1905). No further pelecanid remains

were reported until Miller (1966) reviewed the De

Vis material and described a number of new

specimens collected during the 1950's on the Stirton

expeditions of the University of California and

South Australian Museum, Miller set up a new

species, P. tirarensis, based on a mid-Tertiary fossil

from the Lake Eyre Basin and reterred all of the De

Vis material except the lectotype of P. grandiceps

and the holotype of P. validipes to the living

Australian Pelican, P. conspicillatus. The same year

Scarlett (1966) designated pelican material found in

New Zealand as a new sub-species of the Australian

pelican, P.c. novaezealandiae (see Fig. 1).

Since the mid-1960's more pelican material has

been collected. This paper reports on all the fossil

pelican material from Australasia. Pelecanus tiraren-

sis is quite distinct from all other pelicans, both fossil

and Recent. Most of the Quaternary material,

however, appears to belong to P, conspicillatus, Two

forms, one small, P. cadimurkan, sp. and one large,

P. novaezealandiae from New Zealand, appear ta be

distinct species, The diagnoses of P, tirarensis and

P. cadimurka are based on features of their

tarsometatarsi and of P, novaezealandiae on

features of its synsacrum and femur. We found many

elements to be similar in size and shape in several

species of Pelecanus. The fossi] specimens that

lacked diagnostic features we have referred

therefore tentatively on the bases of size, location

and age to: P, tirarensis, small, central Australia,

Miocene; P. cadimurka n. sp., small, central

Australia, Quaternary; P: novaezealandiae, large

New Zealand, Holocene, and P. conspicillatus,

large, Australia, Quaternary, including the Recent

ABBREVIATIONS

The following abbreviations have been used in this

paper: AM, Australian Museum, Sydney; AMNH,

American Museum of Natural History, Ornithology

Department, New York; AWMM, Auckland War

Memorial Museum, Auckland; CM, Canterbury

Museum, Christchurch; ANWC, Australian

National Wildlife Collection. CSIRO Division of

Wildlife Research, Canberra; NMV, National

Museum of Victoria, Department of Ornithology,

Melbourne; QM, Queensland Museum, Brisbane,

SAM, South Australian Museum, Adelaide;

UCMP, University of California, Museum of

Paleontology, Berkeley; UCMVZ, University of

California, Museum of Vertebrate Zoology; 1.Z..

Institute of Zoology, Chinese Academy of Sciences,

Beijing (Peking). > = greater than, + = slightly

yreater than, @ = approximately, rf = referred.

ACKNOWLEDGEMENTS

Thanks are due to many who aided us in this

study, especially our respective families for their

tolerance throughout this work. Several provided us

with fossil specimens and invaluable comparative

material including:

A, Bartholomai, W. Bock, Cheng Tso-hsin, J. W.

Gregory, P. L. Horn, N. Johnson, L. Kuster, W.

Lanyon, A. McEvey, P. Millener, S. Parker, N.

Pledge, D. E. Savage, R. J. Scarlett, A. Stokes, G.

Turbott, W. J. M. Vestjens, and M. Wade. M.

Canny, C. Armstrong, M. Heslop and R. Sheehan

typed the manuscript. G. Earl drafted the charts.

Frank Knight drew the map and mounted the

photographs which are by Frank Coffa. Thanks are

due to the M. A. Ingram Trust, Danks Trust, Utah

Mining, and the National Science Foundation (BMS

7200102) for support of various aspects of this study.

DIAGNOSIS OF ELEMENTS OF PELECANUS

REPRESENTED BY AUSTRALASIAN FOSSIL

MATERIAL

Fossil material discussed below was assigned to

the Pelecanidae because of the following combina-

tion of characters of these large birds:

Quadrate: Lacks distinct rounded pit for quad-

rato-jugal; entire ventral articular area with little

relief, lacking deep pitting between articular

surfaces for mandible and quadrato-jugal such as

that present in the Phalacrocoracidae; anterior

border of ventral articular surfaces nearly straight

lacking any marked process on ventral end that

projects medial to mandibular articulation; man-

dibular articulation parallel-sided over most of

medial half; squamosal articulation also of low

relief, oriented at 90° to ventral articular surface

being compressed mediolaterally; main shaft com-

pressed anterio-posteriorly and lacking pneumatiza-

tion.

3rd Cervical Vertebra: Prezygapophyses elongate

and narrow; diapophyses about one half total length

of vertebra with short pleurapophyses extending

posteriorly; most dorsally protruding part of neural

spine low, and on posterior one-third of vertebra

lying posterior to end of parapophyses; diapophyses

not bulbous and not extending lateral to prezy-

gapophyses, but continue posterior from ther so

that vertebra remains nearly same width as anterior-

most width to end of pleurapophyses; lateral

margins only slightly concave; deep pit on ventral

surface near anterior end; prominent bulbous ridge

along midline of centrum over posterior half of

centrum ends in two prominent blunt projections on

either side of midline ridge.

REC, S. AUST, MUS. 18(12): 235-264

September, 1981

4th Cervical Vertebra: Differs from 3rd only in

that dorsal-most expansion of neural spine occurs

just slightly posterior to midpoint and at same level

as ends of pleurapophyses; ventral pit near anterior

end deeper and more elongate; two ventral spines

present and very low.

Sternum: (dorso-anterior end only). Lacks dorsal

manubrial spine but broad-based ventral spine

present, extending anteriorly, not dorsally; coracoi-

dal sulci converge upon one another anteriorly,

forming a small obtuse angle; dorsal lips of

coracoidal sulci separated and markedly expanded

medially, tapering to a fine point posterio-laterally;

coracoidal sulci narrow, parallel-sided and deep,

being deepest near postero-lateral end; ventral lips

of coracoidal sulci straight, not curved or indented.

Scapula: Scapular blade not parallel-sided near

proximal end but rapidly narrows distally, becoming

parallel-sided near midpoint without any further

broadening; near distal end blade narrows to nearly

one half of previous width, terminating rapidly in

blunt process, not recurved; entire shaft of blade

nearly straight, not anteriorly concave; proximal

part of shaft quite deep (dorso-ventrally) becoming

more flattened distally, with prominent muscle scar

on dorsal surface well separated from the proximal

end (about one third of the total length of the

blade); furcular articulation on elongate process,

extending proximally beyond glenoid facet.

Coracoid: Coracoidal fenestra small but complete

and well separated from the internal border of the

coracoid; scapular facet deeply excavated; glenoid

facet not flattened but markedly concave latero-

internally; medial and lateral boundaries of shaft

nearly straight, not markedly concave or convex,

nearly perpendicular to long axis of shaft over

medial half and at small obtuse angle over lateral

half.

Humerus: Proximal end differs in palmar view

from other large birds including storks, cranes,

gannets, cormorants, and darters in ligamental

groove not extending from ventral edge as far

dorsally, and in being a pit rather than a groove, A

similar pit occurs also in albatrosses, but the

proximal end is more bulbous and rounded in

Pelecanus, whereas it is thinner and more angular in

Diomedea: Distal edge, as in waterbirds, orientated

at right angles to main axis of shaft and not obliquely

with a wide, protruding entepicondyle as in land

birds, penguins, and auks. Ectepicondylar spur

absent as in most waterbirds except for those in the

Procellariiformes and Charadriiformes. As in other

Pelecaniformes, distal end curved over more in

palmar direction than in other waterbirds. Within

Pelecanus this is more pronounced than in other

THE FOSSIL PELICANS OF AUSTRALASIA

extant genera of the order. The humeri of Pelecanus

are also decidely larger than those of other extant

genera in the order.

Femur: Trochanter low, with both it and head

extending an equal distance proximally; trochanter

and head not grading smoothly into one another but

separated by a distinct, broad channel; head large,

about 60 per cent of the length of the trochanter; in

lateral view, proximal border of trochanter only

slightly, not highly arched with the most proximal

extension lying slightly behind anterio-posterior

midpoint; no pneumatic foramen present near

proximal end on anterior surface.

Tibiotarsus (Distal end only): Internal condyle

extends far distad of external and is much more

elongate than internal; single supratendinal bridge

lacks any ligamental prominences on either medial

or lateral ends and forms small acute angle with long

axis of shaft; shaft lacks external flange just proximal

to external condyle; in distal view, condyles deep,

with internal slightly deeper than external; distal end

slightly broader anteriorly than posteriorly but

lacking any marked constriction near anteropos-

terior midpoint.

Tarsometatarsus (Distal end only): Trochlea I

longer (extending further distally) than trochlea IV;

in medial view, trochlea I] not extending far

posteriorly and lacking medial spike on posteriome-

dial border; trochlea II lacking medial groove

extending over entire trochlea but notched distally

and posteriorly; in distal view, medio-lateral axis

through trochlea II-IV not straight but arched.

SYSTEMATIC POSITIONS OF AUSTRALASIAN

FOSSIL MATERIAL

Australasian fossils referred to Pelecanus by De

Vis (1892, 1905; in Brown, 1894), Archey (1931),

Miller (1966), and Scarlett (1966) are summarized

with our determinations in Table 1 and on Figure 1.

We have re-examined most of this material and have

added to it a number of new specimens collected

over the past decade and a half.

Pelecanus tirarensis

Citation: Miller, A. H., 1966. Mem. Queensland

Museum, 14 (5), 182, Fig. la, c, e.

Holotype: Distal end, right tarsometatarsus SAM

P13858.

Locality and Age: Lake Palankarinna Site 3,

Turtle Quarry (UCMP V-5762), west side of lake,

South Australia, Etadunna Formation, Miocene.

28°48'S, 138°24'E.

Comment: The holotype of P. tirarensis (SAM

P13858) differs from the tarsometatarsi of living

237

pelicans (see Figure 2 and Tables 2, 3 and 4) in

having:

(1) an elongate trough developed on the posterior

margin of the medial surface of trochlea II,

separated from the ligamental pit by a slender ridge

that is orientated nearly due proximo-distally along

trochlea; this ridge trends posteriorly towards distal

end and broadens slightly; in all other species

examined this area is occupied by an elevated,

inflated ridge that continues all the way to the

posterior border of the trochlea, when trochlea I

viewed posteriorly; the presence of the trough in

P. tirarensis produces a more gracile trochlea over

much of its length, as pointed out by Miller (1966);

(2) a decidely narrower anterior margin of

trochlea II than posterior margin (in distal view)

rather than both approaching the same width.

Differs from P. occidentalis and P. erythrorhynchos

in having a trochlea II that extends much further

distally than trochlea IV; in P. occidentalis and P.

erythrorhynchos trochlea II extends only slightly

distad of trochlea IV; P. conspicillatus and P.

onocrotalus have trochlear conditions very similar to

those in P. firarensis.

Miller (1966) noted also that the size of the

ligamental pit on trochlea II is greater than that on

other pelicans; in our sample of P. conspicillatus this

character seems too variable to be of diagnostic

value, and our largest specimen (ANWC BS1202)

had a large ligamental pit approaching that of

P. tirarensis. Miller further noted that the distal

foramen was less elongate on the plantar surface

than that in P, conspicillatus; examination of our

sample of P. conspicillatus (n=20) bore this out, but

in the sample of P. erythrorhynchos (n=5) the

character varied from the condition in P. conspicil-

latus to that in P. tirarensis; P. onocrotalus and P.

occidentalis have distal foramina as in P. tirarensis.

From fragments of SAM P13858 available, we

cannot verify the position of the origin of the plantar

ridge on the posterior surface nor can we see this in

Miller’s illustrations. Thus, comparison of propor-

tions of the shaft (length vs. width) based on the

location of this feature seems impractical.

Referred Material: UCMP 60988: Distal end,

right tarsometatarsus, Lake Palankarinna, Site 6,

UCMP V-5765, 28°46'S, 138°25’E, South Australia,

Etadunna Formation, Miocene.

Comment: UCMP 60988 is very incomplete,

lacking trochlea IJ and with articular surfaces of

trochlea III and IV very worn so that the original

surface cannot be defined on trochlea IV (see Figure

3). Thus, the diagnostic characters defining

P. tirarensis cannot be evaluated sufficiently even

though the distal foramen is small and ovoid, as in

P. tirarensis, and the size is comparable (see Tables 2

238

and 3). We have therefore only tentatively referred

it to this species,

UCMP 113115: Distal end, right tarsometatarsus,

lacking distal end of trochlea [1 and totally lacking

trochlea [V, Leaf Locality, UCMP V-6213 Lake

Ngapakaldi, 28°18'S, 138°16'E, South Australia,

Wipajiri Formation, late Miocene.

Comment? UCMP 113115 lacks those areas

exhibiting characters diagnostic of P. tirarensis. Its

size is comparable to that species (see Figure 3 and

Table 3) and its distal foramen is of moderate size.

We therefore only tentatively refer it ta P. tirarensis.

AMNH_ 11494: Proximal end of right tar-

sometatarsus, lacking much of hypotarsus, Lake

Pinpa, 31°08'S, 140°12'E, southeast af Lake Frome,

Tarkarooloo Basin, South Australia, Namba Forma-

tion, Miocene,

Comment: AMNH 11494 is definitely pelecanid in

that the proximal articular surfaces are deep; the

intercotylar prominence is very prominent and is

restricted to the anterior part of the proximal end;

the hypotarsus viewed posteriorly, is triangular in

shape with the longest portion on the internal side;

there are two well defined calcaneal canals; the

anterior surface of the shaft near the proximal end 1s

deeply excayated with two large vascular foramina in

this deep excavation. AMNH 11494 is smaller than

the tarsometatarsi of Pelecanus conspicillarus, and

differs further in that the lateral caleaneal notch is

only slightly indicated and not deeply incised; the

anterior border of the lateral articular surface does

not form a sharp apex, but is rounded; the whole

lateral articular surface is relatively broader; viewed

antertorly, the external foramen is smaller and not as

enlarged as the internal foramen; the excavation of

the antenor surtace al the proximal end and further

distad is relatively deeper; the tubercle for tbialis

anticus is displaced onto the medial surface of the

excavated area on the anterior surtace rather than

being more or less centrally located: the foramina on

either side of the hypotarsus are relatively smaller

than in P. conspicillatus.

Because AMNH 11494 is badly eroded, only the

following measurements were practical; width across

proximal end @ 18 mm; and depth across lateral

articular surface and lateral caleaneal ridge of

hypotarsus @ 14 mm. In P. conspicillarus these

parameters are width: n = 10, range 23-24 mm,

mean 23 mm, s.d. 1 mm; and depth n = 10, range

17-20 mm, mean 18 mm, ¢.d. 1 mm. Because of its

size and age we refer AMNH 11494 tentatively to

P. tirdrensis.

QM F10703; Distal end of left tarsometatarsus,

Crocpot 1, south end of Lake Palankarina, 28°46’,

138°28'E, South Australia, Etadunna Formation,

Miocene

REC. S. AUST. MUS, 18(12): 235-264

September, 1981

Comment; QM F10703 fits within the above

description of the holotype, except that the

proximodistal ridge does not broaden towards the

distal end; we, therefore, refer it to P. jirarensis.

Pelecanus proayus

Citation:

De Vis. 1892, Proc. Linn. Soc. New South Wales,

2(6): 444 Pl.24,

Holotype:

sometatarsus,

Locahty and Age: Darling Downs, 28°S, 150°.

Queensland, Quaternary.

No number (type lost), left tar-

Comment; Tarsometatarsus, Miller (1966) re-

evaluated this left tarsometatarsus and assigned it to

P. conspicillatus; the illustrations provided by De Vis

(1892) are not helpful in making this assignment,

Based on the illustrations, jt appears that the distal

foramen in Pelecanus proavus is nat rounded, but is

not as elongate as those in our sample of

P. conspicillatus, being more like that in P.

onocrotalus. This may he due to inaccurate

illustration of damage to trochleae, which De Vis

(1892) mentions.

De Vis’ illustrations also indicates that there is

litle flare of the distal end of the tarsometatarsus of

P. proavus, which mast closely approaches the

condition in P. onecrotalus. In most P. conspicillarus,

P. erythrorhynchos, and P. occidentalis this flare,

particularly on the internal margin of the shaft, is

more pronounced.

The measurements made from De Vis’ illustra-

tions of the holotype suggest a medium-sized

pelican, such as P. rufescens and P. philtppensis.

However, we are not entirely convinced that De Vis"

illustrations are accurate because of the discrepancy

between his measurement of the trochlear expansion

and curs of the distal width of the illustration of the

fossil, both of which appear ta be the same

measurement, De Vis (1892) originally recorded the

‘width of the trochleat expansion’ as 16.5 mm, but

our measurements of maximum width of the distal

end made from Figures 6a, 6b, Plate 24 were 18_0-

18.6 mm (see Table 2).

Thus, at present until] the onginal specimen is

available we reserve judgment on the validity of

P. proavus.

Carpometacarpus; De Vis (1892) assigned a

proximal carpometacarpal fragment, QM F1141, to

P. proavus, but we concur with Miller (1966) that itis

nol pelecanid.

Femur; De Vis (1905) assigned a distal right temur

fragment (QM F3752, Lower Cooper Creek, South

THE FOSSIL PELICANS OF AUSTRALASIA

Australia, Quaternary) ta P, proavus, When we

compared it with femora of large pelicans,

?. conspicillatus and P. onocrotalus, we could find no

appreciable differences (see Figure 4 and Table $).

De Vis (1905) mentions that the fibular groove is

more extensive in P. proavus, covering most of the

fibular condyle and making it concave posteriorly

further distad than in P, conspicillatus. Additionally,

the ectepicondylar region in the extitict bird was

supposedly comparatively narrow and rather deeply

sunken between the trochlea and the ectepicondylar

edge, the cavity formed extending distad nearly two-

thirds of the Jength of the trochlea (De Vis, 1905:

17). These characters are so similar in some

specimens of the modern Australian Pelican that this

fossil is best referred to P. conspicillatus,

Tibiotarsus: De Vis (1905) assigned the distal end

of a right fbiotarsus (QM F3753, Lower Cooper

Creek, South Australia, Quaternary), damaged on

the internal condyle, to P. proavus, nating that there

was nothing to distinguish it from the extinct form.

Our comparisons with other large pelicans including

P, conspicillatus, have revealed no diagnostic

characters that set his fossil tibiotarsus apart (see

Figure 4 and Table 6), and thus it should he referred

to P. conspicillatus as Miller (1966) suggested.

Pelecanus. conspicillatus

Citation

Pelecanus conspicillatus Temminck, 1824 Planches

col, Oiseaux 47; pl.276,

New Synonymy of Fossil Forms

Pelecanus validipes. De Vis, 1894 (in Brown,

Proc. Parl. So, Aust,, 2(25): 21, pl. 11, 5-6).

Archagocycnus lacustris (in part) De Vis, 1905

(Ann. Qld, Mus., 6:13).

Pelecanus grandiceps. De Vis, 1905 (Ann. Qld.

Mus., 6: 16, Pl. V, Figs 1-3)

Fossil Material Described by De Vis

Pelecanus validipes

Holotype: Distal end of a right tarsometatarsus,

SAM 18412, Warburton River, South Australia,

Quaternary,

Comment: Contrary to the comments made by De

Vis (in Brown, 1894) we have found. that (see Fig.

2):

(1) the middle trochlea (III) is not broader than

that in P. conspicillatus;

(2) the hallucal depression is no better defined (in

fact ANWC BS1363, P. conspicillatus, matches the

condition in ?. validipes);

239,

(3) the dorsal sulcus is not any deeper nor the

bounding ridges more well defined than in

P. conspicillatus:

(4) the distal end of the plantar ridge is no

straighter than in some P. conspicillarus; and

(5) P. validipes is within the size range of the

living P. conspicillatus that we have measured (sce

Tables 2, 3 and 4).

Thus, we suggest that Pelecanus validipes be

synonymized with P. conspicillarus.

Archagacycnus lacustris (in part)

Relevant Syntype: Fragment of 4th cervical

vettebra (QM F5529), unknown locality,

Comment: Among a series of bones that De Vis

(1905) named Archaeocycnus lacustris, a new genus

and species of swan, was a cervical vertebra that is

not anatid, but pelecanid (see Fig, 3). The vertebra

is quite eroded anteriorly and dorsally and thus

preserves few useful characters. It is, however, of a

pelican within the size range of P, conspicillatus (see

Table 7) to which species it should be referred and

removed from the Anatidae and the syntypes of

Archaeocyenus lacustris.

Pelecanus grandiceps

Lectotype: Miller, 1966: QM F3751, left distal end

of a tarsometatarsus, Lower Coaper Creek, South

Australia Quaternary,

Comment: Miller (1966) in his review of

Australian pelicans chose to retain this species as

distinct from the extant Australian pelican,

P. conspicillatus because of the following features of

the lectotype tarsometatarsus: (1) large size; (2)

‘presence of a deep pit on the plantar surface

between the bases of trochleae [If and Il’ (Miller

1966: 187); and (3) ‘a greater breadth and flattening

of the trochlear ridges on the anterior surface (of

trochlear IT)’ (Miller 1966: 187).

QM F3751 (see Fig. 2 and Tables 3 and 4) only

shghtly exceeds in size the tarsometatarsi of

P. conspicillatus in only one out. of four measure-

ments that are practical on the fossil. We doubt that

this difference is significant. The deep pit on the

plantar surface between the bases of trochleae [] and

If, present on QM F3751 is represented by only a

tiny foramen on a specimen that Miller (1966)

referred to this species (UCMP 56322, Lower

Cooper Creek site 8, UCMP V-5860, South

Australia, Quaternary) (see Fig. 3 and Table 4), In

our sample of recent P. conspicillatus one specimen

(AN WC BS1272) exhibited a distinct pit on the night

tarsometatarsus and a pinhole foramen on the left

tarsometatarsus. Thus, this character seems quite

variable between and even within individuals and

not particularly useful taxonomically for the genus

24U

Pelecanus. The anterior ridges on trochlea [IT

increase in breadth and flattening with size, and our

largest specimens af P. conspicillatus approach or

match that of P, grardiceps

Thus, we suggest that Pelecanits grandiceps be

synonymized with P. conspicillatus,

Quadrate: De Vis (1905) named a fragmentary

left quadrate (QM F3749, Lower Cooper Creek,

South Australia, Quatetnary), P. grandiceps,

because of qualitative differences as well as a

distinctly larger size than P, conspicillatus. We agree

with Miller (1966) that, in fact, this specimen is

within the range of variability of the living

P. conspicillatus (see Fig. 3 and Table 8), and we

therefore refer it ta this species,

Coracoid: De Vis (1905) assigned to P. grandiceps

a left coracoid (QM F37S0, Lower Cooper Creek,

South Australia, Quaternary), lacking both dorsal

and yentral ends but including most of the glenoid

facet and part of the scapular facet, Although not

preserving much that is diagnostic, the position af

the coracaidal fenestra in the fossil form more

closely approaches that in P. onacrotalus, where the

coracaidal fenestra is displaced further !aterad from.

the medial margin and thus is positioned nearer the

centre of the coracoidal shaft than in most of our

sample of P. conspicillatus. Because of the lack of a

more complete bone, it is difficult to interpret the

significance of this character, and because jt js

similar in size to those af modern P. conspicillatus

(see Fig, 4 and Table 9) we refer it to this species as

suggested by Miller (1966).

In conclusion, we can see no yalid reasans for

maintaining Pelecanus grandiceps as a species

distinct from P. conspicillatus.

Additional Material

Quadrate: UCMP 56321, left, Cooper Creek, Site

8, UCMP ¥V-5860, 28°34'S 138°00'E, South Aus-

tralia, Quaternary, Left quadrate nearly complete

except for erosion along some articular surfaces.

Together with those of other living species of

pelicans, this specimen differs from that of

P. occidentalis in that the pneumatic foramen lies on

jnternal rather than external surfaces; the otic

process is short rather than elongate giving the

quadrate a remarkably different shape, Allowing for

variability in modetn populations of pelicans, only

the much larger P, onacrotalus can be ruled out. The

fossil quadrate is similar in size to quadrates of

several living species, including P. conspicillatus, and

thus we follow Miller (1966) 1n referring it to this

species (see Fig. 3 and Table 8).

Palatines: UCMP 60702, fused palatines, Cooper

Creek, Maikuni Waterhale 28°34'S 138°09'E, South

Australia, UCMP V-5382, Quaternary. We follow

REC, 5, AUST. MUS, 18(12): 235-264

September. 198]

Miller (1966) in

P. conspicillaius.

Vertebrae; UICMP 56394, posterior half of 3rd

cervical vertebra, Cooper Creek, Site 7, 28°33'S

138°10'FE UCMP ¥V-5859, South Austraha, Quater-

nary. Miller (1966) assigned this specimen to

P. conspicillatus. tn size it matches measurements of

several of the larger pelicans (see Fig. 3 and Tables

10 and 11), and we can find no diagnostic differences

from any of these forms, including P. conspicillarus,

Thus we refer it ta this species.

referring this specimen to

Sternum: UCMP 61656, anterior fragment of

sternum, Cooper Creek, Site 18, 28°35'S 138°08'E,

UCMP V-6147. South Australia, Quaternary.

Although larger than any of our sample of

P, occidentalis, this specimen falls again in the range

of the larger pelicans, except for perhaps

P, onocrotalus (see Fig.. 3 and Table 12), and is se

fragmentary that definite assignment to any species

is tather senseless. We follow Miller (1966) in

referring it ta P. conspicillaius,

Coracoid: QM F6669, fragment of a left coracoid,

Lower Cooper Creek, South Australia, Quaternary-

Although part of a collection made during the

summer of 1901/2 by J. W. Gregory and his students

during the Melbourne University Lake Eyre

Expedition, we are not aware that De Vis and Miller

have cammented on this specimen in print. We refer

QM F6669 (see Table 9) to P, conspicillarus as it is

within the size range of this species and we can

discern no qualitative differences,

UCMP 60487, fragment of a left coracoid, Cooper

Creek, Site 8, UCMP V-S860, Quaternary. We refer

this specimen to P. conspicillatus in agreement with

Miller (1966).

Ulna: UCMP 56348, distal end of left ulna,

Cooper Creek, Site 16, UCMP V-5868, South

Australia, Quaternary, UCMF 60520, distal end of

left ulna whose articular surfaces are eroded,

Cooper Creek, Sire 8, UCMP V-5860, South

Australia, Quaternary. We agree with Miller (1966)

in referring these specimens to P, conspicillatus (see

Fig 2 and Table 13).

Cuneiform: UUCMP 60503, left cuneiform, Cooper

Creek, Site 8, UCMP ¥V-5860, South Australia,

Quaternary; UCMP 460549, right cuneiform, Lake

Kanunka, Site 2, UCMP V-5773, 28°23'S, 138°17'E,

South Australia Katipiri Sands, Quaternary, We

agree with Miller (1966) in referring these specimens

to P. conspicillatus (see Fig. 2 and Table 14).

Femur: UCMP 604777, proximal fragment of night

fernur, Cooper Creek, Site 8, UCMP V-5860, South

Australia, Quaternary, Miller (1966) assigned this

specimen to P, conspicillatus but because of the Jack

of any diagnostic differences either in morphology or

TELE FOSSIL PELICANS OF AUSTRALASIA

size with most other large pelicans (see Pig, 4 and

Table 15), we can only refer it to this species,

P. rufescens and P, occidentalis appear to be

distinctly smaller, and P. crispus (based on one

specimen) has a flattened, not curved, trochanter

(viewed laterally), which does not slope anterior-

wards as it does in UCMP 6(1477.

Tibtotarsus: UCMP 60521, very fragmentary,

Distal right, tibiotarsus, Cooper Creek, Site 8,

UCMP V-5860, South Australia, Quaternary,

Because of the incompleteness of this specimen,

little can be said of its affinities except ta the family

Pelecanidae, It falls within the size range of P.

conspicillatus and several other large pelicans (see

Fig, 4 and Table 6). Therefore, we refer it

tentatively to P. censpicillatus.

Humerus: UCMP 60640, very fragmentary distal

left humerus with only external and internal

condyles and external margin preserved, Cooper

Creek, Site 18, UCMP V-6147, South Australia,

Quaternary. Miller (1966) placed it in P. canspicil-

latus. In size and shape it is similar to the humeri of

P. conspicillatus and other large species of pelican

(see Fig. 3 and Table 16). We therefore tentatively

refer UCMP 60640 to P. conspicillatus.

Scapula: UCMP 56633, right scapula missing parts

of the glenoid facet and furcular articulation,

Warburton River, Marcus Locality, 27°S5'S,

138°00'E, UCMP ¥V-5569, South Australia, Quater-

nary. The distance between the fureular facets is

very narrow, but this character is extremely variable

even within P. conspicillatus. The angle at which the

two furcular facets meet (in ventral view) is slightly

preater than 90°, but this varies from an acute to

slightly obtuse in P. oonspicillatus, In our small

samples. it is slightly obtuse in P. occidentalis and

P. onocrotalus and acute in P. erythrorhynchos, In

size UCMP 56633 js similar to the scapulae of the

larger species of pelican (see Fig. 3 and Table 17),

and we therefore tentatively refer it to P.

conspieillatus.

Pelecanus novaezealandiae

Citation! Pelecanus conspicillatus novaezealan-

didae. Scarlett, 1966, Notormis 13: 204, Fig. 1 1-11.

Holotype: Large pact of a single skeleton

including mandibular fragments, left quadrate,

pelvis, right and left humerus, right ulna, right and

left radii, right and left carpometacarpi, left scapula,

right and left coracoids, right and left femora, right

tibiotarsus, right and left fibulae, right and left

tarsometatarsi, and several vetebrae, CM AV21355,

Locality and Age: Poukawa, Site I, Hawkes Bay,

New Zealand, (39°45'S, 176°43'E) Quaternary,

between 3500 and 4500 years B,P, The holotype was

found just below the Waimihia Ash, a band of airfall

volcanic ash, that was deposited 3440 + 70 years

B.P. (pers. comm. FP. Horn 1979).

Diagnosis: Very large pelican, Differs from recent

large pelicans in that its pelvis is distinctly broader

and more robust (see Fig, 5 and Tables 2-6, 9, 13.

15-19) particularly across supratrochanteric proces-

ses; iloischiatic foramen not extremely elongate.

thus. differing from P. conspicillatus and more

resembling P. oneerotalus; proximal end of femur

wider than largest of Pelecanus, although most other

measurements overlap. Tarsonietatarsus longer than

femur as in P, conspicillatus and not shorter as in

P. onocrotalus.

Referred Material: Partial skeleton (no number,

now lost), Archey (1931), cave deposits near Lake

Waikaremoana, North Island (38°47'S, 117°06'E);

right coracoid (CM AV15089), right humerus (CM

AV12264), left femur (CM AV12482) and proximal

end, right humerus (CM A¥V13095), Marfell Beach,

Lake Grassmere, South [sland (41°42'S, 174°10'E);

distal left ribiotarsus (AWMM AU5845,1), Puheke

Beach, Kacikari Peninsula, North Island (34°55'S,

173°22'E); left tibiotarsus and fragments of a lefr

ulna (private callectian of Peter Horn), type locality

Poukawa, Site N141/i: part of fused furcula and

sternal keel (CM A¥V32001) type locality, Poukawa,

all Holocene in age.

Comment: Searlett (1966) nated most of the

characters we have utilized to define the species,

Pelecanus novaezealandiae. Because his sample of

modern pelicans was so small, he chose to recognise

the New Zealand form as only a separate subspecies.

With our additional samples, we believe that the

New Zealand form is sufficiently distiact, especially

in the robustness of the pelvis and proximal hind

limb, that it rates separation as a species.

Pelecanus cadimurka n. sp.

Holotype: SAM P22501, left tarsometatarsus

fragment lacking trochlea 1V and with some erosion

of posterior border of trochlea If (see Fig. 6).

Locality and Age: Kuttipirra (Katipir)

Waterhole, Cooper Creek Site 9, UCMP V-S586l,

28°33'S, 138°06'E, South Australia, Quaternary.

Etymology: Cadimurka (Aboriginal), noun in

apposition, Brown (1894:5) in reporting the

presence of bones of extinct animals at several places

along the Warburton River, mainly along the edges

of large waterholes, writes that “The natives account

for the presence of these bones by saying that they

belong to the cadimurka, a large fish which lives in

the bottom of the waterholes, and which has

consequently never been seen by them’, One of the

bones picked up by Brown on the Warburton River

add

is the holotype of Pelecunus validipes, and the

matenal of P. cadimurka has been subsequently

found in the same area,

Diagnosis: Small pelican in the size range of

P. occidentalis and P. rufescens. distinctly smaller

than P. conmspicillarus and all other living species of

pelican (see Fig. 2, 6 and 7 and Tables 2, 3 and 4),

Where comparable publistied tossil material exists,

only P. tirarensis and P. gracilis are of similar size. In

medial view trochlea Il lacks a deep channel on

posterior border of medial surface, thus differing

from P, lirarensis, Trochlea Wf and Ill extend an

equal distance distally, unlike in most other pelicans,

including P. occidentalis and P, conspictilatus where

{Tf is distinctly Jonger, [In distal view, trochlea IT is

relatively narrow with posterior and anterior borders

of nearly equal width, differing from (1)

P, conspicillatus, which tends to be broader overall

and laterally expanded along the posterior border,

(2) P. tirarensis where the anterior border js

decidedly narrower than the posterior, and (3) P.

occidentalis, which tends to be broader, Trochlea [11

is deep and nat shallow as in P. occidentalis, thus

more closely approaching the condition in P.

conspicillatus. Trochlea IIT is relatively slender, end

not as broad as in P, firarensis (see Fig. 2), In

posterior view, articular surface of trochlea Il

terminates in a distinet apex proximally and is not

squared off as in P. occidentalis. The shape of the

distal foramen is not long and slit-like as in P.

conspicillatus but more closely approximates those

of P. onoerotalus and P. occidentalis.

Referred Material: UCMP 60577-60578. Two

distal fragments of right tarsometatarsi with all three

trochlea represented, Lake Kanunka, Site 2. UCMP

V-5773, 28°23'S, 138°17'E, South Australia, Quater-

nary. Both specimens are slightly smaller than any of

our samples of Pelecanus conspicillaius, about the

size of P. tirarensis (See Fig. 3 and Tables 2, 3 and 4),

On trochlea II, the posterior channel that

characterizes P. firarensis is not present, thus

eliminating this species, In UCMP 60578 there is a

small posterior depréssion in this area, but this

condition can be matched in our sample of

P. conspicillatus. The distal foramen is intermediate

in shape between those of P. conspicillatus and

P. firarensis. more like those of P. onocratalus and

P, erythrorhynehos, but this character may quite

likely vary somewhat with broader sampling. The

trochleae of UCMP 60577 do not appear to flare as

broadly distally as they do in P, conspicillatus and

P. tirarensis, more closely approaching the condition

in P. wnocrotalus. Because parts of UCMP 60577

have been eroded, however, a final evaluation of

this character is advisable, In both specimens,

trochlea [Ll extends far distad of trochlea IV as in

other large pelicans including P, tirarensis, distingu-

ishing it from P, eccidenralis and P, erythrorhynchos,

REC. 8, AUST. MUS. 18(2): 235-264

September, 1981

Comment! Although these two specimens are

somewhat intermediate is size berween the holotype

ot P. cadimurka and the tarsonletatarsi of

P. conspicillatus, they appear to be qualitatively

distinct from the latter as well as those ot PF. urarensis

and P. occidenralis. The specimens compare well

with the holotype of P. cadimnurka and thus will be

referred to that new species. Miller (1966) had

previously assigned both the above specimens to P

conspicilatus,

Our measurements trom the De Vis figures of the

holotype tarsometatarsus af P. proapus are within

the ranges of those of the P, cadimurka material but

because of a discrepancy between De Vis’ and our

measurements (see above) we doubt the scale of his

figures and do not feel jusofied in ceferring the

material of P. cadimurka to P. proavus. Further-

more, trochlea Ill is figured substanually longer

than trochlea Ef in P. proawus and thus differs trom

the holotype of P. cadimurka,

UCMP 123382; Nearly complete 4th cervical

vertebra, Cooper Creek, Site 8, V-5860, South

Australia, Quaternary,

Comment: The vertebra is much smaller than any

of P, conspicillatus in our sample (see Table 7)

although very similar in shape. Also it is more

gracile than those al P, occidentalis, which are

anterla-posteriorly compressed and more robust. It

differs further from those of P. occidentalis, and ts

similar to those of P. conspicillatus, in lacking paired

ridges over posterior half of ventral surface and in

having a decidedly shallower pit just posterior to

anterior articular surface of centrum on ventral

surface. The Yertebra may be, relative to total

length, shghtly broader across ventral surface of

centrum ou posterior half than in P. conspteillatus. It

appeats that this represents a species of pelican

smaller than the extant and the Quaternary

P_ conspicillatus, and we have therefore referred it to

P_ cadimurka,

Not Pelecanidae, UCMP 69587. Proximal end,

night humerus, Lake Kanunka, UCMP Y-5773,

South Australia, Quaternary.

Comment: Miller (1966) assigned this specimen to

P conspicillatus. Upon re-examination we find that

the humerus is certainly within the size range of

some living pelicans, but it shows considerable

morphologic differences from all Pelecanidae. In

UCMP 69587 the ligamental furrow is deeper, The

bicipital furrow is deeply excavated instead of flat

In proximal view, the head narrows consideiably

near the external tuberosity, and is not bulbous.

In all of these characters, the fossil so closely

resembles members of the Accipitridae that it should

be transferred to this family. [t appears to represent

@ plant raptor, larger lhan the living Wedge-tailed

THE FOSSIL PELICANS OF AUSTRALASIA

Eagle, Aguila audax and more similar in size to the

extinct New Zealand Eagle, Harpagornis moorei.

COMPARISON OF AUSTRALIAN FORMS WITH

OTHER FOSSIL PELICANS

Brodkorb (1963) lists eight fossil species af

pelicans, and recently Harrison and Walker (1976)

have reported a ninth, Pelecanus aethiopicus from

East Africa. Many of these are not directly

comparable with most of the Australasian material

except P. novaezealandiae. P. intermedius and

P. fraasi, both from the Miocene of Europe, are

known only from cranial material; P. haliews from

the early Pleistocene of North Ametica, a radius

representing a bird the size of P. erythrarhynchas;

Liptornis hesternus from the Miocene af Argentina, a

lower cervies! vertebra, whereas the Australian

material is represented by higher cervicals.

P. aethiapicus from the Middle Pleistacene of East

Africa is nat directly comparable ta Australian

fossils as it is represented only by a proximalend of a

tarsometatarsus and a scapula; scapulae of both

P. aethiopicus and one Australian Pleistocene fossil

pelican are so worn and broken that qualitative

comparisons are futile, although they are clearly

wilhin the same size range,

The remaining fossil pelicans are at least

represented by materials comparable ta Australian

specimens. P, gracilis (proximal humerus, femur,

scapula, partial tarsometatarsus):; P. cautleyi, (distal

ulna); P, sivalensis, (distal ulna); and. P, edessanus

(coracoid, tarsometatarsus).

As Miller (1966) pointed out, P. gracilis from the

early Miocene (Aquitanian) of Europe is decidedly

smaller and more slender-legged than P, conspicil-

latus, Unfortunately, because nothing is known of

the distal part of the tarsometatarsus, it cannot be

compared with P. firarensis or P. cadimurka,

although it is clearly in this size range.

P. cautleyi and P. sivalensis. based on distal ends

of wlnae from the Pliocene of India are smaller than

the living P. conspicillarus, (Miller, 1966) and

although within the size range of P. mrarensis and

P. cadimurka, cannot be compared directly,

P, odessanus from the early Pliocene of Russia is

similar in Size to the living P. conspicillatus, thos

larger than P. tirarensis and P. cadimurka. It difters

from FP. conspicillaius in the shape of the trechlea,

ridges, and muscle scars of the tarsometatarsus

(Miller. 1966),

STRATIGRAPHIC DISTRIBUTION OF

PELECANIDAE WITHIN AUSTRALASTA

Fossil pelicans are known from three parts of the

siratigeaphie column in Australasia, and direct

243

correlation with sediments bearing pelecanid fossils

elsewhere in the world is only approximate.

P. tirarensis accuts in the Etadunna Formation

and possibly the Wipajiri Formation of the Lake

Eyre Basin, Central Australia. The Etadunna

Formation overlies fluviatile rocks dated as

Paleocene-Eocene because of contained plant

remains, Unfortunately, none of the vertebrate

bearing sediments has datable plant remains, and

lithologic correlation must be used ta tie the

vertebrate bearing outcrops with subsurface geologic

sections bearing such plant remains. Further

correlation with plant-bearing marine sections to the

south allow a tentative assignment of the Etadunna

rocks to a mid-Tertiary, Oligo-Miocene age (Stirtan

et al, 1968), Correlation by use of fossil mammal

remains with the Namba Formation, further to the

southeast, which bears. pollen basally, suggests a

post Batefordian—Baleombian age of abour 14-16

million years BP (Tedford, et al., 1977). Based

primarily on marsupial mammalian correlations. the

Wipajitt Formation, which is channelled into the

Etadunna, is thought ta be of mid to late Miocene

age. (See Rich. 1979 for a summary of the

stratigraphy).

The remaining fossil pelicans of Australia have

been collected from a number of sites along Cooper

Creek, at Lake Kanunka (north of Cooper Creek},

and along the Warburton River, all in the Lake Eyre

Basin of northeastern South Australia; and on the

Darling Downs of southeastern Queensland, All of

these have been called Quaternary or Pleistocene tn

age, Locales along the Cooper and Warburton

Rivers include fossils found both as ‘float’ on the

surface and in place and are often associated with

marsupials normally restricted to Pleistocene—aged

sediments within Australia, Because many were

collected out of a stratigraphic context, placement

within the Pleistocene or even within the Recent is

not certain, Dating of the Darling Downs verte-

ebrates is based primarily on correlation using

marsupial mammals, some species of which are

entirely restricted to the Pleistocene, while others

range from Pleistocene into the Recent,

New Zealand pelican fossils are known from

several localities on both North and Sauth Islands.

All are no older than Holocene (see above),

DISCUSSION AND CONCLUSIONS

Pelecanus conspicillatus is the only extant pelican

in Australasia. This has certainly not always been

the case, During the last 2 million years, in the

Quaternary, as many ag 3 species Were present in

this biogeographic province, including: a smiali.

gracile form, P. cadimurka, n. sp.; a very large,

robust species, P. novaezealandiae: as well as a form

244

osteologically indistinguishable from the extant

P. conspicillatus. P. conspicillatus and P. cadimurka

were sympatric and contemporaneous in central

Australia, a situation reflected by the modern broad

geographic overlap of the small P. rufescens and the

large P. onocrotalus in Africa. Likewise, the possible

sympatry of P. conspicillatus and P. novaezealandiae

is mirrored in the modern avifauna by the overlap of

the large pelicans, P. crispus (the more gracile form)

and P. onocrotalus (the more robust form) in the

western Palaearctic. In fact, P. novaezealandiae,

known only as a fossil from New Zealand, was

contemporaneous with P. conspicillatus in Australia,

suggesting probable overlap in distribution for such

large mobile birds.

Only one other fossil pelican species is known

from Australia, Pelecanus tirarensis, of Miocene age.

It was a small form morphologically distinct from all

later species,

Our review of all the Australasian fossil pelicans

and comparison with all the recent pelicans support

the inclusion in the synonomy of P. conspicillatus of:

P, validipes, P. grandiceps and in part Archaeocyc-

nus lacustris, Although we agree with Miller (1966)

that of the three specimens De Vis referred to

P, proavus, one is not of a pelican and the other two

may be referred to P. conspicillatus, the illustrations

of the last holotype suggest a pelican distinct from

P. conspicillatus. Thus, determination of the status

of P. proavus is impracticable unless the holotype is

relocated. Provisionally the mame should be

retained. Our review has also demonstrated the

necessity for recognizing as a new species of small

and gracile pelican, P. cadimurka, and for elevating

to specific status P. noveazealandiae, which Scarlett

had named as a sub-species of P. conspicillatus.

REC. 8. AUST. MUS, 18(12): 235-264

September, 1981

As a result of this study, we have been impressed

by the uniformity in skeletal morphology of the

pelican species we examined, except for that of the

New World Brown Pelican, P. occidentalis, Being

the only diving pelican it is so distinctive in many

aspects of its biology and morphology, that further

study may well indicate it meriting separated generic

status in Leptopelicanus Reichenbach 1852 (1853),

Avium Syst. Nat.: 7,

REFERENCES

ARCHEY, G., 1931. Skeleton of an Australian pelican found

With moa bones, Rec. Auck, Instr. Mus. 1: 116-121.

BRODKORB, P., 1963, Catalogue of fossil birds. Pt, 1 Bul

Florida State Mus,, 7; 179-293,

BROWN, H.Y.L., 1894. Report of the Government Geologist

for year ended June 30, 1894. Government Printer,

Adelaide. 26pp.

DE VIS, C. W., 1892 (1891). Residue of the extinct birds of

Queensland as yet detected. Proc. Linn, Soc. NSW, 2(V1):

437-456,

DE VIS, C. W,, 1905, A contribution to the knowledge of the

extinct avifauna of Australia. Ann. Queenslund Mus,, 6:

1-25,

HARRISON, C. J. O.,and WALKER, C. A.,.1976. A new fossil

pelican from Olduvai. Bull) Br. Mus. Nat. Hist, (Geol.) 27,

4; 315-320,

MILLER, A. H., 1966. The fossil pelicans of Australia. Mem.

Queensland Mus., 14: 181-190,

NORTHCOTE, E. M., 1979. Determination of age and sex of

long bones of Mute Swan Cygnus olor, [bis 121: 74-80.

RICH, P. V., 1979, The Dromornithidae, an extinct family of

Jarge ground birds endemic to Australia. Bur. Min, Res.

Bull. 184: 1-196.

SCARLETT, R,, 1966. A pelican in New Zealand. Notornis

13(4); 204-217.

STIRTON, R. A., TEDFORD, R. H., and WOODBURNE,

M. O., 1968. Australian Tertiary deposits containing

terrestrial mammals. Univ. Calif. Pub. Geol. Sei. 77: 1-30.

TEDFORD, R. H., ARCHER, M., BARTHOLOMAI, A.,

PLANE, M., PLEDGE, N. S., RICH, T., RICH, P., and

WILLS, R. T., 1977. The discovery of Miocene vertebrates,

Lake Frome area, South Australia. Bur. Min. Res. Jour.

Aust, Geol. Geophys. 2: 53-57.

THE FOSSIL PELICANS OF AUSTRALASIA 245

TABLE 1

Fossil specimens assigned to the genus Pelecanus from the Cainozoic of Australasia

Specimens available

to:

PVR

A.H. GFv1

Specimen De Vis Miller (this

Number Element Old Name New Name (1905) (1966) paper)

AMNH 11494 p, R. tarsometatarsus — rf. tirarensis — — ff

Lost, De Vis d. L, tarsometatarsus proavus (holotype) ? proavus 4 _ —

SAM P13858 sd. R. tarsometatarsus tirarensis (holotype) tirarensis — J dV

SAM P18412_ d. R. tarsometatarsus validipes (holotype) rf. conspicillatus Y —_ F

SAM P22501_— d. L. tarsometatarsus — cadimurka, n. sp. (holotype) — - v

QM F1141 p. R. carpometacarpus proavus not pelican f " v

QM F3749 L. quadrate grandiceps rf. conspicillatus Wa J J

QM F3750 L, coracoid grandiceps rf, conspicillatus Vv J V

QM F3751 d. L. tarsometatarsus grandiceps (lectotype) rf. conspicillatus i J if

QM F3752 d. R. femur proavus tf. conspicillatus J J J

QM F3753 d. R. tibiotarsus proavus (syntype) rf. conspicillatus J vf Vf

QM F5529 4th cervical vertebra Archaeocycnus lacustris rf. conspicillatus J = J

QM F6669 L. coracoid _ rf, conspicillatus — — v

QM F10703 d. L. tarsometatarsus — rf. tirarensis — — J

UCMP 56321 —_L. quadrate conspicillatus rf. conspicillatus _— v J

UCMP 56322 d, L, tarsometatarsus grandiceps rf. conspicillatus — Vv Vv

UCMP 56348 d. L. ulna conspicillatus rf. conspicillatus — / J

UCMP 56394 3rd ceryical vertebra conspicillatus rf. conspicillatus — Fi v

UCMP 56633 R, scapula — rf. conspicillatus oe — Z

UCMP 60477 p. R. femur conspicillatus rf. conspicillatus — vf J

UCMP 60487 — L. coracoid conspicillatus rf, conspicillatus _— J J

UCMP 60503 —_L, cuneiform conspicillatus rf. conspicillatus _ J J

UCMP 60520 d. L, ulna conspicillatus rf, conspicillatus — J" if

UCMP 60521 d. R. tibiotarsus conspicillatus rf. conspicillatus = J J

UCMP 60549 — R. cuneiform conspicillatus rf. conspicillatus — Vv J

UCMP 60577. d. R. tarsometatarsus conspicillatus rf. cadimurka, n. sp. — J J

UCMP 60578 = d. R. tarsometatarsus conspicillatus tf, cadimurka, n. sp. i J Vv

UCMP 60640 d. L. humerus conspicillatus tf. conspicillatus _ " J

UCMP 60656 Ant, sternum conspicillatus rf. conspicillatus _ Vv

UCMP 60702 fused palatines conspicillatus rf. conspicillatus — J _—

UCMP 60988 d. R. tarsometatarsus _ tf. tirarensis — — J

UCMP 69587 p. R. humerus conspicillatus Accipitridae — Vv Vv

UCMP 113115 d. R. tarsometatarsus — rf. tirarensis — a J

UCMP 123382 4th cervical vertebra = rf. cadimurka n. sp. — J v

Scar-

Archey leit PVR

(1931) (1966) GFvT

CM AV12264 R, humerus conspicillatus

novaezealandiae tf. novaezealandiae — 4 vl

CM AV12482 d. L. femur conspicillatus

novaezealandiae rf. novaezealandiae — J .

CM AV13095_ p, R. humerus conspicillatus

novaezealandiae tf. novaezealandiae _— Vv .

CM AV15089 — R. coracoid conspicillatus

novaezealandiae rf. novaezealandiae — rd of

CM AV21355 _ partial skeleton conspicillatus

novaezealandiae (holotype) novaezealandiae _ J /

CM AV32001 — part of fused furcula —

and sternum tf. novaezealandiae — Z

Lost, Archey partial skeleton conspicillatus

novaezealandiae tf. novaezealandiae J J —

AWNM

AVS845.1 d. L. tibiotarsus _ rf. novaezealandiae = —

Horn, no No,

L. tibiotarsus

rf. novaezealandiae

246 REC. S, AUST. MUS. 18(12): 235-264 September, 1981

TABLE 2

Measurements in mm of tarsometatarsi of recent and Australasian fossil pelicans

Width of At proximal end of distal foramen

distal end Width of shaft Depth of shaft

n range x sd n range x sd n Tange x sd

FOSSIL

P. tirarensis

SAM P13858 type... 0-22 19,6

UCMP 60988 rf... eee @12.0 8.3

P. cadimurka

LEGMP 60577 £8. eae pete pace Battal 18.5 14.2 8.8

UCMP 60578 rf. 2. oe ee eee @19.3

P. novaezealandiae

CM AV21355 holotype ............... 26.0 18,5 12.0

Archey, Tast,rie.. 3 seas teeters $2 26.9 (Scarlett 1966)

P. proavus from figures of holotype ..... >18.0-18.6 >14.8-15.2

P. conspicillatus;

SAM 18412 (holotype of validipes) rf. -. 26.1 17.9 11.0

RECENT

P. conspicillatus.... 2.0.0.2... 0....02.. 20 21-26 24 1.5 19 15-19 $17 1.4 19 9-12 11 0.8

PV CRISDUS cx fs iittaasgaece «awa sect vleleyee 1 25 9 18 1 11

Phonocrotatus -s 0.5 23:5 ¢eaassssteetas 3 24-30 27 3.3 3 17-22 20 2.5 3 11-16) =«13 2.3

P. erythrorhynchos.................... § 22-25. 23 0.9 5 12-18 15 3.2 5 10-11 10 0.6

Po philippensis oo... 3 thee ean bb bee es 1 25 1 19 1 12

Pr FAlPesCens- v0 fie iveicelecs ile ainee eeMee 2 19-24 2 14-18 2 9-12

P. occidentalis ..... 0.2.00. 02002254. 9 17-21 920 1.4 9 12-16 14 1.0 9 6-9 8 0.8

TABLE 3

Measurements in mm of trochlea III of tarsometatarsi of recent and Australasian fossil pelicans

Internal depth External depth Distal width

n range x” sd n range x sd n range x” sd

FOSSIL

P. tirarensis

SAM P13858 holotype ......-......0.. 11.3 11.2 9.4

TOMDPIOZ0S Te i fea cetacean 9.7 9.8 6.9

UCMP 60988 rf. oe ee >9.9 >10.3 @7.0

UCMP 113115 rf. ...,...--4.- pbrner he @10,0 9.9 6.4

P. cadimurka

SAM P22501 holotype ........-...,.... 9.2 9.6 >6.0

UCMP. 6057756. oss pec cie epee dy peenih >8.9 10.8 7.8

UCMP 60578 rf. 45 tt cele a dep dias 11.5 12.0 7.2

P. novaezealandiae

CM AV21355 holotype ......,.....-.. 14.8 15.2 10,0

P. proavus from figure of holotype -.. . - - >6.8

P. conspicillatus:

SAM 18412 (holotype of validipes) rf... . 15.0 15.3 10.1

QM F37S1 (lectotype of grandiceps) rf... 15.8 17.0 10.2

UCMP'56322). 0 tecaa etsy hesag tees nwt 15.3 10.1

RECENT

P. conspicillatus.......0.0...0225..2-.. 19 12-16 14 1.0 20 13,17 15 1.0 17 8.10 9 0.6

Py CPiSPUS T,. << exh SRS tt eile oo eb we 1 14 1 14 1 10

P. onocratalus . 2... eee eee 2 14-17 3 14-16 15 1.3 2 9-10

P. erythrorhynchos............2...2... 5 13-14 9 13 0.4 5 13-14 14 0.4 2 9

P. philippensis ... 0.0.0.6. 0.000 cue een 1 14 1 14 1 9.5

PL TUPESCENS i el Rael titer de ee 2 11-14 2 12-14 1 7-9

Ppecidenidalisy #& + 5.+) esr Sige 9 8-11 10 0.7 9 8-11 10 0.8 1 6

THE FOSSIL PELICANS OF AUSTRALASIA 247

TABLE 4

Measurements in mm of trochlea II and IV of tarsometatarsi of recent and Australasian fossil pelicans

External depth Anterior width Internal depth

of II of II of IV

n range x sd n range * sd n range x sd

FOSSIL

P. tirarensis

SAM P13858 holotype ........-....... 10.2 4.5

OM FIO703 TE... ee ees 9.1 4.0

P. cadimurka

UGMP 60879 TED ici lity paevacmoncoes 5.7 10.2

UCMP 60578 rf. 2... ec eee 10.6 5.5 11.2

P. novaezealandiae

CM AV21355 holotype ....,.......+-5 12.9 13.0

P. proavus from figure of holotype ...... >5.6

P. conspicillatus:

OM F3751 (lectotype of grandiceps) rf. . . >14.4

SAM 18412 (holotype of validipes) rf. .. . 13.5 7.8 14.0

UCMP 56322 rhe con + bie, cyereirt arian meets 13.9

RECENT

P. conspicillatus...... 2.02.4... ...00.. 20 «11-15 13 1,0 17 6-10 7 0.7 20 =: 12-15 13 0.9

PLCvispuss - boos ss toctss dar tts poneln 1 13 1 1 14

P. onocrotalus 2.0.6.0... eee 3 12-15 13 1.4 2 7-8 3 14-18 15 2.3

P. erythrorhynchos.... 02.0620... eee S 12-13 12 0.4 2 5 12-13 13 0.4

P. philippensis vccsa oc cecsa wees as eeiien ] 13 1 7 1 14

Ps FUPESEOHS oyu, ais inl lle le wooed bla as ohh sities 2 10-13 2 6-7 2 11-14

P occidentalis: ows. eeneyeerersseace 9 8-11 10 0.7 1 5 9 8-11 10 0.7

TABLE 5

Measurements in mm of distal end of femora of recent and Australasian fossil pelicans

Width across internal

Width of distal end and external condyles

n range x sd n range x sd

FOSSILS

P. novaezealandiae

CM AV21355 holotype 2.0.0.0 0 0.000 eee eee eee tees 36.2 28.4

SMA WIZAB2 Tf isi ig eta etch phe aA el be ected tote wert ct a elelhwre dacase bye >35 >28.3

Archey; Ost, rhs. .o cee pss ce Peas ELL La Baas t tae efeilise a army Efe 38 (Scarlett 1966)

P. conspicillatus:

SADE BES TSS TES ps sgeee et yihs bs pean elnfee oa! oleate so rh epajater et olfllsiess nals nah ad PE 3 an @29.0 24.4

RECENT

P. CONSPICH AIS: oye) sp o\ea tht nel odientigeea iene Bins sioeceid y39e $e 15 30-37 =. 332 2.2 15 24-29 26 1.7

PCHSPUS! cs cceg passe p ative es sid qwetttamee ds sadtt tee debra des 2 35-37 2 27-29

P.-QHOCTORTUS.”. i DGGG fe si ajetees tite lee olnegetece si pinaclane on dh Bales Tae ee 2 33-35 2 26-29

PSR ry T TOPPA AE CTO Se ote chan Peete ole fens alain tay ov sh iif so coe «play el ellie leant te jee me 9 28-33 31 1.9 9 24-27 825 0.9

P; piittopensisr. 25.755 cay e200 8 BARR e eae ABO Bate wtih 1 35 1 26

BoPRsesGeH®: sa-rcop een tele g dare eng gis 0 ns carly ln SET Ieee wgup “2 26-32 2 21-25

Pi otcidentalis neney ali aetocscesog i camels bee satan dalels exces. ole Ss 20-26 23 1.8 5 16-19 19 2.1

September, 1981

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249

THE FOSSIL PELICANS OF AUSTRALASIA

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snyoppiaidsuod “gq

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ps x asuel u ps x osuel u ps x osuel u ps x osuel u

saskudodeStiasod dauase setindodsakior fore quinse jo

SSOIOB YIPIA, jo 1otrajsod ssoioe YIPIAA Jolia}sod 0} sashydod

SSOIOB YIDIAA -e3Azoid wo yisusT

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250 REC. S. AUST. MUS. 18(12): 235-264 September, 1981

TABLE 8

Measurements in mm of quadrates of recent and Australasian fossil pelicans

Height from Width across

mandibular to mandibular Width across

squamosal articulation articulation orbital process

n range x sd n range * sd n range x sd

FOSSIL

P. conspicillatus

OM F3749 rf. eet ee een 33.9 27.0

UICIMEP SOS 2042 os ecg order t ne he's doe @31.5 >23.3 >24.6

RECENT

P. conspiciliatus , ..07 70 e 84 o> de oe 92 11 28-36 32 2.3 11 22-27 26 1.9 11 24-30 8=6.28 2.1

PiGrispus’ oo gxscck ee aee yee ei 1 32 1 24 1 26

P. onocrotalus 2.0.0.0... 06 e eee 1 40 1 30 1 30

P. erythrorhynchos...............00445 1 29 1 23 1 24

P. philippensis :o0 emcee cs ga eceig pees es 1 33 1 24 1 27

Pr tUfescens= 06 ice cp cece red cau 2 29-33 2 20-25 2 22-26

P. occidentalis... 0.06... 1 32 1 20 1 19

TABLE 9

Measurements in mm of coracoids of recent and Australasian fossil pelicans

Width from

Width of shaft coracoidal fenestra to

at coracoidal medial margin of shaft

fenestra on posterior surface

n range x sd n range x sd

FOSSILS

P. conspicillatus:

ONGFS TSO The oo sens nies ig aatoote hey fl lg rersaas shite cibaoh Hotuchtch Maki degen hated abyha @19,4 8.4

OM RGG69 TE. cou. dit areca legs areal UUs gare w beat ate aie a gis: 17.4 7.0

P. novaezealandiae

CMA V2135S RORY BE: i. ato es eae eigen OF HT) GF aed orest glelee dagtd a alaty 16.8 9.7

CM AMESGRS TE) sy wetiyerreay Ss hp Hoey o RES A pedhe! aah Sete bebe elated oe opt ERI gk @17.5 8.7

RECENT

P, COnSPictllAtus sc cae cc cee aa Cova dbegg eenes sured be 13 14-19 16 1.7 13 6-9 7 1.0

Pr CHSDUS™ ot se ene SR SUR EES hece gee tovahy damlide praise tet erat sag LE 2 17-20 2 7-9

PY GROCTOTANS. ep ABE dep 4 ayy ne oot + alia ta tel td std te ssew ated d: 2 18-19 2 9

DP. \GPYARTOPRYNCHOS!-s 6%, ols, orod egg 408 || bAace Se As ee lt aa tin eed a dake 9 14-18 16 1.0 9 6-9 7 1.1

Py philippensis. sco 2° tbe ag othe bb rit 18S ee late es polelnlgg oe ee angi 2 18-19 2 8-9

Fis TUPOSCENS TG STE SELECTS DIAL LG Wile BARI dials ol be eu yyy gly 2 13-16 2 5-8

BP. OCCIMERTATIS (fo. cvieeg hay eae ee ty Beene eer ee s tale bs Pl eee ae 5 11-13 11 0.7 5 5-6 5 0.4

THE FOSSIL PELICANS OF AUSTRALASIA 251

TABLE 10

Measurements in mm of third cervical vertebrae of recent and Australasian fossil pelicans

Length of left

Width of anterior side of anterior

Width across articular surface articular surface

prezygapophyses of centrum of centrum

n range x sd n range x sd n range x sd

FOSSIL

P. conspicillatus

TEMP 56394 hg oe cece chee a cetalationss 24.9 16.1 9.8

RECENT

P. conspicillatus ... . - biypeeraayetszan @ 22-26 24 1.4 9 11-14 12 0.9 9 7-11 10 1.5

Bey CHESTS BS wasn ie a caste Shiiin fa abot & jedutena|§ l 23 1 12 1 11

P. onocrotalus ...,...-...---- srr93 2 23-26 2 11-12 2 10

P. erythrorhynchos...- . ceteteleit exeeame Pe 21-23 2 11-12 2 9-11

P. philippensis ............ a cnbe afeten 1 18 1 10 1 8

Prrufestens 2.0% :2¢ 0ti 2a rr uhispe 19-21 2 11-12 2 8-10

P. occidentalis 0.0.0.6... cece ens 1 16 1 9 1 6

TABLE 11

Measurements in mm of left prezygapophyses of third cervical vertebrae of recent and Australasian fossil pelicans

Length Width

n range x sd n range x sd

FOSSIL

P. conspicillatus

DICMP 36394 rE e seescach eld debate occice pais obebalobartoracere peolec’, giaeeazazece pb elefobels atid 11.5 6.3

RECENT

P: COMSPICHIAUS . 5... Be\o.s4.05 canny avery SMPTE TE Peer LE bee? Sista ee, OD 10-1412 1.3 9 5-7 6 0.5

Pi G7ISPUS) os pee ceases eid y phat aqupeee a pac galn ever aneewureait ti, 1 12 1 6

Py CUED oo oss eah YG Son ca tsh ow jaen chee df [acns eft fos tee] ote fe leone] 4 a fave oh Ohi ja wae 2 11 2 6-7

P erythirorWynchoss os\e<es 40-25 bg nesses PSE E esa yea PEED Bene eee 2 11 2 4-5

P. philippensis .-... 2) 6220 0..0225. Et lgeneretewre atoll ae arene acne thse da yet 1 9 1 5

Pmifeseiner are ten ee ot BOR eee eGR Sap s teem tie fob) otitel 2 9-10 2 5

Pu O06tdertAlls: vaqwce eo ot) ae lnjcty erent a teg As paves nla ide eed tect Mle bo pity wd 10 1 4

TABLE 12

Measurements in mm of sterna at coracoidal sulci of recent and Australasian fossil pelicans

Depths across right Depths across left

dorsal and ventral dorsal and ventral Minimum

lips at maximum lips at maximum distance between

expansion expansion sulci

n range x sd 0 range x sd n range x sd

FOSSIL

P. conspicillatus

UCMP 60656 rf. .....,.---5.----44--- 13.9 16.2 4.4

RECENT

P. conspicillatus . .... pesssacesssasaz: 10 13-18 16 1.4 10 «13-19 16 1.7 10 0-7 4.5 2,3

P. onocrotalus ......,. wae eee Ee. 21 1 19 1 10

P. erythrarhynchos. 2.000.066 0 0. 0c eee 5 9-16 12 3.0 4 8-16 12 3.9 5 4-11 7.5 2.7

P. philippensis 2.00.60. ees 1 18 1 17 1 9

P. rufescens ......... =: Lf ‘sescem a 16 1 17 i 6

P, occidentalis. js) icssi. qoccey vests el cnet 9 7-9 8 0.5 9 7-9 8 0.6 9 7-11 9 1.2

September, 1981

REC. S. AUST. MUS. 18(12): 235-264

252

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THE FOSSIL PELICANS OF AUSTRALASIA

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TIISSOA

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anscord douejnanie coakincas sednordseytsoiddo pue

ausoddo jo 1ys1aR Teujn jo yIplA Jeupn jo 1ystoyy uonepnonie reuyn

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254 REC. S. AUST. MUS. 18(12): 235-264 September, 1981

TABLE 15

Measurements in mm of proximal end of femora of recent and Australasian fossil pelicans

Width of proximal end Depth of head Depth of trochanter

n range x sd n range x sd n range x. sd

FOSSIL

P. novaezealandiae

CM AV21355 holotype ............... 35.4 15.1 5.0

Archey, lost, rf. ued piesa eae wes 36.0 (Scarlett 1966)

P. conspicillatus

UCMP 60477 rh. fics edi neeta yp testa ys 29.7 14.5 > 25.5

RECENT

P. conspicillatus.. 0.0.0 14 26-33-29 2.0 14 13-16 14 0.8 14 22-2724 1.5

PPCHISRUS (4-4 pcb oye ett taste e edt 2 31-34 2 15-16 2 25-26

P. onocrotalus .... 0... eee 1 30 1 15 1 24

P. erythrorhynchos.. 0.00... c ee eee 5 29 0.3 5 13-14 14 0.3 5 22-25 24 1.2

P. philippensis .. 06.000 ccc cee ee eee 1 32 1 15 1 23

P, rufescens ........ 0.6. 2 24-29 2 12-14 2 19-24

P. occidentalis ............ 00000 -05- ) 19-25 23 2.0 9 9-12 11 11 9 16-22 19 1.7

TABLE 16

Measurements in mm of distal end of humeri of recent and Australasian fossil pelicans

Length of Depth of Depth of

external condyle external condyle internal condyle

n range x sd n range x sd n range x sd

FOSSIL

P. novaezealandiae

CM AV21355 holotype .............-- 22.7 26.5 14.8

CM AV12264 rf. 2. eee >22 >27 @15

P. conspicillatus

UCMP 60640 rf. 0.0... eee 20.7 >21.1 13.1

RECENT

P. conspicillatus ... 2.22. 17 19-25 22 1.6 18 22.27 25 1.5 18 12-16 14 1,1

PS GVISPUR | Oj RP A coe attr peels 2 = 23-24 1 28 16

P. onocrotalus ...... 0... cece eee jf ae 23-28 2 24-30 2 15-17

P. erythrorhynchos..............2--.-. 5 20-22 21 0.6 5 24.26 24 1.2 5 14-15 15 0.2

PrTUpeseens: 29s pet tec des oly 3 rare cite 2 18-22 2 21-25 2 =12-15

P. occidentalis .......0...00.00 0240... 9 15-20 18 1.5 9 17-22 21 1.4 9 11-13 12 0.8

THE FOSSIL PELICANS OF AUSTRALASIA 255

TABLE 17

Measurements in mm of scapulae of recent and Australasian fossil pelicans

Width of shaft

Width across at distal end of

Total length glenoid facet glenoid facet

n range x sd n range x sd n range x sd

FOSSIL

P. novaezealandiae

CM AV21355 holotype ............... 23.7 14.7

P. conspicillatus

WEMP 56633 Th agstarias ptt teed Wein >101.8 @21.2 @13.8

RECENT

P. conspicillatus.......0......0.00004.. 9 114-140 126 9.2 il =20-25. 0-22 1.2 11 11-1714 1.3

Pe CHSPUS 8.2 ate pes EE bth oD 5. 1 134 1 24 16

Ponocrotalus, M45) S959 2327 AR ad 2 129-137 2 24-25 2 16-17

P. erythrorhynchos...........00..0000% 5 121-133 126 5.5 5 20-22-21 0.8 5 13-20 16 2.9

P. philippensis tes 2. cnt toi sade ceteae : 2 129-137 2 24-25 2 =16-17

Bi FULESCEMS 0 8, PUTMAN Bee toe ae 4 2 101-119 101 1 2 2 16

P. occidentalis ..........0....000004.. 10 81-103 97 6.5 8 14-18 16 1.3 10 8-13 11 LiF

September, 1981

REC. S. AUST. MUS. 18(12): 235-264

256

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tr I ez I 68 I 0Sz [Etter trent ce tne n nese arse e ese nea e cesses suaosafni ‘g

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avipuv]pazavaou ‘q

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sueoted [Issoj uviseyensny pue jusde1 Jo eBioesuAS JO WIU Ul s}UDUTOINsBoy

8. ATaVL

THE FOSSIL PELICANS OF AUSTRALASIA 257

TABLE 19

Logo ratios of width: length of synsacra and of ilio-ischiatic foramina of recent and Australasian fossil pelicans

(cf. Table 18 and Northcote 1979)

Synsacrum Ilio-ischiatic foramen

n range x sd n range x sd

FOSSIL

P. novaezealandiae

CMA NV2I3S5 OILY Pe nF as deste kot a dle edad Sy, aches eM, eR 0.47 0.31

RECENT

P.("CONSPICiAtUS:..,0 eiPinis de 8 sob dk MIN cease tcgapecbsa, whe ae decal a weno ess 12 0.47-0.51 0.49 0.01 20 0.33-0.48 0.40 0.04

P SONOCTOLAIUS sped ros a dk BE ye Pete Sats BARES Se Nit ahs Me | Pela. H o® 2 0.44-0.45 2 0.32-0.35

PERYENTORNYNCHOS Mit niyoe She fewer ees eet TDs anne ae eka ets dM 2 0.42-0.43 2 0.29-0.34

Be TUSESCENS 21). Rose att OA es TR Be! eo OE ee ee iL 0.45 1 0.28

P? OCCIAENIGLIS en Th,.8: PO heats LE TR, Meld: 3. ea Pe haa A 1 0.42 1 0.31

258

REC. S. AUST. MUS. 18(12): 235-264

September, 1981

—

1000 Km

Lake Eyre IN ahr

X 10, 9813

C2 7€

@) 100 Km

FIG. 1. Map of localities where fossil pelicans were found in

Australasia. Map A: Pelecanus proavus, holotype (1); P.

novaezealandiae, holotype (4), referred (2, 3 and 5); Map B:

P. tirarensis, holotype (7), referred (6); P. cadimurka n. sp.

holotype (10), referred (8, 13); P. conspicillatus, referred (8,

9, 11-14).

THE FOSSIL PELICANS OF AUSTRALASIA

FIG. 2. Fossil pelicans of Australia. Pelecanus rf. cadimurka,

UCMP 123382, fourth cervical vertebra (A); Accipitridae,

UCMP 69587, proximal right humerus (B). P. rf.

conspicillatus, UCMP 60503, left cuneiform (C); UCMP

60549, right cuneiform (D); UCMP 60520, distal left ulna

fragment (E); P. validipes, holotype, SAM 18412, distal right

tarsometatarsus (F); P. grandiceps, lectotype, QM F3751,

distal left tarsometatarsus (G); P. tirarensis, holotype, SAM

P13858, distal right tarsometatarsus (H). See scale for size.

REC, 8. AUST, MUS. 18(12); 235-264

FIG, 3. Fossil pelicans of Australia. Pelecanus rf, conspicillatus,

UCMP 60656, anterior sternal fragment (A); UCMP 56394

anterior 3rd cervical vertebra (B); Archaeocycnus lacustris,

syntype, QM F5529, 4th cervical vertebra (C); rf. P.

conspicillatus, QM F3749, left quadrate fragment (D);:

UCMP 56321, left quadrate (E); UCMP 60640, distal left

humerus (F); UCMP 56633, right scapula (G); P. rf.

tirarensis, UCMP 113115, distal right tarsometatarsus (H);

UCMP 60988, distal right tarsometatarsus (I); P. rf.

conspicillatus, UCMP 56322, distal left tarsometatarsus (J);

P. rf. cadimurka, UCMP 60577, distal right tarsometatarsus

(K); UCMP 60578, distal right tarsometatarsus (L). See scale

for size.

September, 1981

THE FOSSIL PELICANS OF AUSTRALASIA

FIG. 4. Fossil pelicans of Australia. Pelecanus rf. conspicillatus,

UCMP 60477, proximal right femur, (A); QM F3752, distal

right femur, (B); QM FF3750, left coracoid fragment, (C);

UCMP 60521, distal right tibiotarsus, (D); QM F3753, distal

right tibiotarsus, (E). See scale for size.

262

Supratrochanteric width (mm)

REC. S. AUST. MUS. 18(12): 235-264 September, 1981

P. onocrotalus i

anaes P. novaezealandiae

P. erythrerhynchos a

)

P. conspicillatus

220

230 240 250 260 270 280 290

Length of synsacrum (mm)

FIG. 5. Comparison of synsacral measurements of the large

species of Pelecanus.

THE FOSSIL PELICANS OF AUSTRALASIA 263

FIG. 6. Holotype of Pelecanus cadimurka, n. sp., SAM P22501,

distal left tarsometatarsus. A. Anterior view. B. Distal view.

264

Distal Width of Trochlea I (mm)

REC. S. AUST. MUS. 18(12): 235-264

September, 1981

ul P. novaezealandiae gat onocrotalus

‘P. validipes’

P cri wi

. Crispus ce P. grandiceps ’

sa ; A ®

10 P. philippensis ¥

o e

Se es

¢ .-t

: P. tirarensis ® oC re

O P. erythrorhynchos

8 e Je =P. conspicillatus

Xe <> ————- P. rufescens

! e

P. occidentalis

P. cadimurka

8 92 Oleh wy <8: te “Tbe -le- VF

External Depth of Trochlea II (mm)

18 19 20

FIG. 7. Comparison of distal tarsometatarsal measurements of

living and extinct species of Pelecanus.

ON SOME OVIPAROUS FILARIAL NEMATODES MAINLY FROM

AUSTRALIAN BIRDS

BY ODILE BAIN AND PATRICIA M. MAWSON

Summary

Three groups of filarial nematodes are represented in this collection.

1. Aproctinae

Four species are described: Pseudaprocta copemani n.sp from Petroica multicolor; Aprocta

boulengeri n.sp. from Strepera graculina; Aprocta bakeri n.sp. from Corvus orru; Lissonema sp.

from Ninox novaeseelandiae.

2. Dicheilonematinae

Serratospiculum guttatum (Schneider, 1866) from Falco longipennis and F. peregrinus, and S.

tendo (Nitzsch, 1819) from F. peregrinus are redescribed. S seurati n.sp. is described.

3. Diplotriaeninae

Three species are described: D. spratti n.sp. from Oreoica gutturalis, D. beveridgei n.sp. from

Corvus orru, and D. smithi n.sp. from Acanthogenys rufogularis. Additional morphology is

given for D. delta Johnston and Mawson, 1940 and D. falconis (Connal, 1912).

ON SOME OVIPAROUS FILARIAL NEMATODES MAINLY FROM AUSTRALIAN BIRDS”

ODILE BAIN* and PATRICIA M, MAWSON**

“Laboratoire de Zoologie-Vers, associé au CNRS, Muséum national d’ Histoire Naturelle, 43 rue

Cuvier, F 75231 Paris Cedex 05

**Department of Zoology, University of Adelaide, G.P.O, Box 498, Adelaide, South Australia

5001

ABSTRACT

BAIN, O,, and MAWSON, P. M. 1981, Some oviparous filarial

nematodes mainly rom Australian birds, Ree, S. Aust, Mus,

18(i3): 265-284.

Three groups of filarial nematodes are rep-

resented in this collection,

1. Aproctinae.

Four species are described: Psewdaprocia cope-

mant n.sp. from FPetroica multicolor: Aprecta

boulengeri n.sp. from Strepera graculina; Aprocta

bakeri n.sp. trom Corvus orru; Lissonema sp, from

Ninox novaeseelandiae.

Pseudaprocta is regarded as being closely linked

with the Seuratoidea.

The species assigned to Apracta are heterogene-

ous, falling into two distinct morphological groups,

one of which corresponds with the genus Lissonema,

which is therefore reinstated, Apracta spp, have a

short ovejector, small caudal papillae in males,

spicules generally subequal with fine distal

extremities, no deirids, small eggs (less than 55 «jm

long), short larvae (around 200 zm long) with short

tail bearing terminal spines. Lissonema spp. have a

long ovejector, at least one large caudal papilla in

male, spicules usually equal, with large blunt distal

extremities, deirids, large eggs (more than 65 1m

long), long larvae (more than 400 jm long) with

long unarmed tail, The genus Squamofilaria is

regarded as a synonym of Lissonema and its species

are shown to belong either to Lissonema or Aprocta.

Lissonema contains 20 spp. (19 n, comb.) and

Aprocta 17 spp. (2 n. comb). The host ranges of

these two yenera are completely different.

2. Dicheilonematinae.

Serratospiculum guttatum (Schneider, 1866) from

Falco longipennis and F. peregrinus, and S. tendo

(Nitesch, 1819) from F. peregrinus are redescribed.

S. seurati n.sp is described,

Serratospiculum, parasites of Falconidac, contains

a total of seven species. These species fall within two

groups depending on the spicule length. (1) Those

with short spicules are separated further, in that S.

turkestanicum Skrjabin, 1916, S. guttatum, S.

kwangsiensis Hsii, 1963, possess specific cuticular

orhamentations in the female; while S. chungi

Hoeppli et Hsii, 1929, S. seurafi n.sp. (= F.

allenuaia Rud,, 1819 sensu Seurat, 1915), S.

congolensis Vuylstecke 1957, are distinguished by

the morphology of the epaulettes and the right

spicule; (2) those with long spicules: S. rendo(=? S.

thoracis Tubangui, 1934; = ? S. lii Ezzat et Tadras,

1958).

3, Diplotriaeninae.

Three species are described: D. spretti nosp. tram

Oreoica gutturalis, D. beveridgei n.sp trom Corviis

orru, and D, smithi n.sp from Acanthogenys

rufogularis. Additional morphology is given for D.

delta Johnston and Mawson, 1940 and D. falconis

(Connal, 1912),

DEPOSITION OF TYPE MATERIAL

Holotypes and allotypes have been deposited in

the South Australian Museum, Adelaide; (SAM in

text); other material has been deposited as noted, in

the Muséum National d’Histoire Nuturelle, Paris

(MNHN) and in the Australian Helminthological

Collection (AHC) at present housed in the South

Australian Museum,

In the species described here, the female has been

designated as holotype, and the male as allotype,

This is because the family, genera and species of

filarial worms are more often identified by the

female than by the male.

DESCRIPTION OF MATERIAL

I, Aproctoidea—Aproctinae

Genus Pseudaprocta Schikhobalova

The morphological characters of the genus

Pseudaprocta are closer to those of Seuratoidea

Ascaridida than of Spirurida (Chabaud, 1974);

mouth large and triangular, six external labial

papillae, situated on the same circle as the cephalic

papillae, oesophagus short and very thick, without

This study was carried out while working under a grant (to O,B,) from the World Health Organisation.

266

differentiation into two parts, spicules equal and

simple, and one pair of caudal papillae situated

laterally (8th pair).

It seems that Pseudaprocta, and possibly the

Aproctinae, evolved from the Seuratoidea.

Pseudaprocta copemani n.sp.

(Figs. 1, 2,)

Material examined

From Petroica multicolor Gmelin (Muscicapidae)

from Maggs Mt, Tasmania. Holotype &, allotype ¢

(SAM V2096, V2097), 12 and 1d (MNHN, Paris,

No. 201 HD); 22 and 3d (AHC 6277).

From Pachycephala pectoralis Latham (Mus-

cicapidae), from Heatherleigh, S. Aust.: 3 pieces of

@ (2 anterior, 1 posterior), 1d, 1 posterior end ¢.

(AHC 6282).

Description

Morphology is shown in Figures 1 and 2. The

specimens, except in one case, are surrounded with a

REC. 8. AUST. MUS. 18(13): 265-284

September, 1981

thin membrane detached from the rest of the cuticle

and forming a vesicle around the body, beginning at

the level of the oesophagus and ending at the level of

the tail.

In the female, the tail ends in a small rounded

knob (Fig. 1, I). In the male, the precloacal unpaired

papilla is particularly big (Fig. 2, B and C). The

spicule sheaths are thickened near the cloaca; these

thickenings apparently assume the role of a

gubernaculum (Fig. 2, E).

Measurements

Holotype 2: body length 16.5 mm, width 500 wm;

nerve ring, deirids and vulva at 160 um, 320 pm and

370 ~m from head; length of buccal capsule 25 .m;

length of oesophagus 585 jim; length of tail 320 wm.

Allotype d: length of body 8.9 mm; width 310

wm; nerve ring 160 wm from head; length of

oesophagus 520 ym; length of left spicule, right

spicule and tail respectively 640 4m, 600 pm, 190

oun

100 Um

150 Jum

Figure 1. Pseudaprocta copemani n.sp., female. A, anterior end,

ventral view; B, C, & D, head in apical, dorsal and ventral

views; E, larva; F & G, head of larva, from hook side and

opposite side; H & I, tail in lateral and ventral views; J, tip of

tail; K, transverse section of body. A, H, I, J, K, to scale

200 »m; B, C, D, to scale 100 pm; E, F, G, to scale 10 pm,

OVIPAROUS FILARIAL NEMATODES

Figure 2. Pseudaprocta copemani n.sp., male. A, anterior end,

lateral view; B & C, caudal region, lateral and ventral views;

D, region of deirid and excretory pore, lateral view; E, distal

extremities of spicules and gubernaculum, lateral view. A, B,

C, to scale 200 pm; D, E, to scale 100 um.

Paratype %: body length 18 mm; width 520 um;

nerve ring, deirids, excretory pore and vulva at 152

wm, 320 wm, 270 wm and 435 wm from head; buccal

capsule 25 um long; egg 52 pm X 42 pm; larva 480

um Xx 8um.

Another 3 from AHC6277: body length 9 mm

long, width 400 jm: nerve ring, deirids and

excretory pore 160 pm, 290 um and 230 um from

head; buccal capsule 22 um long; oesophagus 440

um long; length of left spicule, right spicule and tail

respectively 550, 575 and 200 pm.

Discussion

This material is distinguished by its longer spicules

from P. gubernaculum Schikhobalova, 1930

(Caucasia); P. decorata Li, 1933 (China); P. buckleyi

(Singh, 1949) and P. leiperi (Rasheed, 1960) (India),

and P. ungriai Chabaud et al., 1964 (Madagascar), in

which the spicules are about 400 pm long. It differs

from P. sichotealinensis Oshmarin & Belouss, 1951

from Eastern USSR (Sonin, 1966) and P. skrjabini

(Ali, 1956) from India, by the rather greater spacing,

longitudinally, of the papillae near the cloaca. P.

myzanthae Johnston & Mawson, 1940, from

Myzantha flavigula (Meliphagidae) from South

Australia, is known only by the female, the

measurements of which are very similar to those of

the present specimens with the exception of the body

length which is greater. It seems better to keep the

two species separate, in view of the difference in

hosts, and of the fact that Johnston & Mawson show

a slightly different arrangement of the cephalic

spines (Johnston & Mawson, 1940, fig. 14). We

therefore propose it as a new species, P. copemani,

in honour of our Australian colleague Bruce

Copeman.

The Genera Aprocta Linstow, Lissonema Linstow,

and Squamifilaria Schmerling

The genus Aprocta according to Anderson and

Bain (1976), is large and heterogenous. Some

authors (Boulenger, 1928; Sandground, 1933

etc . . .) had previously proposed to recognize two

genera, Aprocta Linstow, 1883 and Lissonema

Linstow, 1903, separated on the form of the head

(flat or with two lateral elevations), the oesophagus

(divided or not), the caudal papillae of the male (few

and small or large and numerous). As pointed out by

268

Anderson and Chabaud (1958) the two first at these

characters are of doubtful use, because there are

many species in which the division is not clear,

However, since theif work, further descriptions and

redescriptions have been made (Ezzat and Tadros,

1958; Rasheed, 1960; Diaz-Ungria, 1963; Chabaud

et al., 1964; Schmidt and Kuntz, 1970; Yoyotte,

1972; Quentin et al., 1976), so the morphology of the

group is better understood.

Tt appears that the structure of the ovejector ts of

importance, as this can be either of two distinct

types: it may be long (1500-5000 2m) often with the

middle part dilated to form a chamber (e.g, A.

travassosi Caballero, 1938) or very short (200-600

jam) and simple (e.g. A.turgida Stossich, 1902, in

Chabaud and Choquet, 1955), Examination of well

described species shows that the character “ovejec-

tor long" is found in species with large caudal

papillae, and the character ‘‘ovejector short"’ is

found in species with small papillae.

This division is borne out by other characters:

spicules equal or unequal, with tips wide or fine,

presence or absence of deirids, large or small eggs,

larvae with long smooth tails (Chabaud e7 al,, 1959)

or short spiny tails (Quentin er al., 1976).

The two groups represent Linstow’s two genera,

Aprocta and Lissonema, and Lissonema 1s now

reinstated. Aprocta Linstow, 1883: ovejector short;

caudal papillae small; spicules generally subequal

with distal extremity finely pointed; deirids absent:

eggs small (less than 55 yom long); larvae short

(about 200 xm), with short tail with terminal spines.

Type species; A.cylindrica Linstow, 1883 (redes-

cribed by Quentin et al., 1976) trom Passeriformes.

Other species sufficiently known to be allotted to

the genus, with the bird groups in which they occur,

are:

A. angolica (Vuylsteke, 1953); Passeriformes

A. caballerat Ybarra, 1948; Faleoniformes and

Passeriformes

A. cercomelae Sonin. 1966; Passeriformes

A. colaptidis Schuurmans-Stekhoven, 1952; Pas-

seriformes

A. crassa Railliet et Henry, 1910; Ralliformes

A. laevicutis (Sandground, 1933); Passeritormes

A. matronensis Railliet et Henry, 1910; Pas-

seriformes

A. narium Linstow, 1901; Falconitormes

A. ophthalmophaga Stossich, 1902; Palconiformes

A. orbitalis Linstow, 1901; Falconiformes

A. ptiloscelidis

Charadriiformes

Schuurmans-Stekhoven, 1952;

REC. & AUST. MUS. 18(13); 265-284

Seplember, [981

A. suidarikowi Sobolev, 1947*; Charadriiformes et

Ralliformes

A. textori Vuylsteke, 1953, Passeriformes

A, turgida Stossich, 1902 (redeseribed by

Chabaud et Choquet, 1955); Lariformes,

The species A. caudata Mendonca, 1961, of which

only the male is known, seems to belong to the

Splendidoblariinac.

Lissonema Linstaw, 1903: ovejector long; at least

one large caudal papilla (a single precloacal papilla,

0-4 pairs cloacal papillae); spicules generally equal,

with distal extemity wide and blunt; deirids present;

egps large (over 65 wm long), larvae large (over

400 jm long), tail long unarmed. Type species:

L, rotundata Linstow, 1903 (redescrihed by

Rasheed, 1960) parasitic in Cuculiformes.

Other species which are sufficiently known to be

allotted to the genus, listed with the bird groups in

which they occur, are:

L. brevicaudata (Chow, 1939) n.comb,

(Strigiformes).

L. calliderma (Schmidt et Kuntz, 1970) n-comb.

(Cucuhformes),

L. caprimulgi (Kazubski, 1958) n.comb. (Cap-

rimulgiformes)

L, casmetocephala (Yoyotte,

(Cuculiformes)

1972) n.comb.

L, ghesquieri (Ezzat et Tadros, 1958) n-comb-

(Caprimulgiformes).

(Diaz-Ungria,

L. golvani 1963) n.cemb.

(Piciformes)

L, lepidogrammi (Tubangui et Masilungan, 1937)

n.comb, (Cuculiformes.)

L.. noctuae (Spaul, 1927) n.comb., redescribed by

Chabaud (1951) (Strigifarmes),

L. nyctidromi (Caballero et Peregrina, 1938)

o.comb (Caprimulgiformes).

L, obtusa (Dujardin, 1845) n,comb. (= Eucamp-

tus obtusus D,, 1845, Rucampius Cheyr., 1833

preoccupied, Coleoptera) (Caprimulgiformes).

L, papillosa (Chabaud, Anderson et Brygoo,

1959) n.comb, (Cuculiformes),

L proctata (Lent et Freitas, 1948) n.camb.

(Strigiformes).

L. semenovi (Skrjabin, 1934) n,comb. (Caprimul-

giformes),

*A_ sudarikawi Sobolev, 1947 (in Sonin, 1966) is hard ta

classify: according to this author, the spicules are pointed and the

eggs small asin Aprocta spp., bui the caudal papillae are large as

in Lissonema spp. The ovejector cannot be analysed, However,

Sandground (1933) describing analogous material, emphasised

the absence of caudal papillae. IL seems to us thal (his species

belongs to Aprocta.

OVIPAROUS FILARIAL NEMATODES

L, spasskyi (Sonin, 1966) n.comb.

(Cuculiformes),

L. travassosi (Caballero, 1938) n,comb,

(Tragoniformes).

The genus Squamofilaria Schmerling, 1925, which

was reinstated by Anderson and Chabaud (1958) for

Aproctinae with a short well sclerotized buccal

capsule, is itself heterogenous. In these species, the

buccal capsule appears to be a secondary cuticular-

isation of the wall of the buccal cavity, as in

Lissonema sp., described below from Ninox

novaeseelandiae. It is poorly limited externally, as

compared with the true buccal capsule seen in

Dipetalonema spp., Litomosa spp., etc. which are

descended from forms with a buccal capsule formed

from rhabdions.

The species attributed to Squamofilaria fall into

the same two morphological groups found in

Aprocta and Lissonema, and can be allotted to one

or other of these genera as follows:

(a) Aprocta

A, vestibulata (Johnston et Mawson, 1950)

n.comb. (Passeriformes).

A. pillersi (Yorke et Maplestone, 1926)

n.comb, (Passeriformes).

(b) Lissonema

L. coraciae (Gmelin, 1790) n.comb. syn.

Squamofilaria coraciae (Gmelin, 1870), This

is the type species of Squamofilaria which

now falls as a synonym of Lissonema

(Coraciadiformes).

L. sicki (Strachan,

(Strigiformes).

L. nepalensis (Soota et Chaturverdi, 1970)

n.comb. (Coraciadiformes),

L. ungriai (Guerrero,

(Trogoniformes).

1957) n.comb.

1971)* n.comb.

It appears that the host ranges for Lissonema and

Aprocta are mutually exclusive.

Lissonema spp. are known from the Strigiformes,

the Cuculiformest, the Piciformes (Bucconidae), the

Trogoniformes, the Caprimulgiformes and the

Coraciadiformes. [t a5 of interest to note that

Berlioz, in his Classification of Birds, treated these

six groups consecutively (pages 930 to 974, in

Grassé, 1950).

Aprocta spp. are known from the Lariformes, the

Charadriiformes, the Ralliformes (Burrhinidae and

* The eggs measure only 45m x 25 ym, but their concave form

jncheates thar they are not embryonated,

* Among species from Cuculiformes, thoxe from Centropus ure

unusual in having a Jang oesophagus (5-7 mm), clearly divided,

Otididae), the Falconiformes (similarly grouped by

Berlioz, p, 884-929), and the vast group of

Passeriformes.

The Australian species which have been studied

herein are assigned to the two genera Aprocta and

Lissonema.

Aprocta boulengeri n-sp.

(Figs. 3, 4)

Material

Strepera graculina Shaw (Cracticidae) from

Warwick, Qd., from head and nasal cavities.

holotype @, allotype d (SAM V2098 and V2099)

and 1] entire 2 and 2 broken 6 (AHC 5839),

Morphology

The morphology is shown in Figures 3 and 4,

Unpaired portion of ovejector short and simple.

Dimensions

Holotype 2: length 50 pm; width 920 pm:

cephalic papillae form area 110 «xm (laterally) by

70 ~m = (dorsoventrally); nerve ring and vulva

respectively 175 gm and 940 jm from head;

oesophagus 1060 sm long, with muscular part 215

1m long; tail length 385 ym, egg 58 pm by 32 pum.

Allotype d; length 25.6 mm, diameter 600 pn,

cephalic papillae area 110 jm x 55 jam; buccal

capsule 20 44m long; nerve ring and excretory pore

205 pm and 305 pm from head; oesophagus 800 pm

long with muscular part 190 jzm; tail length 320 em;

right and left spicules 400 im and 440 pm long,

22 pm wide.

Other entire female: body length 37 mm, diameter

910 wm; cephalic papillae on area 88 jum x 57 jum:

buccal capsule 38 jum long; nerve ring, excretory

pore and vulva respectively 220 pm, 310 jm and 680

yum from head; oesophagus 965 ym long, with

muscular part 200 «m; unpaired part of ovejector

600 xm long; tail length 240 jum; larvae 190 and 205

um long, 11 pm wide.

Discussion

This species is easily differentiated by the

following characters: presence of a median subtermi-

nal papilla on the male tail, spicules 400-440 jum in

length, with tricuspid distal extremities, vulva

slightly in front of the oesophago-intestinal junction,

anus of the female not terminal. This is proposed as

a new species A. houlengeri-

Aprocta bakeri n.sp.

(Fig. 5)

Material

Corvus orrt Bonaparte (Carvidae) from Emu

Vale, near Warwick, Od,, from nasal cavity:

holotype & (SAM ¥Y21000).

270 REC. 5. AUST. MUS. 18(13): 265-284 September, 1981

Figure 3. Aprocta boulengeri n.sp., female, and male allotype. A, anterior end of female,

lateral view; B, C, & D, head of female, in apical, lateral and dorsal views; E, F, head of

male, ventral and lateral views; G, anterior end of male, lateral view; H, I, tail of female,

lateral and ventral views; J, K, L, tail of male, lateral and ventral views; M, left spicule.

A, G, H, I, J, L, to scale 200 zm; B, C, D, E, F, K, to scale 100 wm; M, to scale 150 um.

ANY

Figure 4. Aprocta boulengeri n.sp. (cont.) A, and B, tips of left and right spicules, ventral

views; C, larva; D, E, head of larvae, with hooks seen in profile and full face; F, tip of tail

of larva, A-E, to scale 10 um; F, free-hand.

OVIPAROUS FILARIAL NEMATODES 271

Description

Morphology is shown in Figure 5. Triangular

buccal cavity. Cloaca opening on conspicuous

protuberance.

Dimensions

Body length 29 mm, diameter 550 4m; buccal

capsule 40 xm long; nerve ring 140 jm from head;

left and right spicules respectively 390 um and

330 ym long; tail length 260 um.

Discussion

The bifid extremity of the testis in these specimens

resembles that of Aprocta laevicutis from Cyornis

banyumas whitei but is distinguished from this

species by the larger body size and by the spicule

length (390 and 330 j1m instead of 210-170 um). It is

also distinguished from all other Aprocta spp. by a

500 Um

200 Um

voluminous protuberance bearing the cloacal open-

ing. It is proposed as a new species, A. bakeri,

dedicated to our Canadian colleague, Michael

Baker.

Lissonema sp.

(Fig. 6)

Material

From Ninox novaeseelandiae Gmelin (Strigidae)

from Berrimah, N.T., 12 (AHC 6261).

Description

The morphology is shown in Fig, 6. Mouth and

buccal capsule narrower dorso-ventrally than later-

ally; buccal capsule stoutly built, of two parts—an-

teriorly, a thickening of the cephalic cuticle,

Figure 5, Aprocta bakeri n.sp,, holotype male, A, anterior end;

B, oesophageal region, lateral view; C, D, head in dorso-

ventral and lateral views; E, F, tail, in lateral and subventral

views; G, H, right and left spicules, ventral views; I, J, distal

extremities of same spicules; A, to scale 500 um; B, E, F, to

scale 200 xm; C, D, to scale 100 4m; G, H, to scale 150 wm;

I, J, to scale 10 pm. v, vulva.

272

posteriorly, a pre-oesophageal thickening, poorly

defined externally, and thicker laterally. Base of

buccal cavity formed by the three prominent and

sclerotised oesophageal lobes. The cuticular ‘scales’

are irregularly distributed over the body length, but

the lateral fields are smooth. Unpaired portion of

ovejector long and complex (Fig. 6 J).

Measurements

Body length 34 mm, diameter 590 ym; buccal

cavity 22 ym long; nerve ring, excretory pore and

deirids at 155 ym, 242 4m and 270 pm from head;

oesophagus 1200 wm long, with muscular part 230

wm long; vulva 700 pm from head; unpaired part of

ovejector 4660 um long; tail length 95 xm; cuticular

REC. S. AUST. MUS. 18(13): 265-284

September, 1981

scales on body 22 pm long, 13 pm wide,

commencing at 1150 xm from head and ending 550

um from posterior end of body. Eggs 66 pm x 38

um; larva 415 jm long, 13.5 xm in diameter.

Discussion

Among the species of this genus in which there is a

buccal capsule, L, sicki, L. nepalensis, L. ungriai and

L. coraciae—only the last named agrees with the

Australian specimens in having ‘scales’ on the body,

but the shape and arrangement of these was not

exactly described. The dimensions of the female are

very close, except the length, which is only half that

of the Australian specimens. In view of this, and of

the different geographical range, no_ specific

identification is made.

Figure 6. Lissonema sp,, female. A, oesophageal region, lateral

view; B, anterior end, median view; C, D, head, lateral and

median views; E, cuticular ornamentation in region of mid-

body, in submedian view; F, deirid, lateral view; G, cuticular

scale in lateral view; H, I, tail, ventral and lateral views, J,

ovejector, distal part; K, larva; L, head of larva; A, B, E, H,

I, to scale 200 um; C, D, F, G, K, to scale 100 ym; J, to

scale, 500 pm; L, to scale 50 pm.

OVIPAROUS FILARIAL NEMATODES 273

IT. Diplotriaenoidea—Dicheilonematinae

Genus Serratospiculum Skrjabin

The genus Serratospiculum contains (a) species

with small spicules (left spicules 600-700 jun, mght

spicule 300-350 ym), which may be considered as

gtouped about S. guttatum and (h) species with large

spicules (left spicule 1100-1300 jim, right spicule

350-500 jm) grouped about S. tendo.

(a) Species with small spicules

S, guttatum (Schneider, 1866): the most character-

istic feature of this species is the cuticular

ornamentation of the female body, in the form of

large bosses, well described by Schneider. The

species is recorded in Australia from Falco berigora

by Schneider, 1866, from F. longipennis and

F. peregrinus (Syn. F. melanogenys) by Johnston &

Mawson, 1941, and in the present paper.

Species grouped with S$. guttatum may have (i)

smooth or (ji) ornamented cuticle,

(i) With smooth cuticle:

S. chungi Hoeppli & Héu, 1929, from Falco sp,,

China.

S. congolensis Vuylsteke, 1957, from Butastur

rufipennis (Sund.) from equatorial Africa*, This

species is distinguished from S. chungi by the

structure of the right spicule; the oblique crests are

present on the distal third of the spicule, not on the

distal half,

S, seurati n.sp. (syn. Filaria attenuata Rud,, 1819

sensu Seurat, 1915) from variaus Falconidae, from

North Africa, is close to S. chiungi but can be

distinguished by the cephalic structure—the epau-

lettes concave with regard to the dorso-ventral axis,

and not rectilinear (Fig. 18 of Hoeppli & H&u, 1929),

relatively narrower—(30 zm wide by 100 wm long in

S. seurati, 50 wm wide by 120 jzm long in S. chungi).

(ii) With cuticular ornamentation:

S. turkestanicum Skrjabin, 1916 (type species of

the genus), trom F. tinnunculus L, from Central

Asia, is well distinguished by the type of cuticular

ornamentation, consisting of points curved back-

wards,

S, kwangsiensis H8u, 1963, from F. columbarius

insignis (Clark) from China, is close to S. guttatum in

the structure of the head and the presence of

rounded bosses on the cuticle of the female, but

these bosses continue to the level of the anus in this

* The apical view shown in Fig. 36 appears ro be in error: the

piece of the head shown is probably seen upside dawn, from (he

posterior aspect.

| Ruilliet & Henry (1916) have resolved clearly the question of

the nomenclature of this species.

Species but only to 1 rom from the posterior

extremity in 8. guttatum; also, the distal half of rhe

spicule is particularly delicate.

The tilanal worms described by Li (1933) from

F. columbarius (syn, Hypotriorchis aesolon), from

China, as S. turkestanicum seems from the character

of the female cuticle, to be S. kwangsiensis.

(6) Species with large spicules

This group contains three species, all without

cuticular ornamentation in the female!

S. tenda Nitasch, 18197, the species is parasitic in

F. peregrinus in Europe (Nitzsch in Rudolphi, 1819;

Dujardin, 1845; Yorke & Maplestone, 1926) and in

Australia (material described here and, as pointed

out by Sonin (1968), some parts of the material seen

by Johnston & Mawson, 1941, and noted as

S. guttatum). The species is also found in other

falcons, and in other parts of the Old World.

S, thoracis Tubangui, 1934, from Falco ernesti in

the Philippines; the dimensions of this species are

similar to those of \S. tendo.

S. lii Ezzat & Tadros, 1958, from F. peregrinus

callidus fram Belgian Congo. In this species the

measurements resemble those of the two preceding

species. The area rugosa and the detailed structure

of the spicules have not been described, and as

pointed out by Sonin (1968), it is possible that these

three species are synonymous.

(t appears that there are more species of

Serratospiculum than allowed by Sonin (1968). It

also appears that S. guttatum is found only in

Australia, and that references to its presence in

other parts of the world are erroneous.

The Australian material from falcons (Faleonidae)

permits the redescription of two species, Serraro-

spiculum guttatum (Schneider, 1866) and S. tendo

(Nitzsch, 1819). Comparison of the Australian

material with specimens in MNHN has led to the

identification of a new species, 8, seurati.

Serratospiculum guttatum (Schneider, 1866)

(Fig. 7)

Material

From Falco longipennis Swainson, from Fleurieu

Peninsula and Yorke Peninsula, South Australia.

93, 5 anterior and 2 posterior ends of ¢, 72,4

anterior and 5 posterior ends ? (AHC 6279); 2¢

and 22 (MNHN, 193 HD).

From Falco peregrinus Tunstall, from Mallala,

S.A, 3d, 1 posteriar end ¢, (AHC 6280).

Description

The morphology is shown in Fig. 7. The shape of

the head is the same in all specimens, , . broader on

lateral axis, epaulettes twice as long as wide (80 pm

274

REC. S. AUST. MUS, 18(13): 265-284

September, 1981

Figure 7. Serratospiculum guttatum A-G, female H-N male, A,

anterior end, lateral view; B, cuticular ornamentation in mid-

body, lateral yiew; C, the same, detail of one boss; D, distal

end of ovejector; E, F, tail, in lateral and ventral view; G,

head, apical view; H, anterior end, ventral view; I, head,

median view; J, K, left and right spicules; L, area rugosa,

ventral view; M, the same, in detail; N, tail, ventral view; A,

scale 500 pm; B, D, H, to seale 200 pm; C, E, F, to scale

150 pm; G, I, L, M, N, to scale 75 wm,

by 40 4m) and in apical view, convex with regard to

the dorso-ventral axis. Cuticular ornamentation

present only in the female. Area rugosa forms two

latero-ventral bands of low cuticular protuberances.

Dimensions

Three females: length 127, 150 and 140 .m, width

700, 790, and 600 um; nerve ring at 175, 180, and

240 rm from head; excretory pore at 205, ?, and 240

uum from head; deirids at 250, 220, and 510 .m from

head; muscular oesophagus 420 xm long, and with

glandular part 13 mm long (measured on one of the

females dissected); vulva at 740, 600, and 1050 4m

from head; Iength of unpaired part of ovejector 2150

pm (in female dissected); tail 90-110 um long;

embryonated eggs 52 by 30 um; cuticular ornamen-

tation starts at 3 mm from the head and ends at 1 mm

from the posterior end.

Two males: length 72 and 93 mm, width 480, 560

p.m; nerve ring and excretory pore 150, 230 44m from

head (measured on one male); deirids at 280 and 340

wm, 212 and 275 «ym from head; muscular

oesophagus 320, 365 wm long, length of glandular

OVIPAROUS FILARIAL NEMATODES 275

oesophagus not determined; tail length 80, 82 wm; This material has been compared with MNHN 510

length of left spicules 650, 670 wm, of right spicules JJ collected in France from the same host species;

350, 360 um; area rugosa lying from 5.4 mm to 450 ‘this consists of one anterior end and two posterior

pm from the posterior end (measured on one male). ends of d and the anterior and posterior ends of 12.

Serratospiculum tendo (Nitzsch, 1819) Description

(Fig. 8) The morphology of the two lots of material is

similar, and is shown in Figure 8. The cephalic

From Falco peregrinus Tunstall, from Port structure is constant: rounded head, epaulette 3-4

Augusta, S.A. 12, 1d (AHC 6273). times as long as wide (110 «4m by 30 ym), concave

Material

Figure 8. Serratospiculum tendo. A, anterior end of female, lateral

view; B, anterior end of male, median view; C, head of

female, apical view; D, E, head of male, median and lateral

views; F, male, area rugosa, ventral view, G, H, female, tail,

ventral and lateral views; I, J, tail of male, ventral and lateral

views; F and J, Australian material, other figures French

material. A, B, G, J, to scale 200 pm; C, D, E, F, I, to scale

100 wm; H, to scale 500 pm.

276

with regard to the dorso-ventral axis, and prolonged

by 4 small additional swellings. Area rugosa in form

of two ventro-lateral bands of small longitudinal

cuticular wrinkles. Spicules not drawn, as studied by

Bain & Vassiliades (1969).

Measurements of Australian specimens

Female: length of body 190 mm, width 825 pm;

nerve ring and deirids 300 wm and 210-220 wm from

head; length of oesophagus 19.7 mm, of muscular

part 600 wm; vulva 1650 wm from head; tail 100 ~m

long.

Male: length of body 148 mm, width 410 um;

nerve ring and deirids 200 4m and 305-310 um from

head; length of oesophagus 11 mm, of muscular part

600 um; length of left spicule 1120 um, of right 505

pm; tail 130 wm long; area rugosa extends from 4650

ym-600 wm from the caudal extremity.

REC. S. AUST. MUS. 18(13): 265-284

September, 1981

Measurements of specimens from France

In female, vulva 1 mm from the head, tail 140 pm

long. In male nerve ring and deirids 165 4m and 230

ym from head; length of left spicule 1150 4m, and

1120 wm, of right spicules 510 wm and 500 wm; tail

145 and 110 pm.

Serratospiculum seurati n.sp.

(Fig. 9)

Syn. Filaria attenuatum Rudolphi,

Seurat, 1915.

1819 sensu

Material

Specimens studied by Seurat, 1915, and deter-

mined as ‘Filaria attenuata Rud., 1819 (= F.

guttatum Schneider, 1866)’. This material, now in

the MNHN, Paris, was taken from Falco biarmicus

erlangeri Kleinschm at Biskra, North Africa; it

comprises 23 of which one is in two pieces, 1

Figure 9. Serratospiculum seurati n.sp. A, anterior end of female,

lateral view; B, head of holotype female, lateral view; C,

head of female, apical view; D, E, tail of holotype female,

ventral and lateral views; F, tail of male, subventral view; G,

H, left and right spicules, lateral view. A, E, to scale 200 um;

B, C, D, F, H, to scale 100 pm; G, to scale 50 pm.

OVIPAROUS FILARIAL NEMATODES 277

anterior end ¢, 2° and 2 posterior ends @, 1 entire

2 and 1 entire ¢ were chosen as holotype and

allotype respectively (MNHWN 110 EJ).

(2) specimens collected by Dollfus (MNHN 114

A) in 1914 from Falco sp., from Rabat, Morocco,

labelled S. guttatum (Schneider, 1866); consisting af

numerous pieces including 2 anterior and 4 posterior

ends of 2, 3 anterior and [ posterior ends ¢.

Morphology

The general morphology, well described by

Seurat, is complemented by Figure 8, The structure

of the head, in both lots of material, is similar, The

position of the vulva is somewhat variable (450-1000

wm from head). An area rugosa appears ta be

absent, The right spicule bears low crests, obliquely

placed.

Measurements

Holotype ¢: length of body 135 mm, width 770

jun; vulva 470 ym from head; tail length 40 pm.

Specimen not sufficiently clear for other measure-

ments,

Allotype ¢: length of bady 42mm, width 480 zm;

nerve ring and deirids 130 pm and 230 wm from

head; length of oesophagus 10 mm, of its muscular

part 271) pm; left spicule and right spicules

respectively 775 jum and 300 jum; tail length 110 wm.

From MNHN 114 A (2 pieces): length unknown;

width 500 wm; nerve ring 170 wm from head;

muscular oesophagus 310 ym) left and right spicules

respectively 650 zm and 350 jvm, tail length 110 pm,

Diplotriaenoidea:. Diplotriaenaidae

Diplotriaena spratti n.sp.

(Fig, 10)

Material

From Oreoica gutturalis Vigors & MHorsfield

(Muscicapidae) from the Petermann Ranges, N.T.

Holotype ¥ and allotype ¢ (SAM, V2101 and

V2102 respectively); 62, 9¢, and numerous pieces

of ends of ? and 1 posterior end 3 (AHC 6272); 19

and 23 paratypes (MNHN, Paris, 204 HD).

Description

Morphological features are shown in Figure 10.

Measurements

Holotype @: length of body 68 mm, width 700

jum; nerve ring 240 jzm from head; tridents 150 4m

and 172 «.m long; length of oesophagus 3720 wm, of

muscular part 370 pm; vulva 500 pm from head,

unpaired portion of ovejector 3300 «2m long; tail

length 110 p.m; eggs, non-fertilized, 32 pm by

18 wm,

Paratype 2s; body length 66-71 mm; in ? length

66 mm, width 500 jam; nerve ring and vulva 290 em

and 425 wm from head; trident 135 2m; oesophagus

4500 um; eggs 45 wm by 35 bm.

Allotype 3: body length 32 mm, width 400 jxm;

nerve ring 200 xm from head; muscular oesophagus

3850 jm long; tail SO pm; left and right spicules

respectively 4500 and 600 jcm long.

Paratype ds: length of body 32-34 mm; of

oesophagus 2600-3275 jzm; left spicule 4200-6300

pm, right spicule SLO-600 pm; in one male 32 mm

long: body width 455 wm, nerve ring 230 jm from

head; tridents 125 sm; length of oesophagus 3400

pm, of tail 55 ums left spicule 4200 pm long, with

calamus 650 jm, right spicule 550 jm.

Discussion

This species can be compared with the others of

the genus in which the left spicule is longer than 2000

pm. These are: D. bharioensis (Parona, 1889)*

redescribed by Anderson (1959) trom Sturnidae and

Turdidae in Asia, has the trident not pointed bur

concave at the apex, a much longer oesophagus (8-

12 mm instead of 2.6-6.3 mm), left spicule not

exceeding 2600 jzm, and the right spicule spiralled

about a strongly curved axis, whereas in the

Australian specimens the axis is almost straight.

In D. ecaudata (Oerley, 1882), from Sturnidae in

Africa the trident is concave at the tip, the left

spicule is not longer than 2800 jm, and the axis of

the right spicule has three strong bends.

D. tricuspis (Fedtschenko, 1874) sensu Singh,

1949, is, as pointed out by Anderson (1959), a

mistaken determination. It is clear from Singh's

paper that though the female is a Diplotriaena sp.,

the male (Fig. 27) is a Hamatospiculum sp. The

Australian species is proposed as D. spratti n.sp.,

named after our colleague David Spratt. The species

is characterised by the pointed apex of the trident,

the long left spicule (over 4000 wm) and the right

spicule spiralled around an almost straight axis.

* B. buckleyi Fotedar & Kaw, 1965, and D, mirzapurensis

Seota & Chaturvedi, 1967, both parasites of Acridotheres tristis,

have no morphological difference from [. bhamoaensis, and are

regarded as synonyms of this species.

278 REC. S. AUST. MUS. 18(13): 265-284

September, 1981

Figure 10. Diplotriaena spratti n.sp. A, anterior end of holotype

female, lateral view; B, head of holotype female, median

view; C, junction of oesophagus and intestine, paratype

female; D, E, tail of paratype female, lateral and ventral

views; F, trident of holotype female; G, to Q, male: G,

anterior end, ventral view; H, head, apical view; I, J, tridents

of two specimens; K, posterior end, lateral view; L, M, the

two extremities of the left spicule; N, right spicule and part of

the left spicule; O, distal end of right spicule, ventral view: P,

Q, tail in lateral and ventral views. A, D, to scale 500 p.m; B,

to scale 150 pm; C, E, G, L, N, to scale 200 pm; F, I, J, M,

O, P, Q, to scale 100 pm.

Diplotriaena beveridgei n.sp.

(Fig. 11)

Material

From Corvus orru Bonaparte (Corvidae) from

Emu Vale, near Warwick Qd: Holotype ° (SAM V

2103) and posterior end (10 mm) of allotype 3

(SAM V 2104); 29, 2 anterior and 3 posterior ends

2 (AHC 6276).

Description

Morphological features are shown in Figure 11.

Measurements

Holotype ?: body length 102 mm, width 1400 zm,

nerve ring 450 4m from head; trident 285 wm long;

oesophagus 9500 ym long, its muscular part 575 um;

vulva 630 zm from head; eggs 50 x 35 wm; tail 200

um long.

Allotype d: the piece is 1.11 mm wide; tail 10 um

long; left spicule 1355 zm long (calamus 250 p.m);

right 1075 «wm.

Paratypes: one anterior end of 2, 1400 ym wide,

with nerve ring 390 ym from head, tridents 250 4m

OVIPAROUS FILARIAL NEMATODES

200 Foal

279

150 jm

100 [Lm

Figure 11. Diplotriaena beveridgei n.sp. A-H, female; I-L, male,

A, anterior end, lateral view; B, oesophago-intestinal

junction; C, head, apical view; D, head, lateral view; E, F,

trident and its apex; G, H, tail, ventral and lateral views; I,

posterior end, ventral view; J, K, right and left spicules; L,

tail, ventral view. A, B, G, H, to scale 500 jm; C, to scale

150 um; D, E, J, K, to scale 200 jum; F, L, to scale 100 um; I,

to scale 1000 ym,

and 270 wm; oesophagus 8950 jm with muscular

part 580 um; vulva 650 um from head, unpaired

ovejector 2900 xm. One posterior end with tail 200

wm. Length of entire female 81 mm.

Discussion

The specimens differ from the other two

Diplotriaena spp. from Australian Corvidae as

follows:

In D. alpha Johnston & Mawson, 1940, the trident

is smaller (120 ym instead of 250-285 um) and

oesophagus twice as long.

In D. beta Johnston & Mawson, 1940, considered

by Anderson (1959) as a synonym of D. flabellata

(Linstow, 1888) the oesophagus is very long, but the

tridents shorter (190-200 1m), more massive, and

with rounded apex, and the right spicule is a

different shape.

280

The right spicule is bent in an ‘L' similar to that of

D. delirae Pinto & Noronha, 1970, from Tyrannidae

in Brazil, however our material is distinguished from

this species by the Janger oesophagus, and the longer

right spicule (1075 jm instead of 610 pm),

This material is therefore proposed as D, bev-

eridgei n.sp., named after the callector, Dr Jan

Beveridge. It is characterised by the large tridents,

the long oesophagus (about 9 mm) and the ‘L’ shape

of the right spicule.

Diplotriaena smithi o.sp.

(Fig. 12)

Material

5 collections from Acanthogenys rufogularis

Gould. From The Bunkers, S. Aust.: 4 birds, as

tollows;

1. holotype 2 (SAM V2105), allotype ¢ (SAM

V2106)

22, 1d, pieces of 2 and & (AHC 6278)

12, 1d, posterior end 1d (MNHAN, Paris 208

HD)

2.12, 1¢ pieces of 2 (AHC 6270)

3. 12%, pieces of and d (AHC 6268)

4. 42, 3d pieces of 2 [AHO 6267)

From Wubin, W. Aust.: 62, 3d (AHC 6264); 3°,

2d (MNHN, 208 HD).

Description

Morphological details are shown in Figure 12.

Dimensions

Holotype 2: body length 50 mm, width 680 ym;

trident length 220 4m; nerve ring and vulva 350 jtm

and 660 4m from head; length of oesophagus 4000

pom, of tail 45m, of unpaired part of ovejector 2100

Lm,

Allotype @: length of body 22 mm, width $30 jum;

trident length 190 jm; nerve ring and excretory pore

310 pm and 350 ym from head; length of oesopha-

gus 2850 pum, of tail 68 xm, of left spicule 800 pm, of

right spicule 650 jm.

Paratype 39s: Length of body 45, 22, 39 mm;

width 650, 610, 520 ,cm; tridents 205, 195, 190 wm;

nerve ring 300, 290, 330 jum from head; length of

oesophagus 2400, 2100 zm, damaged in 3rd; vulva

650, 730, 500 xm from head; tail length 40, 55, 55

p.m.

Paratype d: body length 22 mm, width 520 im,

trident length 165 zm; nerve ring 280 .m from head;

oesophagus 3400 xm long (muscular part 340 pm);

length of tail 80 jxm; of left spicule 785 jum, of right

spicule 500 jum.

REC, S, AUST. MUS. 18(13): 265-284

September, 1981

Variations observed among all the specimens:

tridents 165-190 um in males, 190-220 jm in

females; vulva 480-730 jum from head; oesophagus

2000-3600 «xm long in 9; left spicule 730-850 pm,

right spicule 500-680 zm. One male from AHC 6278

is teratological, with left and right spicules

respectively 260 ym and 310 pm.

Discussion

D. zeta Johnston & Mawsan, 1940, redescribed by

Anderson (1959) was also taken from Acanthagenys

rufogularis; the only temale known is of similar

dimensions to the present specimens, except that the

trident is smaller (150 jtm instead of 190-205 pm),

Moreover, there are two grooves on the head of

tridents in D. zeta not seen on any of the new

specimens, These twa differences, and the absence

of a male of D. zeta, prevent the identification of the

hew material as D_ zeta.

Five other species must be considered:

Two species have similar dimensions and spicule

shape: D. clelandi (Johnston, 1912) from Crac-

ticidae in Australia (male only known) and D,

kennedyi Grewal, 1965, from Pycnonotidae from

India. These two species are of doubtful status, as

their morphology is imperfectly Known, and it is

impossible to compare our specimens with them.

The three other species in which the trident and

spicules are similar to our specimens are disting-

uished as follows: in D, pungens (Schneider, 1866)

from Turdidae in Asia, the oesophagus is very long

(14500 jum in the 2) and three pairs of papillae lie

distinctly in front of the cloaca. In D. flabellata

(Linstow, 1888), from Paradiseidae in the Aru

Islands, the trident is more stoutly built, the body is

particularly wide, the glandular oesophagus very

long (6500-7500 jm in the female)—the left spicule

strongly curved and the male tail rounded,

In D, urocissoides Anderson, 1959, from Corvidae

in India, the head structure is characteristic (‘buccal

eavity surrounded by a clearly defined cuticular

thickening often forming a plug,’ Anderson, 1959)

and the number of papillae is small.

It seems that the material described here is new,

and the name D. smithi is proposed for it, in honour

of our colleague 1. Humphrey Smith. It ts

characterised by the large trident with blunt tip and

without grooves, the oesophagus not very thick and

not particularly long or short (2000-4000 j.m), the

spicules of medium length (730-850 pm and 500-600

j.1m) the tail of male rectangular in ventral view, with

precloacal papillae few in number and situated close

to the cloaca.

OVIPAROUS FILARIAL NEMATODES 281

Figure 12, Diplotriaena smithi n.sp. A, anterior end of female

holotype, lateral view; B, head of female paratype, apical

view; C, D, same female, apex of tridents; E, F, tail of same

female in lateral and ventral views; G, anterior end of male

allotype, lateral view; H, same male, oesophago-intestinal

junction; I, head of same male, lateral view; J, trident of

male paratype; K, tail of male allotype, ventral view; M, N,

male allotype, right and left spicules, ventral views; O, P, two

other views of same right spicule; A, E, G, M, N, O, P, to

scale 200 4m; B, H, F, J, K, L, to scale 100 wm; C, D, to

scale 50 ym; I, to scale 150 pm.

Diplotriaena delta Johnston & Mawson, 1940 tridents 115-120 4m and 100 tm; nerve ring 180,

(Fig. 13 A-O) 190 44m from head; oesophagus length 1850,

2110 wm, its muscular part 330, 160 ~m; vulva 525,

400 4m from head; tail length 70, 32 um; eggs

From Amytornis goyderi (Gould) (Maluridae), 46 x 29 um. The extremes of the oesophagus are

from the Simpson Desert, S. Aust.: 109, 2¢ (AHC 1750 um and 2400 um, in females measuring

Material

6274); 22 and 1d (MNHN 207 HD). respectively 62 mm and 70 mm.

Description

Additional morphological details are given in 29? body length 30, 28 mm, width 400, 380 1m;

Figure 13 A-O. tridents 110, 120 um; nerve ring 170, 160 um from

; ; head; left spicule 950 zm long, (calamus 240 ym),

Dimensions and 1050 ym; right spicule 720, 630 j1m long; tail 50,

22: body length 70, 72 mm, width 460, 400 um; 60 um.

282

Discussion

These specimens are identified as D. delta,

described from Malurus lamberti (Maluridae) in

Australia. This species, placed by Anderson (1959)

as a synonym of D. tridens (Molin, 1858), was

reinstated by Ogden (1965), who had specimens

from M. lamberti from various parts of Australia.

D. delta is recognisable by the simultaneous

presence of the following characters (taken from the

REC. 8. AUST. MUS. 18(13): 265-284

September, 1981

new material and from Ogden): large body size (60-

127 mm in female); tridents small (78-122 4m) with

three small digitations at the tip; oesophagus of

average size (1750-3900 um in female, 1560-

2600 ym in male), with glandular part thin or thick;

left spicule 950-1140 ym, right spicule 620-730 jm,

with triple twist; tail of male with two lateral

swellings.

Figure 13. A-O, Diplotriaena delta, A-E, female: A, anterior end,

subyentral view; B, head, en face; C, D, trident and its apex;

E, tail, ventral view; F-O, male; F, anterior end; G, H,

trident and its apex; I, J, two views of right spicule; K, L,

tight and left spicules of another male, lateral view; M, N,

tips of right and left spicules, ventral views; O, tail, ventral

view.

P-R, D. falconis. P, tail of female, ventral view; Q, profile of

cuticle at mid-body; R, tail of male, ventral view.

A, F, Q, to scale 500 pm; 1, J, K, L, to scale 200 jxm; P, to

scale 125 pm; B, E, G, M,

scale 50 zm.

N, O, R, to scale 100 pm; H, to

OVIPAROUS FILARIAL NEMATODES

Diplotriaena falconis (Connal, 1912)

(Fig, 13 P-R)

Material

From Falco berigora Vigors & Horsfield, from

Yorke Peninsula, S. Aust.: 32, 1d (AHC 6269).

Description

Additional morphological details are given in

Figure 13 P-R.

Measurements

29: body length 70, 72 mm; width 590, 660 ym;

trident length 135, 135 jum; nerve ring 230, 230 ym

from head; vulva 430, 660 4m from head;

oesophagus in second female 2550 jz£m long, with

muscwlar part 400 ym; tail 22, 28 zm; eggs

65 x 30 pm.

3; body length 45 mm, width 550 j.m, trident

length 135 wm, oesophagus length 2700 um, with

muscular part 450 yum; tail 32 xm long; left spicule

broken, 1025 4m long; right 1100 4m long (850 jum

if measured in a straight line).

Discussion

The measurements, general morphology, and the

characteristic anatomy of the left spicule agree with

those of D. falconis, redescribed by Anderson

(1959), from Falconidae in Africa, In the Australian

specimens, there are ten pairs of papillae in the

male, the tail of the female bears papillae, and the

cuticle of the female is marked with regular and

distinct undulations.

ACKNOWLEDGEMENTS

We are grateful to all those who collected the

birds hosts or the parasites which enabled this work

toa be undertaken .. . these are Dr Ian Beveridge,

Mrs Joan Paton, Dr Dom Serventy, Mr Shane

Parker, and officials of the South Australian

Museum and the Northern Territory Administration

(Animal Industry and Agriculture Branch).

APPENDIX

Australian Species Described in this Paper, Arranged under

their Hosts

Corvidae

Corvus arry Bonaparte

Apmeta bakeri n.sp.

Diplotriaena beveridgei n.spy.

Cracticidae

Strepera graculina (Shaw)

Aprocia boulengert sp

Muscicapidae

Petroica multicolor (Gmelin)

Pseudaprocta copemani n.sp.

Pachycephala pectoralis (Latham)

Pseudaprocta copemant n.sp.

Oreoica gutturalis (Vigors and Horsfield)

Diplotriaena gpraii n.sp..

Maluridae

Amytornis goyderi (Gould)

Diplotriaena delia Johnston and Mawson

283

Meliphagidae:

Acanthogenys rufogularis Gould

Diplorriaena smithi n-sp.

Falconidae

Falca berigora (Vigors and Horsfield)

Diplotriaena falconis (Connal)

Serratospiculum guoatum (Schneider)

Faleo peregrinus Tunstall

S. tendo (Nitzsch)

Falco longipennis Swainson

Serratospiculum guiraium (Schneider)

Strigidae

Ninox navaeseelandiae (Gmelin)

Lissonema sp,

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RECORDS OF THE

SOUTH AUSTRALIAN

MUSEUM

VOLUME 18 NUMBERS 14—16 JUNE 1982

No. 14 OBSERVATIONS ON CHYZERIA CANESTRINI AND SOME RELATED

GENERA (ACARINA: TROMBIDIOIDEA) WITH REMARKS ON THE

CLASSIFICATION OF THE SUPERFAMILY AND DESCRIPTION OF A

PYGMEPHORID MITE PHORETIC ON CHYZERIA

by R. V. SOUTHCOTT

No. 15 SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS.

3. ARTHRONOTINA (CRYPTOSTIGMATA)

by DAVID C. LEE

No. 16 POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF

AUSTRALIA

by STEPHEN PRUDHOE

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

OBSERVATIONS ON CHYZERIA CANESTRINI AND SOME RELATED

GENERA (ACARINA: TROMBIDIOIDEA) WITH REMARKS ON THE

CLASSIFICATION OF THE SUPERFAMILY AND DESCRIPTION OF A

PYGMEPHORID MITE PHORETIC ON CHYZERIA

BY R. V. SOUTHCOTT

Summary

The taxonomy of the Australian larvae of the genus Chyzeria Canestrini, 1897, is revised. Two new

species, Chyzeria flindersi sp. nov., from Tasmania, and C. derricki sp. nov., from Queensland, are

described. In addition, the larva of C. hirsti Womersley, 1934 is redescribed from material reared in

South Australia. The species Grossia onychia Womersley, 1954 is redescribed, from the type

specimen from Victoria, and placed in Chyzeria as C. onychia (Wom.), comb. Nov.

OBSERVATIONS ON CHYZERIA CANESTRINI AND SOME RELATED GENERA (ACARINA: TROM-

BIDIOIDEA) WITH REMARKS ON THE CLASSIFICATION OF THE SUPERFAMILY AND DESCRIP-

TION OF A PYGMEPHORID MITE PHORETIC ON CHYZERIA

R, V. SOUTHCOTT

Honorary Associate, South Australian Museum, Adelaide, South Australia 5000

(Manuscript accepted 27 January 1981)

ABSTRACT

SOUTHCOTT, R. V, 1982. Observations on CAyzeria Canestrini

and some related genera (Acarina: Trombidioides) with

remarks on the classification of the superfamily and description

of a pygemphorid mite phoretic on Chyzeria, Rec, S. Aus.

Mfus. 18 (14); 285-326.

The taxonomy of the Australian larvae of the genus

Chyzeria Canestrini, 1897, is revised. Two new spe-

cies, Chyzeria flindersi sp. nov., from Tasmania, and

C. derricki sp. nov., from Queensland, are deseribed.

In addition, the larva of ©. Airsti Womersley, 1934

is redescribed from material reared in South Aus-

iralia. The species Grossia onychia Womersley, 1954

is redescribed, from the type specimen {rom Victoria,

and placed in Chyzeria as C, anychia (Wom.), comb,

nov,

Details of the rearing of larvae of C, Airsti from

adult specimens fre given, The species appears to be

viviparous, and the larvae are carried phoretically on

the mother’s back. Such behaviour allows some allot-

ment of function to the digitations upon the dorsum

of the adult, and the specialized hair characteristics

of the intervening area. Parasitization of larval Chy-

zeria upona host is recorded only in lwo instances,

these being the record of the type series of C. derrick

sp. nov, on an unidentified tettigoniid grasshopper,

and the earlier record of C. onychia (Womersley)

purausilizing an adult identified by Womersley as Chy-

zeria australiensiy First,

A description of the adult of Chyzeria hirsti Wom-

ersley (elevated from a subspecies to a species here)

lrom the holotype is given, and some comparison

made with the Chyzeria australiensis Hirst, 1928,

type series from Western Australia.

The classification of the larvae of Chyzeria, Gros-

sia, Audyana and others is examined. Grossia is

considered a synonym of CApzeria. These genera are

removed from the subfamilies Apoloniinae and Las-

seniinue, where they have been placed by some

authors, to the family Trombellidae Feider 1955 (ear-

lier subfamily Trombellinae Thor, 1935) of the Trom-

bidigidea. Larvae of an Australian species of Mon-

erslevyia Radford are recorded as parasitic upon the

cricket Teleogryllus commodus (Walker), in northern

New South Wales, and nymphs reared experimentally

are placed in Trombella udelajdeae Womersley,

June, 1982

1939, thus allowing the synonymizing of Womers/evia

With Trambella Berlese, 1887. This defines a further

larval form of the Trombellidae,

The relationships of some of the genera, Carmilivs

and subfamilies of the Trombidioidea are. briefly

examined,

A new species of pygmephorid mite (Tarsonemius}

is recorded as phoretic upon the dorsum of an adult

Chyzeria hirsti Wom., and described us Bakerdania

workandae sp, nov,

INTRODLCTION

In this paper the author records a number of obser-

vations on the morphology and biology of the genus

Chyzeria Canestrini, (897, with particular reference

to the larvae. This genus belongs to the family Trom

bidiidue (s. 1.) of the superfamily Trombidiowen

Banks, 1894.* 4 group of prostigmatic mites which

exhibil extreme heteromorphy between the adults

(and mobile nymphs) and the larvae, This hetero

morphy is in fact so great that in the superfarnily the

adults and larvae cannot be correlated with each

other without experimental rearing. As both lurvue

and adults have 1 be described under separate names

until such correlations ure established, classificatory

systems for the post-larval forms may not, and in fact

ollen do not, correspond to thit for the larvae. with

inevitable confusion, and, in a number of cuses.

nomenclatorial difficulties, The present paper attempts

to place the classification of the Jarvae related 1

Chyzerta on u firm basis, with detailed deseripuuns,

and biological data where available. Two new species

of larval Chyzeria will be deseribed, and additionally.

two previously recorded species will be redeseribed.

one being C. eavehia (Womersley, 1954), mn. comb..

previously Grossia anvchia Wom,

“Although this superfamily name was first proposed as such by

Banks in 1894, under Article 36 of the International Code of

Zoological Nomenclature, 1961, all categories in the family-group

(which includes the superfamily category) are of co-ordinate stalus,

As jt is generally accepted that the family name Trombidiidav

was first validly proposed by Leach in 1815 as *Trombidides”,

the superfamily name Trombidioidea is available from the year

1815 (See Oudemans, 1937; Thor and Willmann, 1947; Southcot,

1957d).

The trombidiid mite genus Chyzeria was founded

by Canestrini in 1897, with the adult C. ornata

Canestrini, 1897, as the type species, collected at

Friedrich Wilhelmshafen (later Madang) in New

Guinea, Since that time several further species have

been described from Australia (including Tasmania),

New Zealand, Africa and South America (Hirst

1924, 1928a, b, 1929; Womersley 1934, 1942; Law-

rence 1944: Thor and Willmann 1947, André and

Lelievre-Farjon 1960).

The three #enera Trombella Berlese, 1887, Chy-

ceria and Parachyceria Hirst, 1926, based on adults,

were placed in the subfamily Trombellinae Thor,

1935 by Thor and Willmatin (1947) in their mono-

graphic review of the taxonomy of the fumily Trom-

bidiidae; this subfamily was separiuted from all others

of the Trombidiidae on the ground of its lack of a

orista metopica. Within the subfamily, adults of Chy-

ceria are distinct in possessing & serjes of digitatians

arranged around the rim of the dorsum of the idio-

soma. These digitations ure covered by lwo lypes of

normal body setae (scobalae): one suff and swordlike,

the other flexible, longer and ciliated. In between

these digitations the central dorsal area of the idio-

soma js covered with another type of modilied seob-

alae. In at least two Australian species, CL ausztrel-

jensis Hirst and C. Airsti Womersley, these last

named setae are modified to long, unciliated, highly

convoluted hairs,

The genus Parachyzeria, known from Aftiea and

Asia, is similar to Capzeria, with a central area of

modified setae dorsally upon the idiosoma (Lese

being long and straight), but lacks the peripheral

processes.

So suggestion has been made as lo the possible

funetion of the peculiar structural adaptations of

these mites, although in the case of Chyreria, the

opportunity to do so, to ut Jeast some degree, existed

over 40 years ago,

In }938 | made a series of attempts to correlate

Jarval and postlarval forms of trombidiid mites by

experimental rearings, using South Australian

material. Among genera reared, and made available

to Herbert Womersley, then Entomologist. South

Australian Museum, was Chvzeria. These lurvae were

deseribed by Womersley in 1939, and allotted to

Chyzerla australiensis Hirst, 1928, despite the fact

that all adult Chpzeria from the “Adelaide District”

he had examined earlier (Wormersley, 1934) were

allotted by him to C. australiense (sic) var. hirsti

Wom. 1934,

REC, S. AUST. MUS., 18 (14): 285.320

dune, 1982

] had seen these reared larvae in captivity on the

udult’s back, At times they ventured away to the soil

below and then returned to the protection presumably

allorded by the dorsum of the idiosoma of the parent.

These observations were recounted to Womersley and

offered for incorporation in his article, but although

he was prepared to describe the larvae with their

unusual morphological structure, he chose not to

include the observations, possibly doubting their

accuracy or relevance, or more likcly being little

interested in details of mite behaviour.

Womersley recognized that these larval mites, with

their bizarre complement of dorsal and ventral setae,

as well as unusual palp and (pedi)iarsal claw structure.

were very different trom any previously described

trombidiid mites, They will be placed here as Chyzerra

hirsti Womersley, 1934,

After returning to South Australia front military

duties | made a number of attempts to rear larvae of

Chyzeria, ind on three further oceasions | obtained

lurvae from adults, These experiments tre recorded

below, and confirm the larva-adull correlation origi-

nally seen in 1938.

The larval Chyrerta hirsti has each of its three

pairs of legs 7-segmented (including the coxa); i.e. its

leg Segmental formula is 7, 7, 7. Its leg coxae have

iwo setae upon coxa J, and one each upon coxae Ii

und Ul, i.e. its leg coxal formula is 2, 1, |.

In late 1945 | captured a larval trombidiid mite at

Lake St. Clair, Tasmania, walking up the trunk of a

cucalypi. This mite was very similar to the larval

Chyzeria hirsti, and was clearly referable to the same

genus, It had a trifid palpal tibial claw, 7-segmented

legs, dorsal idiosomal setae placed upon expanded

plates, ley coxal formula 2, 1, 1, and a number of

other resemblances, The most obvious difference lay

im the lack of modification of the scutalac. particu-

larly the AL scutalae. This specimen is described in

the present paper as the holotype of CAyzeria flin-

dersi sp.nov.

In 1954 Womersley (p. 1) 1) deseribed as a new

genus and species Grossia onyehia Wom, 1954, a

larval trambidiid mite which had been collected

altached to an adult identified as Chyzeria ausiral-

fensis Hirst in the Otway Ranges, Victoria, in 1951.

In 1947 Wharton had revised the classification of

the subfamilies of the chigger mites, family Trom-

biculidae Ewing 1929*, separating them on the char-

acters shown in Table I.

*Formally proposed ws such by Ewing (1944).

OBSERVATIONS ON CHYZERI 4A GANESTRINI AND SOME RELATED GENERA o87

TABLE |,

Ley segmental Coxal sétal

CLASSIFICATION OF SUBFAMILIES OF TROMBICLILIDAE, AS PROPOSED BY WIARTON (1947).

Median scowl Anterior scutal

formula formula Tratheae setae projection Sternal setae

Subfamily Leeuwenhuckiinie 6,6, 6 2, Present Frequently (Frequently) No more

Womersley (1944) (= 2 preset (hun 2

Leeuwenhockidae,

Womersley, 1944"

Subfamily Walehiinae 7, 6,8 ier Tiles Absent (1) LAbsentt At least 4

Rwing, 1946

Subfamily Apoloniinue 77,7 1. —. — Present (}-2) Present (4)

Wharton, 1947

Subfamily Trombicu)inae 7,47 Absent | Absent AL leust 4

Ewing, 929

*Formally proposed as such by Womersley (1945),

Womersley’s Grossia onychia has a leg segmental

formula 7, 7, 7, and a leg coxal formula of 2, 1, I.

Ite placed it in the subfamily Apolominac, whose

definition had been enlarged by Wharton and Fuller

in 1952, to accommodate the genus Sauracere/la

Lawrence 1949, which has a leg segmental formula

of 7. 7, 7, but With a coxal setal formula of 1, 12).

! and lacked tracheac, and also has expanded sensil-

lary setae; the earlier criterion of “posterior lateral

scutal setae not on the seutuim” was omitted.

In the sane paper (1954) Womersley revised the

taxonomy of a number of larval mites he placed in

the family Leeuwenhockiidae, subfamily Apoloniinge,

In addition to the larval genera that he deelared had

been placed in the Apoloniinue by previous

authors=-Apolonia Torres and Braga, 1938 and

Womerslevia “Wharton” (actually the genus placed

there by Wharton in 1947 was MWomersia Wharton,

1947) und Savracarella Lawrence, 1949—he included

formally six further genera, these being Cockingsia

Womersley, 1954, Aud vana Womersley, 1954, Muck-

errasiella Womersley. 1954, Neotrombidium Leon-

ard: 1901 (with Monunguis Wharton 1938 as a syn-

onym), Grassia nov., Nothotrombicula Dumbleton,

1947 and Womerslevia Radford, 1946. In his paper

Womersley stated that ‘This is a heterogeneous

assemblage of genera’, which subsequent studies have

amply confirmed. The genus Mackerrasiella has been

shown by Vereammen-Grandjean (1972) to be one of

the Hydrachnellac, genus Midr\phantes Koch, 1841

and falls in synonymy. The genus 4vdyana was

placed alternatively, in the same paper by Womer-

sley, in the subfamily Trombellinae Thor 1935, since

larvae had been reared to nymphs resembling Tron

hella, in Malaya. Additionally it may be commented

that of the genera listed by Womersley, Grovsia,

Nothatrombicula and Womerslevia have leg sezmen-

tal formula 7, 7. 7, while Cockingsia, Andyana and

Mackerrasiella have leg segmental formula 7. 6, 6.

Womersley (/uc cit.) included alsa in his key to the

genera the genera Neotrombidium Leonardi, 1901

and Monunguis Wharton, 1938, stating in error that

they had leg segmental formula 7, 7, 7; actually in

Neotrombidium it is 7. 6 6. and in Monunguts

6. 6, 6, which Womersley eventually recognized (pub-

lished posthumously (Womersley, 1963)),

The resemblance between the larvae of Chyzeria

as deseribed by Womersley in 1939. Grassi onyvelia

Wom. 1954. and Nothorrambicula deinacridae

Dumbletwon, 1947, from New Zealand is striking, but

in this (1954a) paper Womersley made no reference

lo it, Evidently. by 1953-1954, he had forgotten his

1939 description. Another possible fictor may have

been the maccuracies in his account of the Chyzeria

larva, of Which the claws of the tarsi of the legs were

stilted to have “claws three, the lateral ones clavate

“while in Grassfa cach leg tarsus was stated to

Possess “paired spathulate sctae on each larsus at the

base of the single claw’, these lateral claws also being

relerred to as ‘a pair of shorter [than the central

tarsal claw] apically spathulute tenent setae’. Both

Grossia onvehia and Nothotrombicula deinacridue

have been redescribed by Vereammen-Grandjecan

(1972), as possessing spathulate neolateral pedotarsal

claws (see the anatomical terminology in Southeart

196]a).

In 1968 Vercammen-Grandjean and Kolebinova

revised the subfamily Apoloniinac, restricting iv to

seven species, allotted among six genera: these were

arranged in two tribes, the Apoloniinit and the Saur-

acarellini.. They rejected all of the genera included in

the Apoloniinae by Womersley (19544), with the

exception of the type genus Apolenia,

As slated above, Vercammen-Grandjean (1972) re-

examined the status of the mites that Womersley had

placed in his greatly expanded Apoloniinae, He

placed both Grossia and Nothotrombicula in the

subfamily Lasseninae (sic, for Lasseniinae) Newell,

1957, of the family Johnstonianidae Thor, 1935* (for-

merly Johnstonianinae Thor, 1935). Vercammen-

Grandjean made no reference in his paper to the

larval Chyzeria that Womersley had described jn

1939,

* Formally elevated as such by Newell (1957).

IRA

The subfamily Johnsionianinae had been founded

by Thor in 1935 for those (adult) Trombidiidae, in

which a distinet crista metopica is present, wilh two

pairs of dorsal sensilla, together with some minor and

less specific features, This was a distinct group, cle-

vated to family status as Johnstonianidae by Newell

(1957), in a study of a series of trombidioid mites

from Hawaii and North America, on the ground that

these mites differed so profoundly from other terres-

trial Parasitengona that they could not logically be

retained in the old family Trombidiidae. The differ-

ences were, in fact, greater than used by Ewing to

separate off the family Trombiculidae from the

remaining Trombidiidae. In that study Newell

included seven genera in the family, which he divided

into the subfamilies Johnstonianinac Thor and Las-

seniinac Newell, 1957, The latter subfamily was

erected for the general Lassenia Newell, !957, Poly-

discia Methlagl, 1928 and Crossothrombium Wom-

ersley, 1939. Laxssenia was founded on L, lasseni

Newell, 1957 as type genus, an adult mite in which

the principal character was the great reduction of the

anterior sensilla when compared with the posterior

sensilla. As well as two species of adults, Newell (oc.

cit.) described two species of larvae, which he placed

in the genus on morphological characters und field

association, but without the confirmation of experi-

mental rearing. In these larvae the dorsal scutum

carries two pairs of normal setae (scutalae) as well as

the two pairs of sensilla, Newell made no reference

REC, S. AUST, MUS,, 18 (14). 285-326

June, 1982

to any genera of Trombiculidae (Trombiculinae) nor

the Leeuwenhoekiidae (Leeuwenhoekiinae), nor to the

genus Chyzeria or other Trombellinae.

In 1973 Vercammen-Grandjean submitted a clas-

sification of the Trombidioidea, divided into eight

families and a number of subfamilies and tribes.

Among the families considered were listed Trombel-

lidae, Johnstonianidae, Neotrombidiidae noy., and

Leeuwenhoektidac. In that revision he placed Chy-

zeria in the Trombellidae and Grossia in the John-

stonianidae. Womersleyia appears to have been omit-

ted.

As it is not possible in the present paper to deal

with the overall classification of the Trombidioidea,

it is proposed to submit a detailed examination of the

status of the genera Chyzeria and Grossia, and to

make some incidental reference to some other taxa,

tn an effort to clarify the relationships of several

genera of the Trombidioidea.

Table 2 lists a number of genera of larval Trom-

bidioidea which haye been referred to cither the

Trombellinae, Leeuwenhoekiinae (Leeuwenhocki-

idae), Lasseniinae or Apolontinae by various authors,

as well as several genera of other families of the

superfamily Trombidioidea which are included for

purposes of comparison.

TABLE 2. PossiBLy SIGNIFICANT GENERIC CHARACTERS FOR THE LARVAE* IN A NUMBER OF ‘THE SUPERFAMILY

TROMBIDIOIDEA.

Cxi Cx2 Cx3 LI Li LIT AM PL. Nas Claws ss

Chyzeria 2 1 1 7 7 7 0 2 0 3 2

Grassia 2 ! ] 7 7 7 0 2 0 3 2

Nothotromtbleula 2 \ l 7 7 7 0) 2 | 3 2

Chatra 2 ] 5-7 6 6 6 1 2 0 2? 2

Monunguis | 5] ! 6 6 6 2 2 ! ] 2

Neatrombidium 2 l | 7 6 6 2 2 ] 1 2

Cackingsia 2 L | 7 6 6 2 2 | | 2

Leeuwenhoekia 2? \ | 6 6 6 2 2 | 3 2 T

Audyana 2 | | 7 6 6 2 2 0) 1 2

Apalonia | I | 7 7 7? 2 i) | 3 2 at

Womersia ] | ! 7 7 7 l 0 | 3 2 T

Womersleyla 2 1 ] 6 6 6 2 2 | 1,1,2 2

Mackerrasiella 2 1 1 7 7 7 Q 2 0) 1 24

(=Hydryphantes)

Butrombicula ] l 1 7 7 7 \ 2 0 3 2

Babiangia | | | 7 wi T | 2 0 3 2

Odontacarus 2 ] | 6 6 6 2 2 | 4 2 V

Watacarus 1 | \ 7 7 7 i 2 0) 3 2

Preridapus 2 | 1 7 7 7 0 2 (1) 3 4

Palydiseia 2 ! 2 7 7 7 0 2 (1) 4 4

Durenia 2 ) | 6 6 6 Z 2 ] 1,P,2 z T

Taraxithrombium 2 ] l 7 f 6 2 2 I | 2

Podathrombium 2 ] 1 6 4 6 2 2 I 3 2

Ralphaudyna 2 ] ! 7 7 7 (92) 2 | 3 47)

Walchia i a l-3 7 6 6 ) 2 0 3 2

(+)

Lassenla 2 2 3-4 6 6 6 () 2 () 3 4

lassent

Lassenia 2 | 2 6 6 6 0 2 0) 3 4

seutellata

Sauracarella ] (2) ] 7 7 7 2 2 | 3 2

T Tracheae present

jn ne eee ttttIISItSsSSISSSSSS

* The listing of a genus in this Table does not imply that I necessarily accept that these genera are all distinct from cach other: nor

that they are particularly related to each other,

OBSERVATIONS ON CH) ZERIA GANESTRINI AND SOME RELATED GENERA

The abbreviations used are Cx, 2 and 3, the num-

ber of coxalae or setae upon the ventral surface of

the coxae (Le. omilting supracoxalae), L J, If and

LI, the numbers of segments in the legs, including

the coxae (the traditional term coxae being used here,

in place of terms introduced by later authors); AM,

the number of anleromedian setae (scutalae) to the

dorsal scutum; PL the number of posterolateral scu-

talae to the dorsal scutum; Nas, the presence (1) or

absence (0) of a “nasus” or anterior projection ta the

dorsal scutum; 8S, the total number of scutal sensilla.

The information is derived from the most recent or

accurate authorities On these data, as quoted in the

list of references, or from the detailed descriptions

viven in the present paper.

Ii is clear that the classification of the larval Trom-

bidividea is not yet on a firm basis, since we have,

for example, various classifications for a fairly homo-

geneous group of larvae related to Chyreria, these

having been placed in the subfamilies Trombellinac,

Apoloniinae and Lassentinae by different authors.

It is proposed io this paper lo re-examine and

redescribe the larvae of CAvcerfa (including Grossia)

from the further material now availiable, and to

attempt to define the characters of the Trombellinae

(Trombellidae) lor larvae. The relationships of the

genera Neatrantbidium, Monunguis and Cockingsia

have ulready been examined by the writer jn other

papers (Southeott 1954a. 1957c) as well as by others,

and any further consideration will be deferred to later

works, The larva of the genus Audyana Womersley,

1954, with its leg segmentation formula 7, 6. 6 and

single-clawed pedolursi, appears al first sight to be

not particularly related to the Trombellinae considered

here. Nevertheless, it has been correlated by rearing.

One lurther genus was placed by Womersley

(19548) in his expanded Apoloniinae, namely Hoi

erslevia Radford, 1946. In an earlier draft of the

present paper | had written that although this

appeared to be a well-defined genus, there wus nu

strong evidence to connect it with Chizeria, Grossia

and Nothotrombicu/a and had decided to exclude it

from further consideration in the present paper,

Recently, however, | have been sent a batch of larval

orange-coloured miles Laken as ectoparasites on adults

of Teleogrvilus contmadus (Walker), the field cricket,

collected 10 km N of Armidale, New South Wules,

on }2 und 19 March 1980, by Mr Steve Davidson.

These proved to be typical Womerslevia, bul the leg

segmentation formula is 6. 6, 6, and not 7, 7, 7, as

recorded by Womersley (1954a, p. 114). (Vereammnien-

Grandjean (1972) had correeted this for HM’) miinnta

289

Radford.) It has been possible to rear a number of

these larvae to nymphs, which are typical 7rontbella,

keying down (as far as is possible for nymphs) to

Trombella adelaideae Womersley, 1939, in the key

of Womersley (1954b, p, 128),* The further deserip-

tion of these Australian larval Tromibella and their

nymphs will be dealt with in later papers.

A Turther genus referred lo the Trombellidae was

Ralphaudyna Vercammen-Grindjean er al,, 1974, this

action being based on its morphology. I is interesting

lo nate, however, that there are significant resem-

blances to those larvae allotted by Newell (1957) to

Lassenia in the dorsal scutum and other features.

At the present lime it is probably fair to consider

that larval trombidioid mies which can reasonably

be allotted to the Trombellidae are species ascribed

to the genera Chyzeria, Nothetrambicula, Woner-

slevia (=Trombella), Audyana and Ralphaudyna.

*As H consequence. Radfurd's species is here pe-oamed Pronpheller

pitatita (Radtird, 1946), carmb. nay

SUBFAMILY AND FAMILY

CONSIDERATIONS

Within the family Trombidiidae Leach, 1815. us

defined by Thor (1935) and Thor and Willniann

(1947), a number of groups have been separated alt

into families, e.g.

Trombiculidae Ewing, 1929*, Leeuwenhoekiidae

Womersley, 1944*, Vatacaridae Southeott, 1957,

Jobnstonianidae*, Calotrombidjidae (sic) Feider, } 959,

and a number of others, Among these is the family

Trombellidae (foritally proposed as such by Feider

(1955)), based upon the genus Trumbella Berlese,

1887. and erected originally as (he subfamily Trom-

dellinae Thor, }935,

Family Trombellidae Thor, 1935

Syn, Trombellinae Thor, 1935; Thor and Willmann,

1947; Trombellidae Feider, 1955

Definition

‘Trombidioid** mites lacking 2 erista metopica in

the adult und uctive nymphal stages. Propedosuma

eurrics a pair of dorsal sensilla. Definition of Jarva

deferred,

Key to Larval Genera of the Family Trombellidae

| Lee segmental formule 7, 7. 7 2

Leg segmental Tormuli 7,6, 6 or 6, 6. 6 4

2 (1) Dorsal seutum lacking wnreramedian Projection ("nasus |)

Chyzeria Canesten 1897

*See earlier comments in this paper ag the formal dates on which

these numes were proposed,

**For a definition of Trambidivides, see Sourheotl (19570, p )73)

200

Dorsal scutum with interomedian projection (inasus’) 3

3 (2) Dorsal seuruny with # setae

Relphavdvne Vereammen-Crandjeun ef al, 1974

Dorsal scurum with 6 setae

Dumbletun (947

41) leg segmental formula 7, 6.6

Leg segmentil formals 4. 6, 6

[S87 (— Womersteyia Radtopd |946)

Nuthotrambiuha

Andvana Womersley 1954

Trambella Berlese

Genus Chyserta Canestrini

Chy ceria Canestrini 1897, p. 463; 1899, p. 390. Hirst

1924, p, 1077; 1928a p. 563: 1928b p. 1021:

1929 p. 165. Vitzthum 1931 p, 148; 1942 p. 286.

Womersley 1934 p. 182; 1937, pp. 75, 76; 1939,

p. 155: 1942, p. 169, Lawrence 1944 p. 438.

Thor and Willmann 1947 p, 203. Baker and

Wharton 1952 p, 250. Vercammen-Grandjean

1955 p. 260; 1973b p, 110, André and Lelievre-

Farjon 1960 p. 461, Southcott 19614 p. 563,

André 1962 p. 124, 1963 p, 561. Vercammen-

Grandjean and Kolebinbova 1968 p. 250. Robaux

1969 p, 69. Krantz 1978, p. 351.

Grossia (as larva) Womersley 195da p, LI) 1. Audy

1954 p, 166; 1957 p. 442. Vercammen-Grandjean

1967 p. 2; 1971b p, 315; 1972 p. 231; 1973a p.

58: 19734b p. 110, Robaux 1977a p. 666.

Type species: CAyzeria ornata Canestrini, 1897,

(original designation)

Definition of Adult

Trombidijdae without well-defined crista metopica,

but a shield may be present in the central area of

dorsum of propadosoma. A pair of dorsal idiosomal

sensilla may arise upon a protuberance. usually anterior

to the level of the eyes, Eyes 2 + 2, sessile. Dorsum

of idiosoma with a lateral column of finger-like pro-

cesses, covered with setae which may be of two distinet

types, the one stiff and pointed. the other flexible.

Additionally there may be a dorsal anteromedian

projection arising about level with the most anterior

of the dorsolateral projections, and a posteromedial

projection. Area between the dorsal projections of the

idiosoma with numerous fine, long, convoluted setae.

Definition of Larva

Vrombidjid larvae with a single prodorsal scutum,

transversely placed, with AL and PL scutalae placed

towards AL and PL angles respectively of scutum.

No anteromedian scutalac. A pair of seutal sensilla

present, widely separated, placed at about the mid-

level of scutum. Sensillary hairs filiform, ciliated.

Eyes 2 + 2, conjoined, sessile, placed laterally or

posterolaterally to scutum. Intercoxal formula 0, 0,

2. Peducoxal formula 2, 1, 1. Legs with divided

femora, 1c. leg segmental formula 7, 7, 7 (including

coxac). Fedocoxalae may be short, stout, pointed,

blunted, peg-like, smooth or irregular. Pedotarsal claws

REC §. AUST. MLS.. 18 (14), 285-426

June, 1982

all rather slender: anterior terminating in expanded

suction-membrane; middle (neomedian) claw normal,

more or less falciform; posterior claw similar to ante-

rior, Palpal tibial claw strong, divided into three strong

divergent tines. Idiosamal setae may arise [rom small

plates, these being expansions of the normal alveolar

annulus of the seta. Ventral setae may have the basal

part of the shaft (scobillum) thickened.

Key (0 Known Larvae of Chypseria

1 AL, seutalac short, blunt, rebust, somewhut expanded, with

short, strong serrations. about 17-20 um Jong. Pedicoxalae

short, blunted, rhultituberculare, with tubercles present wlong

must of (he setal shaft (scobillum). Posterior dorsal idioso-

Inalae fo wbout 46 um long

Chyzeria hirsti Womersley, 1934 (South Australia)

AL scutalue normal, tapering, moderately pointed. more or

less cihuled, Pedicoxilae cither tapering, potted and ciliated,

or blunted and tuberculate, but in the latter case with (he

(ubereley absent in the proximal half of the seobillum.

Posterior dorsal idigsomual setac in range of 75-140 um 2

2 ()) Pedicoxulae pointed or very slightly blunted. AL. scutalae

wboul | 12 am long; AL/(A+P)> 2.1

Chizerta onvehia (Womersley, 1954) (Victoria)

Pedicoxalue very blunt-ended. AL seutalae in range of 55-

107 wm long: AL /(A-P)< 2.1 3

42) Pedicoxalae weakly tuberculale through (he distal hall of

the setal shalt (seobillum), AL scutalge about 55 wm long;

AL/(A-P) about 1.06, Palpal tibial claw tines only slightly

divergent Chyeeria flinderst sp. nov, (Tasman)

Pedicoxulue weakly (uberculale in wboul the distal one quarter

of the shalt of the seta (scobilum), AL seutahte tn range

of KK-107 pan tong: “WL/(A-P) 19-20. Palpal tibial claw

tines widely divergent

Chyeria derrick sp. nov. (Queensland)

Chyzeria hirsti Womersley

(Figs. [A, B; 2 A-Fy 3; 4 A-D)

Chyzeria australiense (sic) var. firsti Womersley,

1934 p. 182; Womersely 1937, p. 76 (as adult)

Chyzeria australiense (sic) Womersley, 1939, p. 155

(as larva)

Nee C. australiense (sic) Hirst 1928 (1928a) p. 563

(adult)

Description of Larva (based largely upon specimen

ACB 13/14 L.1) (Figs. 1A, B; 2A-G) Colour in life

red, Length of idiosoma (mounted on slide) 285um,

width 190um; Lotal length of animal from tip of

chelicerue to posterior pole of idiosoma 360um,

Dorsal scutum about twice as wide as long, trupe-

zoidal with somewhat rounded angles, the scutum

widest posteriad, Anterior margin somewhat simuous,

posterior margin somewhat convex (a little angulated

in the middle), lateral margins slightly concave. Seutal

sensilla very widely separated, and behind level of

middle of scutum. AL scutalae placed towards AL

angles of scutum, PL scutalae towards PL ungles of

sculum,

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME REI ATED GENERA %|

TABLE 3.

Designated

variate ACB 13/14 ACB 13/14 is isa

1] 12 !

AW S6 83 73

PW 110 116 104

SB 61 66 37

ASBa 9 37 35

ASBp 29 28 2h

I, 68 65 6!

WwW 126 L34 ella

AP 32 3] 3t

Al is WY 20

PL 39 25 29

Sens 59 63 53

DS 29-57 45-57 24-49

mid-DS 39-44 35-42 39-40

PDS 45 44 24-37

Anterior scutalue short, robust, spindle-shaped, cil-

iated, the cilia blunted, outstanding a little and thus

the AL scutala resembles a distorted partly opened

pine-cone with about 9 bracts; posterior scutalae rather

longer, pointed, clongate-spindle-shaped, with proj-

ecting barbed pointed cilia. Scutal sensillary hairs

slender, with distal sparse cilia,

Eyes 2 + 2, sessile, near PL angles of scutum;

corneae oval to ovoid, posterior the larger. Anterior

cornea has maximum diameter |5um, posterior has

maximum diameter |9um.

Dorsal idiosomalue long, very slightly tapering,

pointed, slightly ciliated with barbed cilia. Setae arise

from small plates, oval or somewhat straight-edged

where they impinge upon cach other; plates are

expansions of normal annuli of alveolar sockets; setae

usually arise eccentrically upon the plates. Setae

arranged 6, 6, 6, 6, 2; total 26.

Venter lacking setae on idiosoma between coxae |

and Il. Between coxae II] a pair of tadish-shaped

setae, having an expanded basal part to the shaft

(scobillum), the remainder long, tapering, pointed:

setie 17um long. Behind the level of coxae II] are

about 30 setae (apart from the four anals), similar to

the last, arranged in somewhat irregular transverse

rows across the ventral opisthosoma. These setae

change in character as one proceeds posteriad. The

more anterior setae are short and “radish-like” or

“parsnip-like’, with a thickened basal part of the

scobillum and a maximum thickness at about the

middle of the seta, and are similar to the pair between

coxae IT]. All ventral idiosomal setae arise from small

basal plates (expanded annuli). Posteriad the setae

gradually become more elongate, more ciliated and

STANDARD ASD OTHER DATA FOR SOME LARVAR OF Chyceria hirsty

Specimen

ACB322 AC B468 ACB46% ACBSM

12 L| La LI

79 7K 7 nh?

96 110 107 i)

55 57 sy i)

33 28 2 oh

23 cals 29 Ly)

535 cade s7 64

he 149 138 tan

29 a2 4] a)

\7 18 {9 17

29 29 43 2

50) Ol Ss?

40-52 43-42 34-66) 42-50

40 42 39-66, 43

44 37-46 33-39 3H-d)

stronger, but lose the bulbous expansion. Setac of

anterior row ure 16-2] um long, and the posterior setae

to 37m Jong. Anus with two oval valves, 45um long

by 374m across with the valve lips in apposition.

Each anal valve with two long setae, anterior pointed,

ciliated 22um Jong, posterior blunted, ciliated 28um

Jong. Idiosomal setae lateral to anus tapering, pointed,

clongate, 354m long,

Coxalace 2, 1, 1, All coxalae short, stout, unciliated,

with indefinite coarse tuberculitions in lateral aspect.

und terminally somewhat bifid. resembling an irreg-

ular plum in shape; setae 9-Tlum long by 5-7um

across. The tuberculations number about 7, and ure

placed more or less distally upon the setal shaft or

scobillum.,

Legs normal. with the distal ends of femora. genua

and tibiae tending to expand. Leg [ 375um long, II

380um, Hl 420um (all lengths include coxae and

claws). Leg scobulae normal. curved, ciliated, termi-

nally pointed or blunted.

Tarsus 964m long by 29um high, tibia | 55m

long, genu | 46um (Til/Gel = 1.20); tarsus [1]

1Olum long by 24um high, tibia ITE 55am. genu I

S4um (Ti /Gelll = 1.01). (Tarsal lengths given

exclude claws and pedicle.). Genu. | with specialized

seta VsGel,77pd: tibia | with SoTil.1 4ad. SoTil. Sad,

VsTil.s6pd. Genu [1] with specialized seta

VsGell.79pd. Tibia TI] with specialized seta

SoTi1.22pd. Tarsal setae as figured. Tarsal pedicle

normal. Pedotarsal claws: anterior and posterior slen-

der, spathulate, middle (neomedian) falciform., slen-

der. unciliated, With sharp terminal downturned tip.

Gnathosoma rather small, compact, fattened pas-

292 REC. S. AUST, MUS., 18 (14): 285-326 June, 1982

FIG. 1. Chyzeria hirsti Womersley, larva, specimen ACB13/14

L1. A, Dorsal view, legs on right hand side of figure omitted

beyond trochanters; to scale on left. B, Detail of a dorsal

idiosoma! seta and its basal plate; not to scale.

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 293

oe hy |

eo YI

As sd)\S

2) ——S + A es

E F

FIG. 2. Chyzeria hirsti Womersley, larva, specimen ACB13/14

Li. A, Ventral view, legs on right hand side of figure omitted

beyond trochanters; to scale on left. B, Palpal tarsus, enlarged.

C, Claws of tarsus I, enlarged. D, E, F, Posterior ventral

idiosomal setae, enlarged. G, Pedocoxal seta, enlarged. (B-G

not to scale.)

294 REC, S. AUST. MUS,, 18 (14): 285-326

teriorly. Chelicerae bases 74um long to tip of cheli-

ceral blades, by 55m across conjoined. Cheliceral

blades small, curved, sharp-tipped, with small teeth

along cutting edge and other surfaces. Galeala spini-

form, robust, curved, simple, slightly blade-like, ter-

minally pointed, 17um long. A delicate circular lip

surrounds the tip of the chelicerae. Anterior hypos-

tomala not identified. Posterior hypostomala very

slender, tapering, unciliated, 30um long.

Palpi as figured. Palpal setal formula 0, 1, 1, 3, 9.

Palpal femorala tapering, with one barbed ciliation,

37um long. Palpal genuala tapering, pointed, uncil-

iated, 26um long. Palpal tarsal setae as figured. Pal-

pal tibial claw (odontus) with three strong slightly

divergent tines. Palpal supracoxala a slender blunted

peg, 4um long.

SOURCE OF Chyzeria hirsti LARVAE

Larvae of Chyzeria hirsti have been obtained on

three occasions since 1938 by experimental rearing

from the adults confined in tubes of soil.

(1) Serial ACB13, 14. Two adults were captured

by the author on a hillside above the property of

Birksgate near Glen Osmond, South Australia, on 23

May 1938. These mites were placed in a glass tube

with a small amount of soil from their capture site.

Local natural unsterilized soil* was used, as with all

my similar attempts with many different trombidiids,

on the hypothesis that the mites would find food in

the form of fungi or small arthropods in the soil.

Subsequent notes (abbreviated) were as follows.

13.vi.1938 Both (mites) active. One active and

plump, the other smaller and less plump.

2.vii.1938 Both active and well.

10.vii.1938 As 2.vii.1938.

24.vii.1938 Alive and active. Some fresh [unster-

ilized] soil added.

14.viii.1938 Alive and active.

10.ix.1938 Both active on stimulation. One thin,

one plump. One oribatid (mite) and one parasitid

(mite were) seen in the tube. There were a num-

ber of larval trombid (iid)s seen in the tube. Six

were mounted (on a slide). One (further) one

was lost. The others (were) kept in the tube. The

adults had larvae on their backs, but most of the

larvae were free.

12.ix.1938 One larva put into formalin.

13.ix.1938 Adults alive and active. Not less than

three larval trombid(iid)s were free, running

* The statement by Womersley (1939, p. 156) that the soil was

sterilized was an error on his part.

June, 1982

around inside the tube. One Chyzeria adult had

two larvae on the dorsum; the other adult had

four larvae on the dorsum. The larvae were not

attached by their mouthparts. They ran over the

bodies of their parents, without leaving them. [I

obviously assumed at the time, presumably from

the larval behaviour, that both of the adults were

parents of the mites, although in retrospect that

conclusion may not have been justified.]

(On examination of the slide-mounted adults

one finds that one specimen (ACB13/14A) is

clearly a female, the body being clarified in the

mountant to show it contains a slightly irregular

ovoid egg measuring 295um by 260um. No evi-

dence of internal chitinization in the egg is visible;

a more definite statement cannot be made on

account of the thickness of the setation. In the

case of the other adult (ACB13/14B) the sex is

indeterminate, as the mite’s idiosomal structure

is obscured by attached debris, and no eggs can

be seen; the genitalia are also obscured.)

So there were at least 17 larvae in all (6

mounted on slide initially, one lost 10.1x.1938,

one put in formalin 12.ix.1938, and not less than

9 in the tube),

14.ix.1938 Adults active. Several larvae on dorsum

of each adult. None [was] attached by its mouth-

parts, but all were moving over the surface of

the adult, and keeping to the posterior half of

the adult for the most part. When on the dorsum

of the adults the larvae moved one or two legs

at a time, and quickly, with intervals in between

so they moved forward jerkily.

Several larvae were seen running round [in]

the tube.

One larva was seen to leave the adult by a

leg, and to get several mm [away] from the

adult, and then return to the dorsum by a leg.

(Possibly the larvae reconnoitre [using] the adults

as bases.). One larva was seen dead in the tube

(lost [on] 17.ix.1938).

15.ix.1938 Adults active; one with 4, other with 3

or more larvae on it. None [was] attached by

[its] mouthparts. Two larvae where seen free in

the tube.

17.ix.1938 Two [larvae] were taken out (returned

later) and put on my arm. They were watched

with a strong lens, but neither showed any signs

of wanting to pierce the skin. Each was watched

for about three minutes. They ran quickly over

the skin.

22.ix.1938 Adults active, each with one larva on

the posterior part of the dorsum. Larvae not

attached by mouthparts, moving as before. Four

OBSERVATIONS ON CHYZERLA GANESTRINI AND SOME RELATED GENERA

larvae seen running around tube. One larva seen

dead in the tube.

Observations continued over the succeeding wecks.

By 24 September 1938 all the larvae were dead,

possibly by the tube being made dryer. The tube was

then re-wetted. On & October 1938 the adults were

active, On 22 October they were shrunken and feeble,

and on 29 Octuber were dead. On being slide-mounted

neither was then seen to contain eggs, nor were eggs

seen al any stage in the tube,

(2) Serial ACB322

On | August 1948 a very plump adult Chyzeria

was captured in soil at Workanda Creek, National

Park, Belair, South Australia. It was placed in a iube

with a little soil. Although examined regularly during

the period August to November no epes or larvae

were seen in the tube. On 7 December 1948 two dead

larvae were seen in the tube. The adult mite was

healthy on 28 December but was dead on 28 January

1949,

No. eggs or live larvae were observed in the tube

al any stage.

(3) Serial ACB468

An adult Chypzeria hirsti was captured al the same

site on 3 September, 1950, placed in a tube with

some damp soil, and examined as opportunity offered,

On 29 October | recorded “Worm in tube. One yellow

Xeve in tube, Tube quite damp’. On 14 November

seven larvae were counted in the tube, several being

immobile, trapped in water droplets, The yellow object

noted on 29 October remained unaltered in appear-

ance, and I recorded in my notes “I do not doubt

[that] it has nothing to do with the CAyzeria. To my

mind this tube experiment establishes the viviparity

of Chyzeria,*”

One active larva was then taken out of the tube

and manipulated into repeated encounters with the

adult, these manoeuvres being aided with a fine

brush, Although placed on the back of the adult on

three or four occasions, the larva showed no indication

to remain there, ut once hurrying off. | commented

in my notes that the larva looked “in fact like a

typical larval Trombicula”’ (the term Trombienla

“(When taken in conjunetion with the previous observations).

295

indicating here a larval trombiculid mite of the genus

Eutrombicula and various other genera), On being

replaced in the tube, further encounters occurred

between the adult and the larva, but the larva made

no attempts to climb on its presumed parent's back.

The trapped larvae were in a row as though they had

been recenily laid in a regular fashion, the distance

beiween successive larvae being about 5 or 6 times

the length of an individual larva.

On 15 November a few larvae were still alive and

healthy, but again no altempt for one to climb on the

back of the adult was observed. By 4 December all

larvae were dead, The adult appeared active and

healthy. However, by | January the adult also was

dead.

Thus in this experiment no phoretic relationships

were Observed.

(4) Serial ACB526

On 29 July, 1951 another adult Chyzeria {rom the

same site was captured and confined in a tube with

some soil. Observations were made at intervals. On

9 September no egys or larvae were seen in the tube,

At the next observation, on 24 October, the adult

was recorded as being well, the tube damp. “One

larva is present in the tube. It gives the impression

of walking on liptoes more than the larva of Eremiil-

leria* ete.—in this it resembles Trombicula cte, |

watched jt encounter its parent several times. At one

of these encounters if ran up its parent's leg, over its

back and down another leg, then circled widely and

returned to the moving parent and walked underneath

it. Tt appears to be attracted by movement.....Only

this one larva (was) seen; no egg seen.”

On 20 November the tube was rather dry; the

adult alive and well. Two dead larvae were seen in

the tube.

On 25 November the tube was dry; the adult alive

and well,

The adult remained alive until 29 December, bul

was dead on 27 January, 1952. Only one larva was

enventually recovered from the tube.

The time relationships between the adults and the

larvac in this experiment are summarized in Table 4.

*The larvae of Echinothrombium willungae (Hirst, 1931); see also

u note in Southcott (19574; p, 142),

TABLE 4. SUMMARY OF DATA ON TIME-RELATIONSHIPS OF ADULTS AND LARVAE IN SUCCESSFUL REARING EXPERIMENTS WITH

Chyceria hirsti WOM

Experiment ACB 13,14 ACRI22 ACB468 ACBS22

Adult(s) captured 23.v.1938 Lyi, i948 3.x, 1950 Wyn 19St

Larvate) seen alive {0-22 in. 1938 = 14-15.41.1950 D4.x,195]

Lurva first seen dead 14.1. 1998 T xii 193% 15.47 1950 DOKL IMSL

All larvae deud 23-24.in_ 1938 (7 xi 1948) (G.xi, (950+ 20.81, 1954

4.xii, L950,

Adult(s) died DA-29.x, 1938 29x11, 1948- Sth PYS0- Myxis, 195) -

28.1, 1949 (1951 27.3, 1952

ee See

296

In summary, larvae have appeared on four occa-

sions in tubes containing live adults of Chyzeria hirsti

confined therein, with unsterile soil as a rearing cham-

ber. Eggs have not been observed. Larvae have

appeared alive between September and November

and possibly are restricted mainly to the period Sep-

tember-October. No larva has been captured in the

field, and at present these larvae are the only ones

known of Chyzeria hirsti. No evidence of any poten-

tial host for these presumably parasitic larvae has

been discovered. The adults are used by the larvae in

a phoretic relationship, and there has been no evi-

dence of parasitisation of the parent. Man does not

appear to be a suitable host for the larva, from a

single experiment. (This was expected; it is antici-

pated that eventually the larvae will be found to

parasitise some arthropod host. See also the record

for Chyzeria derricki sp.nov., below of parasitisation

by this species of a tettigoniid host, and also the

record for Chyzeria onychia (Wom.), in which this

species was taken parasitic upon an adult Chyzeria

sp.).

COMMENT ON LARVAL BEHAVIOUR IN Chyzeria hirsti

Although over the years 1938-1980 continued

efforts have been made, as opportunity offered, to

obtain Chyzeria hirsti (and other Trombidiidae) lar-

vae from experimental preparations, the only suc-

cessful attempts with Chyzeria hirsti have been those

recorded above.

In two of the experiments recorded there was evi-

dence of behavioural interaction between the larvae

and the adult(s). In one further instance the obser-

vations were negative, while in the fourth experiment

the larvae were dead when first observed.

There appears to be no doubt, however, that the

larvae of Chyzeria hirsti have an unusual behaviour

among Acarina, in that they have a phoretic relation-

ship with their parents or parent. The evidence given

above clearly indicates that the larvae use the adults

as a means of transportation, and hence dispersion,

this coming within the formal definition of phoresy.

No doubt, in addition, the adults provide shelter and

some degree of protection, in that being able to

retreat to an elevated niche upon a parent represents

some protection from predatory arthropods of a size

capable of preying upon animals the size of the Chy-

zeria larva but not of the adult. This extends the

relationship between the parent and the larva beyond

transportation-utilization or phoresy, into one of pro-

tection. Similar extended relationships occur in other

arachnids, perhaps the best known ones being in the

case of scorpions and lycosid spiders, but such rela-

tionships occur widely among various groups of

arthropods, particularly the Crustacea, and in fact

even more widely and extensively in vertebrates. Thus

not only is this seen frequently among Mammalia,

REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

but is well-known among Amphibia, and in fact the

carrying of juveniles in the mouth or a brood-pouch

is also known widely among the fishes.

Among the Acarina, however, although phoretic

relationships between mites of a number of families

and other arthropods are well-known (see Krantz,

1978), as far as I am aware there has been no previous

record among the Acarina of phoretic behaviour

occurring between an adult mite and the larva of the

same species.

The observations above on maternal care of the

larvae help to provide also some explanation for the

peculiar structure of at least two species of Chyzeria.

The dorsum of the idiosoma of the adult carries a

number of largely peripheral projections. The area

between these projections, in C. australiensise and

C. hirsti, bears a dense mat of fine, convoluted, simple

hairs.

In a recent paper Rovner et al. (1973) have sug-

gested a possible role for the abdominal setation of

the lycosid spiders in terms of maternal-juvenile

behaviour. Adult female lycosids have peculiar

knobbed hairs on the abdomen. Juveniles will not

cling to their mother’s abdomen once these hairs have

been shaved.

Since the Chyzeria hirsti there is an unusual

behaviour pattern in the larvae and an unusual struc-

tural modification of an area of the adult, where the

unusual larval behaviour is manifested, it would be

reasonable to interpret the adult structure as possibly

subserving the larval-adult behavioural modes.

Although in larval Chyzeria the anterior and posterior

(neolateral) pedotarsal claws are provided terminally

with a ‘suction’ membrane, this adaptation cannot

necessarily be considered as related to the adult-larva

behaviour, but could possibly serve some other func-

tion, such as ensuring adhesion to a thick-skinned

host.

It is also reasonable to speculate that the fine

convoluted hairs of the adult allow utilization by the

larva in its rather jerky mode of progression over the

idiosoma. The convoluted hairs appear to be of uni-

form thickness and unciliated, even when viewed

either by phase-contrast or polarized-light micros-

copy. The shaft (scobillum) outer layer is birefringent,

as is usual for any trombidiform normal-type seta

(scobala) (see Southcott 1961b, 1963a for details of

seta terminology). The undulations or coils (and the

terminal hook) on these hairs could serve the same

function as e.g. special knobs (as is the case in the

lycosid spiders mentioned above) i.e. allowing the

larvae better to utilize them in walking or moving

across the adult’s back. For these setae the name

sericala is proposed,

These setae are present in both CAyzeria austral-

OBSERVATIONS. ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 747

iensis and C. hirsti. They are present in some other

species of Australian Chyzerla with various specific

determinations by Womersley, in the South Austra-

lian Museum collection. These inclide specimens

from Tasmania, Victoria (bul not all) and Queens-

land. They are nol present in three specimens in the

collection identified as C. musgravei Hirst by Wom-

ersley. (I have not seen the type of C. musgrave/.).

These three specimens instead possess in the central

dorsal idiosomal space a number of stiff sword-like

setae, arising from small plaques (seta-hases}, as well

as longer, Mexible, ciliated setae, arising similarly.

Details of these three specimens are as. follows:

(1) ACB715. One specimen. Top of Carrington

Drive, Natfional] Pfar]k, N.S,W., 31.%.1944, H.

W [omersley]..

(2) ACB716, Queensland, 1943, No other data.

(3) ACB717 (Two slides) One specimen. In leaf

debris, Brookfield Rd., nr Haven Rd. Brookfield,

5.iv,1961, E. H. Derri [c] k [Queensland].

Other Possible Phoretic Usage of Chy'zeria hirsti

Adult

A further usage of an adult Chyzerfu as a phoretic

host has been observed, In that case an adult female

of Chyzeria firsti, collected at Workanda Creek,

National Park, Belair, South Australia, on | August

194%. was observed to be carrying an adult female

pygmephorid mite. For further details, soe p, 318, and

Fig. 18 in this paper.

NOTES ON THE ADULT OF Chyzeria hirsti WOMERSLEY

As stated above, Womersley separaled this form as

a subspecies in 1934. In his discussion on Chyzeria

australiense (sic), C. occidentalis Hirst, 1929 and C-.

musgravei Hirst, 1928 Womersley (p. 182) included

the following remarks (after reducing C. occidentalis

to a ‘variety’ of ©. australiense (sic) Hirst, 1928)

‘Further, Hirst's C. musgravei must. also be consid-

ered as a variety differing in that the anterior median

plate (this is an error for ‘process’ of

‘tuberosity"—R, V.S,) is developed into a compartively

long process, One specimen amongst the Hirst mate-

rial in Professor Harvey Johnstan’s keeping, and now

in the South Australian Museum, is clearly interme-

diate between the two forms in respect of this char-

acter, the process being shorter and triangular. This

specimen was labelled in pencil by Hirst as C, mus-

gravel. All the specimens collected recently by Mr

M, W. Mules and myself in the Adelaide District

agree with this intermediate form, for which the name

C. australiense yar. hirsti var. nov. is proposed.

‘Loc. Type; Willunga, West, Aust. paratypes:

Woodside, S. Aust., July, 1933 (W.M.); Mt Osmond,

8S. Aust., Sept. 17, 1933 (H-W.),; Glen Osmond, S.

Aust., Oct. 1, 1933 (H-W.)”

Womersley also provided (pp. 183+4) a key to the

Australian and New Zealand species of Chyzeria in

which the ahove differences were restated in separating

C. australiense (sic) var. musgrayei from C. austral-

iense (sic) var, hirsti, The New Zealand species ©,

novaezcadlandiae Hirst, 1924 was misnamed as €-

novae-hollandiae Hirst by Womersley, who was thet

comparatively unfamiliar with the Australian fauna

and history,

Although the characters proposed by Womersley

(1934) in the separation of this taxon as a Subspecies,

and presented by him in the text and the key to the

species, appear comparatively slight, the present

author believes (hai Airsti is worthy of full specific

rank and therefore formally raises i to species status

now, Again, although this paper docs not aim to deal

with the classification of the adult Chyzeri/a, some

redescription of adult Chycerla hirsti and comparison

with Chyzeria australrensis Hirst are required.

DESCRIPTION OF THE HOLOTYPE ADULT OF Chyrera

hirsti’ WOMERSLEY, 1934 (FIGS. 3;.4 A-D)

The specimen (identification ACB718) is slide-

mounted in clear unstained medium, cleared, deco-

lorized, squashed and disrupted, possibly partly dis-

sected, so that it is difficult to give other than impre-

cise details of the soft parts. Scattered around the

mounted specimen are a number of eggs and skins,

without evidence of contained embryos (at least there

is no evidence of internal chitinization). The disrup-

lion of the animal on the slide is such that it is almost

divided into two parts by a line of tedring behind the

coxae Il, and there is a further oblique tear in ihe

opisthosoma (Fig.. 3).

Mite appears to be of normal CAyzeria shape and

charucteristics, with moderately strong legs and gna-

thosoma,, and a somewhat elongate idiosoma with the

usual processes. Legs [ and [V are about equal in

length to the idiosoma, with legs I] and [11] somewhat

shorter,

Length of animal on slide from tip of chelicera to

lip of posterior idiosomal processes 2900 um. (The

length in life is estimated from the degree of damage

to the specimen as about two-thirds of this figure, i.e,

ca 1900-2000 ym. Length of idiosoma in the mount

1950 wm, width (at widest part, including edges of

dorsal processes) 900 pm.

Eyes 2 + 2, sessile, anterior eye 42 ym across,

posterior eye larger, 56 um across, rather more

medial; the two cyes making 4 conjoined mass 116 urn

by 57 um. In the mid-line, somewhat anterior to the

eyes, is a pair of prominent tubercles, each about

46 wm across and high. From each of these sensilla

a fine sensillary hair emerges, about 240 um long.

Between the eye masses and the sensillary tubercles

is the “sensillary area”, which bears about 20 long

cilated setae. A few similar setae arise also along the

298 REC. 8. AUST. MUS., 18 (14); 285-326 June, 1982

“

~o8

FIG. 3. Chyzeria hirsti Womersley, adult. Holotype female, spec-

imen ACB718, shown in transparency (genitalia and anus

omitted), The body and leg setae are mostly omitted. The

specimen has been ruptured in the mounting process, and ova

of various sizes have been released.

OBSERVATIONS ON CHYZERI4 GANESTRINI AND SOME RELATED GENERA

medial edges of each eye mass. There is a weak

indication of a crista running posteriorly from the

sensillary area for about 360 wm; this is thin and has

an irregular margin, becoming vaguer and weaker

posteriorad,

Dorsum of the idiosoma with the characteristic

digitations of projections of the genus. The antero-

median is rather conical, but not well-defined. These

projections are all studded with little bosses carrying

the setal plates. They carry two kinds of setae (scob-

alae), the more prominent. kind being the stiff spine-

like unciliated setae, which project out like spines of

a hedgehog, and range from 86-260 um long. Among

the spine-like scobalae are a number of more norrnal,

slender, cilated setae, which project out beyond the

spine-like setae, The anterolateral and posicrolateral

processes are long, more or less round-ended and

hence sausage-shaped. In between them are less

prominent projections, so that along each side of the

dorsum of the idiosoma is a column of five more or

less distinct projections. The second and third proc-

exses project the least, bul are still distinct. The two

Most posterior projections are bulbous. There appears

also to be a posteromedial projection, probably some-

what ventrally placed, but as in the specimen it is

detached and displaced away from the idiosoma

between the two most posteriar projections, its exact

placing is not determinable in this specimen.

In the central area of the dorsum: of the idiosoma

is 4 large number of long fine convoluted hairs, each

rising fram an expanded seta-base, forming a small

plate, These setae are simple, terminally pointed, but

so heavily convoluted that it is difficult to estimate

their length; the degree of convolution also tends to

give a false eslimate of their numbers. (See Fig. 4C).

Among them are a few of the long spine-lke setae.

Venter of idiosoma is provided with numerous short

ciliated hairs, without any spine-like or sword-like

setae. Genital valves appear normal, wilh the usual

3 + 3 oval or circular “suckers'; the degree of dis-

ruption of the specimen precludes further detail being

given,

Legs fairly robust (see Fig. 3); lengths (trochanter

proximally to tip of tarsus, without claws) I 2.000 um,

Il 1405, UL 1465, IV | 905. Tarsal claws appear

normal, For leg segmental measurements see Table

5 (below),

Gnathosoma as figured. Cheliccrae appear normal,

but not visible in lateral view, as the right chelicera

is at least partly missing (?dissected away). Palp

robust Palpal tibial claw strong, with a strong acces-

sory claw immedialely behind it, reaching to about

24 its length; another accessory claw behind that also.

These two accessory claws are the most distal mem-

bers of a comb of about 12 spines along the lateral

299

face of the palpal tibia. Similar spines are spread

over the lateral face of the palpal tibia, without any

regularity of arrangement. (Note: the customary term

spine is used here, although these projections are not

true spines, but are highly modified normal-type

setae, 1.e. scobalae.) Palpal tarsus ovoid, shorter than

the tibial claw, well-covered with setae, these being

longest peripherally, so that these setac extend ta

reach the level of the tip of the palptibia.

The eggs on the slide are round to oval, or ruptured.

The size 1s quite variable, ranging from about 380 wm

x 345 um, down to 120 um x 120 um; an average

dimension is perhaps 240 um x 185 win.

SITE OF ORIGIN OF THE TYPE OF C. Airsti

Some clarification is required of the details of the

origin of the specimen.

The specimen of an adult in the South Australian

Museum collection is labelled TYPE in red ink in

Womersley’s writing. Other daia on the sole slide

label are “Wondside. SA, July, 1933 M. W. Mules’.

The determination of Womersley's. on the label 1s

‘Chyzeria australiense Hirst y. hirsti an, v | have

added ‘ACB718".

Wamersley (1934, p. 182) stated (hat the Type

came from “Willunga, West. Aust.’, and lists the

Woodside specimen among the paratypes. This is

clearly an error by Womersley, as the slide of the

Woodside specimen js clearly and unequivocally

labelled, and the four slides containing material from

‘Willunga, W. A. have no marking indicating thal

they were ever considered as type material. The four

Willunga slides represent only one specimen, which

Womersley had partially dissected. | have lubelled

these four slides ACB677A-D, for future reference.

Although the slides give no date or name of collector

on their present labels, which are in Womersley’s

writing. one slide (A677B. of a palp) has written upon

it (labelled as C, misgravei by S, H.’, this indicating

it was af Hirst’s collecting and original identification.

No evidence of the original label of Hirst remains.

As each slide bears the notation “Chyzeria austral-

tease Hirst v, hirsei n, v,” (or some abbreviation of

this) it is clear that this specimen is to be correctly

desiznated a Paratype.

Slide ACB677D has written upon the label, in

Womersley’s writing ‘Willunga. W_A./ no date/ from

Univ. Coll./', indicating that this was one of the

specimens deposited by Professor T, Harvey lohnston

in the South Australian Museum, from the Depari-

ment of Zoology, University of Adelaide.

Hirst had worked in ihe Zoology Department while

Professor Johnston was ubsent in the Antarctic with

the British Australian and New Zealand Antarctic

300 REC. S. AUST. MUS., 18 (14): 285-326 June, 1982

100

500

SAS —

0 ‘

100

um 100

FIG. 4. Chyzeria hirsti Womersley, adult. Details of holotype

female specimen ACB718. A, Dorsal idiosomal sensilla, eyes

and neighbouring structures. B, Left palp, in lateral view, and

right chelicera. The left chelicera is missing. The proximal

part of the right palp is also shown. C, Setation of the central

dorsal part of the idiosoma, showing the fine convoluted seri-

calae, and a few of the sword-like setae. D, The most posterior

of the right lateral column of dorsal idiosomal processes,

showing the two types of setae present. (All figures to adjacent

scales.).

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 301

Expedition of 1929/1931, but unfortunately commit-

ted suicide on the return voyage to England, Hirst

had collected at various sites in South Australia and

eastern Australia, one of his species in fact being

named Microtrombidium willungae Hirst, 1931* col-

lected by Hirst at Willunga, South Australia, October

1929.

Reference to the Gazeteer of the United States

Board on Geographic Names, Number 40 (Australia)

shows that the name Willunga occurs only in South

Australia, in the forms of Willunga and Willunga

Hill, which are in fact contiguous, the former being

a township, Both are at 35°17’S lat.; the former al

138°33’E and the latter al 138°34’E long. Womersley

arrived in South Australia only in 1933, and at the

time was not familiar with South Australian place

names (see Southcott, 1963b),

Chyzeria australiense (sic) was described by Hirst

in 1928 (1928a, p. 563) from “Swan River, W.A. In

nest of Ponera lutea.”'

In the South Australian Museum collection are

four relevant slides labelled as originating from Swan

River, W[estern] A[ustralia]. Three of them bear a

label in Hirst's writing, while one bears a label by

Womersley, These represent the TYPE and a Para-

type. | have attached identifying numbers ACB719

and ACB720 to these specimens as follows:

ACB719, Intact specimen mounted in well-slide,

labelled in what | interpret as Hirst’s writing

“Chyzerla — australiense HIRST/Swan

River/W.A./S.A. Mus. Coll./ Nest of Ponera

lutea”

ACB720, A series of three slides, to which [ have

attached the identifying numbers ACB720A, B

and C, as follows:

ACB720A: Heavily dissected specimen, mounted in

well-slide, labelled in same writing as preceding

“Chyzerla australiense HIRST/Swan

River/W.A./S.A, Mus. Coll,/Palp”, the term

“Palp” being crossed out in pencil.

a Ettmuelleria willungae (Hirst); see Southcott 1957a, p.

ACB720B: Slide containing the end of a first leg of

a trombidiid mite, of three segments, labelled as

for preceding slide, but with “Ist Leg” replacing

the term “Palp", Reference to slide ACB720A

shows that three segments are missing from the

left first leg.

ACB720C; Slide bearing a label in Womersley's writ-

ing “Chyzeria australiense Hirst/Palp/ in nest

of Ponera lutea / Swan Riv. W.A./S.A.M. coll.”

and has also the word “TYPE” written on it in

red ink, in Womersley’s writing. This palp

appears to have been the source of Womersley's

(1934) Fig. 2.

There is no doubt that specimen ACB719 is the

Holotype of Chpzeria ausiraliensis Hirst, from the

red slide notation of “Type”, in a writing that does

not appear to have been Womersley’s, and is

presumably Hirst’s. I have attached an additional

label to the slide, identifying it as ACB719, and with

the word “TYPE”, This specimen was not figured by

Hirst, but was apparently the source of Womersley’s

(1934) Fig, |, (The specimen has the palpi flexed,

and not extended as figured by Womersley. Other-

wise, however, there appears to be a general agree-

ment with the figure.)

The three slides | have labelled ACB720 appear to

make up a single specimen, and | have labelled these

ACB720A, B and C with additional slide labels, and

have added to each the word “PARATYPE”. I regard

Womersley’s use of the word “TYPE” on slide

ACB720C as either frankly erroneous, or as a loose

indication that the specimen belonged to the lype

series. | do not know whether there were originally

two slides and a further mount was made by

Womersley, or whether there were originally three,

the label of one being discarded by Womersley in

1933-4.

SOME FURTHER DATA AND COMPARISONS FOR

ADULTS OF Chyzeria hirsti WOMERSLEY AND Chyzeria

australiensis HIRST.

Table 5 gives various measurements for the holo-

types of Chyzeria hirsti Womersley and Chyzerla

australiensis Hirst, as well as for some other speci-

mens of gach of these species.

302

Table 5.

REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

Measurements (um) for some adults of Chyzeria hirsti Wom. and C. australiensis Hirst, including lengths of nominated leg

segments and their ratios, also the lengths of the posterior dorsal swordlike setae.

Chyzeria hirsti

Chyzeria australiensis

Holotype ? Paratype Holotype Paratype ACB721A ACB721B

Specimen ACB718 ACB677 ACB13/14A ACB13/14B ACB719 ACB720 Nyabing, Nyabing,

Woodside, Willunga, 2 Glen Os., Glen Os., Swan R., Swan R., W.A. 2.8.45 W.A. 2.8.45

S.A, S.A. S. Aust. S. Aust W.A. W.A. P.N.F. P.N.F.

Length Ratio Length Ratio Length Ratio Length Ratio|Length Ratio Length Ratio Length Ratio Length Ratio

Segment

Ta | 605 1.00 583 1.00 572 1.00 562 1.00] 482 1.00 482 1.00 580 1.00 576 1.00

Ti | 382 63 367 63 396 69 396 10 | 360 15 396 82 ©6418 712 396 69

Ge I 331 55 324 56 346 60 342 61 310 64 313 65 335 58 353 61

Ta Il 374 1.00 346 1.00 374 1.00 346 1.00} 346 1.00 342 1.00 382 1.00 367 1.00

Ti Il 295 79 259 75 288 17 295 85 | 288 83 288 84 335 88 324 88

Ta Ill 403 1.00 346 1.00 360 1.00 367 100} 353 1.00 363 1.00 382 1.00 385 1.00

Ti Ill 317 719 288 83 0 317 88 317 86 | 328 93 302 83 367 96 382 99

Ta IV 446 1.00 410 1.00 446 1.00 —* — 389 1.00 389 1.00 432 1.00 439 1.00

Ti IV 468 1.05 400 98 482 1.08 ca480* — 482 1.24 493 1.27 562 1.30 522 = 1.19

Posterior dorsal g9 994 88-172 120-195 144-172 86-135 80-143 72-152 86-138

swordlike setae

* Somewhat obscured in the mount, and cannot be measured accurately

Perusal of Table 5 suggests that some of the dimen- ASBa 44 143.0(!) 42

sions could be useful in separating the two species PSB 44 20.0(!) 42

australiensis and hirsti. Of them, the range of the L 88 163.0(!) 84

lengths of the posterior dorsal swordlike setae is the W 167 — —

most useful as the ranges differ considerably, with A-P 52, 52 55.0 50

their maxima ranging from 172-204 um for hirstiand AL 112 112.0 110

135-152 um for australiensis. PL 83 84.0 88

Sens cal80 168.0 162

: ; DS 103-173 163 102-174

Chyzeria onychia (Womersle F

if penta ) mid-DS 110-138 = —

(Figs. 5, 6, 7A-B, 8A-G, 9) PDS 112-138 — —

Synonymy

Grossia onychia Womersley, 1954 (1954a) p. 111;

Audy, 1954, p. 166; Audy 1957 p. 442; Vercam-

men-Grandjean 1972 p. 231.

Redescription of Holotype

Colour in life not recorded by collector G. F. Gross

or by Womersley. Slide-mounted specimen (it has not

been remounted by either Vercammen-Grandjean or

myself) has the idiosoma ca 1750 wm long by ca 1600

um wide; total length of animal from tip of chelicerae

to posterior pole of idiosoma ca 1900 um. (The spec-

imen has been severely compressed on the slide and

ruptured on mounting, hence these measurements are

likely to be significantly larger than the measure-

ments in life.)

The standard (and some other) data of the Holo-

type are as follows (and compared with data pub-

lished by Womersley and Vercammen-Grandjean):

This Womersley Vercammen-

paper (1954) Grandjean

(1972)

AW 96 100.0 97

PW 151 156.0 149

SB 67 73.0 65

Scutalae tapering, somewhat sinuous in the mount,

slightly blunted at tip, with weak, barbed, almost

adpressed ciliations.

Eyes 2 + 2, sessile, conjoined, the eye patch placed

well laterally to the dorsal shield, eyes of about the

same size, anterior more medial, circular, 16 wm

across, posterior oval, 15 um X 17 um.

Dorsal idiosomalae similar to scutalae, tapering,

slightly blunted at tip, with weak, barbed cilia; cilia

only a little outstanding from shaft. Dorsal idioso-

malae have their seta bases expanded to small plates

(see Fig. 8C, D).

Venter: no setae between coxae I, II and III. Behind

coxae III are a small number of opisthosomal ventra-

lae, arranged as figured. (The pattern that exists is

not clear on account of damage that has been done

to the specimen in mounting.) The setae are tapering,

pointed, somewhat (more or less adpressedly) ciliated,

40-77 wm long, the more posterior setae the longer

(these remarks excluding the most posterior setae of

the mount, whose dorsal:ventral relationship cannot

be determined from the mount).

Anal region: anus consists of two hinged valves or

plates, each 61 wm long by 20 um wide, hinged ante-

500

yim

ee)

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 303

Q \

\_ p's

Sf Sa

S36)

FIG. 5. Chyzeria onychia (Womersley). Holotype larva, dorsal

view, to scale on left. The specimen has been distorted and

ruptured by the mounting process. The anus and surrounding

setae cannot be clearly identified as dorsal or ventral in the

mount, hence these are repeated in Fig. 6.

a4 REC,

riarly and posteriorly to make a structure 44 ym

across in the mount, Each anal plate carries two stiff

normal setae, almost straight, tapering, slightly

blunted, anterior seta 40 um long, posterior 64 um.

Surrounding the anus is a group of setae similar to

last, pointed or slightly blunted. These opisthosomal

setae arise from small expanded plates. (The setation

over the posterior end of the idiosoma is contiguous

between the dorsum and venter, but in the mount its

exact posilion on the idiosoma cannot be determined).

Coxalae 2, 1, 1. Coxulae robust, lapering, some-

what spindle-shaped, curved, with a few adpressed

cilia, All coxalae appear to be similar, but in the

Speeinien only the left coxala HI] is intact, the others

being apparently broken, or missing. Coxala LI

38 wi long, terminally tapering to a fine point,

Legs as figured; | 560 xm long, 11 565 wm long, TI

645 pm (from medial point of each coxa to up of

claws). Fenvoral to tibial segments roughly cylindri-

cal, Tarsus | 129m long by 48 um high, tibia |

92 um long, genu | 80 wm (Til/Gel = 1.15). Tarsus

(ll 129 pm long by 34 wm Ingh, tibia ILE 109 pm

long, genu WT 85 pm (Till /Gelll = 1.29), (Tarsal

meisurements exclude claws and pedicle.)

Genu | has specialized seta VsGel.74pd. Tibia |

has specialized setae SoTil-27d, VsTil. 91pd,

SoTil.92d, Genu I has specialized seta VsGell.71 pd.

Tibia Il has specialized setae SoTill.23d and

SoTL s3pd. Tibia lll has specialized seta

SoTill.18d.

Tarsal setae as figured. Tarsal pedicles appear nor-

mul, Each tarsus is provided with three claws, The

middle (neomedian) claw is normal, strong, falciform.

Anterior claw reduced to a slender tapering sinuous

rod, with a spreading terminal adhesive membrane.

Posterior claw is somewhat more robust than the

anterior, and straighter, it terminates in a smaller

adhesive sucker Sel an angle to the preceding claw-

shaft.

Gnathosoma short, robust. Chelae bases 96 um long

by 48 wm across individually, the two combined being

96 um across. Chelie blades short, projecting 32 um

beyond the end of the chelae bases. Chelae bases

more or less pyriform. Blade with small scattered

retrorse teeth on all surfaces, but stronger laterally.

Galeala not identified. Anterior hypostomala taper-

ing, pointed with a few adpressed cilia, 45 um long.

Posterior hypostomala similar, 57 wm long,

Palpal formula 0, 1, 1, 3,9. Palpal femorala robust,

lapering, with strong barbs, 74 um long. Palpal gen-

uala strong, tapering, pointed, terminally filiform,

with barbed cilia, 70 um tong. Palpal tibial claw tri-

fid, the three tines tapering, strong, divergent, slightly

blunted. Additionally the palpal tibia carries a strong

S, AUST, MUS, 1H (14), 245-326

June, 1982

sickle-shaped accessory seta, with a tapering hook-

like end and a small tooth about the middle of its

ventral side; this possibly operates in apposilion ta

the tibial claw. Palpal tarsus as figured. Palpal supea-

coxala present, a slender conical seta 9 um long.

Locality

The specimen is known only from the Holotype,

The collector Dr G. F. Gross, has advised me (pers,

comm., 4 Augusi 1975) that he collected it at Tam-

bryn Junction, Otway Ranges, Victoria in January

1951, The specimen, which was parasilic upon an

adult Chpzerta, was probably brought back alive to

Womersley.

No Chpzerja adult is among the South Australian

Museum slide or spirit collection, with locality data

for the Otway Ranges, Victoria.

Remarks on Biology

Observations upon a phoretic relationship between

Chyzeria hirsti and its larvae have been recorded

earlier in this paper. When C, Ajrsti larvae were

reared in the laboratory, all larvae were small and

clearly unfed. In the case of C. onychia , the larva

is considerably enlarged, and in fact, judging by other

experiences of the author with trombidiid larvae, it

appears to be al a stage not far away from pupation,

even though recorded as attached parasitically, Lt

may seem remarkable that a larva of Chyzeria should

parasitize an adult of the same genus (and conceiv-

ably, of the same species), but among the family

Erythraeidae, in some species the larvae will parasi-

lize Opportunistically a wide range of hosts including

adult erythraeid and anystid mites (see Southcott,

1946 pp. 39, 41)..

Remarks on the Generic Status of Chyzeria onpchia

(Wom.)

There does not appear io be any good reason to

separate C. onvehia generically from the other mem-

bers of the genus Chyzeria (the whole of the known

larvae) considered in this paper, The body setae are

similar to those of other members referred to the

genus, and in fact are less aberrant than, e.g., the AL

seutalae of C, hirst/. The pedocoxalae are in fact the

Most normal in structure for any of the larvae con-

sidered in this paper. The tarsal claws of (he legs are

also quite similar to those of the other members of

the genus. Womersley's erection of Grossia appears

10 have been simply a consequence of his overlooking

his earlier (1939) paper, as remarked above.

Chyzevria flindersi sp. nov.

Fivs. 1OA, B: 1TA-B; 12

Description of Larva (from the Holotype specimen

ACB674)

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 305

ne an

FIG. 6. Chyzeria onychia (Womersley). Holotype larva, ventral

view, to scale on left. Specimen distorted and ruptured (see

also legend to Fig. 5).

306

REC. S. AUST. MUS., 18 (14): 285-326 June, 1982

FIG. 7. Chyzeria onychia (Womersley). Holotype larva. A, Gna-

thosoma, dorsal view. B, Dorsal idiosomal scutum. (The right

AL scutala has been restored in the figure to its correct

position; its position on the slide is shown in outline.) (Both to

scale shown).

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 307

100

FIG. 8. Chyzeria onychia (Womersley). Holotype larva. A, Gna-

thosoma, ventral view; the chelicera shown on the left is

damaged. B, Left eyes and right eyes. C, D, Dorsal idiosomal

setae. E, F, Ventral idiosomal setae. G, Anus and surrounding

setae. (All to scale shown,).

308 REC. S. AUST. MUS., 18 (14): 285-326 June, 1982

p Cy,

wt .

gy a

FIG. 9. Chyzeria onychia (Womersley). Holotype larva. Legs I,

II and III, to scale shown; to standard symbols, (Note: all legs

have been dislocated at the trochantero-femoral joints.). Pedo-

coxala III is also shown, further enlarged.

Colour in life red. Length of idiosoma (mounted on

slide) 410um, width 330 wm; total length of animal

from tip of chelicerae to posterior pole of idiosoma

527 wm.

Dorsal scutum as figured; much wider than long,

more or less oblong or trapezoidal, with concave

anterior and posterior margins, and irregularly convex

lateral margins. One pair of scutal sensilla, well sep-

arated, placed about halfway between the front and

back edges of scutum. AL scutalae at AL angles of

shield, PL somewhat anterior to PL angles.

The standard (and some other) data of the Holo-

type are (um):

139

184

A-P aa

AL 55

SB 93 PL 52

ASBa 57 Sens 96

ASBp 32 DS 52-77

L 89 Mid-DS 67-71

WwW 212 PDS 77

Scutalae normal, tapering, ciliated, slightly blunted

terminally. Scutal sensillary filaments slender, with

a few sparse cilia in distal half.

Eyes 2 + 2, sessile, anterior about 21 wm across,

lateral to the posterior part of the lateral edge of

scutum; posterior eye close behind anterior eye,

17 wm across.

Dorsal idiosomalae similar to scutalae, but more

slender, arranged in rows across dorsum 6, 6, 6, 4, 2,

2; total 26 setae. All dorsal idiosomalae arise from

expanded plates.

100

200

yum

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 309

FIG. 10. Chyzeria flindersi sp. nov. Holotype larva. A, Dorsal

view, to scale on left; legs omitted beyond trochanters. B,

Dorsal idiosomal seta, further enlarged.

310

Venter: no setae between coxae I and II. Between

coxae III a pair of setae, with enlarged basal quarter

of scobillum, and from this enlarged basal part the

distal part arises abruptly, to taper to a smooth point;

setae 43, 47 um long, lacking cilia. Remaining ven-

tralae arise well behind coxae III, about 39 in num-

ber, and are 30-54 um long; they increase in length

posteriad. The more anterior setae are similar to the

pair between coxae III, except that they carry cilia,

these becoming stronger and more outstanding pos-

teriad. The most posterior ventralae tend to resemble

the adjoining dorsal setae.

Anal region not clearly visible in Holotype, but

appears normal. Anus about 47 um long.

Coxalae 2, 1, 1. All coxalae stout, blunted, with a

terminal slight grooving which makes them appear

like small plums projecting from the coxae, 15-17 um

long. Careful microscopy at high power shows an

appearance of distal tuberosities (see Fig. 11C),

Legs normal: I 700 um long, II 655 um, HI 735 um

(all lengths include coxae and claws). Femoral to

tibial segments more or less cylindrical. Femora

divided. Scobalae normal. Trochanteralae 1, 1, 1,

basifemoralae | (2), 2 (3), 1, telofemoralae 5, 4, 4.

Tarsus I 174 wm long by 50 um high; tibia I 108 wm

long, genu I 94 uv long (Til/Gel = 1.15). Tarsus III

176 wm long by 40 um across; tibia HI 119um long

genu IIT 94 wm long (TilI[/Gell] = 1.27). (All

tarsal lengths without claws and pedicle.)

Genu I has specialized seta VsGel.74pd. Tibia |

has specialized setae SoTil.23d, VsTil.88pd,

SoTil.92d (and hence the distal SoTil is distal to the

VsTil). Genu II has specialized seta VsGell.79pd.

Tibia Il has specialized setae SoTill.23pd and

SoTill.88pd. Tibia III has specialized seta

SoTil11.26d.

Tarsal setae as figured. Tarsal pedicles short.

Pedotarsal claws: anterior curved, slender, two thirds

as long as middle claw, with a terminal flattened

piece which is more or less at right angles to the

claw; middle (neomedian) claw falciform, strong,

unciliated; posterior claw curved, slender, shorter

than anterior claw, and with a similar flattened trans-

verse piece at the tip (see Fig. 12).

Gnathosoma robust, normal, pear-shaped, chelicera

122 um long to tip of cheliceral blade, by 108 um

across. Cheliceral blades strong, falciform, 44 um

long. In the Holotype (the only specimen available)

the left blade is quite smooth, while the right blade

bears ventrally a number of small retrorse adnate

teeth; the concave upper surface of the right fang has

7 small teeth, the left fang, none. Galeala spiniform,

curved, 33 um long. Anterior hypostomala short, peg-

like, 7 um long. Posterior hypostomala slender, ca

69 um long, unciliated.

REC. 8S. AUST, MUS., 18 (14): 285-326

June, 1982

Palpal formula 0, 1, 1, 3, 9. Palpal femorala taper-

ing, slightly ciliated, very slightly blunted, 52 um

long. Palpal genuala slender, unciliated, 71 um long.

Palpal tibial claw (odontus) with three strong, slightly

divergent prongs. Palpal tarsalae as figured. Palpal

supracoxala a blunted peg, 8 um long.

Locality

Tasmania: Lake St. Clair, 28 December 1945, R.

V. Southcott. One larva, ACB674, Holotype, running

up trunk of Eucalyptus sp., at about 1.5m above

ground. The site was in fairly open cleared ground,

near the shore of the Lake. Type to be deposited in

the South Australian Museum.

Nomenclature

This species is dedicated to Matthew Flinders, R.

N., 1774-1814, who was the first to survey the major

part of the Tasmanian coastline.

Remarks on Asymmetry in the Chelicerae

In the above description an asymmetry in structure

of the cheliceral fangs was recorded, one fang (the

presumably normal one) being denticulate, and the

other smooth. The present author is unaware of a

similar asymmetry of the mouthparts having been

recorded previously in at least the Trombidioidea, or

Erythraeoidea. Among the Erythraeoidea differences

in larval shield setation occur at times; for example,

see the discussion on this under the account of Char-

letonia venus Southcott in Southcott (1966, p. 709)

and for Charletonia feideri Southcott (see ibid., Fig.

32, p. 756, and the cases recorded under “Material

examined” on pp. 754-8). However, as these setae are

presumably responsible for only the sensory input,

and some degree of protection, minor abnormalities

are unlikely to be a major significance. Major struc-

tural abnormalities affecting e.g. locomotory capacity

or feeding ability, are more likely to be such a hand-

icap as to threaten survival, and hence are less likely

to be encountered in field-collected material.

The specimen was apparently unfed, and whether

the asymmetry recorded would be a handicap to

feeding is a matter of speculation.

Even among better known animals, major asym-

metry in external structures is comparitively rare.

Possibly gynandromorphy in insects and _ hetero-

chromia in mammals are comparable examples. In

some groups, notably the fiddler crabs, asymmetry

has become part of the genetic inheritance of the

group, and is either sex-linked, or hormonally related.

Chyzeria derricki sp. nov.

(Figs. 13A-H, 14A-B, 15)

Description of Larva (from the Holotype Larva

ACB633A)

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 311

VIG. 11 Chyzerfa flindersi sp. nov, Holotype larva. A, Ventral

view, to scale on left; legs omitted beyond trochanters. B,

Ventral opisthosomal seta, further enlarged, C, Pedocoxal seta

(x in A), further enlarged. D, Palp tarsus, further enlarged.

REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

FIG. 12 Chyzeria flindersi sp. nov, Holotype larva. Legs 1, 1] and

LIT, to scale on left; to standard symbols,

Colour in life not recorded, presumably red. Length

of idiosoma (mounted on slide) 1055 um, width 610

um. Idiosoma rather elongate, slightly constricted

behind level of coxae II, and somewhat pointed at

posterior pole. Total length of animal from tip of

galeal seta to posterior pole of idiosoma 1165um

(chelicerae damaged in Holotype, and fangs missing;

Paratype ACB 633B lacks gnathosoma, etc.).

Dorsal scutum more or less trapezoidal, with

rounded angles, wider posteriorly; anterior and pos-

terior edges slightly sinuous (scutum tends to project

a little in the middle posteriorly). Scutum is lightly

punctate, particularly in its anterior and more central

part, in the area between the sensilla and the AL

scutalae. Scutal sensilla normal, the setae with slight,

sparse, terminal cilia. Scutalae tapering, pointed, slen-

der, lightly blunted at tip, with adpressed cilia.

The standard (and some other) data of the Holo-

type (A) and the Paratype (B) are:

A B A B

AW 102 99 A-P 56,54 53,57

PW 152 152 AL 88 (7+) 107

SB 61 61 PL 88 94,89

ASB 55 55 Sens 153 -—

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 313

500 : ge N PAG IO TT 4

um . ; ?

eh,

100

FIG. 13. Chyzeria derricki sp. nov. Holotype larva. A, Dorsal

aspect, legs omitted beyond trochanters. B, Ventral aspect,

legs omitted beyond trochanters. C, D, E, Dorsal idiosomal

setae: C, anterior dorsal idiosomala; D, mid-dorsal idiosomala;

E, posterior dorsal idiosomala. F, G, ventral idiosomal setae:

F, anterior opisthosomala, G more posterior ventral opistho-

somala, H, Anal region of venter. (A, B to top scale; C-H to

bottom scale.)

314

A B A B

PSB 37 39 DS 85-120 74-120

I, 92 94 ADS 83-118 77-114

W 178 180 MDS 93-98 74-96

PDS 85-120 79-120

PVS 96-114 42-744

Eyes 2 + 2, sessile, subequal, circular, conjoined,

antero-posterior in orientation, lenses 26 um across.

Dorsal idiosomalae long, tapering, curved, with

adpressed cilia, and slightly blunted at the tip;

arranged 6, 6, 6, 6, 4, 2. Each seta arises, usually

eccentrically, from a small plate, suboval, an expan-

sion of a normal alveolar annulus.

Venter: No setae between coxae I or coxae II,

Somewhat anterior to level of coxae III is a pair of

tapering, pointed, simple setae, 57 wm long. Behind

coxae III is a patch of about 40 similar setae which

arise from small plates that increase gradually in size

posteriad, with the more posterior setae the longer.

The chitinized shaft (scobillum) of the seta also

becomes thicker at its proximal end, as one proceeds

posteriorad. Anus externally consists of two hinged

valves or plates, with a longitudinal opening, the

conjoined two making an oval structure 55 um long

by 49 um wide. Each valve has two setae, tapering,

slightly blunted, with adpressed cilia; anterior 48 wm

long, posterior 74 um.

Coxalae 2, 2, 1. All coxalae short, blunted, smooth

except for faint distal tubercles (see Figs. 14B, 15).

coxalae 15-20 um long.

Legs comparatively long and thin for a trombidiid

larva, compared with body size; tarsi long and ter-

minally attenuated. Leg lengths: I 705 um, II 655 um,

III 795 um (all lengths include coxae and claws).

Femora divided. Femoral to tibial segments more or

less cylindrical. Scobalae of legs tapering, with

barbed cilia, and are terminally blunted. Scobalae of

telofemora and tibiae tend to be long and particularly

outstanding.

Tarsus I 224ym long by 36um high. Tibia |

118 wm long, genu I 112 um long (Til/Gel = 1.05).

Tarsus III] 210 wm long by 24 um high (at proximal

end). Tibia III 141 um long, genu III 106 um long

(TillI/Gelll = 1.33). (All tarsal lengths exclude

claws and pedicle.)

Genu | with specialized seta VsGel.82pd. Tibia I

with specialized setae SoTil.10d, SoTil.91d and

VsTil.92pd. Genu II with specialized seta

VsGell.81pd. Tibia II with specialized setae

SoTill.20d, SoTil[.90d. Genu II] without specialized

setae. Tibia II] with SoTilll.16d.

Tarsal setae as figured. Pedotarsal claws three in

number: anterior slender, falciform with ill-defined

REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

terminal spreading suction pad; middle claw falci-

form, over-reaching others; posterior slender, sinuous,

with obliquely placed terminal suction pad.

Gnathosoma short, robust. Conjoined cheliceral

bases form a squat pyriform mass 110 um long (to

anteriormost chitinization, except setae) by 105 um

wide. Cheliceral blades missing in Type and Paratype.

Galeala normal, pointed, tapering with filiform tip,

32 wm long. Anterior hypostomala not available. Pos-

terior hypostomala long, fine, pointed, simple, about

100 wm long. Palpal setal formula 0, 1, 1, 3, 29.

Palpal femorala tapering, terminally blunted, with

outstanding sparse cilia, 123 um long. Palpal genuala

tapering, pointed, unciliated, 96 wm long. Palpal tibi-

alae tapering, pointed. Palpal tibial claw trifid, the

three curved tines strongly divergent (see Fig. 14B).

Palpal tarsalae as figured, including one relatively

huge scythe-like seta, which reaches as far back as

the bases of the posterior hypostomalae. Palpal supra-

coxala short, narrow, pointed, 5 um long.

Locality

Queensland: Mt Tamborine, 23 February 1954, E.

H. Derrick, two specimens taken parasitic upon uni-

dentified tettigoniid grasshopper (identification A294).

Holotype ACB633A, Paratype ACB633B (two slides

B1 and B2). Specimens forwarded by Dr R. Domrow,

Queensland Institute of Medical Research. Type and

Paratype to be deposited in South Australian Museum

collection. The Paratype has been badly damaged in

mounting, and lacks gnathosoma.

Nomenclature

The species is dedicated to the collector, Edward

Holbrook Derrick, 1898-1976, medical research worker

(see obituary notice in Medical Journal of Australia

6 November 1976).

Remarks on Biology of Chyzeria derricki

Parasitization of an insect host is not unusual for

a trombidioid larval mite. The only other record of

parasitization by a larval Chyzeria upon a host is in

the case of C. onychia (q. v., discussion). The capture

of C. derricki larva upon a tettigoniid host allows

some degree of speculation upon the functional sig-

nificance of the various morphological adaptations

seen in this genus. It is reasonable to suggest, in the

case of mites which live as ectoparasites upon mobile

insects with wide areas of heavy, smooth chitinization

(e.g. Acridoidea and Tettigonioidea), that long legs

and development of suction pads upon the tarsi would

be useful, as would also strong mouthparts to pierce

thickened integuments. On the other hand, it is appar-

ent from a study of at least the erythraeoid mites,

that there may be a considerable degree of host

specificity, e.g. Smaris prominens (Banks) larvae upon

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA

FIG, 14. Chyzeria derricki sp. nov. Holotype larva, A, Gnathosoma

and adjacent part of idiosoma, and trochanters I, dorsal view.

B, Gnathosoma, adjacent idiosoma, with coxae I and part of

trochanters I, ventral view. Both to scale shown. In B, a

pedocoxala is shown further enlarged.

316 REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

FIG. 15. Chyzeria derricki sp. nov. Holotype larva. Legs I, II and

III, to standard symbols; to scale shown.

small Psocoptera and Erythrites osmondensis (South-

cott) larvae upon Thysanoptera. There may also be

site specificity upon a host, e.g. Callidosoma wom-

ersleyi (Southcott) preferring a site under the wings

of psyllids, as does also the larva of Rainbowia imper-

ator (Hirst, 1928) (Southcott 1961a, p. 477). On the

other hand the larvae of other erythraeid mites, Ery-

thrites reginae (Hirst, 1928) and Erythrites urrbrae

(Womersley, 1934), utilize only the external surfaces

of the thorax and abdomen and not the under-wing

surfaces when parasitizing psyllids and other small

insects.

With regard to those members of the Trombidioidea

whose larvae utilize insect hosts, in general, there are

less systematic data on the range of hosts and the

OBSERVATIONS ON CHYZERIA GANUSTRINI AND SOME RELATED GENERA 417

sites Of parasitic attachment than in the case of the

Erythraeoidea. This means that at present it is not

possible to offer any detailed discussion of the mor-

phological and functional adaptations of these larvae

and their adults. This is particularly so where there

is some unusual structural modification, such as, for

instance, the anterior scutalae of the larva of CAyzeria

Airsti, or the hind tarsi in the larvae of Ettorvelleria

Oudemans, 1911 and Eutrombidium Verdun, 1909,

CHARACTERISTICS OF LARVAL TROMBEL-

LIDAE

Earlier an attempt at a formal definition for larval

Trombellidae was deferred. With the redescription or

description of four species of Chyvzeria in the present

paper plus the rearing of Audyana Womersley 1954,

and with the knowledge that 7rombella Berlese 1887

and Womers/eyia Radford 1946 are synonyms, it is

proposed to examine what characteristics separate

them and the genera Nothotrombicula Dumbleton,

1947 and Ralphaudyna Vercammen-Grandjean et al,,

1974, based on larvac, and also presumed to belong

to the Trombellidae, from other trombidioid (or

related) mites.

This group of larval trombidioid miles has ihe

following characteristics

(1) coxal formula 2, |, |

(2) leg segmental formula;

fa) 7,7, 7 Chyzeria; Nothotronibicula;

Ralphaudyna

(b) 7, 6, 6 Audpana

(3) AM scutalae:

(a) absent Chyzeria; Nothotrombicula

(b) present as such Tronibella; Audvana

(4) sternal formula 0, 0, 2

(5) vestigiotibialae |, 0, 0

(6) vestigiogenualae |, 1, 0

(7) solenotibialae:

fa) 0, 0, O Chyzeria; or

(b) 2, 2, 2 Audyana; or

($) solenogenualac:

(a) 0, O. O Chyzeria: or

(b) 11,1 Trombella, Nothratrambicula; ot

(ce) 2, 1, | Audyana; or

(d) 2, 1, 2 Ralphaudyna

(9) trucheae absent

* Vercammen-Grandjean et al. (1974, p. 246)

(10) pedotarsal claws:

(a) 1,1, 2 Trombella; or

(b) 2, 2, 2 Audyana; or

(ce) 3, 3, 3 Chyzeria; Nothotromhicula;

Ralphaudyna

(11) palpal tibial claw:

(a) trifid Chyzeria,; Nothotrombicula; or

(hb) bifid Ralphaudyna; Trombella; or

(12) idiosoma seta-bases:

(a) expanded Chyzeria; Ralphaudyna; or

(b) normal Tromhella; Norhotrombiteula:

Audyana

(13) seutum “nasus’;

(a) absent Chyzeria; Audyana; or

(bh) present Trombella; Nothotrombicula;

Ralphaudyna

Taking. these characteristics in order:

(1) This is a characteristic of Leeuwenhoekia,

Odontacarus, and Neotrombidium (Leeuwenhoeki-

idac); it Lhus cannot be taken as a definitive character

of a trombellid larva, but includes also the leeuwen-

hoekiids,

(2) Subcharacter 2 /a) is possessed by many trom-

biculid genera, together with Apolonia, Womersia,

and in fact members of the Erythracoidea and

Hydrachnellae have this characteristic of all femora

divided.

Subcharacter 2 (), of having only the first femur

divided is possessed by other trombiculids, e.g, Wa/-

chia (Walchiinac), also Neotrambidium.

Subcharacter 2 (c), of having all femora undivided.

is possessed by the genera Leeuwenhoekia, Odonta-

carus, Monunguis and various others of the Leeu-

wenhockiidae; also by Lassenia (Johnstonianidae) (see

Table 1).

Clearly this character is of no great value al the

family classification level.

(3) This is another characteristic which may be

present or absent, or present in some degree, and the

sume occurs with other trombidioids (see Table 1).

Similar remarks apply.

(4) This appears to be a constant characteristic. It

is also a characteristic of Leewwenhoekla, Odentu-

earus and Neotrombidium (Lecuwenhoekiidae),

(5) Although this character is present uniformly

among the genera, it also applies widely among

related mites. e.g. Leplotrombidium, Babiangiu

(Trombiculidae), Leeuwenhoekia and Neotrombi-

dium (Leeuwenhoekiidae), Preridopus, etc.

It is thus not of much diagnostic value,

(6) This character is also uniform, but occurs also

318

e.g. in Leeuwenhoekia and Neotrombidium (Leeu-

wenhoekiidae), and also in Pteridopus.

It also is not of much diagnostic value.

(7) Character 7 (a) is shared by Leptotrombidium,

Babiangia, Neotrombidium, for example. Character

7 (c) is shared by Pteridopus, Polydiscia.

It is difficult to see how such a widespread and

variable character can be considered of much diag-

nostic use.

(8) It is clear also that character (8), similarly

widespread and variable, can be considered of little

diagnostic usefulness.

(9) This character is uniform, but also applies to

the majority of the trombidioid mites; see Table 1.

(10) It is clear from inspection that the number of

pedotarsal claws is in no way diagnostic of the trom-

bellids.

It is apparent also that characters (11), (12) and

(13) cannot be used to separate trombellids from

other trombidioids.

In a proposal to separate the Trombellinae from

the Neotrombidiinae Vercammen-Grandjean (1972,

p. 239) proposed the following supposedly differen-

tiating characters for the Trombellinae:

1, a single but trifurcate claw on each anterior and

mid leg, two claws on posterior leg tarsus.

2. no microspurs [=vestigialae here] on anterior

and mid genu.

3. round urstigma between contiguous anterior and

mic [sic, for mid] coxae.

4. uniform porosity on coxae [as against a mosaic

pattern over the lateral part of some of the

coxae]’.

In 1973 (1973b, p. 109) he used the term Trom-

bellidae, ascribed to Feider (1955), although the term

as such appears to have been introduced by Vercam-

men-Grandjean there. As no formal redefinition

appears to have been proposed, it appears reasonable

to accept that the characters listed by Vercammen-

Grandjean (1972, p. 239) remained as his concept of

the differentiating characters of the family Trombel-

lidae, for the larvae. Larval genera accepted by Ver-

cammen-Grandjean (l.c.) as belonging to the Trom-

bellinae were Womersleyia and Audyana.

If we compare the characters listed above with

those discussed in the preceding text of the present

article, we find, with respect to character (1) of

Vercammen-Grandjean, that with the addition of

Chyzeria alone to the list the pedotarsal claw pattern

may be 1, 1, 2 or 2, 2, 2 or 3, 3, 3 (see my character

(10) above).

With respect to Vercammen-Grandjean’s character

REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

(2), of the possession of a vestigiogenual formula of

0, 0, [0], the formula recorded here is of 1, 1, 0 for

all the genera of the Trombellidae considered here,

which is the same as the formula for the Neotrom-

bidiinae proposed by Vercammen-Grandjean (/.c.),

also for Nothotrombicula.

With respect to character (3) of Vercammen-

Grandjean, above, unless some quantitative measure

of ellipticity is to be applied it is difficult to see how

such a character can be used. The problem here is

not to differentiate the trombellids from the genus

Neotrombidum and its near relations, but to find one

or more criteria which will separate the trombellids

from the other trombidioids, at least in the first

instance.

For character (4), the presence of a mosaic pattern

in the outer parts of coxae I and III in Neotrombidium

and related genera, this appears to be confined to

that group, and is not seen elsewhere among the

Trombidioidea.

It would appear therefore that an examination of

the available data has not yielded criteria which will

allow separation of a larval trombellid with any cer-

tainty, although in some cases, where no rearing cor-

relation has been achieved, e.g. for the genera

Nothotrombicula and Ralphaudyna, the resemblances

to known larval trombellids are so great that they

may be allotted to this family with confidence. At

present therefore it is probably justifiable to retain a

family Trombellidae, but this has to be based on the

adult definition only, and no formal definition of

larval Trombellidae can be proposed with any relia-

bility.

This amounts to confirming the view of Robaux

(1977b) that, in order to work out the phylogenetic

relationships of the Trombidioidea, one cannot rely

solely on one aspect of the morphology of these mites,

such as that of the larvae or of the adults. In fact

Robaux has proposed that biological and ecological

factors must also be considered.

DESCRIPTION OF A PYGMEPHORID MITE PHORETIC

UPON AN ADULT Chyzeria.

We now proceed to a description of a species of

mite, of the family Pygmephoridae, recorded above

as phoretic upon an adult Chyzeria hirsti Wom.

Bakerdania workandae sp. nov.

(Figs. 16, 17, 18)

Description of Non-Gravid Adult Female, Holotype

ACC226, Mounted.

Colour in life light brown. Length of idiosoma

260um; length of gnathosoma from tip of chelicerae

to posterior superior margin of the cheliceral bases

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA ay

FLG. 16. Bakerdania workandae sp. nov. Holotype female, A,

Dorsal view. B, Dorsal view left leg II (setae of tarsus II in-

complete in B, from obscurity of mount), A, B to scale shown.

28um; total length of mite from tip of chelicerae to

posterior pole of idiosoma 288um. Width of idiosoma

at level of legs IIL 170um.

Gnathosoma small, free, subquadrate, 33um wide.

li carries 4 simple spiniform setae dorsally on the

cheliceral bases, lateral 23um long, originate somewhat

posterior to medial, which are 20um long. Ventrally

the gnathosoma carries a short spiniform seta (palpal

coxala) 144m long, originating immediately behind

the proximal free palpal segment. Both dorsally and

ventrally a midline chitinous raphe separates the twa

gnathobases, Chelicerae small, the stylets not

extruded, but confined in the specimen in a small

conical cap.

320

Palpi of two mobile segments. Proximal segment

(femur-genu) with a dorsolateral spiniform seta, 11m

long, and distal segment (tibiotarsus) with a similar

dorsolateral spiniform seta, 15um long. Palpal tibi-

otarsus bears distally a strong broad-tipped tooth,

with three or four weakly defined cusps, the lateral-

most projecting farthest. Distal segment bears ven-

trally two solenoidalae, expanded, projecting somewhat

anterolaterally. The more posterolateral (solenoidala

“1"), about 4 um long by lum wide, is thinner,

somewhat clavate; the more medial (solenoidala ‘*2’’)

is larger, 8um long by 3um wide, approximately

thumb-shaped with an excavated base.

Propodosoma dorsally trapezoidal, widest poste-

riorly, and with the anterolateral angles rounded;

about half as long as wide. Laterally, at about the

middle level on each side, is a large circular peritremal

opening, 9um wide, with the centres 37um apart.

Behind and lateral to each peritremal opening is the

short anterolateral propodosomal setae, spiniform, 7um

long. Behind and slightly lateral to this seta is the

larger posterior propodosomal seta, tapering, pointed,

lightly barbed, 29um long. Behind this, on each side,

is a clavate sensillum, 25um long, by 7um wide at its

thickest part.

Ventral propodosomal setae consist of two more or

less transverse rows of 4 setae, total 8; these are

tapering, pointed and barbed. The anterior of these

arise from the coxal plates (‘‘ventrites”) of the first

pair of legs. The medial pair of these setae strong,

4ium long, arising at about the middle of the coxal

segment; the lateral pair 30um long, and arise from

the cusp between the two leg trochanters (“coxae”

of some authors), almost directly below the alveoli of

the posterolateral dorsal propodosomal setae. These

setae appear as normal type setae, i.e. scobalae, and

they are not split or modified in any unusual way.

The second row of propodosomal ventralae are similar

to the first row; they originate laterally upon a small

triangular section of the second coxal plate, at its

anterolateral angle, and juxtaposed to the acetabular

socket of leg II; medial setae 50um long, lateral

(which is slightly posterior) 52um long (these two

setae are the internal and external ventral setae II of

authors). The margins of the circum-gnathosomal for-

amen are only moderately thickened. Apodemes II

transverse. The posterior marginal apodeme of the

propodosoma somewhat thickened, more or less linear,

running anterolaterally, the two lateral members

meeting in a blunted obtuse angle.

Hysterosoma, dorsum: setae long, strong, tapering,

pointed, with barbed ciliations. First dorsal pair of

setae level with first lateral pair of hysterosomal

setae, dorsals 53um long, laterals 78um long; dorsals

I are 91um apart. Hind margin of anteromost dorsal

plate weakly emarginate. Dorsals II 734m long, 72um

apart. Dorsals II] 91um long, 59um apart: laterals

REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

II] somewhat anterolateral to the dorsals, and 43 um

long. Posterior margin of segment III with distinct

smooth median excavation. Dorsals TV ca 90um long

(broken at tip), and centres 28um apart; laterals 1V

63um long with centres 80um apart, and a little

anterior to dorsals IV.

Hysterosoma, venter: all setae of posterior ventral

plate tapering, pointed, with weak barbed ciliations.

Medial presternal setae 414m long with centres 30um

apart, somewhat anterior to lateral presternals which

are 53um long with bases 714m apart. The latter are

a little anterior to the axillary I setae, which are 35um

long with bases 102m apart. Lateral presternals arise

well anterior to apodemes IV, and well lateral to

medial presternals. Axillary setae II (arising between

the origins of legs HI and IV) 53um long by 98um

apart. Medial poststernal setae are 44um long by

22um apart; lateral poststernal setae are 73um long

by 50um apart. Thus the lateral poststernals are the

longest of these setae; and the axillary I setae are the

shortest. Apodemes III not clearly defined in the

specimen. Apodemes IV are only slightly flexed pos-

teriorly to meet edges of acetabular sockets of legs

II]. Apodemes V not clearly defined. The posterior

margin of the posterior ventral plate is more or less

transverse, only a little sinuous. Opisthosoma ventrally

entire, with only a minute clefi indicated at posterior

pole. The most posterior opisthosomal setae (“‘cau-

dals’’) consist of a transverse row of 6 pointed, ciliated

setae: lateral pair 19um long, bases 37um apart

between centres, as usual; the other 4 setae in 2 groups

of 2, each lateral group adjoined, the lateral of these

26um long, medials 324m long, centres of bases of

each pair 3um apart, the medial of these setae with

bases 164m apart. From the posterior opisthosomal

cleft two small ‘“‘apodemes” diverge in a wishbone-

like manner, 17m long.

Legs as figured. The lengths of the non-fixed part

of the legs, to the tip of the tarsal claws, as follows:

I 106um, I] 133um, HT 152um, TV 226um.

The lengths of the mobile leg segments are as

follows (um) (excluding pedicle and claws in tarsus):

Leg ] II Il IV

trochanter* 19 25 33 59

femur 28 34 44 52

genu 18 15 15 14

tibia 56 18 21 27

tarsus —_** 41 37 73

* Cross (1964, 1965) calls this the coxa, and labels subsequent

segments accordingly. ;

**In leg I tibia and tarsus are fused to a single segment.

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA 321

¢ ‘

i N =o

~ ‘p if .

piv \

ey bo JX ee

es eee —*:

Zea ee

\, = Hy

Tm,

y i, ~

ras

FIG, 17. Bakerdania workandue sp. noy. Holotype female. A,

Ventral view. B, Ventral view of left leg II (setae of tarsus II

incomplete in B, from obscurity of mount). (A, B to scale

shown.) C Distal segment of left palp, further enlarged.

322

The setation formula of the legs is (see Figs. 16,

17):

Leg I I Ill IV

trochanter ld 0 lv lv

femur ld, lv, 3d, Id, lv Id, lv

ll (‘c’?) 0-lv

genu 2d, 2v 2d, lv 2v Ip

tibia 8d, 7v 1d, 11, 2d, 2v, 2d, 2v

+3So0 2v+So +So

tarsus — 4d, 4v 4-5d, 2v 4d, 2v

+So

Details of leg specialized setae are as follows:

Leg I. The tip of ‘seta c’ of femur I (a modified

scobala) somewhat falciform and sharply proflexed.

On dorsum of tibiotarsus I are two expanded sole-

noidalae, and more distally one attenuated solenoi-

dala, and more distally again a retroflexed sensory

seta; at the “proral situation” a further seta, probably

a solenoidala.

Leg II. On tibia II the proximal end is overlain

dorsally by a stout clavate solenoidala. On tarsus II

a similar clavate solenoidala lies dorsally and proxi-

mally.

Leg III. On tibia HII a clavate solenoidala arises

proximally and dorsally.

Tarsal claws: On leg I single, on a short pedicle,

and the tibiotarsus I distally and ventrally carries a

short “‘thumb-process”. On tarsi II-IV the claws are

double, each claw strong, falciform, and with a basal

tuberosity or “cushion”; the middle (neomedian) claw

has a chitinized wishbone-like basal piece, from which

an adhesive membrane spreads out fanwise, lobate.

Material Examined

Holotype specimen, ACC226, to be deposited in

South Australian Museum collection. Taken attached

to central dorsal area of adult Chyzeria hirsti Wom.,

specimen ACB320, from under stone, Workanda

Creek, National Park, Belair, South Australia,

1 August 1948, R. V. Southcott.

Notes on Behaviour

On capture the adult Chyzeria hirsti was examined

under a binocular microscope to see if it carried any

larval Chyzeria. None was observed. I recorded “To

my surprise I found a small brown mite in the cavity

of its back as shown [see Fig. 18]... It was cleaning

the back part of the abdomen by hooking its last pair

of legs over, in the same way as a fly does”.

The relationship between the adult female pygme-

phorid mite and the adult Chyzeria hirsti is presum-

ably best described as a phoretic one. The relationship

is unlikely to be a parasitic one in view of the small

REC. S. AUST. MUS., 18 (14): 285-326

June, 1982

size of the cheliceral stylets of pygmephorids, and the

fact that the pygmephorids, in a restricted sense, are

not known to be parasitic; a number of species are

now known to be fungus-feeders (see e.g. Wicht (1970),

Kosir (1975) and the further references they quote).

B'S

Urns

FIG. 18. Adult male Chyzeria hirsti Womersley carrying an adult

female pygmephorid mite, the holotype of Bakerdania wor-

kandae sp. nov., in the space on the dorsum between the

dorsal idiosomal processes. From life, redrawn from a sketch

made at the time.

As a phoretic host, it would appear that the adult

Chyzeria hirsti is not likely to be efficient. Although

Chyzeria hirsti adults are, among similar-sized soil-

inhabiting Australian adult trombidoid mites, rela-

tively fast-moving, it would seem unlikely that they

would disperse over other than small distances.

Remarks on Classification

In the systematic placing of Bakerdania workandae

I have been initially guided by the works of Krezal

(1959) and Cross (1964, 1965). Cross produced a

monographic revision of the genera of the Pyemotidae

(sensu lato), and this was to have been followed by

a first application of the revision to localized faunas.

The large monographic revision was published as

Cross (1965) and contained a division of the old genus

Pygmephorus Kramer, 1877 into the new genera

Acinogaster Cross 1965, Parapygmephorus Cross

1965, Pseudopygmephorus Cross 1965, and Neopyg-

mephorus Cross 1965, as well as a number of subge-

nera. Cross applied these names in his paper studying

OBSERVATIONS ON CHYZERIA GANESTRINI AND SOME RELATED GENERA

the relevant mite fauna of Macquarie Island, which

appeared earlier as Cross (1964), By then he had

reached the conclusion that the separation of the

genera named might not be entirely valid, as the

Macquarie Island survey immediately revealed species

with characters belonging ta one or more of the

genera proposed. In response to this he placed the

five Macquarie Island species in Neopyymephorus,

with the comment that “In the above paper [Cross,

1965], | divided the tribe Neopygmephorini into four

new genera .-.. Acquisition of new material since

then indicates. that this grouping is probably unjus-

tified, ard that it is more realistic to join at least

Pseudopygmepharus and Neopygmephorus ... 2. OF

the two last-named generic names, Pseudopyemepho-

rus has. page priority, but Cross (1964) presumably

chose to use the name Neopygmephorus on the ground

that a tribal name was based upon it, although this

is not stated.

Subsequently Cross (1970) has shown ihat /Veo-

pvegmephorus is a junior synonym of Bakerdania Sasa

1961, and this has been accepted by other workers

on the Pygmephoridae and Pyemotidae, e.g, Mahunka

(1973a, b), Rack (1974).

A further complication has since been discovered

which may have an important bearing on the generic

placing of “pyemotid’’ mites. This is ihe observation

that in certain of the pygmephorid and pyemotid

mites the adult females may appear in two forms,

the phoretomorph and the non-phoretomorphic lorms,

which have different generic placernents in existing

classifications, e.g. Siteroptes and Pediculaster. or

Siteroptes und Pygmephorellus (Rack 1974, Cross

and Moser 1965; Moser and Cross 1975), So far this

phenonienon does not appear to have been recorded

with those pygmephorid mites allotted Lo the generic

forms proposed by Cross in 1965, but clearly this

possibility cannot be excluded.

In recent years no full revision of the Pygmephor-

idae and Pyemotidae has been published*, and at

present the only general work on the pygmephorids

and pyemotids of the world is by Krezal (1959). In

this latter the old genus Pygmephorus Kramer 1877

was not divided into subgenera. By Krezal’s key the

species described above comes to the caption of Pyg-

mephorus kachi Krezal 1959, from which there are

a number of morphological differences. Among these

are the more. spindle-shaped tibiotarsi I, the presence

of a strong opposition-piece (anvil, or incws) ta the

claw of tibiotarsus 1, the far more elongate tarsus FY

“In recent years further taxonumie changes at the family and

suprafainifial levels have been introduced for these mites. Thus tho

superfamily names Pyemotoidea and Pygmephoroidea have been

tised by some authors (see further in Krantz, 1978). These taxonomic

changes are, however, outside the soupe of the present work,

323

(three times the length of tibia IV, as against twice

in the case of P. Kochi) and a number of other factors.

P. kochi was one of the 16 species. or subspecies

placed by Cross (1965, p. 224) tentatively in his genus

Pseudopyamephorus

By the generic table of Cross (1965) the species

described aboye comes within Neupygmephoris Cross

1965, subg, Neopyemephorus, Cross (1965, p. 230)

gave a list of the already described “pyemotids”

which he believed should be placed in this subgenus,

amounting to 17 species and one subspecies, but did

nol submit a diagnostic key to separate them, The

only recent key to a group of related pyemotid mites

from the Australian region known to the present

author is Cross’s (1964) study of the fauna fram

Macquarie Island, allotted to Neapygmephorus. These

indicate the wide-spread distribution of these mites,

as species known hitherto from Europe, Japan and

North America were represented,

Accepting that the species described above belongs

to Neopygmephorus as used by Cross in his 1964

paper, Le. to Bakerdania Sasa 1961, and using that

key, one finds that B. workandae cames closest to B

arvorunt (Jacot 1936), originally described from North

America by Jacot (/. c.) but how known to occur alsu

in Macquarie Island (Cross 1964), Germany (Rack

1967), Jugoslavia (Mahunka L975b) and Tunisia

(Mahunka 1978) and to & fogata (Willmann 1942),

originally described frorn Germany, but now also

recorded from Macquarie Island (Cross 1964). Hong

Kong (Mahunka 1975a) and Africa (Mahunka 1976).

Rack (1967 p.10) places P. arvorum Jacot, 1936 in

Pseudopygmephorus Cross, as did alsa Cross (1965,

p. 224).

From these two species 8. workandae may be sep-

arated as follows:

B. workandae has 6 caudal setac instead of the 4

which oceur in 8. arverum.

B. workandae has the lateral presternal setae well

lateral to the medial presternals, while in B. logaia

the lateral presternal setae are only slightly lateral 10

the medial presternals. In B. workandae the lateral

poststernal setae are significantly longer than trochan-

ter IV, while in B. rogata the lateral poststernals are

significantly shorter than trochanter IV (see further

in Cross 1964; Willmann 1942),

A number al papers have been written on the

pyematid fauna af the world since the acceptance of

the status of Bakerdania as a senior synonym of

Neopyemephorus; some of these describe new species

uf Bakerdania. Of the new species described, jn the

literature available to me from the extensive reprint

library of the South Australian Museum, and in my

own and other libraries, there is only one which

appears to have a significant resemblance to B. wer-

314

kandae sp, nov, This 1s B, diserepata Mahunka, 1973,

from Ghana (1973b, p, 318). Both of these species

are known from only a solilary female; males are

linknown for them. These two females may be sepa-

rated as follaws:

Tarsus IV (excluding claws and pedicle) less

than twice as long as tibia LV. Tibiotarsus I short,

ellipsoidal, Distal solenoidala on tibiotarsus I

short, about half as long as the immediately

proximal thick solenvidala, ,.B, discrepata

Mahunka 1973

Tarsus TV (excluding claws and pedicle) more

than twice as long as tibia IV. Tibiotarsus [

spindle-shaped, Distal solenoidala of tibiotarsus

I long and attenuated, longer than the proximal

thick solenoidala... 8. workandae sp, nov.

Acknowledgement

Tam indebted to Evert E, Lindquist, Senior Acar-

ologist, Biasystematics Research Institute, Ottawa,

Canada, for helpful criticism of this paper.

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SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

3. ARTHRONOTINA (CRYPTOSTIGMATA)

BY DAVID C. LEE

Summary

A study of sarcoptiform mites from surface soil (usually greatest depth = 4cm) at 9 florally diverse

sites in South Australia is continued. Notations for hysteronotal fissures, kinds of gnathosoma,

solenidia, form of notal setae and shield sculpturing are presented. Comments are made on the

systematics of Macropylides in which the disbanding of the cohort Ptycimma is supported. The

included Arthronotina are divided into 3 new subcohorts: Monofissurae, Retrofissurae and

Profissurae. A new family, Trichthoniidae, is established. A new name, Neoliochthonius, is given to

Paraliochthonius Moritz not the pseudoscorpion genus Paraliochthonius Beier. New diagnoses are

given for some superfamilies, families and genera. In this study, 11 species were collected.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS.

3. ARTHRONOTINA (CRYPTOSTIGMATA)

DAVID C. LEE

South Australian Museum, Adelaide, South Australia 5000

(Manuscript accepted 3 July 1981)

ABSTRACT

LEE, D.C, 1982. Sarcopliformes (Acari) of South Australian soils.

1, Arthronouna (Cryptosuigmata), Ree 5S. dus Mus, 18 (15):

327-359,

A study of sarcoptiform mites from surface soil

(usually greatest depth = 4 cm) al 9 florally diverse

sites in South Australia is continued. Notations for

hysteronotal fissures, kinds of gnathosoma, solenidia,

form of notal setae and shicld sculpturing are pre-

sented, Comments are made on the systematics of

Macropylides in which the disbanding of the cohort

Ptyclimina is supported, The included Arthronotina

are divided into 3 new subcohorts: Monofissurae,

Retrofissurae and Profissurae, A new family, Tni-

chthoniidae, is established. A new name, Nealiovh-

thonius, is given to Paraliochthonius Moritz not the

pseudoscorpion genus Paraliochthonius Beier. New

diagnoses are given for some superfamilies, families

and genera. In this study, 11 species were collected.

Two species are new: Gehypochthoniuy yirenckei, Wer-

achthonius movritzi. Six species are newly recorded

from Australia: //ypochthoniella minutissima Berlese,

Brachychoehthenius elsosneadensis (Hammer), 8. cri-

coides Weis-Fogh, Trichthonius pulcherrimus (Ham-

mer), Phylozetes emmae (Berlese), Sphaerochthonius

splendidus (Berlese), Three species were previously

recorded from South Australia: Cosmochihonius aus-

tralicus Womersley and 2 which were misidentified:

Liochthonius simplex (Forsslund) as “'Brauchychthonius

ch perpusillus’, Liochthonius finibriatissimus (Hani-

mer) as “'Brachyvehthonius ef. horridus’'. The deserip-

lions of 4 species not collected in this study are

extended: Gehypachthonius rhadamanthus Jacat,

Hypochthontella borealis Jacot, Poecilochthonius par-

allelus (Womersley), Liochthonius longipilus (Wom-

ersley), Caosmochthonius wallworki and

Sphaerochthonius wallworki ave established for "'C.

sp.” Wallwork and “'S. sp." Wallwork, The synonymy

of Casmochthonius domesticus under C. lanatus by

van der Hammen is revoked. Three subspecies of

Coysmochthonius are given species status. New com-

binations are Gehypochthonius urticinus (Berlese) (ex

Parhypochtheamus), Poecilachthonius parallelus

(Womersley) (ex Brachychthonius), Liochthonius lon-

gipilus (Womersley) (ex Brachyehthonius), Perach-

thontus montanus (Hammer) (ex Kobrachychthontus)

Besides 2 new identifications above, Liochthonius

ocellaius: Hammer, 1958 is identified as L. /ongipilus

(Womersley).

INTRODUCTION

This publication is part of a previously introduced

study (Lee 1981) in which | indicated that | would

fallow Balogh’s (1972) classification. However,

although that classification 1s valuable because of its

comprehensive nature, in dealing with the Macropy-

lides or “Primitive Oribatei” it ignores parts of the

published work of Grandjean and van der Hammen,

In doing so it retreats from maturing the classifica-

lion. Therefore, although ihis paper is primarily con-

cerned with the Arthronotina, if includes a syste-

matics section on the Macropylides in order to explain

the higher classification used here, which is a modi-

fication of that presented by Grandjean (1969). Fur-

thermore, higher taxa are given endings similar to

those of Krantz (1978) in order to standardize the

differentiation of each level. It should also be noted

thai, contrary to common practice amongst acaralo-

gists, other zoologists follow Simpson (1945) in

regurding a cohort as a category between class and

order rather than between order and family.

In order to examine the optical activity of some

hairs, the Leitz microscope and interference contrast

device T normally used was modified for use as a

polarizing microscope by having a bright-lield con-

denser dise and objectives so that no Wollaston prisms

were in the beam path.

All material collected in this study is deposited in

the South Australian Museum, Adelaide, unless

stated otherwise.

NOTATION FOR MORPHOLOGY

The notation. followed is as previously presented

(Lee 1981), but comments are made below on the

notation for hysteronotal fissures and shields. kinds

of gnathosoma, solenidia, form of notal setae and

shield sculpturing.

Hysteronotal Fissures and Shields

On most adult Cryptostizmata the hysteronotum is

covered by a single continuous shield. The Arthro-

notina are unusual in that this integunient is broken

up into a series of shields separated by striated cuticle

which when it forms a narrow strip is referred to as

a fissure, In a description there is often a bias towards

deseribing one or the other. Descriptions of the dorsal

integument of Arthronotina are mainly biased towards

328

describing the transverse fissures. On the other hand,

the lateral integument is usually described in terms

of its shields. The latter can be confusing because a

shield may be described as absent either because it

is replaced by striated cuticle or because it is still

present but is not separated from the rest of the notal

shield by a fissure. The notation to be used is consid-

ered under two headings: (a) fissures, (b) shields (see

also Fig. 1).

(a) Fissures. The signature “B” (initial letter of

“break”, “F” having already been allotted) will be

used for a line separating one part of a shield from

another part that is either a strip of striated cuticle

or is regarded as representing what was a strip of

striated cuticle in some allied species. Fissures are

regarded as belonging to 5 categories: (i) acutely

hinged fissures, with articulated sides capable of

movement from a level position to enclose an angle

of 80° to 140°; (ii) Ainged fissures, with articulated

sides only capable of movement to enclose an angle

of 140° to 175°; (ili) complete fissures, with immovy-

able sides completely dividing a shield into separate

parts; (iv) partial fissures, not completely dividing

shield into separate parts; (v) relict fissures, no strip

of striated cuticle, but some demarcation furrow on

shield representing a fissure. The taxonomically

important fissures are the transverse hysteronotal

fissures of which there is a maximum of 3 (TBI,

TB2, TB3) each lying behind the same numbered

seta in hysteronotal file J. There may be transverse

lines behind seta J4 but these are not regarded as

representing strips of striated cuticle. Only one lateral

hysteronotal fissure is referred to: the dorsolateral

longitudinal fissure (LB). This separates what are

called pleural shields from the notal shields.

(b) Shields. On the hysteronotum there are a max-

imum four notal shields (NS1-4) and four subnotal

shields (SNS1-4) numbered from the anterior. The

subnotal shields have been referred to as “‘supra-

pleural” shields (Moritz 1976) but, since they some-

times merge with the notal shields as for example

shield SNS1 which bears seta S1, I have preferred

to relate them to the notal shield. Ventral to the

dorsolateral longitudinal fissure are the pleural

shields (PS1, 2) and subpleural shields (SPS1, 2).

Kinds of Gnathosternum

The 3 parts of the gnathosternum which contribute

to its general shape are the palp coxites, external

malae and mentum. In a few groups there is no

mentocoxal fissure so that the precise border between

the mentum and the palp coxites is not known. Grand-

jean (1957) regarded gnathosterna as belonging to 3

basic kinds (anarthric, stenarthric and diarthric), but

I have found it more useful to directly describe the

structures involved.

REC. S. AUST. MUS., 18 (15): 327-359

June, 1982

The following notation is introduced to describe

the external malae and mentum. Disjunct external

malae are well separated and lie either laterad or

dorsolaterad to the internal malae and the bases of

the preoral setae. Conjunct external malae lie

ventrolaterad to the internal malae and the bases of

the preoral setae, whilst a ridge indicates that the

protruding parts of the palp coxites from which they

originate come together midventrally behind the

internal malae. Coarctate external malae abut ven-

trad to the preoral setae, obscuring them from view.

An undelineated mentum has no distinct border with

the palp coxites because there is no mentocoxal fis-

sure, although, when this is true for members of the

primitive beetle mites, the mentum is probably trian-

gulate and sometimes ridges suggest its outline. A

triangulate mentum is almost triangular in outline

coming gradually to an anterior apex. The lateral

margins lie well behind both this apex and the cox-

otrochanteral joint of the palps. A quadrangulate

mentum is almost square or rectangular in outline,

usually with a straight, transverse anterior margin.

This shape results from a thickening of the lateral

margins of the mentum so that its anterior shoulders

move forward close to the coxotrochanteral articula-

tion of the palps.

A general statement can be made that the gna-

thosterna of Cryptostigmata have evolved from those

with disjunct external malae and an undelineated

mentum to those with coarctate external malae and

a quadrangulate mentum. The Arthronotina have

either disjunct or conjunct external malae and an

undelineated mentum.

Solenidia

Three of Grandjean’s established terms for soleni-

dia of certain shapes (baculiform, ceratiform and

piliform) (see Norton 1977) are used below. The term

“tactile” is replaced by flagelliform and the term

spiniform is used for stout evenly tapering solenidia.

Some solenidia are coupled with a seta, occupying

the same alveolus, whilst others are either associated

with a seta or well separated from any seta.

Although solenidia are usually dorsal, either | or

2 ventral solenidia have been recorded (Grandjean

1954b 1963, Covarrubias 1968, Reeves and Marshall

1971) on the tarsus I of some Brachychthoniidae,

Protoplophoridae and Sphaerochthoniidae. In describ-

ing a new species of Verachthonius below these solen-

idia were examined with polarized light and appeared

to fit the definition of solenidia, but are spiniform,

which is not a shape found amongst dorsal solenidia.

The symbol sov is used for these solenidia, and if in

a pair the anterior one is numbered first.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

Form of Notal Setae and Shield Sculpturing

In identifying the Cosmochthonius species col-

lected in this study it was found that some attributes

of the form of the notal setae and shield sculpturing

had to be carefully delineated since these characters,

which may be trivial and inaccurately described in

some previous descriptions, had to be used because

few other data were available.

In describing the long hysteronotal setae, seta J3 is

always referred to and it is assumed that setae J4, Z3

and 24 are similar unless otherwise indicated. The

important attributes are the size and spacing of the

marginal files of cilia on the setae. Cilia are regarded

as even if each cilium is intermediate or equal in

length to the cilia on either side of it, and uneven if

this is not so. Cilia are short if less than diameter of

setal mid rib, medium-lengthed if * 1-x 1.25 and

long if more than x 1.25 this diameter. Cilia are

dense if distance apart is less than diameter of setal

mid rib, medium-spaced if * 1-x 1.25 and sparse if

more than X 1.25 this diameter.

The notal sculpturing consists of pits or puncta

which are described as punctations if circular or oval

and reticulations if polygonal, the straight edges

forming a network. Puncta are /arge if similar or

greater in size to setal base J3, or small. Puncta are

abutting if the space between them is not greater

than X 0.5 each one’s diameter, close if between

x 0.5-x 1 and well separated if greater than this

diameter.

SYSTEMATICS

Supercohort MACROPYLIDES

Remarks: Balogh (1972: 32) stated in referring to

his classification “The . . . Primitive Oribatei or

Macropylina is in complete accordance with van der

Hammen’s system (1959).” This is inaccurate. Van

der Hammen (1959) has 7 groups as follows (with

Balogh’s grouping in parentheses): Palaeacaroidea

(= Bifemoratina), Parhypochthonioidea (included in

Arthronotina), Enarthronota including Protoplophor-

idae (Arthronotina and part of Ptyctimina), Mesoplo-

phoroidea (= part of Ptyctimina), Phthiracaroidea

(= part of Ptyctimina), Perlohmannioidea (= part

of Holonotina) and Nothroidea including Nanher-

manniidae and Hermanniidae (= part of Holonotina

and part of the Apterogasterina in the Brachypylides

or “Higher Oribatids”’). Therefore, Balogh’s classifi-

cation differs in 2 important respects from that of

van der Hammen: it ignores the rejection of the

Ptyctimina as a valid group and changes the point of

delineation of the Macropylides from the Brachypy-

lides. Van der Hammen’s classification reflects that

of Grandjean who established both the Enarthronota

(Grandjean 1947a: 215) with the Protoplophora

329

included in it, and a classification (Grandjean 1954a:

428-431) which is a detailed precursor of van der

Hammen’s classification.

It is hard to understand how the Ptyctimina (recog-

nisable by the adult being ptychoid i.e. having an

acutely hinged prehysteronotal fissure) has survived

as a taxon for so long. Grandjean (1933: 319) pointed

out that Mesoplophora (ptychoid) has an adult gna-

thosternum remarkably like that of Hypochthoniidae

(non-ptychoid) and should be grouped with that fam-

ily rather than with the Phthiracaroidea (ptychoid).

It appears that the belief that Grandjean’s work is

based on characters of immature stages which pro-

duce a “natural” but difficult classification to follow

has inhibited the application of his work. For the

Macropylides this had led to an unnecessary stagna-

tion since much of Grandjean’s classification is based

on adult characters. Not that there is any basis for

regarding adult characters as producers of an “arti-

ficial” classification. The disadvantage of Balogh’s

currently accepted classification of the Cryptostig-

mata seems to be that it was based on descriptions

of adults which in many cases included too few char-

acters.

Although the classification by Balogh (1972) is

widely used and is included in a recent manual on

mites (Krantz 1978) some specialists do follow Grand-

jean’s work. Even then, there may be uncertainty as

when Norton (1980), in referring to Mesoplophora

and other ptychoid mites, states “these and several

other apparently unrelated oribatid mites” (my ital-

ics).

The classification used in this publication follows

the trend of Grandjean’s (1969) work. The Ptyctimina

are not regarded as a valid group. The Protoplopho-

roidea and Mesoplophoroidea (ex Arthroptyctima)

are grouped in 2 different new subcohorts within the

Arthronotina, The remaining majority within the dis-

banded Ptyctimina, the Phthiracaroidea and

Euphthiracaroidea (ex Euptyctima), is grouped in the

Holonotina which are not considered below.

Cohort ARTHRONOTINA

Diagnosis: Macropylides. Often pale, minute to

medium-sized (790 or less) adults. One, 2 or 3 trans-

verse hysteronotal fissures present (Mesoplophoridae

appear to lack these fissures, but the presence of only

8 pairs of setae on notal shield indicates that shield’s

posterior edge represents fissure 783, while posterior

and pleural shields inconspicuously merged with ven-

tral shields). External malae either disjunct or con-

junct. Mentum undelineated. Femora not divided into

2 separate parts. Immatures similar to adults except

sometimes in disposition of hysterosomal shields and

always in chaetotaxy and nature of genital shields.

330 REC. S. AUST. MUS., 18 (15): 327-359

Remarks: The most conspicuous attribute of mem-

bers of this cohort is the presence of transverse hys-

teronotal fissures breaking up the rigid integument

into a number of separate shields. Just as the form

of these fissures varies from a vague line to a hinged

break in the soma, so do their functions probably also

vary. In the case of Elliptochthonius profundus a

fissure probably allows bending of the soma in order

to negotiate narrow pore spaces in the deeper soil

layers, whilst for Protoplophora palpalis it allows the

back as well as the front of the mite to be folded

down to form a protective ball. The advantage of the

fissure is not so obvious for mites such as Hypo-

chthoniella minutissimus and Sphaerochthonius

splendidus, but possibly it allows expansion of the

hysterosoma to accommodate a large egg.

The attributes of morphological characters in the

Arthronotina vary considerably compared with those

of the other 2 cohorts of Macropylides. This is the

main reason for my establishing 3 new subcohorts

within it. One, the Monofissurae, represents a well

established taxon in that it includes only the Parhy-

pochthonioidea. On the other hand, the Retrofissurae

and Profissurae are entirely new taxa. It may prove

preferable to downgrade these 3 subcohorts to super-

families (Parhypochthonioidea, Mesoplophoroidea,

Protoplophoroidea). However, for the time being, the

novelty of the grouping, the strangeness of the 2 ex-

Ptyctimina superfamily names being applied to much

larger and diverse taxa, and my uncertainty as to the

importance of the differences between Phyllochthon-

ioidea and the other Retrofissurae, make my classi-

fication a reasonable stage in the maturing of the

systematics of the cohort.

If the grouping into taxa that I have called the

Retrofissurae and Profissurae survives, it will suggest

a number of cases of parallel evolution: the folding-

up-soma of the Mesoplophoridae and Protoplophori-

dae; the short, simple somal setae of the Brachy-

chthoniidae and Haplochthoniidae; the long erectile

mid-dorsal setae of the Trichthoniidae and Cosmo-

chthoniidae; and the large toadstool-like dorsal setae

of the Phyllochthoniidae and Pterochthoniidae.

The Arthronotina exhibit primitive attributes. On

the other hand, I regard some attributes as specialised,

such as slim setose external malae (e.g. Hetero-

chthonius gibbus) and comb-like fixed cheliceral digits

(e.g. Pterochthonius angelus), which are considered

by Grandjean (1947b 1957) as reflections of the setal

origins of these structures. Also, although the presence

of hysteronotal fissures on all species reflects the

primitive nature of the cohort, a greater number of

fissures may not be an indication of a more primitive

taxon.

June, 1982°

The classification of the Arthronotina that I propose

to use is outlined below.

A, Subcohort: Monofissurae

1. Superfamily: Parhypochthonioidea

Families: Parhypochthontidae

Gehypochthontidae

Elliptochthonidae

B. Subcohort: Retrofissurae

1. Superfamily: Hypochthonioidea

Families: Hypochthoniidae

Eniochthoniidae

Brachychthoniidae

Heterochthoniidae

Trichthoniidae

2. Superfamily: Mesoplophoroidea

Families: Mesoplophoridae

Archoplophoridae

3. Superfamily: Phyllochthonioidea

Families: Phyllochthoniidae

Atopochthoniidae

C. Subcohort: Profissurae

1. Superfamily: Cosmochthonioidea

Families: Cosmochthoniidae

Sphaerochthoniidae

Pterochthoniidae

Haplochthoniidae

2. Superfamily: Protoplophoroidea

Families: Protoplophoridae

KEY TO SUBCOHORTS OF ARTHRONOTINA (ADULTS)

1. Hysteronotal gland present. Only single transverse hyster-

onotal fissure (7B2) present. External malae conjunct. Both

pairs of adanal pores (Jaf, Zaf) present. Genua of legs with

| or 2 solenidia. Pretarsi with 3 claws (shorter central claw

sometimes rudimentary) Fits _ Monofissurae

Hysteronotal gland absent. More than | transverse hysteronotal

fissure (if only | is conspicuous it is either TB1 or TB3).

External malae either disjunct or conjunct. Either | pair of

adanal pores (Zaf) present or both pairs absent, Genua may

have 0, | or 2 solenidia. Pretarsi with 1, 2 or 3 claws (if 3

claws then central | most robust) ; pe:

2. Two (rarely only 783) transverse hysteronotal fissures (7B2,

TB3) so that both setae JI and J2 on first shield (NS1/2)

(an exception is Phyllochthonioidea, with toadstool-like dorsal

setae, broad areas of striated cuticle on hysteronotum and

SARCOPTIFORMES (ACARI) OF SOLUTE AUSTRALIAN SOILS Ba

narrow first Shield nit bearing setu./2), External malae disjunct.

Fither | pair (4a// of adanal pores present or both pairs

absent, At Toust genua of legs Tand YW have | or 2 solenidia,

Pretarsi with enher | or 2 subequal claws Retrofissurae

Three (rarely only 7AI obvious: 782 obscure, 7'BS absent)

hysteronotal fissures (TBI, 7B2 TB4) so that-seta J) but not

Jon Sirst shield (NSE), External matic conjuncl Na adanal

pores, All genua without solenidia, Prelurs) with 1, 2-or 3

cluws but ib 2 elaws present. | usually much slimmer and

recly on pretarsis FV Profissurue

Subcohort MONOFISSURAE an.

Diagnosis: Arthronotina. Single transverse hyster-

onotal fissure (7'B2) complete ar hinged. Hysteronotal

gland with egress pore between setac 24 and 75.

Cowl absent or rudimentary, enclosing only part of

retracted chelicerac. Cheliceral digits unmodified,

dentate. Cheliceral setae both (cht, ef2) present and

setose, Distal plasmie setae on palp tarsus simple not

bifureate. External malie conjurct, Two pairs of

adanal pores (Jaf, Zaf) present. On tarsus 1, seta d/

never simple, at least bifureate. Genua of all legs

with | or 2 solenidia. Pretarsi with 3 claws of which

shorter central claw may be: rudimentary.

Remarks; The Monofissurae is not a mew taxon

since it includes only the Parhypochthonwwidea van

der Hammen, 1959, with the following 3 monogeneri¢

families: Parhypochthontidae, Gehypochthoniidae,

EMiptochthoniidac. Although Grandjean (19544) sep-

arated the Parhypochthonigidea from what is here

regarded as the rest of the Arthronotina, | prefer to

follow Balogh (1972) in placing itia the Arthronotina.

especially since the Elliptochthonudae partly “closes

the gap” be|ween the 2 taxa. As | gehypochthoniid

ayinph was collected in the present study, that family

15 considered in more detuil below. The 2 other families

aire considered brietly as Sollaws. i) Parhypochthoni-

due Grandjean, 1932b. Similur to Gehypochthoniidae

especially in the general appearance of the gnatho-

soma. However it is casily distinguished [rom other

Monofissurie by the egress pore of the hysteronotal

eland being on an apophysis and by the presence of

a setal file Sa which includes 4 setac, Parhypochihon-

fuy does have attributes in common with Ellipte-

chthonius thal do not accur on Gehypochthonius,

such as 3 palpocoxal setae, hysteronotal seta 24 pres-

ent and a solenidium on tibia IV, but these are

considered at the most to distinguish families. it)

Elliptochthunjidae Norton, 1975, When established it

was suggested that this family may in the future have

lo be grouped in a separate superfamily. Amongst

the combination of attributes used in the diagnosis,

it should be noled that the hinged transverse hyster-

Onotal fissure (Which merges laterally with the ventral

fissure immediately behind coxite LV so that the

posterior part of the body 1s movable in relation (o

the unterior part) oeeurs on members of the rhada-

monihussomplex in Gelhypochthonius, Also used to

diagnose the family was the gnathosoma which has a

number of substantially modified characters consid-

ering that the Monofissurae occupies a primitive pasi-

tion in the classification. The external malae ventrally

obscure much of the internal malaec and their palpo-

coxal bases abut onto each other for almost their

entire length. There is only one pair of adoral setae,

The palp is reduced in that the trochanter and femur

are fused and it carries few setae (1-0-2-7). The cowl

is extensive, although not completely enclosing the

retracted chelicerae. These gnathosomal altribules

alone support the suggestion that this family may

have Lo be grouped in a separate superfamily,

Family GEHYPOCHTHONIIDAE Strenzke

Gehypochthoniidae Strenzke, 1963; 251.

Type-genus: Gehypochthonius Jacot, 1936.

Remarks: The Gehypochthoniidae currently in

cludes only Gehypochthonius so that most further

points are made under that generic heading. Gel

pochthonius and Parhvpochthonius are closely allied.

I would group Gehypochthoniidae under Parhypo-

chthoniidae, but for the presence on Parhypochthon-

ius of 4 setae in file Sa. which is regarded as impor-

tant in representing a “preanal segment” (Grandjean

1954a) and is usually absent amongst the Arthrone

tina (exceptions: Pterochthoniidae and possibly Phyl-

lachthonioidea).

GEHYPOCHTHONIUS Jacot

Gehypochthonius Jacot, 1936; 22, Type designation

(original): “Gehvpochthonius rhadamanthus sp.nov.”

Type-species: Gehypochthonius rhadamanthus Jacot,

|936: 22.

Diagnosis: Monatissurae. Cowl absent. External

malae conjunct. Adoral setal file includes 3 pairs, of

which aol is. either bifurcate or spreads out distally

into-a triangulate fap. Palpocoxal setal file includes

2 pairs. Adanal setal file Sa absent.. Transverse lys-

teronotal fissure may or may not be hinged and merge

with a ventral fissure. Mysteronotal seta 24 absent

(present on larva, see Strenzke 1963; fig. 20). No

selae in adanal file Sia. Palp wiih 5 free segments

and 2 setae on femur. On tarsus [, bifurcate seta q!

with | branch tapering and 1 expanded into an oval

flap. Tibia TV without solenidia,

Morpholagy: Minute (250-380), pale ivory-winte

and someiimes partly straw-coloured, chelicerae light

brown. Soma bulbous behind seta /2 with clear,

medium-lengthed refractile setae sticking out like

spines. Legs | and IV longer than II and IU, |

stoutest,

Gnathosoma with conjunct external malae, internal

malae not partially obscured ventrally. Leg coxites

completely delineated by apodemes, although apo-

deme separating coxite 11] from IV sometimes bro-

ken.

332

REC. S. AUST. MUS., 18 (15): 327-359

June, 1982

FIGS. 1-4—Arthronotina: 1, signatures for hysteronotal shields and

fissures; 2, Gehypochthonius rhadamanthus Jacot, adult,

notum; 3, Gehypochthonius strenzkei n.sp., tritonymph, notum;

4, Hypochthoniella minutissimus Berlese, adult notum.

When transverse hysteronotal fissure hinged, ante-

rior hysteronotum narrower at the fissure and fits

into posterior hysteronotum. Similar to structure in

Elliptochthonius, but not to equivalent fissures of

most Arthronotina, where the posterior fits into ante-

rior hysteronotum.

Chaetotaxy. Soma: 2/, 2z, 2s; 6J, 5Z, 4S; 3ao, 2c,

led; 31, \Id, 217, 3111, 41V; 5 to 8Jg, 3Zg, 28g; 2Ja,

3Za, OSa. Appendages: ch(2), pa (0-2-0 or 1-2-11),

I (1-4 or 6-5-6-22), 7/7 (1-5-3 or 5-5 or 6-17), T/T (2-2

or 3-2 or 3-3 or 4-13 or 15), JV (1 or 2-2-1 or 2-3 or

4-11 or 13). Solenidia: pa (0-0-1), 7 (1 or 2-1-3), I/

(1-1-1 or 2), //7 (1-1-0), IV (1-0-0).

Hairs generally setose. Adoral seta aol either bifur-

cate or spreads out distally into triangulate flap, ao2

sometimes stout, blunt rod. Some somal and appen-

dage setae with inconspicuous cilia; plasmic seta z2

has conspicuous cilia and take | of 2 forms described

below under diagnoses of the 2 species-complexes.

Posterior hysteronotal setae always tapering distally,

but in some cases medially and proximally dilated

into a refractile cylinder around a central canal. Some

dorsal leg setae (as G. xarifae) lanceolate with lateral

flaps. Tarsus I, with seta dl short and bifurcate, 1

branch tapering and | ovate. If solenidia present on

a tarsus or tibia at least 1 is ceratiform or baculiform.

Solenidia of genua either piliform (sometimes slightly

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

blunter than surrounding setae and so approaching

ceratiform) or flagelliform,

Distribution: Possibly cosmopolitan—Canada (Nn),

Carolina and Florida (Na), France (Pe), Japan (Pc),

Maldive Islands (Or), Australia (Aa).

Remarks; The only species af Gehypachthonius

described extensively enough to refer Lo the gnatho-

soma and appendages is G. xarifae Strenzke, 1963,

Since the lateral view of the type-species (G. rhada-

manthus Jacot, 1936: fig. 1) is similar to that of

Elliprochthonius, 1 needed to establish whether or

not these 2 species were congeneric. because, although

the single iritonymph collected during this study is

congeneric with G. xarifae, it was not certain if either

species should be grouped in Gehypochthonius.

Strenzke (1963: 24%) has compared G. xarifae and

G, rhadamanthus, referring to Grandjean's unpub-

lished extensions of the description of G, rhadaman-

thus based on specimens collected in France, bul at

that time Elliptochthonius was not known.

To clarify the situation | examined both described

and undescribed mites not collected in the present

study. These, together with the material belore me,

confirm that. G. xarifae and rhadamanthus are sim-

ilar. However whilst regarding them as congeneric, |

place them in separate species-complexes, pending

upgrading or abandonment.

rhadamanthus-complex

Diagnosis: Gehypochthonius. Smaller (250-275),

Transverse hysteronotal fissure merges with ventral

fissure and appears hinged, Plasmic seta 22 with short

cilia (length »% 2 or less of setal diameter). Hyster-

onotal seta 72 more than * O.5 length of 82. Hys-

terondtal seta J5 less than * 2 length of J3 and

diameter never mare than that of outer rim of alveolus.

Setal file Jg with 5 or 6 setae. Adoral seta ao2

tapered as eo3. Appendage chaetotaxy reduced, for

example no setae on genu of palp and only 3 setae

on both tibiae [f1 and [V. One solenidium on genu

{, Solenidium on tibia TIL Jonger (at least % 1.1)

than the tibia. Solenidium on geno JV about twice

(x 1,9-2,1) Jength of the genu,

Remarks, As only | species is included, further

information 1s under the species heading. As noted

there, the species may be composite.

Gehypochthonius rhadamanthus Jacot

(Fig. 2)

Gehypochthonius rhadamanihus Jacot, 1936: 22.

Gehvpochthonius rhadamanthus Jacot: Aoki, 1975:

55.

Adult

Idiosomal length 250-275 (Carolina—255,

N.S.W.—250, Japan—262-275). Main cheliceral axis

334

apparently al less than 45° angle from horizontal

somal axis even in unsquashed specimens, Apodeme

separating coxite LI] from IV broken, Specimen from

Japan atypical (a8 original description) in that seta

Z2 longest in second rank. Seta J5 slimmest on Car-

olina specimens and stoutest (equal in diameter io its

alveolus) on N.S.W. specimens, Specimen (Fig. 2)

from Carolina differs compared with those from ather

localities in that setae /6 and 26 displaved forward.

Chaetotaxy of soma varies in that file Je has 6

setac On Carolina and Japanese specimens and 4 setae

on N.S.W. specimens. Chaetolaxy of appendages

(NLS.W. adults, and same on femora, genua and

tibiae of Carolina adults): pa (0-2-2-11), f (l-4-5--

22), HH (1-5-3-5-17), TM (2-2-2-3-13), FV (1 -2-1-3-1 1}.

Marerlal examined; Twenty-five svntypes

(“cotypes”, 34F23.2-5, Smithsonian Institution,

U.S.A.) from Carolina on single slide. One adult

(Ac8636, National Science Museum, Tokyo, Japan)

from Japan, Three adults (N19794.N 19796), litter and

soil. open £uca/pptus forest, Cordeaux (34° 08°S, 150°

43°E), New South Wales, Austraha, 9.6.1978, T.

Moulton.

Specimens from N.S.W. well cleared but squashed,

and characters of gnathosoma and appendages (some

presented under genus heading) clearly discernible,

Syntypes in good condition yet poorly cleared and

sometimes shrivelled. Most characters of gnathosoma

and appendages discernible, bul chaeiataxy of appen-

dages only confirmed on femur, genu and tibia, and

for solenidia alone on tarsi. Specimen from Japan

poorly cleared and shrivelled. Many characters of

gnathosoma and appendage chaetotaxy not discernible.

but attributes of transverse hysteronotal fissure, sete

72 and solenidia on tibia [II and genu LV justify its

inclusion in this. species-complex,

Distribution: Widespread—Carolina (Na), France

(Pe), Japan (Pc), Australia (Aa), The majority of

specimens examined by Jacot (1936) were from 5-

12.5 cms deep in the soil. Records of other specimens

are {fom soil of plant litter under shrubs or trees.

Remarks: G. rhadamanthus, with a hinged trans-

verse hysteronotal fissure and simpler setae, appears

adapted to live in deeper soil layers than its congeners

and therefore superficially resembles E//iptochthonius

species. If the morphological variations of this wide-

spread species later prove to reflect distinct species,

and the above diagnosis for the rhadamanthus-com-

plex remains valid, } would recommend that the genus

be limited to this complex alone.

xarifae-complex

Diagnosis: Gehypochthonius. Bigger (290-380).

Transverse hysteronotal fissure not merging with a

ventral fissure and not hinged. Plasmic seta 22 with

long cilia Vength of longest *% 5 or more of setal

334 REC. S. AUST. MUS., 18 (15): 327-359

diameter). Hysteronotal seta Z2 less than xX 0.5

length of S2. Hysteronotal seta J5 either more than

2 length of J3 or its diameter in parts greater than

that of outer rim of its alveolus, or both attributes

present. Setal file J/g with 7 or 8 setae. Adoral seta

ao2 blunt, spindle-shaped or parallel-sided rod.

Appendage chaetotaxy not so reduced, for example

| seta on palp genu and 4 setae on both tibiae Il

and IV. Two solenidia on genu I. Solenidium on tibia

Ill shorter (at most & 0.9) than tibia. Solenidium on

genu IV conspicuously shorter than twice (x 1.5 or

less) length of genu.

Material examined: Specimens of new species

below. Holotype of G. frondifer from Japan (Ac8567,

National Science Museum, Tokyo, Japan); notum

uppermost, slightly shrivelled, only partially cleared

and including 2 substantial opaque food boli, some

attributes difficult to discern.

Distribution: Widespread—Florida (Na), Maldive

Islands (Oc), Australia (Aa). From moss or plant

litter under shrubs or trees.

Remarks: Although little is known about Parhy-

pochthonius urticinus Berlese 1910a: 219 and fig. 43,

the dorsum of the female illustrated with egg is so

similar to that of members of this complex that it is

here included in Gehypochthonius.

Four species are included in the xarifae-complex:

G. frondifer Aoki, 1975; G. strenzkei nsp., G. urti-

cinus (Berlese, 1910a) n.comb., G. xarifae Strenzke,

1963.

Gehypochthonius strenzkei n.sp.

(Fig. 3)

Tritonymph

Idiosomal length 290 (1); appendage lengths—ch

27.5, pa 55, I 125, I 95, HT 95, IV 120; femur

breadths—pa 12.5, J 25, I7 15, IIT 15, IV 15, Trans-

verse hysteronotal fissure (782) indistinct, bearing

setae J2 and Z2, (fissure 782 of adults set behind

these setae). Indistinct disc around the hysteronotal

gland pore (AG) not illustrated. No indication of

vestigial presence of seta Z4. Five hysteronotal setae

(J4, J5, J6, Z3, Z5) have swollen, refractile zone,

but not setose seta Z6. Four of these 5 setae‘(all but

J6), on right side only, bear a ring-like refractile

structure, only present on seta J5 on left side. Opis-

thosternal chaetotaxy: 5/g, 3Zg, 2S'g, 2/a, 3Za, 0Sa.

Gnathosoma similar to that of adult G. xarifae except

adoral seta aol bifurcate with 2 tapering branches

that lack joining hyaline flap. Appendage chaetotaxy

and solenidiotaxy includes 6 setae on femur | and 4

setae on genu II but otherwise matches that of tri-

tonymph of G. xarifae. On genu IV, dorsal seta

simple and short (x 0.5 length of genu).

June, 1982

Adult

Idiosomal length about 350 (2—very squashed).

Similar to tritonymph except adult attributes as in G.

xarifae. There are further differences. Plasmic seta

z2 with 11 cilia in long-cilia file as adult G. xarifae,

but not its own tritonymph and G. frondifer which

have 14 or 15 such cilia. Some posterior hysteronotal

setae swollen and refractile as tritonymph, but lack

ring-like structures and have relatively longer slim,

tapering tips on some setae (J5, J6, Z5). Appendage

chaetotaxy and solenidiotaxy as adult G. xarifae

except only | seta on trochanter IV as tritonymph of

G. strenzke and xarifae. Differs from its own trito-

nymph in having 5 setae on genu IJ. Dorsal setae on

legs tend to be short and simple compared with long

lanceolate equivalent setae on adults of G. xarifae,

for example, on genu IV where solenidium about

x 1.2 length of genu, associated seta about x 0.5

length of genu (on G. xarifae solenidium about equal

to length of genu, associated seta about X 1.6 length

of genu with hyaline flap so that lanceolate).

Material examined: Holotype tritonymph (N19797),

litter and sparse grass, under Acacia sophorae, Pic-

caninnie Ponds, 3.7.1974, D. C. Lee. Two females

(N19798 and N19799), litter and soil, open Eucalyptus

forest, Cordeaux (34° 08’S, 150° 43’E), New South

Wales, Australia, 9.6.1978, T. Moulton.

Distribution: Australia (Aa). New South Wales:

sclerophyll forest, Great Dividing Range. South Aus-

tralia: Piccaninnie Ponds, coastal closed-scrubland,

ITN (-/8).

Remarks: G. strenzkei is based on 1 tritonymph

collected as part of this study, supported by what are

regarded as conspecific adults from N.S.W. Certain

characters, such as the number of cilia on plasmic

seta z2, suggest that the adults may belong to a

separate closely allied species. But the ring-like

refractile structures, which are not bilaterally sym-

metrical in their distribution amongst some hyster-

onotal setae of the tritonymph, are regarded as a non-

inherited attribute.

The shape of the posterior hysteronotal setae

clearly distinguishes G. strenzkei from urticinus and

xarifae on which such setae are much longer and

slimmer. Because G. xarifae is fully described, dif-

ferences in leg chaetotaxy and shape of hairs on leg

IV have been referred to above, but this is not pos-

sible for G. urticinus. The hysteronotal setae are

much more similar to those of G. frondifer, for which

I was unable to establish the leg chaetotaxy or shape

of hairs on leg IV. The main attribute I use to

distinguish this species from G. frondifer, is the setose

hysteronotal seta Z6. On G. frondifer seta Z6,

SARCOPTIFORMES (ACARI) OF

uthough smaller, is distinctly swollen like J6. Fur-

thermore, although not conspicuous, the swollen hys-

teronatal setae such as J5 on G. frondifer are more

curved in outline with a shorter tapered Lip.

Subcohort RETROFISSURAE a.

Diagnosis: Arihronotina. Usually 2 transverse hys-

teronotal fissures (7R2, THF), sametimes obscure as

when hysteronotal setae so far towards anterior that

seta J4 set well within anterior half af hysteronotum

(Heterochthoniidae) or when notal setae broadly lan-

ceolate, spatulate or umbellate and extent of notal

shields considerably reduced (Phyllochthoninidea), or

only | fissure (7B3) present when prehysteronotal

fissure acutely hinged (Mesoplophorvidea), No hys-

teronotal gland with egress pore between setae 74

und 25, Cowl present, enclosing retracted chelicerae,

Cheliceral digits usually unmodified, sometimes

edentate. Cheliceral setae both usually present and

setose, sometimes ch] absent, rarely ch? enlarged,

lanceolate. One distal plasmic seta on palp tarsus

usually bifurcate, no such seta longer than tarsus.

External malae disjunct. Either | pair (Za/) of adanal

pores present or both pairs absent, On tarsus 1, seta

d1 simple or bifureate. At leasi genua [ and U1 with

1 or 2 solenidia. Peetarsi with cither | claw or 2

subequal claws.

Remarks; The Retrofissurae is a new taxon, Et is

very diverse in form and, as pointed out above in the

remarks on the cohort, the variations reflect similar

variations in the Profissurae suggesting the parallel

evolution of the 2 groups. The name Retrofissurac is

intended to indicate a main attribute thal contrasis

with the Profissurae, i.e, that the transverse hyster-

onotal fissures are usually more posterior, fissure TBI

not being represented, The most heavily weighted

attribute in this study is the presence of disjunct

external malae, a possibly primitive candition alsa

found in the Bifemoratina, but not elsewhere amongst

the Cryptostigmata, On the other hand, certain spe-

cies (see Trichthonius pulcherrintus below) have :

conspievously modified gnathosoma. The presence of

solemidia on jeg genud is an attribute common

amongst Cryptostigmata, contrasting with their

absence on members of the Protissurac. Attributes of

the adult claws also appear lo be a reliable diagnostic

feature although there are exceptions and they diag-

nose lower taxa amongst some other groups of Cryp-

lastigmata, The correlation of the attributes. of these

4 characters, with few exceptions, is considered here

us a good case fora new splitting of a large part of

the Arthronotina into the Retrolissurae and Prolis-

surae.

The Retrofissurae includes 3 superfamilies: Hypo-

chthonioidea Balogh, 1961; Mesoplophorojdea van

der Hammen, 1959; Phyllochthonioidea Traye’, 1967.

The Hypochthonioidea include most members of this

SOUTH ALSTRATIANS Sots j45

subcohort, and since some were callected iq this

study, they are considered further below. The Meso-

plophoraidea. which only include 2 genera and about

7 species, were considered briefly by myself (Lee

1981) under the now disbanded Arthroptyctima. The

Phyllochthonioidea are considered below in these

remarks, allhough none were collected in this study.

The superfamily Phyllochthonioidea Trave, 1967

LO3 is 4 group of minute (length 200-340) pale mites

with large dilated umbellate dorsal setue. Arope-r

chthonius artiodactvius Grandjean, 1948 was

described as the type und only species of Alopo-

chthoniidae with a Holaretic distribution across

southern Canada (No), Mediterranean region (Pm)

and forest zones of Western Europe (Pe), as fur us

the Altai Mountains in Mongolia (Ps). A second spe-

cies (Phylochrhonins aourii Trave, 1967) from the

Ivory Coast (Ew) was established us the type of

Phyllochthontidae. This family has since been recorded

fram broad-leaved forests in southern Eurasia (Fs)

and the far east of the U.S.S.R. (Ps), The 2 genera

ure similar in appearance although their chaetotaxy

differs. For example, 4. artiodacty/us las a gnatho-

soma very similar to that of P. aourti, but lacks 2

setal pairs (aol, clp). The hysterosainal chaetotaxy

is confused because Grandjean (1948) regards seta

J/2 as absent so that the large erectile seta (third in

file J) is labelled “/V. whilst Travé (1967) labels it

“el, There are more hysterosomal setue on P. aowtii

and, if Grandjean (1948) is correct in considering the

second hysteronolal rank to be absent, then setul ile

Sais present on this species. [t may prove preferable

to group these 2 genera in 1 family, bul for the Lime

being | have retained both families because the pres-

ence or absence of setal file Sa has in the pust been

given considéruble weight on the basis that it reflects

the presence or absence of a “preanal segment’ (see

above remarks on Gehypochthoniidae).

The Phyllovhthonjoidea are unusual jn their attri

butes. of 2 of the 4 characters which are Weighted in

the diagnosis of Reirofissurae and this makes its

grouping with the other 2 superfamilies questionuble,

The 2 atiributes are that there is a fissure (? TBI)

close behind seta /| and the pretarsus has 2 claws

rather than | chiw, Bul it should be noted that the

hysteronotal shields are considerably reduced and that

the 2 claws are stout and symmetrical, whilst, when

2 eluws are present on Profissurae species, they (are

known exeeption) are assymmetrical with | hyaline

and slim as for the lateral claws of allied species and

the obhér refractile and stout us for the central chkiw

(rarely the 2 claws are more similar in girth, in which

case the slimmer claw is cilute),

Superfamily HYPOCHTHONIOIDEA

Diagnosis: Retrofissurae. Prehysteronotal fissure rot

aculely hinged, Two narrow Lransverse fissures (7 B82.

336

TB3) dividing hysteronotal shield into 3 parts. Dor-

solateral longitudinal fissure (LB) anterior to seta J5

delineates dorsal margin of pleural shield, ventrally

separate from aggenital and adanal shields. Notal

setae rarely broadly umbellate with J3 extending

beyond posterior edge of soma, if so then ciliate.

Pretarsus with 1 claw.

Remarks: The Hypochthonioidea as used here is a

more extensive superfamily than as delineated by

Balogh (1972) and includes 5 families: Hypochthon-

iidae Berlese, 1910a; Eniochthoniidae Grandjean,

1947a; Brachychthontidae Thor, 1934; Heterochthon-

iidae Grandjean, 1954a; Trichthoniidae n.f. The

Hypochthoniidae and Heterochthoniidae were not

represented in collections for this study and are not

further considered.

Family ENIOCHTHONIIDAE Grandjean

Eniochthoniinae Grandjean, 1947a: 223.

Type-genus: Hypochthoniella Berlese, 1910a.

Remarks: The Eniochthoniidae currently include

only Hypochthoniella so that most further points are

made under that generic heading. The family is closely

allied to the Hypochthoniidae. These 2 families are

usually regarded as the only members of the Hypo-

chthonioidea.

HYPOCHTHONIELLA Berlese

Hypochthoniella Berlese, 1910a: 218. Type designa-

tion (original): “H. (Hypochthoniella) pallidulus

K.”, misidentification of Hypochthonius minu-

tissimus Berlese, 1903 subsequently indicated as

type (van der Hammen 1959: 17).

Arthrochthonius Ewing, 1917: 130. Type designation

(original): ‘“Hypochthonius pallidulus Koch”,

misidentification as type designation for Hypo-

chthoniella,

Eniochthonius Grandjean, 1933: 32. Type designation

(original): ‘“Eniochthonius pallidulus (Mich.)”,

Hypochthonius pallidulus Koch: Michael, 1888:

537 syn. with Hypochthonius minutissimus

Berlese, 1903 (see van der Hammen 1959: 17).

Type-species: Hypochthoniella minutissima (Berlese,

1903: 252),

Diagnosis: Hypochthonioidea. Minute (280-385),

pale mites. Anterior of 2 transverse hysteronotal fis-

sures (7B2) relict, not continuous across mid-notal

line and conspicuously closer to seta /2, than seta J3,

posterior fissure (7'B3) about halfway between seta

J\ and J6. Anterior part of pleural shield (PS1) not

separated off by vertical fissure level with transverse

hysteronotal fissure 7B3. Hysteronotal setae J3 and

Z3 subequal in length to setae J2 and Z2, setose and

never longer than distance between setal bases J/1

and J3, Hysteronotal seta S2 not as near to mid-notal

REC. S. AUST. MUS., 18 (15): 327-359

June, 1982

line as Z1. Pair of merged coxites II] and IV also

merge with each other across mid-sternal line. No

proteronotal eye or eyes in zone between setae j1 and

zl. On tarsus I, bifurcate seta dl with 1 branch

tapering and | expanded into small disc. Adoral file

includes 3 setae of which ao2 has distal teeth.

Material examined: Other than the specimens of

H. minutissima, | syntype of H. borealis (“cotype”,

37F1w-4, Smithsonian Institution, U.S.A.) from New

Hampshire on a slide. This specimen slightly curled

up, dorsal surface uppermost, well cleared and despite

granular integument most attributes of dorsal and

ventral surfaces distinguishable except on gnathosoma.

Probably female since positor has long distal setae.

Distribution: Widespread in temperate regions (Nn,

Na; NTc; Em; Pe, Pm, Ps, Pc; Os; Aa, An) with

some indication that its members commoner in fer-

mantation or deeper soil layers, where there is a

substantial leaf-litter layer as in forests, woodland or

scrubland.

Remarks: The most data on Hypochthoniella is to

be gleaned from Grandjean’s work (see below) on the

type-species. The brief description of H. borealis

clearly distinguishes it from the type-species, stating

that it is larger (about 385) and has a notal plasmic

seta z2 with only 4 or 5 short cilia (their length is

about half that of the broadest width of the seta). I

was uncertain as to whether or not H. borealis was

grouped in the correct genus, but the disposition of

somal shields is very much as on H. minutissima and

both species have a large pore posterior to the ace-

tabulum for leg IV.

Two species are included in Hypochthoniella: H.

minutissima (Berlese, 1903); H. borealis (Jacot, 1939).

Hypochthoniella minutissima (Berlese)

(Fig. 4)

Hypochthonius pallidulus Koch: Michael, 1888: 537.

Hypochthonius minutissimus Berlese, 1903: 252.

Hypochthonius (Hypochthoniella) pallidulus Koch:

Berlese, 1910a: 218

Eniochthonius pallidulus: Michael not Koch: Grand-

jean, 1933: 32,

Eniochthonius grandjeani van der Hammen, 1952:

13.

Female

Dull ochre, becoming orange at shield edges. Inte-

gument appears granular even in well cleared speci-

mens. Refractile parts (external malae, cheliceral

extremities, setae and claws) at least as pale as general

integument. Idiosomal length 315 (25 ex Piccaninnie

Ponds, 280-360); appendage lengths (for 320 not con-

taining an egg)—ch 20, pa 50, J 115, 77 110, 7/7 95,

IV 115; femur breadths—pa 10, J 17.5, If 17.5, IT

17.5, 7 15, IV 15. Prehysteronotal fissure appears

SARCOPTIFORMES (ACARI) OP SOUTH AUSTRALIAN SOILS 337

hinged and expandable so that, in relation to hyster-

osomna, propodasama can move up and down through

about 10°, and backwards and forwards over about

20um. Anterior transverse hysteronotal fissure {7B2)

relict, appearing to be shallow furrow on inner surface

of integument, Posterior transverse hysteronotal fissure

(TB3) hinged. In specimen illustrated (Fig. 4), pos-

terior edge of anterior shield consists of both dorsal

complete line and ventral broken, line level with seta

J3, whilst anterior edge of the posterior shield (which

can be in forward position, level with seta J3) in

backward position level with dorsal postenor edge of

anterior shield. Specimens contain up to 3 food boli,

anterior | usually diffuse, including what appears to

be fungal hyphae and spores. All positors obseryed

have stout long distal setae as an females with eggs,

50 all adults regarded as female, Amongst 25 regis-

tered females examined in detail, 15 without eggs,

whilst 10 contain single large (length about | 30)

ellipsoidal egg. Average length of females with eggs

1() greater than those without.

Material examined: Twenty-five females

(N1979150-N1979174), litter and sparse grass, under

Acacia sophorae, Piccaninnie Ponds, 37.1974 and

208.1975, D. C, Lee, One female (N1979175), litter

and sparse moss, under Eucalyptus incrassata, Fer-

ries-McDonald, 206.1974, D. C. Lee. One female

(N19791 76), litter and sparse moss, under Eucalyptus

obliqua, Mi Lofty, 9.5.1974, D. C. Lee.

Distribution: Widespread, as for genus, South Aus-

tralia: Ferrries-MeDonald, mallee-broombush open-

scrubland, | (1/8); Piccaninnie Ponds, ¢oastal closed-

scrubland, 60 (6/8), Mt Lofty, sclerophyll open-for-

est, | (1/8).

Remarks: Confusion over the nomenclature relat-

ing to this species and its genus results from Mypo-

chthonius paltlidulus Koch, 1836; 3 (20) being the

nymph of H, rufulus Koch, 1836: 3 (19) (syn, Grand-

jean 1933: 32), whilst Hl. pallidulus Koch; Michael,

1888; 537 is Hypochthoniella minutissima.

The most informative descriptions of A. miinutis-

sima (as Eniochthonius pallidulus) are by Grandjean:

development of ventral region of soma (Grandjean

1933) development of lateral region of soma (Grand-

jean 1934): structure of seta d1 on tarsus | (Grand-

jean 1941); setation of genua (Grandjean 1942); gna-

thosonya (Grandjean 1957); solenidiotaxy (Grandjean

1964). The external morphology of this species, and

so this family, is still incomplete in that for example,

the leg chaetotaxy is not fully described.

Family BRACHYCHTHONUDAE Thor

Brachychthoniidae Thor, 1934: 115.

Type-genus: Brachychthonius Berlese, 1910a-

Diagnosis; Hypochthonioidea. Minute (130-270),

pale mites. Two complete transverse hysteronotal fis-

sures (782, 783), anterior fissure (7B2) never con

spicuously closer to seta J2 than ta J3, posterior

fissure (783) about halfway between seta J1 and J6

or posterior to that position. Anterior part of pleural

shield (PSL) separated off by vertical fissure level

with transverse hysteronotal fissure 763 Hysterono-

tal setae J3 and Z3, not more than * 1,5 length of

setae J2 ot Z2, setose, ciliate or lanceolate and never

longer than distance between setal bases J1 and J3.

Hysteronotal seta 52 as near to mid-notal line as Z1

or nearer, Pair of merged coxites IT and TV do not

merged with each other across mid-sternal line, No

proiernotal eye or eyes jn zane between setae /! and

21. On tarsus |, seta d1 simple, Adoral file usually

includes | seta (a02), reduced seta ao3 rarely present

(Neabrachyehthonius).

Material examined: Other than species collected

during present study. brachychthoniid mites previ-

ausly recorded from South Australia (Womersley

1945) also considered. Two species represented in

both collections, 2 in Womersley’s work alone and 3

in just this study, making 7 species altogether.

Distribution. Widespread, probably cosmopolitan

(Niedbala 1972b 1977, Chinone 1978) and if wide

variety of habitats especially moss, plant litter, and sail

Remarks: The Brachychthoniidae are by far the

largest family in the Arthronotina whilst exhibiting

relatively little morphological diversity. Balogh (1972)

included 5 genera. | fallow a classification of European

brachychthoniids (Moritz 1976a, [976b) including 10

genera which, although it may be excessively divided,

works Well in classifying the material before me. One

problem in classifying brachychthoniids is that the

descriptions of many included species are toa brief.

For example, the leg chaetotaxy is known for very

few Species, as tabulated with the redeseription of

Brachychovhthonius lydiae by Reeves and Marshall

(1971), which refers to all of the more compleie

descriptions of brachychthoniids.

Niedbala (1972a, 1973) recognised 7 genera

amongst which he attempted to establish phylogenetic

relationships. Eobrachychthonius was regarded as (he

Most primitive because of its 4 subnotal shields, whilst

other genera Were tore or less advanced depending

on the degree of either loss or fusion of these shields

with the notal shield or each other. Furthermore, the

family was regarded as having evolved along 2

branches: Poecilachthonjus and Svnchthonius consti-

tuting a minor branch in which 4 Jurrow indicates

the line of merging of 2 subnotal shields (SNS2 and

SNS3), The similarities of Poecilochthonius to Bra-

vhychachthonius and of Synachthonius to Liochthonius

were regarded as “convergent similarity” and not due

ia “ways of evolution”

338 REC. S. AUST. MUS.,

| have found it useful to newly group the 10 genera

of Brachychthoniidae in 3 genus-complexes as follows:

1) Brachychthonius-complex— Brachychochth-

ochnius Jacot, 1938; Brachychthonius Berlese,

1910a; Eobrachychthonius Jacot, 1936; Poecilo-

chthonius Balogh, 1943,

2) Liochthonius-complex—Liochthonius van der

Hammen, 1959; Mixochthonius Niedbala, 1972a;

Spachthonius van der Hammen, 1952; Verach-

thonius Mortitz, 1976a.

3) Neobrachychthonius-complex— Neobrachy-

chthonius Moritz, 1976b; Neoliochthonius n. name

(for Paraliochthonius Moritz, 1976a).

The genera are considered below only if their rep-

resentatives have been collected in South Australia,

on the other hand all the genus-complexes are con-

sidered. The 2 characters heavily weighted in diag-

nosing the genus-complexes are the presence or

absence of a fissure separating off the anterior subnotal

shield (SNS1) and the position of hysteronotal seta

Z2. Normally the position of the anterior 2 ranks of

hysteronotal setae of Arthronotina ts that the 3 files

are well spaced and sub-parallel, but in the Brachy-

chthoniidae, seta S2 is always displaced towards mid-

notal line, whilst Z2 is sometimes similarly displaced.

Key to Genera of Brachychthoniidae

|. Hysteronotal seta Z2 displaced towards mid-notal line, base at

least X 2 its own diameter away from line joining setae S2

and Z3. Anterior subnotal shield (SNS1) may or may not be

separate from hysteronotal shield 2

Hysteronotal seta Z2 not displaced towards mid-notal line, base

on or close to line joining setae S2 and Z3. Anterior subnotal

shield (SNS1) separate from hysteronotal shield. Brachy-

chthonius-complex 3

2. Anterior subnotal shield (SNS1) separate from hysteronotal

shield. Neobrachychthonius-complex 6

Anterior subnotal shield (SN.S1) merged with hysteronotal shield.

Liochthonius-complex 7

3. Longitudinal furrow delineates, without separating off, marginal

parts of hysteronotal shield bearing setae Z1, S2, Z2, 23, Z4,

Four separate subnotal shields present, Adanal seta Za2 con-

spicuously stouter than Zal and Za3. Pair of merged coxites

Il] and IV also merge with each other across mid-sternal

line Eobrachychthonius

No such longitudinal fissure on hysteronotal shield as above.

Two or 3 separate subnotal shields present. Adanal setae Zal

and Za2 similar, either both slim like Za3 or both stout. Pair

of merged coxites II] and IV separated by fissure along mid-

sternal line 4

4. Adanal setae in file Za similarly slim, although Za2 may be

slightly longer than other 2 setae Brachychthonius

Anterior 2 adanal setae in file Za conspicuously longer and/or

stouter than Za3 5

5. Three subnotal shields (SNS4 present). Adanal seta Jal on

striated cuticle or narrow oval shield Brachychochthonius

Two subnotal shields (S/'V.S4 absent). Adanal seta Jal on broadly

triangulate shield Poecilochthonius

6. Two palpocoxal setae (clp absent). Adoral seta ao2 tapers to

distal end. Proteronotal plasmic seta z2 broadens distally into

18 (15): 327-359 June, 1982

a subspherical ciliate ball. Subnotal shield SNS3 merges with

hysteronotal shield Neoliochthonius

Three palpocoxal setae (clp present). Adoral seta ao2 stout and

distally dilated. Seta z2 broadens distally into oval ciliate ball

with longitudinal axis at least X 2 greatest transverse axis.

Subnotal shield SNS3 separate from hysteronotal

shield Neobrachychthonius

7. Subnotal shield SNS3 merges with hysteronotal shield or absent.

Posterior setae in hysteronotal file J spaced normally, /6

nearly level with Z6 in relation to posterior margin of hyster-

onotal shield. Adoral seta ao2 slim and tapering

Liochthonius

Subnotal shield SNS3 separate from hysteronotal shield. Posterior

setae in hysteronotal file J almost as Liochthonius or compacted

forward, J6 level with point midway between Z5 and Z6 in

relation to posterior margin of hysteronotal shield. Adoral seta

ao2 \anceolate or parallel sided with blunt distal end 8

8. Hysteronotal setae conspicuously ciliate. Seta /6 more nearly

level with seta Z6 in relation to posterior margin of hysteronotal

shield than to point midway between Z5 and Z6. Adoral seta

ao2 broadly lanceolate Mixochthonius

Hysteronotal setae never conspicuously ciliate. Seta /6 nearly

level to point midway between ZS and Z6 in relation to

posterior margin of hysteronotal shield. Adoral seta ao2 parallel

sided with blunt distal end 9

9. Adanal seta Za2 conspicuously longer and stouter than Zal and

Za3. Three setae in aggenital setal file Jg (/Jg4 absent). Two

setae on both genu III and IV Synchthonius

Adanal seta Za2 and Zal similar, either both slim or both stout,

although Za2 may be slightly longer. Four setae in aggenital

setal file J/g (Jg4 present), Three setae on both genu III and

lV Verachthonius

BRACHYCHTHONIUS-complex

Diagnosis: Brachychthoniidae. Anterior subnotal

shield (SNS1), bearing seta S1, separate from hys-

teronotal shield. Hysteronotal setae Z1, S2, Z2 andZ3

in nearly straight line parallel to lateral margin of

notal shield because seta Z2 not displaced towards

mid-notal line. Subnotal shield SNS3 separate from

hysteronotal shield. Adoral seta ao2 parallel-sided

with blunt distal end. Three setae on both genu III

and IV (1 ventral seta present). Coxite IV bearing 4

setae.

Remarks: The Brachychthonius-complex is made

up of 2 groups: Eobrachychthonius and the other 3

genera. Eobrachychthonius is not closely allied to the

other genera, but it is useful to group them together

because of the similar position of hysteronontal seta

Z2, 1 of the most important characters referred to in

the briefer descriptions. Eobrachychthonius is hardly

sculptured at all on its notum, but its hysteronotal

shields have a marginal strip delineated by a furrow.

The other 3 genera usually have strongly sculptured

hysteronotal shields with a marginal ridge. Possibly

these 2 types of marginal zonation are reflected in

the similar lack of displacement of seta Z2 towards

the mid-notal line so that it remains on the margin.

The marginal ridges also appear in this genus-complex

to be correlated with the position of the posterior

setae in file J, since they often end posteriorly in a

SARCOPTIPORMES (ACARI) OF SOUTH ALSTRALEAS SOILS 4

pair of tubercles bearing setae /6, which in turn are

usually more antertorad, file J being compacted for-

ward,

Eohrachychthoanius montanus Hammer, 1952 does

not (it into that genus wor the rest of this penus-

complex, but is tentatively newly combined below

with Ferachthonius,

Pour genera are included in the Brachychthonius-

complex: Brachychthanius Berlese, 1910a, Brachyp-

chochihonius Jacot, 1938; Eobrachyehthonius Jacot,

1936: Poecilachthonius Balogh, 1943 Only Brachy-

chochthonius and Poecilochthonius fave been col-

lected in South Australia und ure considered below.

BRACHYCHOCHTHONIUS SJacot

Brachychochthonius Jacot, 1938: 130. Type desig-

nation (original): “Brachvehochthonius jugatus

sp.nov.”

Selinickochthonius Krivolutsky, 1964; 935, Type des-

ignation (original): “Brachyehthonius celawaten-

vis Sellnick’',

Brachvehthonius Berlese: Chinone and Aoki, 1972:

223 (in part). Type indication, “Brachychthonius

berlesei Willmann, 1928", correct identification

of “A, brevis (Mich): Berlese 1910a (sce van

der Hammen 1959; 19),

Type-species: Brachychochthonius jugalus Jacot,

1938; 130,

Diagnosis: Brachychthonins-complex. — Hystero

notal shields without marginal longitudinal furrows,

so that shield bearing setae Z!, 82, 22, 73 and 24

not delineated from more central part. Three subnotal

shields (SNS1, SNS3, SNS4) separate from hyster-

onotal shields. Pair of merged coxites II] and IV

separated by fissure along mid-sternal line. Adanal

seta Jal on striated cuticle or narrow oval shield.

Anterior 2 adanal setae in file Za conspicously longer

and/or stouter than Za3. Posterior setae in hyster-

onotal file J compacted forward so that /6 level with

seta 25 in relation to posterior margin of hysteronotal

shield.

Distribution: Widespread, probably cosmopolitan.

Remarks: Brachychochthonius species separate out

best under Sellnickochthanius in the keys of Balogh

(1972), It is difficult to establish how many nominal

species should be included in this genus because a

number of possible members are known only by char-

acters of their notal surface, which is insufficient to

distinguish them from Brachychthonius and Poecil-

ochthonius, Also, in the comprehensive list by Nied-

bala (1972), species of these 3 genera are all grouped

in Brachychthonius. When Moritz (1976b) established

the concept off this taxon as used here, he included

12 speeies from Europe and later Chinone (1978)

included a further 5 species from Japan also Brachy'-

chochthonius lydiae Jacot from North America ts

returned to this genus. Therefore, with at least 15

species. il may be the largest genus in the Brachy-

chthonius-complex. Two species (8. el/sesneadensis

and A. ericoides) in Brachvehochthonius have been

collected in South Australia and are considered below.

Brachychochthonius elsosneadensis (Hammer)

(Fig. nane)

Brachychthonius elsosneadensiy Hammer, 20,

Brachyehthonius elsosneadensis Wammer: Chinone

und Aoki, 1972: 229.

Female

Shields either dull orange or almost straw-coloured

or pale ivory white. Durker orange specimens some-

times with adhesive, dirly exudate conspreuous al

posterior end of soma, Idjosomal length 165 (25 ex

Piccaninnie Ponds, 142.5-177.5); appendage lengths

(for 165 nul containing egy)—ch 5. pu 20, 7 47.5, 1

40, 111 45, 7 S75: femur breadths—pa 5S, J 14. 1

Vl. 7 11. 7 13, Adults from Met. Lotty and Katt

Hill with idiosomal length within 162.5-182,5, All

posilors observed appeared similar to those of speeci-

mens containing ceg, therefore all adults regarded as

female. Amongst 25 females (ex Piccaninnic Ponds),

8 included 1 or 2 boli, Boli light or dark brown,

homogencously granular, level with or posterior to

genital shields. Nine included single ellipsoidal cee

with length within 70-92.5 Females with epgs

uppeared futter but not longer than those without

cee.

Material examined: Twenty-five females

(N1979177-N1979201), litter and sparse grass. under

Acacia sophorae, coastal scrubland, Piceaninnic

Ponds, 3.7.1974 and 20.8.1975, D. C. Lee Five

females (N1979202-1979206) litter und sparse moss,

under Eucalyptus obliqua, Mt. Lofty, 95.1974, D.

C. Lee, Five females (N1979207-N1979211) liver,

under Pinus pinea, Knowl Hill, 225.1974, D.C, Lee.

Distribution, Argentina (NT¢), Japan (Pe), South

Australia (Aa) Sauth Australia: Pidcaninnie Ponds.

coastal clased-scrubland, 35 (8/8), ML, Lofty, selero-

phyllopen-forest, 13 (4/8); Knott Hill, oultivated pine

forest, 6 (2/2).

Remarks: As the hysteronotal setae JT and /2 are

selose, they differ from the larger seta /3 that has

lateral hyaline flaps giving it a lanceolate shape,

which distinguishes &. e/yosneadensis from all other

possibly congeners except 8. elisabethae (Mahunka,

1974: 211) from Rhodesia (Ee). 8. e/sosneadensis is

distinguishable from &. elfsabethae in having & cou-

spicuous hysteronotal pare Af] and narrower

(breadth:length=0,3 or less:]) seta ./3.

340

Brachychochthonius cricoides (Weis-Fogh)

(Fig. none)

Brachychthonius cricoides Weis-Fogh: 1948: 269.

Brachychthonius cricoides Weis-Fogh: Evans, 1952:

236.

Brachychochthonius cricoides (Weis-Fogh: Moritz,

1976b: 287).

Female

Shields pale straw-coloured. Idiosomal length 142.5

(5 ex Mt. Lofty and Knott Hill, 135-147.5); appendage

lengths (for 145 ex Mt. Lofty)—ch 7.5, pa 20, 1 45,

1 40, IH 42.5, IV 47.5; femur breadths—pa 6.25, J

10, 77 8.75, JIT 8.75, IV 10. Specimens from Knott

Hill have weak notal sculpturing, only a faint ridge

associated with setal file J, a scalloped transverse

ridge between tubercles bearing setae /6, and a line

between setae zl and sl. Specimens from Mt. Lofty

have more complex notal sculpturing as illustrated

for this species by Moritz, 1976b, but not as conspic-

uous as for B. elsosneadensis. Hysteronotal pore hf

large, with conspicuous encircling refractile ring. One

pale homogeneous granular bolus observed. Positors

relatively small, involuted tubes forming a ring (diam-

eter 10-12.5). No eggs seen, all adults assumed female.

Materials examined: Three females (N1979212-

N1979214); litter and = sparse moss, under

Eucalyptus obliqua, sclerophyll forest, Mt. Lofty,

9.5.1974, D. C. Lee. Two females (N1979215,

N1979216), litter, under Pinus pinea, Knott Hill,

22.5.1974, D. C. Lee.

Distribution; Denmark, Czechoslovakia, England,

Germany, Italy, Sweden, Poland (Pe); Asiatic USSR

(Ps); South Australia (Aa). South Australia: Mt.

Lofty, scleropyll open-forest, 3 (2/8); Knott Hill, cul-

tivated pine forest, 2 (1/2).

Remarks: B. cricoides is minute and pale, with

faint sculpturing, short setose notal setae and a very

conspicuous hysteronotal pore (Af1). The following 5

species are similar: B. aokti (Chinone, 1974); B. jacoti

(Evans, 1952); B. mivauchii Chinone, 1978; B. rotun-

datus Hammer, 1958; B. suecicus Forsslund, 1942.

B. cricoides is more precisely distinguishable in that

it has rostral teeth, lacks fine punctuations in the

median delineated areas of the proteronotal sculptur-

ing, proteronotal seta z1 is without cilia and hyster-

onotal setae are short (J2 is about 0.35 & the distance

between its bases; /3 is about 0.75 & the distance

between its bases).

POECILOCHTHONIUS Balogh

Poecilochthonius Balogh, 1943: 22. Type designation

(original): ““Poecilochthonius italicus (Berl.)”.

Poecilochthonius Balogh: Moritz, 1975b: 248.

Type-species: Poecilochthonius italicus (Berlese,

1910a: 220).

REC, S. AUST, MUS., 18 (15): 327-359

June, 1982

Diagnosis; Brachychthonius-complex. Hysteronotal

shields without marginal longitudinal furrows, so that

shield bearing setae Zl, $2, Z2, Z3 and Z4 not

delineated from more central part. Two subnotal

shields (SNS1, SNS3) separate from hysteronotal

shields. Pair of merged coxites III] and IV separated

by fissure along mid-sternal line. Adanal seta Jal on

broadly triangulate shield (width of broad anterior

end =X 2.5 length of Jal), Anterior 2 adanal setae

in file Za conspicuously longer and/or stouter than

Za3. Posterior setae in hysteronotal file J compacted

forward, so that seta J6 in zone (breadth =x 0.5

distance Z5-Z6) centred on point midway between

ZS and Z6 in relation to posterior margin of hyster-

onotal shield.

Distribution: Widespread, Greenland (Nn); Argen-

tina (NTc); Austria, Germany, Hungary (Pe); Italy,

Yugoslavia (Pm); Japan (Pc); South Australia (Aa).

Remarks: Poecilochthonius species separate out to

the couplet including Brachychthonius and Syn-

chthonius in the key of Balogh (1972). No mites in

the present study are grouped in this genus, but a

species has been recorded from South Australia. Mor-

itz (1976b) excludes 1 species, Brachychochthonius

hungaricus (Balogh, 1943), originally included in this

genus.

Three species are included in Poecilochthonius: P.

italicus (Berlese, 1910a); P. parallelus (Womersley,

1945) n. comb.; P. spiciger (Berlese, 1910a).

Of the 2 specimens of P. parallelus from South

Australia referred to by Womersley (1945), only the

holotype (N1979297) has been found. Even after

remounting, it remains strongly squashed around an

axis that cuts through the region of setal file S on |

side and Za and Zg on the other side. Despite this,

3 characters were observed on | side: subnotal shield

SNS4 is absent: shield bearing seta Jal is broadly

triangulate; adanal setae Zal and Za2 are similar to

each other and conspicuously stouter than Za3. Attri-

butes of the hysteronotal sculpturing which gave rise

to the name parallelus (i.e. the straight median and

lateral ridges almost devoid of sculpturing in between)

distinguish this species from its congeners.

LIOCHTHONIUS-complex

Diagnosis: Brachychthoniidae. Anterior subnotal

shield (SNS1), bearing seta S1, merged with hyster-

onotal shield. Hysteronotal setae Z1, $2, Z2 and Z3

not in nearly straight line parallel to lateral margin

of notal shield because seta Z2 displaced towards

mid-notal line. Subnotal shield SNVS3 separate, merged

with hysteronotal shield or absent. Adoral seta ao2

either parallel sided with blunt distal end, setose or

lanceolate. Two or 3 setae on either genu III or IV

(ventral seta present or absent). Coxite IV bearing 4

setae,

SARCOPTIFORMES (ACARI) OF SOUTIL AUSTRALIAN SOILS 441

Remarks; Exeept for Syachthonius, members of

the Liechthonius-complex are hardly sculptured at

all on their dorsal integument. Svachthonius does

have seta 42 displaced towards mid-notal line as on

the unsculptured genera, but seta J6 is displaced

anteriorad as for the sculptured genera of the Bra-

chychthonius-complex. Verachihonius also has an

anteriorad displacement of seta /6, although species

have an almost smooth dorsal integument, Yet, there

is a lateral ridge posteriorly around setae 75 and 95,

and sometimes a fainter 1 reaching J6. Possibly this

ridge indicates that ancestors of this genus were more

sculptured with the anteriorad displaced seta J6 on

a tubercle,

Four genera are included in the Liochthonius-com-

plex: Liachthonius van der Hammen, 1959; Mixo-

chthonius Niedbala, 1972a, Syrehthonius yan der

Mammen. 1952: Verachthonius Moritz, 1976a. Only

Lochthonius and Ferachthonjuy have been collected

in South Australia and are considered below.

LIOCHTHONIUS van der Wammen

Brachychthonius Berlese: Strenzke, 195}; 235. Type

indication; none, but & of 12 included species

grouped in Liovhrhanius by Moritz (1976a), other

4 species not in Brachyehrhonius, Assume original

designation of “8, brevis Mich.”, although mis-

identification (Berlese. 1910a) of Brachyehthon-

ius berlesei Willmann, 1928 (Type of

Brachvehthonius Berlese, 19100).

Livehthonius yan der Hammen, 1959; 19. Type des-

ignation (original); “Brachychthonius perpusillis

(redescription by Forsslund, 1942, p. 4, fig. 4).

Livchthonius van der Hammen: Moritz, 1976u; 38.

Type indication: “H\poehthonius brevis Michael,

1888 non Berlese 1910" (syn. Bravhychthanius

perpusillus Berlese, 1910a),

Type-species: Liovhthonius brevis (Michael, [888:

539).

Diagnosis; Liochthanjus-complex, No subnotal

shields separate from hysteronotal shields. Hysteron-

otal setae never conspicuously ciliate, Posterior setae

in hysteronatal file / not conspicuously compacted

forward, seta /6 nearer level of seta Z6 than point

midway between 25 and 26 in relation to posterior

murgin af hysteronotal shield, Notal shields relatively

smooth with few seattered circular markings. Only

udoral seta (av2) setose, tapering to distal point.

Adanal seta Za2 conspicuously longer and stouter

than other 2 setae (Za), 2a3), Three setae on both

genu II] and IV (ventral seta present), Aggenital file

Ja with 4 setae (/24 present).

Distributions Widespread, probably cosmopolitan.

-Reniarks: Liochthonius separates out correctly in

the key of Balogh (1972). Species of all genera without

a sculptured notum and with a centralward displaced

seta Z2 have until recently (1972 and alter) been

grouped in Livch/honius despite the fact that, amongst

a vumber of differences fram the Lype of thal genus.

they have at least | separate subnotal shield: either

SNS\ (Neoliochthonius) or SNS3 (Mixochthanius,

Verachthonius) or both (Neobrachpehrhonits). This

illustrates the fact thal many brachychthoniids have

until recently been classified on the basis of only a

dorsal view of their soma. On the other hand, because

of the strongly sculptured notal shields, the type of

Spnehthonius was previous grouped in Brachychech-

thonius, despite a centralward displaced seta 22.

Niedbala (1972b) included 40 species in Livch-

thonius, but it is difficult to establish for certain haw

inany nominal species should now be included in this

genus, Moritz (19762) includes 22 species from Europe

and Chinone (1978) includes some olf these plus a

further & species from Japan. So that, with a minimum

of 30 nominal species al the moment, if ts likely that

this is the largest genus in the Brachyehthoniidae.

Niedbala (1977) used 37 morphological charucters

to calculate valucs of similarity amongst 19 species

of Liechthonius. Whilst these species clustered into

4 separate groups, Niedbala concluded that establish-

ing subgeneric taxa would be excessive splitting of

the penus.

Moritz (1976a) divided Livehthoniis into 4 specics-

complexes, which do nol correspond to the groups

noted by Nicdbala (1977) and are mainly delineated

by attributes of the notal setac, | have found it useful

to use these species-complexes and a derivation from

Moritz’s key is presented below.

Key to species-compleaes of Liochihoniuy

|. Hysteronotal setae broad, similar breadth ta plasmic proteronpti|

yet 22 (Jl = % 0.75-% 125 23) forridus-coniplex

Hysteronotal setac setose, conspicuously slimmer than dilated

distul half of plasmie proteronotal seta 22 iJ] =leys than x

OTS 22) a

2, Plasmte proteronutal seta c2 distally spindle-shaped. with single

terminal point and bearing 3 to 5 lateral files of 4 to 8 cilia

hrevi-conyples

Plasmic proteronotal setn 22 distally bearing 2 lamellac, each

Will separate leciinal puiqk wad 16 to 20 small maryyoul eis

3, Notal setae mostly (always 2) and J3) setose, rounded in crans-

verse section penduncularis-complex

Notal setae mostly (always ot and J3) laieeolate, with niprow

lateral lamellae, V-shaped iq trinsverse section

lappanicuscompley

Members of the /apponicus-complex have not been

collected in South Australia, Species of the brevis-

complex and horridus-complex have been collected

in the present study and are considered below, |

species under each species-complex heading. A single

species in the penduncularis-complex was collected

previously from South Australia and is considered in

the remarks on that species-complex,

342 REC. 8. AUST, MUS., 18 (15): 327-359

brevis-complex

Liochthonius simplex (Forsslund)

(Fig. none)

Brachychthonius simplex Forsslund, 1942: 7.

Brachychthonius cf. perpusillus Berl.: Womersley,

1945: 219.

Liochthonius simplex (Forsslund): Moritz, 1976a: 50.

Female

Shields straw-coloured. Idiosomal length 162.5 (25

ex Knott Hill, 155-170); appendage lengths (for

162.5)—ch 14, pa 35, 185, 1155, I] 50, IV 75; femur

breadths—pa 8, / 13, 17 11, W711, IV 13. Specimens

from Mt. Lofty similar in size, but 5 specimens from

Normanville (Womersley 1945) sometimes larger

(idiosomal length 152.5-205, previously recorded as

235, Womersley 1945), probably because squashed.

No specimens contain eggs. Proteronotal plasmic setae

z2 bears 5 lateral files of 4 to 6 cilia.

Material examined: Twenty-five females

(N1979272-N1979296), litter, under Pinus pinea,

Knott Hill, 22.5.1974; D. C. Lee. Five females

(N1979298-N 1979302) litter and sparse moss, under

Eucalyptus obliqua, Mt. Lofty, 9.5.1974, D. C. Lee.

Five females (N1979303-N1979307) form Norman-

ville (Womersley 1945).

Distribution: Greenland (Nn); Czechoslovakia,

England, Finland, Germany, Hungary, Poland, Sweden

(Pe); Bulgaria (Pm); Asiatic USSR (Ps); Japan (Pc);

South Australia (Aa). South Australia: Mt. Lofty,

sclerophyll open-forest, 61 (6/8); Knott Hill, cultivated

pine forest, 113 (2/2); Normanville (Womersley 1945).

Remarks: Moritz (1976a: 41) comments on L. cf.

perpusillus (Berlese: Womersley, 1945) under Lioch-

thonius brevis (Michael, 1888) stating that it is larger

than the latter and with distinctly shorter dorsal setae.

While I agree that this material should be grouped

in the brevis-complex, Womersley’s (1945) measure-

ments for it are misleading and it fits better under

L. simplex than L. brevis according to Moritz’s

(1976a) own descriptions. It should be noted that the

proteronotal plasmic seta has to be examined carefully,

since at certain angles it can appear to have the

attributes of the /apponicus-complex, in which case

this species might be grouped under L mollis (Ham-

mer, 1958). Regarding the description of L. simplex

by Chinone (1974), this appears to fit the description

of L. leptaleus Moritz, 1976a rather than that of L.

simplex.

horridus-complex

Liochthonius fimbriatissimus (Hammer)

(Fig. none)

Brachychthonius cf horridus Sellnick: Womersley,

1945: 220.

June, 1982

Brachychthonius fimbriatus Hammer, 1958: 14 (not

Jacot, 1936: 248).

Liochthonius fimbriatissimus Hammer, 1962: 15.

Liochthonius fimbriatissimus Hammer: Hammer,

1966: 13.

Female

Shields straw-coloured. Idiosomal length 170 (25

ex Piccaninnie Ponds, 162.5-185); appendage lengths

(for 177.5)—ch 13, pa 35, 1 72.5, 11 60, 1/1 60, IV

75; femur breadths—pa 10, / 17, // 14, 17714, IV 16.

Form of some notal setae difficult to assess. Some

notal setae (/1, z1, /6, 76, S6) lanceolate with smooth-

edged hyaline flaps. Other setae (e.g. /2-J5) appear

similar, but hyaline flap sometimes wrinkled so that

edge looks serrate. In contrast, seta S1 has hyaline

flap of | side broken up into file of 6 distinct cilia.

Other setae lanceolate in outline, but uncertain if

edge ciliate or wrinkled. Positors present in all spec-

imens examined and assumed females although no

eggs seen. Of 25 specimens ex Piccaninnie Ponds, 16

contain | or 2 boli made up of material including

distinct cellular forms.

Material examined: Twenty-five females

(N1979218-N1979242), litter and sparse grass, under

Acacia sophorae, coastal scrubland, Piccaninnie

Ponds, 3.7.1974, D. C. Lee. Five females (N1979243-

N1979247), litter and sparse moss, under Eucalyptus

incrassata, Ferries-McDonald, 20.6.1974, D. C. Lee.

Five females (N1979248-N1979252), litter, under

Banksia ornata, Tamboore, 4.7.1974 D. C. Lee. Five

females (N1979253-N1979257) litter and sparse moss,

under Eucalyptus obliqua, Mt. Lofty, 9.5.1974, D.

C. Lee. Five females (N1979258-N1979262), litter,

under Pinus pinea, Knott Hill, 22.5.1974, D. C. Lee.

Seven females (N1979263-N1979271) from Norman-

ville (Womersley 1945).

Distribution: Argentina (NTa), Tierra del Fuego

(Sm), New Zealand (An), South Australia (Aa). South

Australia: Ferries-McDonald, mallee-broombush open-

scrubland, 6 (3/8); Tamboore, mallee-heath tall open-”

shrubland, 68 (6/8); Piccaninnie Ponds, coastal closed-

scrubland, 658 (8/8); Mt. Lofty, sclerophyll open-

forest, 45 (4/8); Knott Hill, cultivated pine forest, 216

(2/2); Normanville (Womersley 1945).

Remarks: The leg I of L. fimbriatissimus is shorter

and stouter than that of L. simplex described above.

The top end of the range of idiosomal lengths given

for L. fimbriatissimus (ex Piccaninnie Ponds) is prob-

ably too high since the specimens all belonged to a

group which had been in lactic acid for a long time

and were distended with extended chelicerae.

Moritz (1976a: 99) pointed out that L. cf horridus

(Sellnick: Womersley, 1945) has shorter and stouter

setae than L. horridus (Sellnick, 1928) and he

excluded if from L. horridus. This material has been

newly identified here as L. fimbriatissimus and the

SARCOPTIPORMES (ACARL QF SOUTH ALSTRALIAN SOILS 545

many specimens from the present study have been

similarly identified. They all look yery much more

like the illustration of this species (Hammer 1958:

fig. 2) from Argentina rather than Womersley’s ilus-

tration (Womersley 1945; fig, [D-G), [lammer (1958)

does figure the notal setae as almost parallel-sided

compared lo the more convex edges of those of the

South Australian specimens, but this may result from

the lateral hyaline Plaps being angled up at their

edges,

pedunculuris-complex

Remarks: Members of the peduncilaris-complex

were not collected in this study, On the other hand,

Livchthonius longipilus (Womersiey, 1945: 220)

mcotb. from Normanville, South Australia (holotype,

N1979308; paratypes N1979309-N1979314) is

grouped jn this species-complex. The illustration

(Womersley 1945: fig 10) of the proteronotal plasmic

selu 22 18 inaccurate; 1b should be as for this species-

complex, The following 2 more reeently described

species are very similar and | or both may be syn-

onyimous With £. loagipilus.

Livohthonins ovellatus, (Hammer, 1958: 19) (newly

identified as 2. longipilus) from Argentina is not L.

ocellatus (Hammer, 1952: 19) (syn. L. Ayiserieinus by

Moniz, 1976a: 66) from Cunada. The subcircular

markings and other characters on the natal surface

ol L. dongipilus are similar to thoge illustrated for

the Argentinian specimens. Therefore, since the

Argentinian material has no valid name it is pro-

visionally grouped under ft. Jongipilys,

Livehthonius perfusorius Moritz, |976a: 93 from

Germany is very similar lo &. lengini/us although

there are fewer notal markings illustrated.

VERACHTHONIUS Moritz

Ferachthonius Moritz, 1976a: 112. Type designation

(original): “Brachyehrhonius laticeps Sitrenzke,

1951”,

Type-species: Verachthonius laticeps (Strenzke, 1951:

240).

Diugnosis, Liachthonius-complex. Qne subnotal

shield (SNS3)separate from hysteronotal shield. Hys-

tcronotal setae never conspicuously ciliate. Posterior

setae in hysteronotal file / conspicuously compacted

forward seta, /6 nearly level to point midway between

Zi and Z6 in relation to posterjor margin of hyster-

onotal shield. Notal shields relatively smooth with

few ridges or circles, usually | ridge runs [rom anterior

notch between posterior notal and pleural shields back

to region of seta 45. Only adoral seta (ae2) stout,

almost parallel-sided with blunt tip. Adanal setae Zal

and Za2 similar, cither setose like seta Zu3 or con-

spicuously stouter than Za3. Three setac on both genu

Wt and IV (ventral seta present), Aggenital file /g¢

with 4 setae (/g4 present).

Distribution: Canada (Nn), Argentina (NTe), Ger-

many (Pe), Japan (Pc), Australia (Aa),

Remarks: Werachihonius species separale oui Lo

the couplet including Brachyehrthonius and Syn-

chthonius in the key of Balogh (1972), But, because

the descriptions of many speicies are mainly based

on a dorsal view of the sama, any such incorrectly

placed species of Verachthonius would probably be

currently included ip Livehthanius rather than Bra-

chychthonius or Synchihonius.

Ferachthanius montanus (Hammer, 1952; 17) fram

Canada and Argentine (Hammer 1958: 19), was

included in Eabrachychthonius but on criteria used

here it clearly belongs in the Liochthonius-complex

and jn either Liochthonius or Verachthonius. My

most heavily weighted attribute for supporting its

inclusion in Verachthanius is the forward displacement

of seta /6,

Five species are included in Ferachthontus: ¥. con-

eruus Moritz, 1976a; V. diversus Moritz, |976a: ¥-

laticeps (Strenzke, 1951); FV. monianus (tlammer,

1952), ncomb.; b wroritc’ m.sp.

Verachthontus moritzé n.sp.

(Figs. 5-13)

Female

General appearance; Shields straw-coloured, |dio-

somal length 157.5 (20, 145-165); appendage lengths

(for holotype, 152.5)—ch 10, pa 32.5, f 62.5, 1747.5,

7/1-47.5, 7 62.5; femur breadths—pa 9. £ 14, ff 11,

HAO, TVA,

Prosternune Gnathosoma illustrated cither dorsay-

entrally squashed (Fig, 9) or laterally squashed so

chelicerae unnaturally extended (Fig, 6). Single adoral

seta (wo2) nearly parallel-sided for entire length,

slightly spread out at blunt distal ip. External malae

refractile and colourless. Coxites Il] and tV of cach

side clearly separated by mid-sternal fissure, but not

established whether or not trough between coxites |

and Il occupied by striated cuticle.

Proteronatuny: Single refractile recess under cow!

shows through anterior part of proteronotal shield of

cleared specimens. Seta 42 present, extremely minute

and lying mid-way between a pore and si, Plassnic

seta 22 bears 5 files of 7 or 8 cilia on distal swollen

part.

Opisthosternum: Genital and anal shields illustrated

(Fig, 7) have lifted away fram mid-sternal! line, appear

narrower, Adanal shields (bearing setal file Za) not

posteriorly merged together, Transverse ridge on shield

between setae Za2 and Za3, Ovipositors involuted,

chaelolaxy dilficull to observe. Ovipasilor of hulotype

(contains egg) tlustrated (Fig. 10) only 5 pairs of

genital setae observed, possibly more present,

344 REC. S. AUST. MUS., 18 (15): 327-359 June, 1982

ch2 aoees

FIGS. 5-13—Verachthonius moritzi n.sp., female: 5, notum; 6,

pleura; 7, idiosternum; 8, proteronotal cowl; 9, gnathosternum;

10, left genital shield and involuted ovipositor; 11, left chelicera,

anterior surface; 12, part left palp, anterior surface; 13, legs,

dorsal setae on genua, tibiae and tarsi.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

Hysteronotum: Weak tidges on shields appear

stronger as illustrated (Fig. 5) than they should be.

Outlines of pleural and subnotal shields faint. Seta

55 on low tubercle.

Appendages: Chelicerae pale, not refractile, weakly

dentate. Setae: ch (1), pa (0-2-1-3-9), [ (0-3-3-4-17), IT

(0-4-3-3-14), 777 (2-3-3-3-1 1), PV C1-2-3-4-11). Solenidia:

pa (0-0-1), J (2-1-3), 17 (1-1-1), 27 (1-1-0), 2 (1-0-0).

On tarsus [, only | solenidium in normal dorsal

position, other 2 solenidia (soy) and larger sov2) ven-

tral and spiniform. Solenidia on all genua and on

tibia Tl] coupled with setae. other dorsal solenidia

separate, Absence of solenidium on tibia [V not known

on other brachychthoniids,

Somal inclustons: Amongst 20 registered rites, 4

(2 from cach site) contain single egg, rest without

epgs. Eggs about 80 (70-82.5) long, ellipsaid with

uniform smooth surface. Two specimens from Piccan-

innie Ponds contain 1] and 18 small (diameter 20-

25) spherical objects in hysterosoma which showed

no internal structure and became very faint in gum

chloral, Possibility that they were spermatophores or

cysticercoids of anoplocephalid tapeworms considered

and latter, if either, seemed more likely, Six specimens

contained | or 2 boli, mainly granular, included some

rectangular cellular forms,

Male

Unknown

Material examined: Holotype female (N1979315)

and 13 paratype females (N1979316-N1979328), litter

and sparse grass, under Acacia sophorae, coastal

scrubland, Piccaninnie Ponds, 20.8.1975, D. C, Lee.

Six females (N1979329-N ]979334), litter, under

Pinus pinea, Knott Hill, 22.5.1974, D. C. Lee.

Distribution: South Australia (Aa), South Australia:

Piccaninnie Ponds, coastal closed-scrubland, 14 (4/

%); Knott Hill, cultivated pine forest, 7 (1/2).

Remarks; Besides having more notal ridges than

the other 4 nominal species, ¥. morirzi has conspi-

cously' longer notal selae, with seta J] longer than

half the distance between J] and J2.

The centralward displacement of hysteronotal seta

72 is not conspicuous for a member of the Lioch-

thonins-complex, but it is more obvious if viewed at

righi-angles to the integument in the region of that

sela or on squashed specimens.

NEOBRACHYCHTHONITUS -camplex

Diagnosis’ Brachychthoniidac. Anterior subnotal

shield (SNS1), bearing seta S|, separate [rom hys-

teronotal shield. Hysteronotal setae #4], $2, 272 and

Z3 nal in nearly straight line parallel to lateral margin

of notal shield because seta Z2 displaced towards

mid-notal line. Subnotal shield SNS3 separate or

445

merged with hysteronotal shield. Adoral seta ao2

cither blunt or stout with swollen distal end. Chae-

totaxy of genu II and TV unknown, Coxite 1V bearing

3 setae,

Remarks: If the 2 genera included in this genus-

complex had not been established, the species would

probably be included in Liochthonius because of the

position of 22. So far they are only known from

Europe (Pe), Paraliochthonius has w be renamed

because its use for a pseudoscorpion genus has priority.

Two genera are included in the Neobrachychthon-

ius-complex; Neobrachychthonius Moritz, 1976b;

Neoliochthonius name for Paraliocehthanius Muritz.

1976a not Paraliochthonius Beier, 1956,

Family TRICHTHONIIDAE n.fam.

Type-genus: Trichthenius Hammer, 1961.

Diagnosis: Hypochthomioidea. Minute to medium -

sized (155-790), pale mites. Two complete transverse

hysteronotal fissures (72, 783); anterior fissure

(7B2) never conspicuously closer to seta J2 than to

J3, posterior fissure (7B3) about halfway between

sela J1 and J6 or posterior to that level. Antertar

part of pleural shield (PS1) either partially (less than

* O.6 breadth) separated off by vertical fissure or

completely merged with posierior pleural shield (P82),

Hysteronotal setae vary considerably in length, setae

J3 and Z3 more than & 2 length of setae J2 or 72.

Longer setae (J3, J4, Z3, 24) setose with inconspi-

cuous lateral cilia or linguiform with cilia and some-

times fringe of balloon-like structures, at least as long

as distance between setal bases J] and J4. Hyster-

onotal seta $2 not as near to mid-notal line as 71,

Each coxite of legs Il and IV completely separate

ar only separate along mid-sternal line with [11 and

IV merged on cach side. No proteronotal eye or eyes

irl Zone between setae /1] and cl, On tarsus |, seta d]

simple, setose or peg-like Adoral file includes | (a2)

or 2 (@o2, an3) setac.

Remarks; Trichthonius was grouped in the Cos-

mochthoniidae by Balogh (1972), But, in the classi-

fication presented in thus paper, Trichthonius has 1o

be grouped in the Retrofissurae rather than the Pral-

issurae which includes Cosmochthoniidae, If the

remarks on the Retrofissurae are studied, it can be

seen by characters used there for the diagnosis of

superfamiles, that Trichthonius must be included in

the Hypochthonioidea, Within this superfamily, the

Heterochthoniidae is the most suitable to include

Trichthonius, on the other hand, the presence of a

proteronotal eye or eyes in members of the Hetero-

chthoniidae and the forward compression of the Jarge

spine-like hysteronotal setae and their shields, has

rade me reluctantly prefer to establish this new

family, the Trichthoniidae,

346

Three genera are included in Trichthoniidae: Mar-

shallia Gordeeva, 1980; Nipponiella Gordeeva, 1980;

Trichthonius Hammer, 1961. Trichthonius has been

collected in South Australia and is considered below.

Marshallia includes 2 species: M. majestus (Marshall

and Reeves, 1970) from North America; M. golosovae

Gordeeva, 1980 from U.S.S.R. Nipponiella includes

| species: NV. simplex (Aoki, 1966) from Japan,

TRICHTHONIUS Hammer

Trichthonius Hammer, 1961; 15. Type designation

(by monotypy); “Cosmochthonius pulcherrimus

Hammer”.

Type-species: Trichthonius pulcherrimus (Hammer,

1958: 22).

Diagnosis: Trichthoniidae. Most notal setae

(including j2, J1, /2, J3, /4,) enlarged, linguiform,

ciliate without fringe of balloon-like structures, and

able to cover most of notum. Fifth rank of hyster-

onotal setae (J5, Z5, S'5) conspicuously nearer to 6th

rank than to zone midway between 4th and 6th ranks.

Cheliceral seta ch2 conspicuous longer (x 1.2 or

more) than fixed cheliceral digit. Two pairs of adoral

setae (ao2,ao03). External malae bifurcate. Each cox-

ite of legs HI] and IV completely separate. Chaetotaxy

of genua: setae /-6, J/-3, IIJ-2, IV-3; solenidia J-1, [I-

1, HTI-1, IV-0.

Remarks: The single Trichthonius species is super-

ficially very similar to members of the Cosmochthon-

iidae, especially to Phyllozetes, so it is understanda-

ble that it has been grouped in that family. Even the

gnathosoma is considerably modified (a feature of

Cosmochthoniidae), although not in a similar way

and there is no conspicuous chitinous structure sup-

porting enlarged pharyngeal muscles. The similarity

is mainly due to the enlarged, erectile hysteronotal

setae (J3, J4, Z3, Z4) which can lie close to the body

or spread upwards like a peacock’s tail. If the weight-

ing of characters presented here is accepted, the

similar characters must be regarded as resulting from

convergent evolution. In contrast, Trichthonius looks

rather different from the much larger, confamilial

Nipponiella, with its almost simple notal setae and

unmodified gnathosoma.

One species is included in Trichthonius: T. pul-

cherrimus (Hammer, 1958).

Trichthonius pulcherrimus (Hammer)

(Figs. 14-21)

Cosmochthonius pulcherrimus Hammer, 1958: 22.

Trichthonius pulcherrimus (Hammer): Hammer,

1961: 15.

Trichthonius pulcherrimus (Hammer): Hammer,

1962: 16.

REC. S. AUST, MUS., 18

(15); 327-359 June, 1982

Female

Shields ochre, covered by granulate adhesive layer.

Idiosomal length 187.5, (4, 180-190); appendage

lengths (for 185)—ch 13, pa 40, I 67.5, IJ 60, I/I 70,

IV 77.5; tibial breadths—/ 14, 17 13, TT 11, IV 13;

femur breadths—/ 14, V7 13 III 14 IV 16. Tibial

breadths given because trochanter and femur of legs

III and IV disproportionately stout compared with

rest of leg. Most notal setae subsemicircular in cross-

section and possibly hollow with branched struts

radiating from ventral midrib to convex dorsal sur-

face. Only dorsal setal surface ciliate, seta J4 has

dorsal bald patch which seta J3 rests on when col-

lapsed down close to body. Erectile setae (J3, /4, Z3,

and Z4) illustrated (Fig. 14) in intermediate position

between lying close to body and being at right-angles

to it. Many notal setae on tubercles with large spines.

For seta s2, spine on tubercle larger than minute seta.

Tubercles of 4 pairs of erectile setae carry 2 large,

bifurcate spines, apparently articulating with spines

of next erectile seta in same rank. Curiously, outward

pointing spines of Z3 and Z4 fit this description

although no other spines with which to articulate.

Posteriorly, hysteronotal shield has central keel which

setae J4 rest beside when lowered close to notum. On

gnathosoma, adoral setae have rigid curved shaped,

seta ao2 also has hyaline flap, external malae bifur-

cate with 2 slim, tapering branches. Chelicerae unu-

sual. Movable digit appears foreshortened with anter-

olateral groove and large, hooked posterolateral

process. Fixed digit edentate, may be movable. Seta

ci2 large, curved with ventral hyaline flap. chelicerae

has groove along anterolateral surface from the region

of the mouth to base of movable digit. Pharynx

appears simple and without conspicuous muscles sup

ported by chitinous struts. On legs HI and IV, tro-

chanter, and to lesser extent femur, large and wrin-

kled.

Appendage setae: ch (2), pa (0-2-1-3-10), J (1-3-6-

6-19), I (0-4-3-5-17), IIT (2-3-2-4-15), IV (2-3-3-4-13).

Solenidia: pa (0-0-1), 7 (1-1-1), 77 (1-1-2), T/T (1-1-0),

IV (0-1-0). Solenidial form: baculiform on tapa, tal,

u//, tall(sol + so2); ceratiform on ge//, gellI llI,

n/V; flagelliform with coupled seta on ge/, til.

No somal inclusions (food boli or eggs) observed.

Material examined: One female (N1979335), litter

or grass and moss, under Eucalyptus viminalis,

Chambers Gully, 12.6.1974, D. C. Lee. Three females

(N1979336-N1979338), litter and sparse moss, under

Eucalyptus obliqua, sclerophyll forest, Mt. Lofty,

9.5.1974, D. C. Lee.

Distribution: Argentina, Bolivia, Chile, Peru (NTc);

South Australia (Aa). South Australia: Chambers

Gully, savannah woodland, 1(1/8); Mt. Lofty, scler-

ophyll open-forest, 3(2/8).

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 347

15,17, 19, 20,21

FIGS. 14-21—Trichthonius pulcherrimus (Hammer), female: 14,

pleura; 15, right chelicera, anterior surface; 16, idiosternum;

17, gnathosternum; 18, left trochanter and femur IV, anterior

and ventral surfaces; 19, right leg I, dorsal setae on genu,

tibia and tarsus; 20, left palp tarsus, anterior surface; 21, left

genital shield and ovipositor.

348

Remarks: There are differences between the

description of the type material and the South Aus-

tralian specimens, such as larger (230), proteronotal

plasmic setz z2 not strongly dilated distally, hyster-

onotal seta J4 without dorsal bald patch, tibia I

solenidium about half length of genu I solenidium.

For the time being, these are regarded as either

intraspecific variations or reflecting inaccurate illus-

tration.

The gnathosoma is interesting in having primitive

(only present in Bifemoratina and Retrofissurae) dis-

junct external malae and yet being considerably mod-

ified. The adoral setae and external malae appear to

be modified to hold a globule of liquid and the

chelicerae to provide a tube to suck up liquids. Unfor-

tunately, there were no boli in the 4 specimens col-

lected to give an indication of the nature of the food.

The shape of trochanter III and IV has not been

used in the diagnoses of higher taxa. But the simple

cylinder (Fig. 18), possibly given more mobility in T.

pulcherrimus by being wrinkled, could be regarded

as primitive, and within the Arthronotina be diagnostic

of Retrofissurae. In contrast, Cosmochthonius has a

modified trochanter (Fig. 24) which must give much

more mobility to the leg, and could be diagnostic of

Cosmochthonioidea.

Subcohort PROFISSURAE n.

Diagnosis: Arthronotina. Usually 3 transverse hys-

teronotal fissures (7B1, 7B2, TB3), sometimes obscure

as when notal setae umbellate (Pterochthoniidae), or

when notum folded so that second fissure (7B2) cov-

ered by first hysteronotal shield (Sphaerochthoniidae),

sometimes posterior fissures (TB2, 7B3) partial (some

Protoplophoridae), sometimes posterior fissure (7B3)

absent (Sphaerochthoniidae). No hysteronotal gland

with egress pore between setae Z4 and Z5. Cowl

present, enclosing retracted chelicerae. Chelicerae

often modified so that either fixed digit or seta ch2

comb-like; rarely seta chl absent in which case ch2

simple, Palp tarsus without distal bifurcate plasmic

seta and often with 2 ribbon-like dorsal setae con-

spicuously longer than itself, External malae conjunct.

Often conspicuous internal chitinous structure sup-

porting enlarged pharyngeal muscles. One pair (Zaf)

of adanal pores present. On tarsus I, seta dl never

bifurcate. No genua bear solenidia. Pretarsi usually

with either 1 claw, 2 unequal claws (1 stout simple

claw, 1 slim or ciliate claw) or 3 unequal claws

(central stout claw, lateral slim claws), very rarely 2

subequal claws.

Remarks: The Profissurae is a new taxon. It has a

diversity of form that reflects that of the Retrofissurae

(see remarks on parallel evolution under Arthronotina).

The name, Profissurae, is intended to indicate the

main attribute which contrasts with Retrofissurae, i.e.

that the anterior hysteronotal fissure (7B1) is always

present.

REC. 8S. AUST, MUS,, 18 (15): 327-359

June, 1982

The comb-like structure of the fixed cheliceral digit

has been considered primitive in Pterochthonius, sug-

gesting the origin of chelate chelicerae amongst Acari.

This is unlikely. Probably, both it, the comb-like seta

ch2 and other specializations of gnathosoma amongst

Profissurae, function by sifting single-celled organisms

from liquids. What is peculiar about these attributes

is that they occur regularly amongst the Profissurae

(whilst being absent elsewhere amongst Cryptostig-

mata) yet they may be either present or absent in

very similar genera. This might lead to the conclusion

that there were polymorphic forms or stages of the

same species, but there is no support for this concept.

The attributes of pretarsal claws are used in the

diagnosis although they are complex. The model is

that, amongst Profissurae, pretarsi have a stout central

claw, with slim lateral claws | or both of which may

be lost especially on anterior legs, so that there should

not be 2 subequal claws as on some other Arthronotina.

The | known exception is Cosmochthonius bengalensis

with 2 equally thick and curved claws on all pretarsi.

The form of the claws of Krivolutskiella (Cosmo-

chthoniidae) is not known, although they are unusual

for the Profissurae in there being 2 claws on pretarsus

IV.

The Profissurae includes 2 superfamilies: Cosmo-

chthonioidea Grandjean, 1969; Protoplophoroidea

Grandjean, 1965. The Protoplophoroidea includes 1

family as listed by Balogh (1972) and in relation to

this study has been referred to (Lee 1981) under the

now disbanded Arthroptyctima. The Cosmochthon-

ioidea, some of which were also collected in this

study, are considered further below.

Superfamily COSMOCHTHONIOIDEA

Diagnosis: Profissurae. Prehysteronotal fissure

usually not hinged and never acutely hinged.

Remarks: The genera of Cosmochthonioidea as

delineated here are very much as listed by Balogh

(1972), except that the Pterochthoniidae are included.

Grandjean (1954b: 334) drew attention to the remark-

able similarity between members of this superfamily,

particularly Sphaerochthonius, and the Protoplophor-

idae. In fact, whilst the Protoplophoridae was given

quite a substantial diagnosis in my previous paper

(Lee 1981: 212), if it was diagnosed here as a super-

family to be delineated from the Cosmochthonioidea,

I would be confident only of giving its acutely hinged

prehysteronotal fissure as a diagnostic attribute.

Mahunka (1977b) established a new protoplophorid

genus, Hauseroplophora from Kenya, which, because

of its superficial similarity to Sphaerochthonius, he

regarded as further supporting Grandjean’s supposi-

tion. But in fact, although the form of its setae is

similar to Sphaerochthonius, Hauseroplophora is not

as similar as Aedoplophora and Cryptoplophora

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 349

(2 other genera of Protoplophoridae) are to Sphaer-

ochthonius in the shape and disposition of the somal

shields. Certainly, further study of established taxa

within the Profissurae may indicate a broader basis

for distinguishing the Protoplophoroidea from this

superfamily. For example, Sphaerochthonius appears

to share with Cosmochthonius the proximal construc-

tion of femur and trochanter IV, whilst these segments

on members of the Protoplophoroidea are without

such a constriction. On the other hand, a more thor-

ough examination of Sphaerochthonius may indicate

that it can, or originates from mites that could, fold

up its soma around hinged fissures. This or other

attributes may make the superfamily groups here

redundant. Possibly, the present diverse Protophopho-

roidea represents a convergence from different evo-

lutionary trends within the Cosmochthonioidea, but

at this stage, | am adopting a conservative approach

and maintaining it as in the classification of Balogh

(1972).

The Cosmochthonioidea includes 4 families: Cos-

mochthoniidae Grandjean, 1946; Haplochthoniidae

van der Hammen, 1959; Pterochthoniidae Grandjean,

1950b; Sphaerochthoniidae Grandjean, 1947a. The

Haplochthoniidae and Pterochthoniidae were not rep-

resented in collections for this study and are not

further considered.

Family COSMOCHTHONIIDAE Grandjean

Cosmochthoniidae Grandjean, 1946: 315.

Type-genus: Cosmochthonius Berlese, 1910.

Diagnosis: Cosmochthonioidea. Minute (240-360),

ivory white to brown mites. Three complete, unob-

scured hysteronotal fissures (7B1, TB2, TB3). Four

pairs of hysteronotal setae (J3, J4, 73, Z4) long (at

least as long as distance between setal bases /1 and

J4) and erectile. Cowl broken up by 2 to 4 ranks of

4-9 longitudinal slits. Anterior subpleural shield

(SPS1) has protruding ventral wing-like process, whilst

subpleural shield (SPS2) has ventral extension to

large pore. Genital tracheae present. Anterior cheli-

ceral seta (ch2) pectinate with, 5 or 6 prongs. Tarsus

I bears | solenidium, Tarsi I-IV with 2 or 3 claws.

Remarks: This widespread family, the Cosmo-

chthoniidae, has very similar members grouped in 3

genera: Cosmochthonius Berlese, 1910; Krivolut-

skiella Gordeeva, 1980; Phyllozetes Gordeeva, 1978.

Both Cosmochthonius and Phyllozetes have been col-

lected in South Australia and are considered below.

Krivolutskiella includes 1 species: K. pubescens Gor-

deeva, 1980 from the Canary Islands.

COSMOCHTHONIUS Berlese

Cosmochthonius Berlese, 1910a: 221. Type designation

(original); “C. /anatus (Mich.)”.

Type-species: Cosmochthonius lanatus (Michael,

1885: 396).

Diagnosis: Cosmochthoniidae. Some notal setae

(J3, J4, Z3, Z4) long, stout, ciliate without broad

leaf-like lateral membrane. Pretarsus | with 2 claws,

pretarsi Il, [1] and IV with 2 or 3 claws (2 subequal

claws or stout central claw with | or 2 slim lateral

claws). Coxites II] and IV separate from each other.

More setae on femur II and tarsi ill and IV (6, 16

and 14).

Distribution: Widespread—Canada (Nn), Califor-

nia (Nc), Nevada (Nr); Bolivia, Argentina, Peru

(NTc): Ghana (Ew); England, Germany, France, Rus-

sia (Pe); Italy (Pm); Russia (Ps): East Java (Om);

Australia (Aa), New Zealand (An).

Remarks: Cosmochthonius is an easily recognised

genus, but 2 species originally grouped in it are now

combined in other genera. One, Trichthonius pul-

cherrimus (Hammer), see above, is here regarded as

only superficially similiar, and is now in another sub-

cohort (Retrofissurae). The other, Phy//ozetes emmae

(Berlese), see below, is similiar but was excluded by

a restriction in the extent of the genus. The value of

restricting the genus is questionable since the discovery

of taxa with 2 claws on all legs: Krivolutskiella with

some leaf-like hysteronotal setae (/3, Z3) like those

of Phyllozetes; Cosmochthonius bengalensis with hys-

teronotal setae characteristic of Cosmochthonius.

Most descriptions of Cosmochthonius species are

almost confined to the notal surface: exceptions are

those of C. reticulatus by Grandjean (1962) and C.

fovealatus as originally described. Having given new

status to 3 subspecies, revoked a synonymy and named

a previously described but unnamed species there are

now listed below 15 species of Cosmochthonius. Gor-

deeva (1980) provides a key for 7 palaearctic species,

which still leaves a problem for those wishing to

identify Cosmochthonius from elsewhere. Since, in

the past, notal sculpturing and the ciliation of the 4

large erectile setae (J3, J4, Z3, Z4) are the main

diagnostic characters, | have used a unified termi-

nology (see above under “Notation for Morphology’)

to briefly describe such attributes for all species.

Some characters that differ from C. australicus (as

described below) are also referred to. For convenience,

the species are grouped in 4 species-complexes: reti-

culatus-complex, foveolatus-complex, plumatus-com-

plex, asiaticus-complex.

reticulatus-complex

Diagnosis: Reticulate sculpturing covering posterior

hysteronotal shield (NS4y. Cilia on hysteronotal seta

J3 even, usually short and sparse, rarely longer or

medium spaced, between 14-38 on | side.

350

Remarks: Five species are included in the reticu-

latus-complex. The inclusion of C. lanatus is debat-

able.

Cosmochthonius bengalensis Chakrabarti et al.

Cosmochthonius bengalensis Chakrabarti, Bhaduri

and Raychaudhuri, 1972: 86.

Length, 288-290 (ex West Bengal—Oi). Seta zl

unbranched. Seta s2 long, ciliate as jl. Seta J2 not

on anterior ridge of second hysteronotal shield (NS2).

Seta /3 with about 36 short sparse cilia on each side.

Coxites I, I], II, IV with 3, 1, 3, 3 setae. Pretarsi I,

Il, IL, IV with 2, 2, 2, 2 equally thick claws.

Cosmochthonius lanatus (Michael)

Hypochthonius lanatus Michael, 1885: 396.

Hypochthonius lanatus Michael, 1888: 541.

Cosmochthonius lanatus: Willmann, 1931: 101.

Length, 330 (ex England—Pe), 290-320 (ex Ger-

many—Pe). Seta z1 unbranched. Seta J3 with more

than 25 medium lengthed and medium spaced cilia

on each side.

Remarks: Michael (1888) illustrates C. /anatus

with large abutting puncta on the notum, but in the

text states “notogaster is strongly and coarsely reti-

culated”, also Willmann’s (1931) illustration shows

dorsal pits with distinctly straight edges. On the other

hand, in Gordeeva’s (1980) key (couplet 9/10), for

“Cosmochthonius sp. (lanatus?)”’ the notal surface is

described as having rounded abutting pits, and van

der Hammen (1952) regards C. domesticus as syn-

onymous with C. /anatus. Therefore the position of

this poorly described species is uncertain.

Cosmochthonius reticulatus Grandjean

Cosmochthonius reticulatus Grandjean, 1947b: 354.

Cosmochthonius reticulatus Grandjean, 1962: 404.

Length, 300-340 (ex France—Pe). Cheliceral fixed

digit with 5 well developed teeth, seta ch2 with 6

prongs. Transverse ridge between setal pair jl. Seta

s2 long, ciliate (compared with C. australicus)

although shorter and slimmer than other proteronotal

setae. Plasmic seta 22 with distal half much stouter

(< 6) than proximal half. Seta J2 not on anterior

ridge of second hysteronotal shield (NS2). Seta J3

with 14 long sparse cilia on each side. Appendage

setae and solenidia similar to C. australicus except

tibia II] with 5 and tarsus IIT with 15 setae.

Cosmochthonius sublanatus Mahunka

Cosmochthonius sublanatus Mahunka, 1977a: 254.

Length, 273-294 (ex eastern Java—Om). Seta zl

unbranched. Seta s2 long, ciliate as j2. Seta j3 with

about 30 short medium spaced cilia on each side.

REC. S. AUST. MUS., 18 (15): 327-359

June, 1982

Seta z3 with sparse cilia, /4 and Z4 shorter (x 0.5)

than 3rd setal rank.

Cosmochthonius wallworki n.sp.

Cosmochthonius spec. (C. lanatus?) Wallwork, 1960:

386.

Length, 291 (ex Ghana—Ew). Seta s2 long, ciliate

(compared with C. australicus) although shorter than

other proteronotal setae. Plasmic seta z2 with distal

half slightly stouter (* 1.5) than proximal half. Seta

J2 on anterior ridge of second hysteronotal shield

(NS2). Seta J3 with 18 short sparse cilia on each

side.

foveolatus-complex

Diagnosis; Punctate sculpturing covering posterior

hysteronotal shield (N.S4). Cilia on hysteronotal seta

J3 even, long and either sparse or dense, between 5-

60 on | side.

Remarks: Six species are included in the foveola-

tus-complex. Three species (C. domesticus, C. fov-

eolatus, semiareolatus) are similar to each other in

having denser cilia and closer spaced puncta, and

have been referred to as similar to C. lanatus (reti-

culatus-complex).

Cosmochthonius domesticus Grandjean

Cosmochthonius domesticus Grandjean, 1947b: 354.

Length, 245-285 (ex France—Pe). Cheliceral fixed

digit with 4 teeth, central 2 inconspicuous, seta ch2

with 6 prongs. Form of cilia on hysteronotal seta J3

unknown. Puncta on posterior hysteronotal shield

(NS4) large, circular or oval, never polygonal, and

abutting.

Remarks: The synonomy of C. domesticus with C.

lanatus (van der Hammen, 1952: 22) is revoked on

the basis that it is not yet satisfactorily established.

Cosmochthonius foveolatus Beck n. status

Cosmochthonius lanatus foveolatus Beck, 1962: 232.

Length, 320-360 (ex Peru—NTc). Cheliceral fixed

digit with 4 conspicuous teeth, seta ch2 with 5 prongs.

Seta zl unbranched. Ridge and tubercles connecting

posterior margin of bothridia of plasmic setae 22.

Seta J2 not on anterior ridge of second hysteronotal

shield (NS2). Seta J3 with about 35 long, dense cilia

on each side. Puncta on posterior hysteronotal shield

(NS4) large, circular and abutting.

Cosmochthonius ponticus Gordeeva

Cosmochthonius ponticus Gordeeva, 1980: 844.

Length, 245 (ex Crimea—Pe). Seta s2 long, ciliate

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS 35]

(compared with C. australicus) although shorter than

other proteronotal setae. Seta J3 with 5 or 6 medium

lengthed cilia on each side near base. Puncta on

posterior hysteronotal shield (V.S4) large, irregular in

outline and well separated.

Cosmochthonius semiareolatus Hammer

Cosmochthonius semiareolatus Hammer, 1966: 14.

Length, 285 (New Zealand—An). Seta s2 long,

ciliate as j2. Seta J3 with about 60 long (longest in

genus), dense cilia on each side. Puncta on posterior

hysteronotal shield (NS4) large, circular and abut-

ting.

Cosmochthonius tenuisetus Gordeeva

Cosmochthonius tenuisetus Gordeeva, 1980: 844.

Length, 245 (ex Crimea—Pe). Seta s2 long, ciliate

(compared with C. australicus) although shorter than

other proteronotal setae. Seta J2 long (reaches back

to setal base J4). Seta J3 with 6 or 7 long, sparse

cilia on each side, evenly spaced along seta. Puncta

on posterior hysteronotal shield (N.S4) small, circular

and well separated.

Cosmochthonius ugamaensis Gordeeva

Cosmochthonius ugamaensis Gordeeva, 1980: 846.

Length, 290 (ex Uzbekistan—Ps). Seta /1 bifurcate,

Y-shaped. Seta s2 long, ciliate as j2. Seta J2 not on

anterior ridge of second hysteronotal shield (NS2).

Seta J3 with 6 long, sparse cilia on each side, evenly

spaced along seta. Puncta on posterior hysteronotal

shield (NS4) large, star-shaped in outline and close

although not abutting.

plumatus-complex

Diagnosis: Punctate sculpturing covering posterior

hysteronotal shield (NS4). Cilia on hysteronotal seta

J3 uneven with both sparse long cilia and dense short

(3 or 4 to each long cilium) cilia.

Remarks: Two species are included in the p/umatus-

complex, | of which is C. australicus collected in

this study.

Cosmochthonius australicus Womersley n.stat.

(Figs. 22-29)

Cosmochthonius plumatus australicus Womersley,

1945; 222.

Female

Dull, ochre to orange. Cuticle covered in granulate

hyaline adhesive layer which easily peels off. Refrac-

tile parts (external malae, cheliceral extremities, setae

and claws) paler than general integument. Idiosomal

length 280 (20 ex Ferries-McDonald, 275-285: others

within this range); appendage lengths (for 280)—ch

15, pa 60, f 120, 17 95, TT 100, 1V 115; femur

breadths—pa 14, J 24, I7 22.5, 1/7 20, IV 20.

Puncta on posterior hysteronotal shield (NS4) large,

circular and close although not abutting. On next

shield (NS3), few very faint puncta not represented

in illustration (Fig. 22). Few distinct puncta on second

shield (NS2). On anterior shield (NMS1), few rectan-

gular pits. Setae J3, Z3, with about 24 long cilia and

about 100 short cilia on each side. Setae J4, 24

shorter with longer sparser long cilia. Proteronotal

seta z] bifurcate and T-shaped, with shorter posterior

branch.

Cowl sometimes has 3 ranks of slits, posterior rank

with 9 slits, middle rank with 7 slits, anterior rank

with 5 slits, but arrangement may be more haphazard.

Bases of conjunct external malae with central tri-

angular flaps covering bases of internal malae and,

with external malae, encompass oval aperture through

which preoral setae protrude. Mentum undelineated,

transverse ridge not regarded as mentocoxal fissure.

Three wing-like processes on pleural and subpleural

shields, lying above coxae III and IV. Posterior sub-

pleural shield (SPS2) with ventral extension that

becomes groove, lined with processes or hairs, leading

to conspicuous pore behind coxa IV,

Chelicerae pale, fixed digit with 4 well developed

refractile teeth. Trochanter IV and femur TV, to lesser

extent same segments on leg III, unusual for primitive

oribatids (possibly true for all Cosmochthonioidea) in

being constricted proximally. These segments look as

if they may be able to rotate around their proximal

articulation, rather than rock backwards and forwards.

Appendage setae: ch (2), pa (0-2-1-3-11), I (0-5-5-

6-19), IT (1-6-5-6-17), III (2-3-4-4-14). Solenidia: pa

(0-0-1), 7 (0-0-1), 7 (0-1-1), H/T (0-1-0), JV (0-1-0).

Claws: I-2; I], II], 1V-3. Cheliceral seta ch 2 with 6

prongs. Some setae on palp tarsus long and at least

seta d4 ribbon-like and possibly plasmic. On tarsus I,

plasmic seta d1 dilates slightly distally, curling around

seta pd3. On tibia I, solenidium and coupled seta on

conspicuous tubercle. Solenidia on posterior 3 pairs

of legs ceratiform, those on tibiae (especially tibia

IV) small. Positor short, but too large to be male,

although all genital setae appear similar in size.

Conspicuous internal chitinous structures encompass

pharynx between level of preoral and postoral setae,

also ovoid refractile membrane (not illustrated) arises

in region of palp coxite and extends backwards to

coxite Il, probably equivalent of “vpx’’ (Grandjean

1948: fig. 3A). Genital tracheae extend forward from

anterior end of genital orifice. Gut contents consist

entirely of dark green granular material and refractile

spheres (diameter, 2-3). Narrow pharynx expands

about level of seta J2 into fat horseshoe-shaped pair

of dorsal diverticulae that loop backwards around

352 REC. S. AUST. MUS., 18 (15): 327-359 June, 1982

AONv00S

GLE PAA

50m

23,24,26,27 28,29

22,25

FIGS. 22-30—Cosmochthonioidea: 22-29, Cosmochthonius aus-

tralicus Womersley, female; 22, pleura; 23, right chelicera,

anterior surface; 24, left trochanter and femur IV, anterior

and ventral surfaces; 25, idiosternum; 26, left palp tarsus,

anterior surface; 27, gnathosternum; 28, right leg I; 29, left

genital shield and ovipositor; 30, Sphaerochthonius splendidus

(Berlese), notum.

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOTLS

lateral margins of hysteronotum, Dorsal diverticulac

often uniformly full of food. At anterior end of horse-

shoe-shape (between level of setae J2 and J3) gut

inclines ventralwards to anus. This strip of straight

gut usually contains. 2 or 3 boli, anterior bolus being

diffuse and larger whilst posterior bolus smaller

(diameler, about 25) and dense. No eggs observed

inside mites.

Material examined: Lectotype (NJ979378) and 2

paralectotypes (N1979379-N1979380), moss, Mt.

Arden, southern Flinders Ranges, 11,1943, H.M,

Cooper. Twenty females (N1979339-N1979358), lit-

ter and sparse moss, under Eucalyptus incrassata,

mallee scrubland, Ferries-McDonald, 20.6.1974, D.C.

Lee. Six females (N1I979359-N 1979364), grass and

moss or Jitter under Eucalyptus virrinalis, savannah

woodland, Chambers Gully, 12.6.1974, D.C. Lee,

Twelve females (N1979365-N1979376), litter and

sparse moss, under Eucalyptus obliqua, sclerophyll

forest, Mt. Lofty, 9.5.1974, D.C. Lee. One female

(posterior half) (N1979377), grass and plantain, Glen-

thorne, 12,6,1974, D.C, Lee,

Distribution: South Australia (Aa). South Aus-

\ralia; Ferries-McDonald, mallee-broombush open-

scrubland, 20(3/8); Chambers Gully, savannah wood-

land, 6(3/8); Mt. Lofty, sclerophyll open-forest,

12(4/8); Glenthorne, cultivated pasture, 1(1/8); Mt.

Arden and Waterfall Gully (Womersley, 1945).

Remarks: Regarding the collection data, the record

of | specimen from Glenthorne is based on a hyster-

osoma alone, but from its condition it is assumed that

the mite was damaged after collection. The original

6 specimens from Waterfall Gully cannot be found,

Instead of the 2 specimens, as originally recorded

from Mt. Arden, there are 3 specimens with exactly

the same labels. The specimen designated as the

lectotype is the specimen drawn (Womersley, 1945;

fig. 2) on the basis of the position of hysteronotal

setae. One specimen designated as a paralectotype

cannot belong ta the published syntype series, but,

since it is not known which one, this has to be dis-

regarded,

The original description (Womersley, 1945) is

inaccurate for a number of attributes, for example

the form of the cilia on hysteronotal setae J3, J4, Z3,

#4 and the presence of punctuations on the third

hysteronotal segment (VS3). Since these inaccuracies

apply to the main 2 groups of characters used in

diagnosis, it highlights the need for this genus to be

reviewed by someone with type specimens before him

rather than only descriptions in the literature.

Cosmochthonius australicus can be distinguished

from C. plumatus, the only other member of the

species-complex, by the form of the hysteronotal

puncta. On the other hand, these puncta are similar

A53

to those of some members of the foveolatus-complex

which also have seta J3 with even, dense, long cilia.

The gnathosoma of this species, as that of a number

of other members of the Profissurae, is unusual, as

initially described by Grandjean (1948 and 1954b),

The ribbon-like palp setae and the comb-like cheli-

cera] seta suggest that the food is swept out of a

liquid, possibly with the slits in the cowl also acting

as a sieve. Whilst the structures at the anterior end

of the pharynx may be a crushing or suctorial mech-

anism. On the other hand, although in South Aus-

tralia C. australicus is recorded from moister regions,

mallee scrubland is not an environment in which free

water is often present.

Cosmochthonius plumatus Berlese

Cosmoachthonius plumatus Berlese, 1910: 221.

Casmochthonius plumatus: Grandjean, 1950a: 78.

Length, 300 (ex Italy—Pm), Cheliceral fixed digit

with 4 well developed teeth, seta ch2 with 6 prongs,

Seta s2 with long cilia. Seta J3 form only partially

known, but cilia uneven with sparse Jong cilia and

dense short cilia. Puncta on posterior hysteronotal

shield (NS4) small, varying in size and well separated,

asiaticus-complex

Diagnosis: No sculpturing covering posterior hys-

teronotal shield (VS4). Cilia on hysteronotal seta J7

even, long and sparse, between 8-22 on 1 side,

Remarks: Two species are included in the asiaticus-

complex. One species (C. suramericanus), because it

apparently has only | claw on each of its pretarsi,

may be based on a nymph

Cosmochthonias asiaticus Gordeeva

Cosmochthonius asiaticns Gordeeva, )980: 846.

Length, 319 (ex Tadzhikstan—Ps)}, Seta s2 long,

ciliate as 72. Seta J2 long (reaches back almost Lo

setal base /4). Seta J3 with 8 long, sparse cilia on

each side, evenly spaced along seta.

Cosmochthonius suramericanas. Harmer n.stat,

Coxsmochthonius pluntatus suramericanus Hammer,

[958: 23.

Length, 220 (ex Argentina—NTe).. Seta 92 long,

ciliate as j2. Seta J3 with about 25 long, sparse cilia

on each side.

PHYLLOZETES Gordeeva

Phyllozetes Gordgeva, 1978; 1099, Type designation

(original): “Cosmochthonius entmae Berl., 1910"

Type-species; Phyllozeres emmae (Berlese, 1910: 222)

4 REC, S, AUST, MUS,, 14 (1S): 327-759

Diagnosis: Cosmochthoniidac. Some notal setae (/3,

J4, Z3, Z4) long, stout, leal-like with broud lateral

membrune bearing marginal cilia, Pretarsi 1, I} and

HI with 2 claws (stout central claw, slim anterior

claw), pretarsus TV with 3 claws. Coxites I] and IV

merged together. Fewer selae. on femur IL and tarsi

HW} and IV (5, 13 and 12).

Distribution: Widespread—Texas (Nr); France, east

Crimea (Pe); Algeria, Austria, Greece, Hungary, Italy

(Pm); Japan (Pe); India (Ol); Kormodo Islands (Am):

South Australia (Aa).

Some |iteraiure suggests that species are found in

humus imongst trees or in moist ground even if poor

in organic material. But PAvilozetes is also recorded

from rapid draining sandy soil with depleted vegeta-

won. In South Australia it was collected in a fairly

dry sandy area, as well as from moister sites, | having

a substantial humus layer.

Remarks. Phyllozetes is certainly closely allied to

Cosinochthonius and Krivelutskiella. Wt may in the

future prove preferable to group these genera together.

The name “Ovechthonius’ is used in cambination

with P. evmiae by Mahunka (1980: 110) with the

authority of Gordeeva 1978: 1099. This is presumed

to result from confused communication, since Ovo-

chthonius Ryabinin in Ryabinin and Krivolutzky, 1977

is grouped in the Heterochthoniidae.

Five species are included in Phyplluzeres: P. emmiae

(Berlese, 1910); P; Aypoquercus McDaniel and Bolen,

)980), P. latifolius Gordeeva, 1980; P. osithchnjukovi

Gordeeva, 1980; PF. tauricus Gordeeva, 1978.

The specimens collected in this study are grouped

in FP, emmiae, Because it is difficult to delineate this

species, the other 4 species ure briefly described. P.

Aypoquercus: length, 285 (ex Texas—Nr); seta jl Y-

shaped; setae J2-/2. close (*% 0.5 J2-72); setae J3,

£3 similur to /4, 24, with short, dense marginal cilia

(over 50 on | Side), comes I, 11, HL, VY with 3,3,

3, 3 setae. P. latifolius: length, 150 (ex Canary

Islands— Pm, Crimea-—Pe); seta /1 unbranched; setae

J2-J2 close (& OF J2-272); setae J3, Z3 narrower,

with short, medium spaced marginal cilia (about 32

on | side); setue J4. 274 broader, with long, medium

spaced marginal cilia (about 32 on | side); number

uf setae on coxiles unknown. #. osi/hchnjukavi. length,

160 (ex Russia—Pe); seta jl unbranched, setac J2-

J/2 with bases touching (less than x 0.) /2-772); setae

J3, 23 narrower, with short, dense marginal cilia

(about 55 on 1 side); setae /4, 24 broader, with

medium lengthed and long. medium spaced marginal

cilia (about 35 on | side); number of setae on coxiles

unknown. P. fauricus: length, 285 (ex Crimea—Pe);

seta 7] Y-shaped, seta J2-J2 widely spaced (x 0.8

or more J2-22); setae J3, 73 narrower, with long

sparse marginal cilia (about 'S on | side); 4, 24

June, 1Y82

broader, with long, sparse marginal cilia (about 17

on | side), coxites 1, TH, ILL, FV with 3, 3, 3, 3 setae.

Phyllozetes emmae (Berlcse)

(Fig. none)

Cosmochthanius emmae Berlese, 1910: 222.

Cosmochthonius enmae Berlese; Mahunka, 1977:

254,

Ovochthonius emntae (Berlese): Mahunka, 1980; 110,

Phyllozetes emmae (Berlese): Gordeeva, 1980: 849,

Female

lyory-white to pale straw-coloured, Cuticle covered

by dirty, hyaline, adhesive exudate. Idiosomal Jength

200 (4, 187.5-210); appendage lengths (for 187.5 ex

Piccaninnie Ponds)—ch 11, pa 40, 1 72.5, 17 57.5, LU

60, #F 72.5; femur breadths—pa 10, J 19, 77 18, IT

15, /F AS.

Hysteronotal shields appear to lack seulpturing,

Proteronotal seta jl bifurcate (Y-shaped), seta z]

bifurcate (T-shaped) with posterior branch about

X0.5 length of anterior branch. Hysteronotal setal

pair J2 almost (X0.8 or more) as far apart as distance

between sctal bases. J2-22, Anterior erectile setae

(J3, 23) with lateral lamellae gradually tapering, and

long, dense to medium spaced cilia along margin

(about 30 on | side), Posterior erectile setae (/4, 24)

with broader gradually tapering lateral lamellae, and

long, dense to medium spaced cilia along margin (32-

35 on | side). Shape of most other somal setae similar

to that of P. tauricus, Coxites 1, 1, LI, TY with 3,

2, 3, 4 setae, Central margin of genital shield with 6

Jg setae. Chaetotaxy and solenidiotaxy of appendages

similar to C. australicus, except femur 11 has only 5

setae (sela a absent), tarsus Il] has only 13 setae

(setae p2, py2, v3 absent), tarsus 1V has only |2 setae

(selae pv2, v3 absent). Claws: I, I], I[1-2; P'V-3.

Gnathosoma, internal structures around pharyns,

genital tracheae, structure of alimentary canal and

contained food, ovipositor, all similar to those of C.

australicus.

Material examined: Two females (N1979383 and

N1979384), litter and sparse moss, under Eucalyptus

incrassata, mallee scrubland, Ferries-McDonald,

20.6.1974, D. C. Lee. One female (N1979382), litter,

under Banksia ornata, mallee-heath shrubland,

Tamboore, 4,7.1974, BD. C. Lee. One female

(N1979381), litter and sparse prass, under Acacia

sophorae, coastal scrubland, Piceaninnie Ponds,

20.8.1975, DC. Lee.

Pistrihution: Uncertain because of difficulty of

identification. Possibly Palaearctic and Australian

regions. South Australia; Ferries-McDonald, mallee-

broombush open-scrubland, 2 (1/8); Tamboore, mal-

lee-heath tall open-shrubland, | (1/8); Piccaninnie

Ponds, coastal closed-scrubland, | (1/8).

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOILS

Remarks: Specimens regarded as P. emmae are

described from 3 localities other than the type mate-

rial and that from South Australia, These descriptions

are almost limited to 2 erectile hysteronotal setae (J3

and/or J4) and they show considerable variation. On

the original illustration of the type (Berlese 1910: fig.

49) from Italy (Pm), the erectile hysteronotal setae

have marginal cilia which are so short and dense they

are hardly discernable. On the other hand Mahunka’s

(1980: figs. 11, 12) illustrations of these setae shows

the marginal cilia as long and medium-spaced to

sparse (22-27 on 1 side) with seta J4 slightly bigger

but similar to /3. Mahunka’s (1977: fig. 9) illustration

of J3 or J4 on a specimen from Vienna (Pe) is similar

to this latter description of the type. Seta J3 or J4

on a specimen from Komodo Island (Am) (Mahunka

1977: fig. 8) has short dense marginal cilia (about 56

on | side) and is almost parallel sided with an acutely

tapering tip, therefore, it is excluded from P. emmae.

Specimens from the Crimea (Pe) (Gordeeva 1980:

figs. 5, 6) have short medium spaced marginal cilia

(29-31 on | side) and should possibly also be excluded

from P. emmae.

In identifying the South Australian specimens,

Mahunka’s (1980) redescription of the type is

accepted, and the fact that there are more and denser

marginal cilia on setae J3 and J4 is regarded as an

intraspecific variation. A more comprehensive redes-

cription of the type may require that a new species

be established for this material.

Family SPHAEROCHTHONIIDAE Grandjean

Sphaerochthoniidae Grandjean, 1947a: 224.

Type-genus: Sphaerochthonius Berlese, 1910

Diagnosis: Cosmochthonioidea. Minute (237.5-375),

whitish to brown mites. One complete hinged hyster-

onotal fissure (TB1) present; second hysteronotal fis-

sure (7B2) just posterior to seta J2, but obscured by

anterior hysteronotal shield (VS1); 3rd hysteronotal

fissure (T7B3) present on larva or some nymphs in

region of seta J3, rarely represented on adult, although

sometimes transverse dorsal ridges near setae /3 and

J4, No hysteronotal setae erectile or long (not as long

as distance between setal bases J/ and J/4) and,

except setae J2 and Z2, all ciliate and seta 21 T-

shaped. Setae J2 and Z2 simple, shorter than distance

between setal bases J2-J2, and normally under anterior

hysteronotal shield. Cowl without longitudinal slits.

Anterior cheliceral seta (ch2) pectinate with 3 prongs.

Tarsus I bearing 3 solenidia. Tarsi J-IV with 3 claws.

Remarks: Despite the superficial lack of similarity

between the Sphaerochthoniidae and the Cosmo-

chthoniidae, they have usually been regarded as closely

allied. Also (see above in the remarks on the Cos-

mochthonioidea) Sphaerochthoniidae has long been

regarded as allied to the Protoplophoridae although

355

this is not reflected in the classification by Balogh

(1972). Regarding the relationship of the Sphacro-

chthoniidae and Protoplophoridae, the nature of the

prehysteronotal fissure is an important character com-

mented on in the remarks on Sphaerochthonius splen-

didus.

The Sphaerochthoniidae includes only a single

genus: Sphaerochthonius Berlese, 1910a.

SPHAEROCHTHONIUS Berlese

Sphaerochthonius Berlese, 1910: 223. Type designation

(original): “Hypochthonius splendidus Berl.”.

Type-species: Sphaerochtonius splendidus (Berlese,

1904: 26).

Diagnosis: Sphaerochthoniidae (monogeneric).

Distribution: Widespread—Canada (Nn), California

(Nc); Brazil (NTc); Ghana (Ew); Somali (Ee); Cau-

casus, Crimea (Pe); Algeria, Greece, Italy, Morocco

(Pm); Japan (Pc); Java (Om); Komodo Island (Am);

South Australia (Aa).

Remarks: Sphaerochthonius is an easily recognis-

able genus, but delineating the included species is

difficult. The unusually small specimen before me

has been placed in the type-species, S. splendidus,

which was originally described from the Mediterra-

nean region. The reason for this placement is partly

because the poor descriptions of a long established

species make it less exclusive. In order to give some

idea of the degree of similarity of the South Australian

specimen to other species, | have given a short

description of the other species.

In describing S. suzukii, Aoki (1977) has reviewed

the genus, especially with regard to the form of the

somal setae. In referring to hysteronotal setae J2 and

Z2 (as dl and d2), he regards the fact that they are

short and simple as diagnostic of S. suzukii. In fact

this attribute is true for the South Australian speci-

men, the larva of an unnamed species (Grandjean

1934: fig. 2) and for S. suwblanatus (Mahunka 1977a:

fig. 7). In the case of S. swblanatus and S. suzukii,

the mite drawn is slightly squashed and the anterior

hysteronotal fissure (7B1) has opened up so that setae

J2 and Z2 are no longer obscured by the anterior

hysteronotal shield. On the illustrated specimen of S,

suzukii, the second hysteronotal shield (N.S2) appears

fused to shield NS3, whilst for S. swblanatus it is

attached to shield NS1. It is likely that in other

species where setae J2 and Z2 are obscured they

have not been described and so what Aoki (1977)

would regard as setae J2 and Z2 are in fact J3 and

Z3.

The 7 species included in Sphaerochthonius are

considered below mainly in alphabetical order but

356

with S. splendidus, in which the South Australian

specimens are grouped, last.

Sphaerochthonius gemma (Oudemans)

Hypochthonius gemma Oudemans, 1909: 319.

Sphaerochthonius elegans Berlese, 1910: 266.

Cosmochthonius gemma (Oudemans): Oudemans,

1917: 25.

Sphaerochthonius gemma (Oudemans): van der

Hammen, 1959: 26,

All specimens regarded as nymphs since few genital

setae and only | claw on each of its pretarsi. Length,

268 (ex Java—Om). All illustrated notal setae, except

z2, T-shaped, ciliate and stick-like. Two branches of

hysteronotal setae subequal in length. Setae in hys-

teronotal ranks, 3, 4 and 5 with horizontal branches

directed transversely. Two heavy lines in illustration

(Oudemans 1917: fig. 51) run through setae J3 and

J4 and I agree with Grandjean (1932a: 34) that they

represent dorsal ridges, because posterior line would

otherwise have to be a 4th fissure.

Sphaerochthonius longisetus Mahunka

Sphaerochthonius longisetus Mahunka, 1977a: 256.

Length, 360-374 (ex Komodo Island—Am). Many

dorsal setae (jl, j2, JI, Z1, S1, S2, 24, J5, S5)T-

shaped, papillose and leaf-like. Seta j1 similar but

ciliate. Setae s1, ?s2, J3 and Z3 unbranched, papil-

lose and stick-like; seta J3 unusually long (about

distance between setal bases Z1-Z2).

Sphaerochthonius phyllophorus Balogh and Mahunka

Sphaerochthonius phyllophorus Balogh and

Mahunka, 1969: 32.

Length, 289-319 (ex Brazil—NTb). Setae J1, zl,

j2 T-shaped and possible intermediate between stick-

like and leaf-like. “Anterior” hysteronotal setae T-

shaped, papillose and leaf-like, “Posterior” hysteron-

otal setae unbranched, papillose and leaf-like.

Sphaerochthonius suzukii Aoki

Sphaerochthonius suzukii Aoki, 1977: 85.

Length, 310 (ex Japan—Pc). Most notal setae cil-

iate (short, blunt cilia) and stick-like. Setae jl, z1,

J1, Z1, Sl, $2, J5, Z5, SS, J6 T-shaped, whilst s1,

j2, 82, J3, Z3, Z4 unbranched. Setae Z6 and S6, like

setae in file Za, unbranched, slim with long, tapering

cilia. Transverse dorsal ridge passes just posterior to

seta J3.

Sphaerochthonius transversus Wallwork

Sphaerochthonius transversus Wallwork, 1960: 377,

Length, 256-277 (ex Ghana—Ew). Form of notal

setae similar to S. gemma , especially since horizontal

REC. S. AUST. MUS., 18 (15): 327-359

June, 1982

branches of setae J3 and Z3 directed transversally

(not an artifact, occurs on living specimens) and 2

branches of each seta similar in length. Different,

since setae s1 and j2 unbranched. Apparently dorsal

ridge associated with 3rd hysteronotal setal rank, no

such ridge associated with 4th rank as on S. gemma.

Nymphs grouped in this species have long seta J3

like S. /ongisetus and may not be conspecific.

Sphaerochthonius wallworki n.sp.

Sphaerochthonius spec. Wallwork, 1960: 382.

Length, 298 (ex Ghana-Ew). As pointed out in

original description, this species very similar to S.

gemma, having 2 transverse dorsal ridges passing

along 3rd and 4th hysteronotal setal ranks, and mainly

T-shaped, ciliate, stick-like setae. On the other hand,

seta j2 unbranched and horizontal branches of setae

J3 and Z3 directed longitudinally with shorter ante-

rior branch. Nymph grouped in this species with

horizontal branches of setae /3 and Z3 directed trans-

versely as S. gemma. Despite 4 setae in file Za, not

certain that this nymph belongs to S. wallworki.

Sphaerochthonius splendidus (Berlese)

(Fig. 30)

Hypochthonius splendidus Berlese, 1904: 26, fig. 37.

Sphaerochthonius splendidus (Berlese): van der

Hammen, 1959: 25.

Adult

Length, 237.5 (1). Hysteronotum light brown, pro-

teronotum mid-brown. Cuticle covered by granulated

hyaline adhesive layer. On soma, network of lines

(with granules clustered around) encloses 5-sided or

6-sided areas. Granules also clustered around notal

setae. Refractile parts (external malae, cheliceral

extremities, setae and claws) paler than general inte-

gument.

Single specimen illustrated (Fig. 30), slightly

squashed by coverslip, but natural body-form laterally

compressed like PAthiracarus (Holonotina) or Pro-

toplophora (this subcohort). Anterior 2 pairs of legs

curl up under edges of proteronotum whilst posterior

2 pairs of legs curl up under flaps from pleural

shields. Adanal shields may be capable of closing

over anal shields. No evidence confirming acutely

hinged prehysteronotal fissure, but soma looks capa-

ble of folding so that cowl covers genital shield.

Anterior transverse hysteronotal fissure (7B1) also

appears hinged. By squashing mite, second hyster-

onotal shield (NS2—narrow band bearing setae /2

and Z2) moves backwards from under anterior hys-

teronotal shield (VS1) near level of setae JI and Z1

to be exposed at almost level of setae J3 and Z3. In

illustration (Fig. 30), posterior of 4 broken transverse

lines appears to represent posterior edge of folded

SARCOPTIFORMES (ACARI) OF SOUTH AUSTRALIAN SOLLS 487

hyaline membrane between venter of first byster-

onatal shield (VS1) and anterior edge of second hys-

teronotal shield (NS2), Reticulate hyaline adhesive

layer passes unbroken across anterior transverse hys-

teronotal fissure so that any movement of shields

could mean rupturing this. layer.

Legs difficult to observe because folded up, Tro-

chanter and femur [II and IV similar to Cosmto-

chthonius. Pretarsus 1V with 3 claws (stout central

claw, 2 slim lateral claws). Adanal setae Za and

hysleronotal setae J6, Z6 and S6 simular io J4. File

Za includes 4 setae. Single, dense granular bolus

present, but no eggs observed inside specimen.

Material examined: One adult (N1979385), grass

and moss or litter under Eucalyptus viminalis, savan-

nah woodland, Chambers Gully, 12.6.1974, D. C.

Lee.

Distribution- Medilerranean-type climate: Italy,

Greece (Pm); South Australia (Aa). South Australia:

Chambers Gully, savannah woodland, 1 (1/8),

Remarks: The specimen from South Australia,

although much smaller than the type (310), can be

grouped in S. splendidus. Because [ have not wished

to damage the single specimen, the description is

unusually limited. Earlier work on the hysterosomal

shields of ihe adult (Grandjean 1932a) and larva

(Grandjean 1934) of a Sphaerochthonius species,

indicating a hinged first transverse hysteronotal is-

sure that is folded forward under the anterior shield,

is confirmed, which clarifies some of the confusion

about hysteronotal fissures resulting from more recent

descriptions of Sphaerachthonius. Furthermore, the

presence or absence of small simple setae in the

second hysteronotal rank as allribules to distinguish

species can now be seen as resulling from these setae

sometimes being hidden, Unfortunately, I have not

established whether or not the transverse prehyster-

onotal fissure is acutely hinged or not. Lf the proter-

onotum can be folded right down, distinguishing the

Cosmochthonioidea from the Protoplophoroidea may

not be tenable.

ACKNOWLEDGEMENTS

] am indebted to the Interim Council of the Aus-

tralian Biological Resources Study for funds for

equipment, and to the Mark Mitchell Research Foun-

dation and to the Science and Industry Endowment

Fund for funds to travel in connection with this pro-

ject,

| also appreciate the making available of certain

specimens for study by Dr J, Aoki (Yokohama

National University, Japan), Dr R. E, Crabill (Smith-

sonian Institution, U.S.A.) and Mr T, Moulton (Mac-

quarie University, N.S.W,). Thanks are due to Dr B.

G, M. Jamieson, University of Queensland, Brisbane

for commenting on the manuscript,

My greatest debt of gratitude is to Ms Jenni

Thurmer for her excellent illustrations and to Mra

Debbie Melloy for typmg this manuscript.

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POLYCLAD TURBELLARIANS FROM THE

SOUTHERN COASTS OF AUSTRALIA

BY STEVEN PRUDHOE

Summary

Eighteen species of polyclad turbellarians are recorded from the southern coasts of Australia, 14 of

which are from South Australia and 4 from Victoria. Of the South Australian species, 2 are also

recorded from Tasmania, and | of the Victorian species has been recorded hitherto only from South

Australia. Of the species dealt with, 3 are regarded as new to polyclad systematics, and these are:

Notoplana distincta sp.nov., and Pseudoceros lividus sp.nov., from South Australia and

Candimboides cuneiformis gen. et sp.nov. from Victoria. The genus Ancoratheca is diagnosed. The

descriptions are extended for the following 5 species: Ancoratheca australiensis, Notoplana

longicrumena, Planocera edmondsi, Cestoplana meridionalis and Cycloporus australis. Three

known species, Euplana gracilis, Echinoplana tenerrima and Pseudoceros reticulatus show very

wide gaps in their geographical distribution, suggesting that polyclads may be artificially

transported from one part of the world to another, where they may settle, if conditions are suitable.

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUSTRALIA

STEPHEN PRUDHOE

British Museum (Natural History), London, England

(Manuscript accepted 14 July 1981)

ABSTRACT

PRUDHOE, 8. 1981. Polyclad turbellarians from the southern

coasts of Australia. Rec. 5, dust, Mus, 18 (16): 361-384,

Eighteen species of polyclad turbellarians are

recorded from the southern coasts of Australia, 14 of

which are from South Australia and 4 from Victoria.

Of the South Australian species, 2 are also recorded

from Tasmania, and | of the Victorian species has

been recorded hitherto only from South Australia, OF

the species dealt with, 3 are regarded as new to

polyclad systematics, and these are: Notaplana dis-

lineta sp.nov,, and Pseudoceros lividus sp,nov., from

South Australia and Candimboides cuneiformis gen.

et sp.nov. from Victoria. The genus Ancoratheca is

diagnosed. The descriptions are extended for the fal-

lowing 5 species: Amcorarheca australiensis, Nota-

plana longicrumena, Planocera edmondsi, Cestoplana

meridtonalis and Cycloporus australis, Three known

species, Euplana gracilis, Echinoplana tenerrima and

Pseudoceras reticulatus show very wide gaps in their

geographical distribution, suggesting that polyclads

may be artificially transported from one part of the

world to another, where they may settle, if conditions

are suitable.

INTRODUCTION

Background, Methods and Acknowledgements

The following report is based chiefly on a collection

at polyclad turbellarians housed in the South Austra-

lian Museum, and for the pleasure of studying this

material the writer is indebted to Mr David C. Lee

and Mr Ifor M, Thomas. Grateful acknowledgement

is made to Dr 8. J. Edmonds, formerly the University

of Adelaide, Mr S, A, Shepherd of the Department

of Fisheries, Adelaide, Dr G. C, B. Poore of the

Minisiry of Conservation, Melbourne, and Mr Neville

Coleman of the Marine Photographic Index, Caring-

bah, N.S.W,, for sending specimens to the writer at

different times during the pasi 25 years.

Little has been published on polyclads of the coasts

of South Australia and Victona and it is therefore

not surprising that of the 18 dealt with here only

Notoplana australis (Schmarda) and Haploplana rosea

Prudhoe have been recorded previously from these

coasts. Fourteen species are here recorded from South

Australia for the first time, and of these 7 are new.

Of the known species, 5 have beet recorded from the

vicinily of Part Jackson, N.S.W., 2 from the coast of

Victoria, and 1, Thvsanozaon skoitshergi Bock, from

the coast of Wester Australia. Of the 4 species now

recorded from Victoria, only 1 has been recorded

from other Australian coasts, bul whether or not this

relationship denotes any zoogeographical significance

requires further investigation of the polyclad fauna of

Victoria, especially as Bennett and Pope (1960) indi-

cate the Victorian coast as a cold-lemperature zone

situated between 2 warm-temperate regions. Hitherto,

only 2 species of polyclads have been recorded from

Tasmania, namely, Notoplana australis and Microcelis

schauinslandt Plehn, bul the number is now increased

to 4 with the records af Planocera edmonadsi Prudhoe.

and Pseudaceras reticulatus Yer and Kaburaki,

The present study and a review of the literature,

sugpest that Noroplana australis is probably the com-

monest species of polyclad inhabiting the southern

and south-eastern coasts of Australia. Moreover, the

species has alsu been recorded from New Zealand

and Chile.

The occurrence of Euplana gracilis, Echinoplana

lenerrime and Pseudoceros reticulatus, inhabitants

of other regions of the oceans, in the present collections

reveals an important problem in the study of the

geographical distribution of polyclad turbellarians,

for it now seems that some species are or have been

transported artificially by some means or other from

their endemic areas to very distant localities offering

suitable habitats. This observation gives some credence

to Laidlaw's (1904) record of the north-eastern Atlan-

tic boreal polyclad Cryptocelides loveni Bergendal

occurring in Port Phillip Bay, Victoria—the authen-

ticity of this record has always been doubted by

subsequent writers, even by Laidlaw himself.

The examination of the above-mentioned material

has been carried out on whole specimens cleared in

methyl saheylate and on longitudinal serial-sections

of copulatory complexes cut at 10 wm and double-

stained with haemalum and cosin, For the preparation

of the serial sections, the writer would like to thank

Mr D. W. Cooper of the Histology Section of the

British Museum (Natural History)..

Deposition is in the Australian Museum, Sydney,

Australia (AM); British Museum (Natural History),

London, England (BM); National Museum of Victoria,

Melbourne, Australia (NMY); and South Austrahan

Museum, Adelaide, Australia (SAM).

362

List of Species

From South Australia

Discocelis australis Hyman

Enterogonia orbicularis (Schmarda)

Ancoratheca australiensis Prudhoe

Notoplana australis (Schmarda)

Notoplana longicrumena Prudhoe

Planocera edmondsi Prudhoe

Echinoplana tenerrima Haswell

Cestoplana meridionalis Prudhoe

Pseudoceros reticulatus Yeri and Kaburaki

Pseudoceros lividus sp.nov.

Notoplana distincta sp.nov.

Tripylocelis typica Haswell

Thysanozoon skottsbergi Bock

Cycloporus australis Prudhoe

From Victoria

Latocestus sp.innom.

Euplana gracilis (Girard)

Candimboides cuneiformis gen. et sp.nov.

Hoploplana rosea Prudhoe

From Tasmania

Planocera edmondsi Prudhoe

Pseudoceros reticulatus Yeri and Kaburaki

SYSTEMATICS

Family DISCOCELIDIDAE Laidlaw, 1903, emended

Poche, 1926

Discocelis australis Hyman, 1959

Locality: South Australia—one specimen (acci-

dently destroyed) under rocks, West Is., S.A., Shep-

herd, 29.v.73.

Morphology: Broadly oval in outline, firm consist-

ency, about 18 mm long and about 12 mm maximum

width, which occurs in cephalic region. Ground colour

greyish white with tinge of brown along median line

on the dorsal surface. Surface also bears many scat-

tered dark brown spots, so evident in most species of

Discocelis Ehrenberg. Cerebral organ situated at about

4mm from anterior margin of the body and near

anterior end of pharynx. Eyes in region of cerebral

organ, arranged more or less in two elongate clusters

representing cerebral eyes. Tentacular eyes lie more

dorsally than cerebral eyes and disposed in two

rounded groups, laterally to cerebral eye-clusters.

Numerous very small anterior marginal eyes distrib-

uted in two or three irregular rows which extend

posteriorly to level of tentacular eyes. Distribution of

eyes precisely as figured by Hyman (1959). Mouth

opens into posterior region of the pharyngeal chamber,

at about 7 mm from hinder margin of body; common

genital pore about 3 mm posterior to mouth.

REC. S. AUST. MUS., 18 (16): 361-384

June, 1982

As seen in the whole specimen cleared in methyl

salicylate, the copulatory complexes, although not

fully developed, closely resemble those described and

figured by Hyman for Discocelis australis, originally

collected at Long Reef, near Sydney, N.S.W.

Family LATOCESTIDAE Laidlaw, 1903

Latocestus sp.innom.

Locality; Victoria—one fragmentary juvenile spec-

imen (NMV/G3292), Crib Point Benthic Survey; Stn.

41 N/5, Westernport, date unknown.

Morphology. Elongate, about 5 mm long, brownish

dorsally and ventrally. Eyes spread fanwise over

cephalic region and marginal eyes form complete

series round body.

Remarks: The genus Latocestus Plehn has not been

recorded hitherto from Australia, but it is widely

distributed in the tropical and temperate regions of

the Atlantic and Indo-Pacific Oceans. Of those species

recorded from the Indo-Pacific region, the present

form bears some resemblance to L. argus Laidlaw

and L. maldivensis (Laidlaw) in the coloration of the

body and in the distribution of the eyes.

Family STYLOCHIDAE Stimpson, 1857

Enterogonia Haswell, 1907

Diagnosis: Stylochidae with rudimentary tentacles.

Marginal eyes in complete series around body; cerebral

eyes merge with frontal eyes to spread fanwise ante-

riorly and join with marginal eyes; tentacular eyes

located nearer dorsal surface than cerebro-frontal eyes.

Genital pores well separated. Pair of vasa deferentia

unite to form ejaculatory duct; no seminal vesicle.

Prostatic organ very small and “free”. Penis-papilla

thick, unarmed. Vagina long, forming anteriorly-

directed loop from female antrum. Vagina terminating

in genito-intestinal canal.

Type-species: E. orbicularis (Schmarda, 1859)

Stummer-Traunfels, 1933.

Type-locality: although originally recorded from the

Chilean coast, Stummer-Traunfels states that

there is evidence that Schmarda’s specimens were

collected in New Zealand.

Remarks: The diagnosis of Enterogonia by Hyman

(1959) is modified mainly because of a reinterpretation

of the structure of the reproductive system (see mor-

phology of E. orbicularis).

Enterogonia orbicularis (Schmarda, 1859)

Synonymy: Polycelis orbicularis Schmarda, 1859;

Enterogonia pigrans Haswell, 1907; Enterogonia

pigrans var. novaezealandiae Bock, 1925; Enterogonia

orbicularis Stummer-Traunfels, 1933; Enterogonia

pigrans Hyman, 1959; Enterogonia orbicularis orbi-

POLYCLAD TURBEL] ARLANS FROM THE SOUTHERN COASTS OF AUST.

culariy Wyman, 1959: Enterogonia erbicularis pigrans

Hyman, 1959,

Diagnosis; Fleshy forms, oval or elliptical in outline,

varying from 10-34 mm long and 5-23 mm wide.

Dorsal surface greenish, pale olive-green, dark grey

or light brown, ofien with numerous greenish brown,

dark olive or brownish spots, Nuchal tentacles searcely

noticeable, Antetiorly-extending cerebral eyes com-

mence behind cerebral organ. Mouth situated cen-

trully; elongate pharynx in middle third of body, with

10-12 pairs of lateral folds. Long, narrow, ejaculatory

duct opening into ndrraw male antrum through

unarmed penis-papilla. Epithelial lining of “shell”-

chamber thrown into a spiral fold,

Localities: South Ausiralisa—2 specimens (SAM/

V243\-V2637 consisting of uncut portions and serial

sections), shore line, Ceduna. B. L. Baker, Nov. 1940;

| specimen (SAM/V¥2625-¥2630 consisting of an

uncut partion and serial sections), Whiltlebee Point.

W, Zeidler, 2417.75; specimens (BM/1980.5.1.25),

under stones, upper tidal zone, Port Noarlunga, St

Vineent Guld. 5, J. Edmonds, 24,x.53 and on Aldinga

Reef, St Vincent Gulf, S.A. Shepherd, 2.ix,72; spee-

imens (BM/1980.5.1.29), Japanese oysters. Caffin

Bay, near Port Lincoln S.A. Shepherd, date unknown:

specimen (BM/1980.5.1.30), south side of Salt Creek,

Tumby Bay, Spencer Gulf, S. J, Edinonds, date

unknown.

Morphology: The above specimens [rom South Aus-

tralia, measuring 9-22 mim in length, possess a very

small “free” prostatic organ and agree exactly wilh

that described by Bock (1925) and Stummer-Trauofels

(1933) in the New Zealand nvaterial. This organ may

be interpreted as a small swelling arising from the

dorsal wall of the ejaculatory duct, where the duct

turns ventrally to enter the penis-papilla. It is lined

with a glandulir epithelium provided with long cilia

and coaied with a musculature distinctly thicker than

that in the wall of the ejaculaiory duct. Neither

Haswell (1907) nor Hyman (1959) appear to have

recognized this organ in specimens from N.S,W.

Hyman (Fig. i8) depicts a dilation of the ejaculatory

duct and a thickening of its muscular wall (“a curved

chamber”) immediately before the duct turns ven-

trally, and it seems possible that she mistook the

Insignificant prostatic organ merely as a swelling of

the ejaculatory duct. Morcoyer, in the present mate-

rial, Where the prostatic organ opens into the ejacu-

lutory duct, the latter is provided with a relatively

thick musculature and further suggests that the male

complex in the material examined by Flaswell and

Hyman might not have been fully developed.

Remarks: These worms appear to associate with

colamivs of bivalve molluscs, but there is no direct

evidence that they are predatory on the colonics, even

theugh they undoubtedly scavenge among them by

464

feeding on dead or injured ammals. &. orbicularis

seems to be one of the better known Australian poly-

clads, for Haswell (1907) and Hyman (1959) des¢ribed

it from N.S.W, under the name Enterogenia pigrans,

and Bock (1925) deseribed specimens trom New Zea-

land and considered them to be so very closely related

to Maswell’s form that he gave them the name #arer-

ogonia pigrans var. novaezealaddiae, Stummer-

Traunfels (1933) redescribed Schmarda’s (1859) ong-

inal material of Polycelis orbicularis [rom New Zca-

land and considered it to be identical with &. pigrans

var. novaezealandiae Bock. The main differences

between the specimens from New Zealand and those

from Australia are found in the greater size (26-

34 mm in length of the former and 10-15 mim in the

latter) and the supposed absence of prostatic organ

in the latter. In addition (o these features, Hyman

(1959) stated that the New Zealand form differed

from the Australian one in ils less roomy male antrum,

in the larger penis-papilla, and in the lack of an

expanded lumen in the distal region of the vagina,

Bock (1925) rightly discounts the difference in size

as a specific character. Concerning the differences

listed by Hyman, with the exception of the prostatic

organ, (hey are merely variations influenced to a more

ar less degree by size and may be considered as

superficial and of no diagnostic importance,

ANCORATHECA Prudhoe, 1982

Diagnosis: Siylochidae with aval body. From anie-

rior end of pharynx, cerebro-frontal cyes spread [an-

wise over cephalic region; tentacular eyes in two sinall

groups among cerebro-lranlal eyes; marginal eyes var-

iably distributed, Genital pores adjacent. Accessory

and true seminal vesicles together forming, trilobed

or adnchor-shaped structure: prostatic organ muscular,

vlongale, with narrow lumen and smooth epithetial

lining; penis-papilla bearing short stylet, lying in penis-

pocket opening into male antrum through shallow

penisssheath. Vayina simple, terminating in Lang’s

vesicle sometimes opening on ventral surface of bady.

Type-species} Anceratheca australiensis Prudhoe,

1932

Other species: 4. pacifica (Bock, 1923).

Remarks: The single specimen of 4. austealienss

described below is grouped in the Slylochidae because

of the distribution of the eyes, the long pharynx «ind

the posterior positions of the copulatory complexcs.

li bears a sirong resemblance to Neesrylochus paci-

fieus Boek 1923, from Juan Fernandez ts. off the

coast of Chile. This similarity with N. pacificus seems

to indicate that the latter 1s not congencric with Lhe

type species of Neostylochus (N. fulvopunctatus Yeri

and Kaburaki 1918) from Japan. Por instance, N.

Julvopunctatiiy possesses a simple. clongate seminal

vesicle and 4 vagina bulbosa, whereas V, pacificus

364 REC. 8. AUST. MUS., 18 (16): 361-384

and A, australiensis have a trilobed seminal vesicle

and no vagina bulbosa, differences which are here

regarded as generically important. Therefore, the latter

2 species are grouped in Ancoratheca.

Ancoratheca australiensis Prudhoe, 1982

(Fig. 1)

Locality: South Australia—holotype (SAM/V2608-

V2612, consisting of an uncut portion and serial sec-

tions), 15 metres deep, Upper Spencer Gulf, S.A.

Shepherd, 5.ix.73.

Morphology: Fleshy, somewhat oval in outline,

rounded anteriorly, tapering posteriorly, and of firm

consistency. About 10 mm long and up to 6 mm wide.

Dorsal surface of body (when preserved in alcohol)

buff-coloured, dappled with dark spots, due to under-

lying gonads, except anteriorly and marginally, where

body is somewhat lighter in colour. No tentacles. Very

small marginal eyes disposed in three to five irregular

rows and confined to about anterior third of total

length of body. Cerebral organ somewhat bilobed. At

about 3.5 mm from anterior margin, further minute

eyes spread fanwise over the cephalic region to merge

with marginal eyes and may be regarded as cerebro-

frontal eyes. Among these, a short distance anteriorly

to the cerebral organ, there is a cluster of a few eyes

on either side of the median line. These are nearer

to dorsal surface of the body and may be regarded

as small clusters of tentacular eyes.

Pharynx narrow, about 6 mm long, mainly in pos-

terior half of body; shows no lateral folds, only a

slight rippling of lateral margins. Mouth opens into

hinder region of pharyngeal chamber. Numerous

intestinal branches extend to the periphery of body

and do not appear to anastomose.

Genital pores very close together and appear to

open into a short narrow genital atrium or a depression

in the body-wall, lying at about 2.5 mm from posterior

margin of the body. Near posterior end of the pharynx,

the vasa deferentia become exceedingly muscular and

form a pair of elongate accessory seminal vesicles

which are directed postero-dorsally above ‘genital

atrium.’ These open directly into a muscular seminal

vesicle directed antero-ventrally, parallel with, and to

left side of, accessory vesicles. Prostatic organ elongate

and lies parallel with, and close to right side of,

accessory seminal vesicles. It is invested with very

thick musculature consisting of an outer layer of

circular fibres and a doubly thick inner layer of

longitudinal fibres. Through these layers the efferent

ducts of extracapsular gland-cells pass to open into

narrow lumen of prostatic organ. Latter lined with a

shallow glandular epithelium, which shows no radial

or longitudinal folding. Short muscular ejaculatory

duct extends from seminal vesicle uniting with a

muscular prostatic duct and enters very small mus-

June, 1982

cular penis-papilla, armed with a short stylet. This

lies in a small penis-pocket, which opens into a small

antrum masculinum through a shallow penis-sheath.

Female antrum short and narrow, leading into a

relatively wide vagina lined with an exceptionally tall

ciliated epithelium. Numerous ‘shell’-glands open into

a short portion of distal half of vagina to form a

‘shell’ chamber, epithelium of which appears to be

thrown into four or five spiral folds. From ‘shell’-

chamber, vagina extends posteriorly for a short dis-

tance to receive short common duct of uterine canals.

From this union, vagina interna continues posteriorly

to open into a spacious arcuate Lang’s vesicle through

a sphincter. Inner end of vesicle narrows considerably

to open to exterior through a tiny pore situated at

about 0.5 mm posterior to common genital opening.

The epithelium of the proximal region of the vagina

interna is thrown into radial folds, a feature commonly

associated with the ‘stalk’ of Lang’s vesicle. Lang’s

vesicle is in poor histological condition but it appears

as a membranous structure lined with large gland-

cells containing a secretion staining weakly with eosin.

An interesting feature is that it contains small packets

of spermatozoa, one of which is clustered around the

sphincter at the union of the vagina and the vesicle.

A few individual spermatozoa are also present in the

proximal region of the vagina interna suggesting that

fertilization may take place in the uterine canals.

Further, it might be thought that the packets of

sperm had been deposited through the body-wall by

hypodermic impregnation with a penial stylet of

another worm, but this does not seem to be so because

the wall of Lang’s vesicle is fused to the tiny external

aperture.

Remarks: A. australiensis may be distinguished from

A. pacifica by the marginal eyes not completely encir-

cling the body and by Lang’s vesicle opening on the

ventral surface of the body.

The fusion of the musculature of the seminal vesicle

(which consists principally of circular fibres) with its

accessory vesicles appears to produce a structure sim-

ilar to the trilobed or tripartite seminal vesicle of the

genus Stylochus (Imogine). The tight clustering

together of the organs of the male copulatory complex

may not be normal but due to contraction of the

hinder region of the body.

The opening of Lang’s vesicle directly to the exterior

is very interesting, because it could be regarded as

the original condition of the ductus vaginalis found

in several acotylean polyclads, particularly among

species belonging to the family Stylochidae, Bock

(1925a), however, assumes that the ductus vaginalis

has been derived from the ‘stalk’ of Lang’s vesicle,

and he presents evidence to support this assumption,

with particular references to Discocoelides langi Ber-

gendal, where an accessory canal connects the ‘stalk’

directly to the antrum femininum. Whilst Bock’s

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST.

assumption may be justified in some instances, it

should not be overlooked that in other instances the

ductus vaginalis might also have arisen solely from

Lang’s vesicle, which having made contact with the

exterior, as in the present form, eventually lost its

365

function solely as a receptaculum seminis and became

reduced to a narrow canal for the discharge of waste

reproductive materials, as well as for copulation, thus

being comparable with Laurer’s canal of trematodes

and the vagina of cestodes.

asv sv

sc Lv

FIG.1. Ancoratheca australiensis. A. Dorsal view. B. Sagittal section of copulatory complexes. asv., accessory seminal vesicles; Lv.,

Lang’s vesicle; po., prostatic organ; pp., penis-pocket; ps., penis-sheath; s., stylet; sc., “shell”-chamber; sv., seminal vesicle; uc.,

common uterine canal

Family LEPTOPLANIDAE Ehrenberg, 1831,

emended Stimpson, 1857

Euplana gracilis (Girard, 1850); Girard, 1895

(Fig. 2)

Locality: Victoria—one damaged specimen and

several fragments (BM/1980.5.1.11 and NMV/G3290

and G3291), Port Phillip Bay Environmental Study;

Stn. 137/3 and 137/5.

Morphology: Body delicate, elongate-oval; 3.25-

5.0 mm long, 0.9-1.75 mm maximum width (measure-

ments are estimates since no specimens complete). In

alcohol, dorsal surface yellowish brown with dark

brown spots from underlying ovaries; ventral surface

greyish. No tentacles. Cerebral organ 0.25 mm from

anterior end. Eyes few, in two rows each of 4-5 on

either side of cerebral organ. Mouth opens into middle

of pharyngeal chamber which lies in anterior half of

body closely behind cerebral organ. Pharynx has a

few pairs of shallow lateral pockets. Median intestinal

trunk extends from near cerebral organ to posterior

region giving off several pairs of lateral branches;

these anastomose initially but are free marginally.

Male and female genital pores separate and lie in

middle region well behind pharynx.

Remarks: Unfortunately, the histological condition

of the material does not permit of a precise exami-

nation for description, but sufficient has been made

out of the reproductive system to show that it is

undoubtedly comparable with that described by

Hyman (1939) in American specimens.

Euplana gracilis was known hitherto to occur only

on the Atlantic coast of North America, where,

according to Hyman (1940), it is common in the

benthic fauna of the boreal/warm temperate region

and ranges from Prince Edward Island, Canada, to

the western coast of Florida. Its occurrence in Port

Phillip Bay is therefore surprising, and it must be

assumed that the worm had somehow been transported

from America, possibly when eggs attached to the

366

hull of a ship hatched in Victorian waters. There 4s

also the possibility that they were carricd with oysters

or other animals and deposited in these waters. Bennett

and Pope (1960) apparently regard the Victorian coast

as a cold-temperate region, and it seems therefore to

be a suitable atea for Enplana gracilis-—clearly a

eurythermic species to establish itself,

Notoplana australis (Schmarda, 1859) Bock, 19]3

Synonymy: Polvoells australis Schmurda. 1859;

Leptaplana australis Laidlaw, 1904; Lepraplune

suleri Jucubowa, 1906; Noroplana australis

forma Asa Mareus, 1954,

Diagnosis: Body elongate oval, of variable size and

coloration. Nuchal tenlacles rudimentary or not

upparent. Byes of the cerebral groups smaller than

the tentacular eyes, Mouth in middle region of body;

pharynx with 12-17 pairs of lateral folds. Gemital

pores well separated. Seminal vesicle smaller than

prostatic organ, which has sia epithelial longitudinal

chambers, Penjs-papillu yery small and provided with

4 long stylet enclosed in elongate muscular penis-

pocket. Anirum masculinum narrow. Vagina bulbosa

large; Lang's vesicle clongute,

Localities. South Australia—specimen (SAM/

V¥2638-V2645— consisting of uncul portions dnd seria}

seclions). near Jones Island, Buird Bay,

W. Zeidler, 27.1).75; specimen (SAM/¥2640) under

tucks at Jow tide, Daly Head, Yorke Pernimsula,

W. Zeidler, 9.xi,76; specimen (SAM/¥V2647) Brown

Point, W. Zeidler, 9.viii.74; 2 specimens (SAM/V2648

and V2hd9), Wittelbee Point, neur Ceduna, W

Zeidler, 2.11.75; specimen (SAM /V2650) under inter-

tidal rocks, Pondalowie Bay, W. Zeidler, 9-xi. 7h: spee-

imen (BM/1980.5.1.8), Wittelbce Point, near Ceduna,

W. Zeidler, 2.111,75; speeitnen (BM/1980.5.1,9. in

part), under rocks, upper tidal zone, American River

Inlet, Kangaroo Island, S. J. Edmonds, Aug. 1946;

speeamen (BM/1980.5.1.9, in part) under rocks, upper

tidal zone, Port Noarlunga, St, Vincent Gull. Ss. J,

Edmonds, November 1950; specimen (BM/

1980.5.1.10), under stones, 5 metres of waler, Tipara

Reef. Spencer Gull. S.A. Shepherd, 25.v.73.

Morphology: This species has been adequately

described by Haswell (1907), Bock (1913) and Stunme

mer-Traunfels (1933), seems lo be exceedingly

variable in size, Existing records give the size of

preserved adult worms as 19-37mm_ long und

920 mim wide, bul Haswell records specimens up to

about 76mm in length when alive. Present adult

specimens are 15-32 mm long and 7-15 mm wide; one

immature specimen is 20mm long and 9 mm wide,

Haswell also records small sexually mature specimens

under [Onmm in length from New Zealand and sup-

Bests that they may be a dwarl varicly of N. wustralis.

Among the present material, a specimen from Baird

REC. S AUST. MUS. IS (16) 401384

June, IY82

Bay measures only 9 tom an length und 4mm in

maximum width, but although it has both sets of

copulalory complexes Lhere is no indication ol’ devel-

oping ova. Marcus (1954) recorded a similar form

from the coast of Chile and named it Noteplana

australis forma Atina, which he regarded as distinct

fron the dwarf form of Haswell and listed two or

(three morphological differences, but these appear to

have no systematic importance,

The coloration of the dorsal surface of this polyelad

varies greatly from almost black, through differing

shades of brown, to orange; Hyman (1959) records

specimens of vlive-grey or greenish grey. The present

adult specimens are greyish with an elongyte-oval

central area of pale brown; immature specimens a

dull white with a tinge of brown in the pharyngeal

region. The yentral surface of nature and immuture

worms is always somewhal lighter. The specimen

from the Tipara Reel was originally orange coloured,

inoltled with white above and belyw, The dorsal sur

face also hud a dark brown median band, due to the

underlying pharyax, and the shallow nuchal tentacles

were also of un orange colour with « whitish apex and

a band of white about midway along cach tentacle.

The area uround the base of each tentacle is without

colour, After prolonged unmersion in an aleohol pre-

servalive, his specimen has lost such coloration,

On the dorsal surface, at about the junction of the

first and second quarters of the total length of the

body, Haswell mentions the presenee of ‘iwo clear

culuurless rounded knobs om which tentacular groups

nf eyes are borne,” In the present specimens one may

uecasionally distinguish a pair ul very shallow bosses,

representing rudimentary tentacles, but in their place

in other specimens there may be a pair of clear

roundish areus in the region of the cerebral organ

dnd in them lie tentacular eyes. Laidlaw (1904) and

Jagubowa (1906) found rudimentary jentacles in

specimens from Australia, In Chilean specimens,

Marcus fuund rounded bosses as nuchal tentacles,

which were transparent in the living aninoal.

The pharynx lies in the middle third of the body

and possesses from $ to 15 pairs of lateral folds. The

main intestine hus 12 to 15 pairs of lateral branches,

whieh vamify towards the periphery of the body and

appear 16 anastomose Only in Ure pharyngeal region at

feature to be seen in other species of the pemis

Notoplana Laidlaw. In the posterior region of the

pharynx, the main intestine bifurcates and ihe arms

pass round the copulatory complexes, similarly, the

anterior end of the intestine bifureates and the limbs

pass round the cerebral organ.

The siructure of the copulatory complexes has been

sulficently deseribed by Haswell (1907) and well

figured by Bock (1913), Mareus (1954) and Hynian

()959) in sagittal or longitudifal section, The pras-

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST. 367

tatic organ in the present material agrees with that

described by Laidlaw (1904) in having its epithelial

lining modified into six longitudinal chambers.

Remarks: Notoplana australis is evidently a common

and widely distributed species on the southern coasts

of Australia. In addition to the present material from

various localities on the South Australian coast, this

species has been recorded from Port Phillip, Victoria,

by Laidlaw (1904), from Tasmania by Haswell (1907)

and from N.S.W. by Schmarda (1859), Haswell

(1907), Stead (1907), Bock (1913) and by Hyman

(1959). It has also been recorded from New Zealand

in various localities extending from Lyttleton Harbour

northwards to Auckland Harbour by Schmarda

(1859), Jacubowa (1906) and by Haswell (1907).

Latterly, it has been recorded from the coast of Chile

between 20°S and 41°S by Marcus (1954).

Stead (1907) reported the destruction of oyster-

beds by “Leptoplana”’ australis in N.S.W. While

circumstantial evidence might point to this, it is more

likely that some other factor was reponsible, and that

the polyclads were merely engaged in cleaning up by

devouring dead and injured animals. In fact, Stead

speaks of the abundance of a polychaete predator and

the oyster-drill Urosalpinx among the oysters.

FIG, 2. Euplana gracilis. Anterior half of body, dorsal view

(bar-scale=0.5 mm.)

Notoplana distincta sp.nov.

(Fig. 3)

Diagnosis: Body elongate oval, 21.5 mm long and

pale yellowish brown. Eyes in two elongate cerebral

clusters. Mouth anterior to middle of body; pharynx

with 8 or 9 pairs of lateral folds. Genital pores well

separated. Seminal vesicle larger than prostatic

organ, which has four longitudinal epithelial cham-

bers. Penis-papilla in the form of a long stylet entirely

enclosed in penis-pocket. Antrum masculinum long

and narrow, lined with tall glandular epithelium

thrown into longitudinal folds. Antrum femininum

narrow, lined with glandular epithelium; Lang’s vesi-

cle small and bulbous.

Locality: South Australia—holotype

(BM/1980.5.1.5), under rock, upper tidal zone, Port

Noarlunga, Gulf St Vincent, S. J. Edmonds, Nov.

1950.

Morphology: Rounded anteriorly and tapering pos-

teriorly. Length, 21.5 mm; maximum width, 6 mm,

which occurs in the mid-region of the body. Dorsal

surface is pale yellowish brown. The cerebral organ

lies at about 4 mm from the anterior margin of the

body. Eyes are arranged in two elongate groups on

either side of the median line in the region of the

cerebral organ. There are 36 to 50 eyes in each group,

and in the hinder region of each group, on the postero-

lateral borders of the cerebral organ, lie 12 to 16

larger eyes more dorsally situated and represent the

tentacular eyes, although tentacles are absent.

The dorsal musculature of the body-wall is, as usual

in polyclads, less well developed than the ventral

musculature, but the basement membrane of the dor-

sal wall is about twice as thick and the epithelium

taller than that of the ventral wall.

The mouth lies at about 5 mm posteriorly to the

cerebral organ and a little behind the middle of the

pharyngeal chamber. The pharynx is situated a little

anteriorly to the middle of the body and measures

3.5 mm long. It possesses 8 or 9 pairs of lateral folds.

The intestinal branches appear to anastomose in the

pharyngeal region and terminate trichotomously in

the marginal zones of the body.

The male genital pore is situated in the central

region of the body, 3.5 mm posteriorly to the mouth,

and the female pore | mm behind the male and about

8 mm from the posterior end of the body.

The testes are scattered in the ventral parenchyma,

as far as the submarginal zones of the body. The vasa

deferentia arise on either side of the pharynx and

extend posteriorly to a level close behind the female

genital pore, where they turn inwardly to unite in the

median line to form a posterior loop. About midway

along their length, each vas deferens gives off a

medially-directed branch, which becomes much con-

voluted and swollen with sperm. These two branches

open independently into a muscular pyriform seminal

vesicle, the narrow end of which is directed anteriorly.

From the narrower end of the seminal vesicle, a short

ejaculatory duct runs dorsally to lead into a globular

prostatic organ provided with a relatively thick mus-

cular wall. The ejaculatory duct passes well into the

368

prostatic organ and is surrounded by a tall glandular

epithelium; in which there are four tubes lying par-

allel with the ejaculatory duct. The epithelial tubes

and the ejaculatory duct open into a small chamber

in the distal region of the prostatic organ. From this

organ, the ejaculatory duct turns posteriorly to enter

an elongate penis-papilla: The papilla is covered with

a membrane or perhaps a thin cuticle and lies entirely

REC. S. AUST. MUS., 18 (16): 361-384

June, 1982

within a penis-pocket. The proximal half of the penis-

pocket encloses the papilla and is lined with a mem-

brane or cuticle, similar to that covering the penis-

papilla, whereas the distal half of the penis-pocket is

lined with tall cilia. Through a shallow penis-sheath,

the pocket opens into a long narrow antrum mascu-

linum lined with a tall glandular epithelium thrown

into longitudinal folds.

vn of

e@ oP

29°® 9

FIG. 3. Notoplana distincta. A. Dorsal view. B. Arrangement of eyes (bar-scale=0.2 mm.) C. Sagittal section of copulatory complexes.

af., antrum femininum; am., antrum masculinum; Lyv., Lang’s vesicle; po., prostatic organ; pp., penis-pocket; s., stylet; sc., “shell”-

chamber; st., stalk of Lang’s vesicle; sv., seminal vesicle; uc., common uterine canal.

The dorsally-disposed ovaries are numerous and

widely scattered, extending farther into the submar-

ginal zones of the body than the testes. The uterine

canal on the right side of the pharynx extends ante-

riorly a little beyond the pharynx, whereas that on

the left side appears to be malformed, for it is thrown

into several coils alongside the hinder margin of the

pharynx. The posterior ends of the two uterine canals

unite to form a short common duct opening into the

vagina at the union of the vagina interna and the

“stalk” of Lang’s vesicle. The epithelium of the “‘stalk”

is thrown into radial folds, so that the lumen appears

moniliform in longitudinal section. Lang’s vesicle is

small and lies ata level a little posteriorly to the

female genital pore. It is lined with a highly glandular

epithelium and coated with a relatively thick mus-

culature. From the uterine opening, a short vagina

interna extends ‘anteriorly to enter a narrow “shell”

chamber, which soon turns ventrally to run posteriorly

to a point ventrally to the union of the uterine canals,

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST.

where it curves ventrally to open into the antrum

femininum through a distinct conical papilla. The

“shell”-chamber is heavily invested with “‘shell”-glands

and lined with an epithelium bearing tall cilia. The

antrum femininum is narrow and lined with a tall

glandular epithelium.

Remarks: The features of this new species are the

forward position of the pharynx, the wide separation

of the male copulatory complex from the posterior

end of the pharynx, the widely-separated genital pores

in the middle third of the body and the glandular

nature of the male and female antra. A further feature

is the conical papilla through which the “shell”-cham-

ber opens into the antrum femininum, but this struc-

ture is unusual among polyclads, and it needs to be

confirmed as a characteristic of the species.

Notoplana longicrumena Prudhoe, 1982

(Fig. 4)

Diagnosis: Body elongate oval in outline, up to

25 mm in length; dorsal surface yellowish grey, tinged

with brown along median line; ventral surface greyish.

No nuchal tentacles. Eyes in two elongate groups.

Mouth centrally situated; pharynx with about 9 pairs

of lateral folds. Genital pores separated. Seminal vesi-

cle more or less as large as prostatic vesicle, which

has five longitudinal epithelial chambers; long penis-

papilla in form of stylet projecting well into male

antrum, which is narrow and deep. Antrum femininum

narrow; Lang’s vesicle bulbous.

Locality: South Australia—holotype (SAM/

V2651—V2665—consisting of uncut portion and 14

slides of _ serial sections), Point Brown,

W. Zeidler, 9.viii.74.

Morphology: It is elongate oval in outline, rounded

at both ends and measuring 25 mm long and 10 mm

wide. Its dorsal surface is yellowish grey, with a tinge

of brown in the middle region, while its ventral surface

is greyish. There is no indication of nuchal tentacles.

The mouth is centrally placed and opens into the

middle region of the pharyngeal chamber, which holds

a pharynx measuring about 6 mm in length and pos-

sessing about 9 pairs of lateral folds.

Eyes are disposed in two elongate groups extending

anteriorly for about 1 mm from the cerebral organ,

which is situated at about 5mm from the anterior

margin of the body. The two groups of eyes tend to

converge gradually as they move forward from the

cerebral organ, and each group consists of cerebral

and tentacular eyes, the latter being larger, nearer to

the dorsal wall of the body and laterally to the cerebral

organ and hinder cerebral eyes.

The dorsal epithelium of the body is higher than

the ventral epithelium, and the musculature of the

ventral wall is two to three times deeper than that in

the dorsal wall.

369

The male genital pore is situated 4 mm posteriorly

to the pharynx and the female pore 0.8 mm behind

the male. A pair of convoluted vasa deferentia orig-

inate in the ventral parenchyma on either side of the

median line, close behind the pharynx. They extend

posteriorly to about level with the male genital pore,

where they turn inwardly to open quite independently

of each other, into the seminal vesicle. There is no

indication of branches to the vasa deferentia extend-

ing posteriorly and uniting behind the vagina, as

found in other species of Notoplana. The seminal

vesicle is pyriform and possesses a well-developed

musculature of circular fibres. Its narrow end is

directed anteriorly. The globular prostatic organ lies

dorsally to the distal end of the seminal vesicle and

is thickly coated with circular and mainly longitudinal

muscle fibres, investing a tall glandular epithelium.

From the distal end of the seminal vesicle, a short

ejaculatory duct extends dorsally to penetrate some

way into the prostatic organ, the epithelium of which

envelops the duct and contains five elongate com-

partments lying parallel with the duct. The compart-

ments and the duct open together into a small cham-

ber in the distal part of the prostatic organ. From the

latter, the ejaculatory duct leads into a relatively long

slender penis-papilla coated with a cuticle and may

be regarded as a stylet, which projects into the male

antrum through a long convoluted penis-pocket with

no sign of a penis-sheath. The penis-pocket is lined

with a ciliated epithelium coated with a thin mus-

culature invested by a thickened parenchyma. The

male antrum is deep and narrow, and lined with an

epithelium continuous with that of the ventral wall

of the body.

The female genital pore leads into a narrow and

shallow antrum femininum or vagina externa, which

extends dorsally to open into the vagina media. The

inner two-thirds of the vagina media is surrounded by

numerous “shell”-glands and functions as a “shell”-

chamber. This chamber extends to the dorsal wall of

the body, where it turns posteriorly to open into the

vagina interna. This region of the vagina continues

posteriorly to open into a bulbous Lang’s vesicle.

Shortly before opening into Lang’s vesicle, the vagina

receives the common duct of the two uterine canals.

Anteriorly, the uterine canals are confluent in front

of the pharynx. The vagina is coated with a muscu-

lature of longitudinal and circular fibres and lined

with a ciliated epithelium, whereas Lang’s vesicle is

lined with a shallow glandular epithelium.

Remarks: Many species have been assigned to the

genus Notoplana Laidlaw and about 10 of these have,

like the present form, no tentacles, a relatively large

Lang’s vesicle. and a long penis-pocket enclosing a

long slender penial stylet. N. longicrumena differs

370

REC. S. AUST. MUS., 18 (16): 361-384

May, 1982

FIG. 4. Notoplana longicrumena. A. Dorsal view. B. Arrangement of eyes (bar-scale=0.5 mm.) C. Sagittal section of copulatory

complexes. af., antrum femininum; am., antrum masculinum, Lyv., Lang’s vesicle; po., prostatic organ; pp., penis-pocket; s., stylet;

sc., “shell”-chamber; st., stalk of Lang’s vesicle; sv., seminal vesicle; uc., common uterine canal.

principally in the comparatively long penis-pocket,

which measures about the combined length of the

seminal vesicle and prostatic organ, whereas in other

species the length of the penis-pocket is only as long

as one or other of these organs. Of the species of

Notoplana possessing a long slender penis-papilla cov-

ered with a thin cuticle and lying in a long penis-

pocket, the present form closely resembles N. sawayai

Marcus, 1947, from Brazil and N. parvula Palombi,

1924, from Indonesia. Both of these latter forms,

however, possess a vagina bulbosa, which is absent in

N. longicrumena. This species also closely resembles

the foregoing N. distincta. It differs, however, in the

arrangement of the eyes, in the position of the mouth

and pharynx, in the relative length of the penial stylet

and in one or two other less important features.

Tripylocelis typica Haswell, 1907

(Fig. 5)

Locality: South Australia—a mature and an imma-

ture specimen (SAM/V2667), Wittelbee Point, West

Coast, W. Zeidler, 2.i11.75.

Morphology: The mature specimen is elongate oval

and measures 12 mm long by 5 mm wide. Dorsally,

the body is light brown, ramified with a darker col-

ouration, due to the dark green lining of the gut

branches, but in the immature specimen the greenish

ramification is apparent only in the pharyngeal region.

The ventral surface is of a lighter colour.

A pair of nuchal tentacles occurs at about 1.5 mm

from the anterior extremity of the body. About 15

eyes lie on one tentacle and about 19 in the other.

The cerebral organ lies in the median line, between

the tentacles, and around it cerebral eyes are arranged

in two groups.

The mouth is situated in the middle region of the

body. The pharynx is about 4 mm long and lies in

the middle third of the body. It possesses 9 to 10

pairs of lateral folds. The intestinal trunk is about as

long as the pharynx and has 9 or 10 pairs of lateral

limbs, which branch towards the margins of the body

and do not anastomose. The trunk bifurcates anteriorly

and posteriorly.

The male pore is situated at 3.5 mm from the

posterior margin of the body, and the female pore

about 1 mm behind the male. As is usual among

polyclads, the testes and ovaries are disposed ventrally

and dorsally, respectively. The vasa deferentia appear

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST. 371

on either side of the pharynx in the region of the

mouth and proceed posteriorly in the ventral paren-

chyma to a little beyond the level of the hinder end

of the pharynx, where they turn inwardly to meet in

the median line and lead into a short convoluted

ejaculatory duct, directed anteriorly. The duct opens

into the inner end of a small, dorso-ventrally elongate,

seminal vesicle, differentiated from the ejaculatory

duct by a somewhat thicker musculature. The dorsal

end of the seminal vesicle narrows and bends pos-

teriorly as the ejaculatory duct is thrown into several

coils. This latter portion of the ejaculatory duct is

lined with a relatively tall ciliated epithelium and has

a very narrow lumen. Finally, the duct enters a well-

developed unarmed penis-papilla containing numerous

gland-cells, and, as noted by Haswell (1907), this

Suggested that the distal region of the ejaculatory

duct might have a prostatic function. The penis-papilla

appears as an evagination of the inner end of the

long, convoluted, posteriorly directed antrum mas-

culinum, for its epithelial covering is similar to the

lining of the male antrum. This epithilium is highly

glandular and bears very long cilia. Almost throughout

its length, the lumen of the male antrum holds accu-

mulations of an eosinophilous granular material, an

indication that the gland cells of the epithelial lining

possibly produce a prostatic substance.

The female pore leads into a short narrow antrum

femininum, directed dorsally. The antrum opens into

a narrow vagina media extending anteriorly to near

the male antrum, where it loops dorsally and runs

posteriorly to a region above the female antrum. In

this region the vagina turns ventrally to receive the

common duct of the 2 uterine canals. The vagina

media represents that portion of the vagina extending

from the female antrum to near the entrance of the

common uterine duct. This portion has plicate walls

lined with a ciliated epithelium, and into its lumen

open the ducts of innumerable extracapsular “‘shell’’-

glands investing its entire length, and for this reason

the vagina media may be regarded as a long narrow

“shell”-chamber. From the latter, the vagina interna,

lined with well-developed ciliated cells, continues to

the point where it turns ventrally. At this point, after

receiving the common uterine canal, the vagina interna

gives rise to a ductus vaginalis which extends ventrally

to open into the ectal region of the female antrum.

The ductus vaginalis is lined with a tall ciliated epi-

thelium, similar to that lining the vagina externa and

the uterine canals. On leaving the vagina interna, the

uterine canals run alongside the pharynx to near its

anterior extremity, where their ends remain separated.

From its union with the vagina interna, the lumen of

the ductus vaginalis gradually narrows, presumably

contracting to prevent the flow of eggs not covered

with “shell” material, but used for the expulsion of

excess reproductive materials and as a copulatory

organ, corresponding to Laurer’s canal in trematodes

and the vagina in cestodes, as suggested by Haswell,

Remarks: The mature specimen described above

agrees very well with Tripylocelis typica as described

by Haswell (1907) from N.S.W., but differs principally

in one feature, for the ductus vaginalis opens into the

antrum femininum and not independently on the ven-

tral surface of the body, as Haswell states. This

difference is here not regarded as diagnostically

important and the present specimen is determined as

Tripylocelis typica Haswell, 1907. On the other hand,

if it were found that the difference in the position of

the opening of the ductus vaginalis was constant in

specimens from South Australia and N.S.W., then

this feature could be regarded as one of systematic

interest. Moreover, there are differences in the struc-

ture of the male copulatory complex, but these are

probably no more than varying stages of development.

CANDIMBOIDES gen.nov

Diagnosis: Leptoplanidae. Body elongate, without

tentacles. Eyes in two elongate clusters in region of

cerebral organ. Pharynx in middle third of body. Male

and female genital pores approximate and well sep-

arated from posterior margin of body. Male copulatory

complex immediately posterior to pharynx. Vasa

deferentia unite to form common duct opening into

muscular seminal vesicle. Prostatic organ muscular

and lined with tall, smooth, glandular epithelium.

Penis-stylet lying in penis-pocket. Female copulatory

complex includes bursa copulatrix and Lang’s vesicle.

Type-species: C. rabita (du B. R. Marcus & Marcus,

1968) comb.nov.

Other species: C. cuneiformis sp.nov.

Remarks: Two specimens of this genus are available.

Although their copulatory complexes are not yet fully

developed and the gonads are not yet ready to begin

their reproductive processes, it is possible to recognise

that they are closely related morphologically to Can-

dimba rabita du B. R. Marcus and Marcus 1968

from the Caribbean island of Curacao. However, the

two forms differ from one another in several features,

principally in the structure of the copulatory bursa

and in the relative positions of the seminal vesicle

and the prostatic organ. When du B. R. Marcus and

Marcus (1968) considered the species C. rabita to be

congeneric with the type-species of Candimba

(C. divae Marcus, 1949, from Brazil), they regarded

the absence of a bursa copulatrix in the latter to be

of no generic importance. There are however, other

important features by which the two forms may be

differentiated. In C. divae the pharynx lies in the

anterior third of the body, the genital pores are widely

separated, a penis-stylet and a penis-pocket are absent,

whereas in C. rabita the pharynx lies in the middle

third of the body, the genital pores are close to each

other, a penis-stylet and a penis-pocket are present.

372

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REC. S. AUST. MUS., 18 (16): 361-384

June, 1982

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FIG. 5. Tripylocelis typica. A. Arrangement of eyes. B. Sagittal section of copulatory complexes. af., antrum femininum; am., antrum

masculinum; dv., ductus vaginalis; e., ejaculatory duct; p., penis-papilla; sc., “shell”-chamber; sv., seminal vesticle; uc., common

uterine canal.

C. cuneiformis agrees with C. rabita in all of these

features, and they are both dissimilar from Candimba

divae, which is the reason for establishing Candim-

boides.

Candimboides cuneiformis sp.nov.

(Fig. 6)

Localities: Victoria—holotype (NMV/G3294),

sample M7/1-3, Westernport Bay Environmental

Study, Jan., 1974; paratype (BM/1980-5.1.15),

sample 32N, Crib Point Benthic Survey, Aug. 1966.

Morphology: The specimen from Westernport Bay

is rather distorted and slightly macerated, and that

from Crib Point is also in poor histological condition,

but apparently both have a somewhat cuneiform out-

line, rounded anteriorly and tapering posteriorly, and

measure about 6.5 mm long and about 2.5 mm wide.

Dorsally, the body is brownish, speckled with dark

spots, due to underlying ovaries, which are widely

scattered, leaving only the marginal regions free.

The cerebral organ is situated at 1 mm from the

anterior margin of the body and bounded by an elon-

gate cluster of eyes on either side. The clusters are

short and tend to converge anteriorly. In the specimen

from Crib Point there are about 40 eyes in each

cluster, whereas in the younger worm from Western-

port Bay there are only about 20 eyes in each cluster.

The mouth is situated at about 3 mm from the

anterior margin of the body and opens into the middle

region of the pharyngeal chamber, which is about

1.5 mm long and contains a short pharynx provided

with six pairs of lateral folds. Details of the intestinal

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST, 373

system have not been made out, but the posterior gut

branches of the younger specimen appear to be

infested with many gregarines.

The hinder half of the specimen from Crib Point

has been serially sectioned, and in this specimen the

male genital pore is situated at 2.5 mm from the

hinder margin of the body, and the female genital

pore at 0.3 mm posteriorly to the male pore.

As is usual in polyclads, the testes lie in the ventral

parenchyma and the ovaries in the dorsal, with a

tendency for the ovaries to move ventrally and mingle

with the testes. It has been difficult to make out the

vasa deferentia, but they appear to arise in the pha-

ryngeal region and run posteriorly to about the level

of the male pore, where they turn inwardly to unite

in the median line and form a short anteriorly

directed ejaculatory duct. This duct opens into the

posterior end of an oval seminal vesicle lying close to

the dorsal wall of the body and directed obliquely

ventrally. The vesicle is very muscular, its wall con-

sisting of a relatively thick layer of longitudinal and

circular fibres. Presumably, the male copulatory com-

plex is not yet fully functional, because the seminal

vesicle contains no sperm, but an eosinophilous mat-

erial similar to that found in the prostatic organ and

had probably been forced into the vesicle by contrac-

tion of the body at fixation. The anterior end of the

seminal vesicle narrows to open into an ejaculatory

duct, which curves posteriorly and ventrally to open

into the anterior end of the prostatic organ through

a shallow papilla. The prostatic organ is oval and lies

ventrally to the seminal vesicle, which is of similar

size. It is invested with a thick layer of mainly longi-

tudinal muscles, but the portion of the ejaculatory

duct passing through the wall of the prostrate is

enveloped by a layer of circular muscles, probably

functioning as a sphincter. The prostatic organ is

lined with a tall, vacuolate, glandular epithelium

which produces a granular material. The posterior

end of the prostatic organ opens directly into a long

slender penis-papilla, the distal two-thirds of which,

about 0.35 mm long, is covered with a thick cuticle

and may be regarded as a penis-stylet. This stylet

appears to be slightly flattened dorso-ventrally to

produce an unusual shape, as illustrated (Fig. 6B). It

lies in a penis-pocket that opens into a narrow male

antrum through a shallow penis-sheath.

The ovaries are widely distributed, extending from

level with the eyes to the posterior region of the body.

The female pore leads into a narrow antrum. Through

the anterior wall of the antrum, near its external

aperture, opens a relatively large bursa copulatrix.

This bursa is directed dorsally and lies to one side of

the median line. The efferent canal of the bursa is

exceedingly narrow and invested with compact par-

enchymatous tissue. Dorsally, it opens into a swollen

muscular structure lying above the male pore. In the

present specimen there appears to be little or no

lumen to the bursa, until it opens into its bulbous

portion, which is provided with a thick muscular wall

and lined with a tall ciliated epithelium. From the

opening of the bursa copulatrix into the antrum fem-

ininum, the vagina extends to near the dorsal wall of

the body, where it turns posteriorly and curves ven-

trally to receive the common duct of the uterine

canals. From this union, the vagina continues ventrally

to open into a small globular Lang’s vesicle. “Shell’’-

glands are not yet developed, hence a “‘shell”-chamber

is not yet apparent.

Family HOPLOPLANIDAE Stummer-Traunfels,

1933

Hoploplana rosea Prudhoe, 1977

Localities: Victoria—one specimen (NMV/G3293),

sample 23S/5, Crib Point Benthic Survey, 1965; one

specimen (BM/1980.5.1.21), sample W/1, Western-

port Bay Environmental Study, 29.xi.73.

Morphology. The two specimens are discoid and

have a diameter of 5-6 mm. The colour of the papillate

dorsal surface of the specimen from Crib Point is

brownish, instead of pinkish as in the type-specimen

of the species from South Australia, while the spec-

imen from Westernport Bay is greyish after preser-

vation in alcohol. The morphology of both specimens

appears to be identical with that of the type-specimen.

Family PLANOCERIDAE Stimpson, 1857

Planocera edmondsi Prudhoe, 1982

(Figs. 7 and &)

Localities: South Australia—holotype (SAM/

V2668), under rocks, upper tidal zone, Port Noarlunga,

St Vincent Gulf, S. J. Edmonds, Nov., 1950.

Tasmania—2 paratypes (BM/1980.5.1.13-14), on

Sonderophycus in 30 metres, Little Squally Cove,

Deal Island, Bass Strait, S.A. Shepherd, 4.v.74.

Morphology: Strongly leaf-like and rounded oval

in outline. The holotype measures 27 mm by 19.0

mm. Dorsal surface of the living worm reddish or

reddish brown. Preserved specimens still retain patches

of such pigment, particularly on tentacles. On the

dorsal surface, there is a pair of conspicuous, non-

retractile, nuchal tentacles lying 1.5 mm apart and

situated at about the junction of the first and second

thirds of the total length of the body. Between the

tentacles lies the cerebral organ. At the base of each

tentacle there is a ring of 40 or 50 tentacular eyes,

and around the cerebral organ numerous cerebral

eyes are scattered. The cerebral eyes show indistinctly

a division into two lateral clusters, commonly found

among species of the genus Planocera Blainville, and

each cluster showing some indication of a division

374 REC. S. AUST. MUS., 18 (16): 361-384

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FIG 6. Candimboides cuneiformis. A. Arrangement of eyes; B. Penis-stylet (bar-scale=0.1 mm.) (ventral view). C. Sagittal section of

copulatory complexes. af., antrum femininum; am., antrum masculinum; bc., bursa copulatrix; e., ejaculatory duct; Lv., Lang’s

vesicle; po., prostatic organ; s., stylet; sv. seminal vesicle; uc., common uterine canal; vm., vagina media.

into an anterior and a posterior group in relation to

the cerebral organ (Fig. 7B).

The mouth occurs in the middle region of the body,

and in the largest specimen at about 5 mm behind

the cerebral organ. It opens into a pharyngeal cham-

ber, about 5 mm long, containing a well-developed

pharynx possessing four pairs of deep lateral folds.

The intestinal trunk is about as long as the pharyngeal

chamber and has six or seven pairs of deep lateral

branches and a forwardly-directed branch passing

over the cerebral organ. The dendriform offshoots of

the intestinal branches do not anastomose.

The male pore is situated at 3.5-4 mm posteriorly

to the mouth, and the female pore 2.0-.25 mm behind

the male. The male copulatory complex lies close

behind the pharynx and is directed anteriorly from

its ventral aperture. The testes are widely distributed,

ventrally to the intestinal branches. The vasa defer-

entia arise on either side of the female copulatory

complex, close to the ventral wall of the body, and

take a convoluted course anteriorly to near a level

with the hinder end of the pharynx, where they turn

medially to open together into a seminal vesicle. The

latter appears to be a structure of two compartments,

one being vesicular and the other wholely muscular.

The union of the vasa deferentia gives rise to a narrow

ejaculatory duct which passes through the muscular

compartment of the seminal vesicle to open into the

vesicular compartment containing a mass of sperm.

From the sperm-containing section, a narrow duct

passes through the muscular section, skirting the pos-

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST.

375

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FIG 7. Planocera edmondsi. A. Dorsal view. B. Arrangement of eyes (bar-scale = 1 mm.) C. Copulatory complexes, with their respective

apertures (ventral view).

terior and ventral walls of the prostatic organ, through

a pocket surrounding the proximal region of the cirrus

cavity, to unite with the prostatic duct. The prostatic

organ is a globular structure lying antero-dorsally to

the seminal vesicle. As in other species of Planocera,

the prostatic organ is lined with a very tall, much-

folded, glandular epithelium. The prostatic duct

springs from the posterior wall of the prostatic organ

and soon opens into the ejaculatory duct, which enters

the anterior end of the cirrus-sac or intromittent organ

through a broad projection that is probably no more

than a slight eversion of the inner or proximal end of

the cirrus-sac, since it is covered with spines. The

cirrus-sac has a narrow lumen and is lined with strong

spines that gradually increase in size towards the

outer or distal end. Around the opening of the cirrus-

sac, as it opens into a very shallow male antrum,

there are five very large spines or teeth. The cirrus-

sac and the prostatic organ are invested with a thick

muscular sheath, in which there is a cavity around

the proximal half of the cirrus-sac and the ejaculatory

duct.

The ovaries are numerous and distributed widely

in the dorsal parenchyma. The female genital pore

opens directly into a large bursa copulatrix provided

with an exceedingly thick musculature of circular,

longitudinal and radial fibres. The bursa appears to

contain three indistinct sections. The distal and prox-

imal sections are lined with a ciliated epithelium,

whereas the central section is lined with a cuticle

thrown into longitudinal ridges. The bursa is directed

anteriorly and leads into a wide vagina media, much

of which functions as a “shell” chamber, being

invested with innumerable “shell” glands. From the

bursa, the vagina media makes a wide posteriorly

directed curve to receive the common uterine canal.

From this union, the vagina continued posteriorly as

a narrow rudimentary Lang’s vesicle.

Remarks: This species is named after Dr S. J.

Edmonds, formerly of the University of Adelaide,

and is readily differentiated from all known species

of the genus Planocera by the unusual structure of

the seminal vesicle and by the presence of five thorn-

like spines embedded in the wall around the opening

of the cirrus-sac.

Family GNESIOCEROTIDAE du Bois-Reymond

Marcus and Marcus, 1966, emend.

Gnesiocerinae du Bois—Reymond Marcus and Mar-

cus, 1966

Diagnosis: Planoceroidea with elongate-oval body.

Eyes in two elongate clusters alongside cerebral organ,

or in paired cerebral and tentacular clusters. Pharynx

376

REC. S. AUST. MUS., 18 (16): 361-384

June, 1982

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FIG, 8. Planocera edmondsi. A. Male copulatory complex. am., antrum masculinum; cs., Cirrus-sac; ct., cirrus-teeth; e., ejaculatory

duct; po., prostatic organ; sv., seminal vesicle; vd., vas deferens; B. female copulatory complex. af., antrum femininum; bc., bursa

copulatrix; Lv., Lang’s vesicle; sc., “‘shell”-chamber; uc., common uterine canal.

much folded, situated in middle region of body, or

somewhat anteriorly. Genital pores separate. Vasa

deferentia may form pair of accessory seminal vesicles

or open into true seminal vesicle. Prostatic organ well

developed, interpolated between sperm reservoir and

cirrus sac. Epithelium of prostatic organ tall and

thrown into folds, distal or ectal region of prostatic

organ narrows to open into cirrus-sac invested with

thick musculature. Cirrus-sac contains well-developed

cuticularized papilla or lined with thick cuticle, often

covered with stiff bristles or spines. Vagina simple,

but may possess vagina bulbosa and Lang’s vesicle.

Remarks; The known genera constituting this family

are listed by du Bois—Reymond Marcus and Marcus

(1968) under the subfamily “Gnesiocerinae”. They

divided the Planoceridae into two subfamilies: the

Planocerinae and the Gnesiocerinae; based on the

presence of an interpolated prostatic organ or of an

independent or so-called “free” prostatic organ. The

Gnesiocerinae is characterised by an interpolated

prostatic organ, through which sperm passes during

copulation, and the Planocerinae by an independent

prostatic organ. The position of this organ in relation

to the ejaculatory duct is of family importance among

POLYCLAD TURBELLARIANS FROM TILE SGUTUERN COASTS UF AUS]

acotylean polyclads and differentiates the family Lep-

toplanidae from the Callioplanidae, and the Stylo-

chidae Irom the Cryptocelididae. Therefore, ta be

consistent in classifying acotylean polyclads, il seems

necessary to raise the subfamily Gnesiocerotinge

(emended) to family rank.

ECHINGPLANA Haswell, 1907

Diagnosis; Gnesiocerotidae without tentacles. Eyes

disposed in two elongate groups, Pharynx mainly in

anterior half of body, provided with 10-15 pairs of

shallow lateral folds: intestinal branches ramify and

anastomose. Male and female genital pores well sep-

arated by genital sucker with corrugated surlace.

Seminal vesicle elongate. Prostatic organ muscular

and clongate, united with cirrus-sae by convoluted

ejaculatory duct enclosed in mass of muscle fibres,

Cirrus-sac with thick musculature and lined with

spines gradually increasing in size towards the

exterior, No male antrum. Vagina narrow, directed

anteriorly, invested with “shell’-glands over much of

its length, Lang’s. vesicle rudimentary, Uterine canals

confluent in median line anteriorly to pharynx.

Type-species: E, tenerrinia Haswell, 1907,

Type-locality; Port Jackson, N.S.W., Australia.

Remarks: Although Echinoplana has been defined

by Haswell (1907), by Bock (1913) and by Gualleni

(1978), for one reason or another it was necessary to

redefine it, Haswell regarded Eehinoplana asa tnem-

ber of the Leptoplanidae, but Bock (1913) placed it

in an appendix of the Planoceridae, and it has sub-

sequently been generally accepted as a member of

the latter family. En fact, Bresslau (1928) placed iL

in the subfamily Planocerinae, but du Bois—Reymond

Marcus and Marcus (1968) referred it to a new

planocerid subfamily Gnesiacerimae (sic).

Echinoplana tenervima Haswell, 1907

Locality: South Australia—l specimen (SAM/

¥2623), Point Hrown, W. Zeidler, 9 viii.74:

| specimen (SAM/V2624), west coast of Wittelbee

Point. W. Zeidler, 2.11.75; | specimen (SAM/V2622),

under rocks, intertidal zone, Pondalowie Bay,

W. Zeidler, 7.x1.76; 1 specimen (SAM/V2621), under

rocks, low tide, Daly Heads, Yorke Peninsula,

W. Zeidler, 9.xi.76-

Morphology; The elongate-oval body in the present

material measures 12-22 mm long and 5-9 mm wide.

and of a very light brown, with a dark reticulation.

No nuchal tentacles. Eyes are arranged in two elongate

groups, disposed laterally to the cerebral organ, each

consisting of 30-50 cyes and tending to converge

anteriorly. The tentacular eyes are situated in the

Posterior region of each group and are larger and

appear nearer to the dorsal surface than the remaining

(cerebral) eyes. The cerebral organ lies. between the

pharynx and the anterior margin of the body, bul

rather nearer to the pharynx, In fact, the arrangement

af the eyes in relation to the cerebral organ agrees

well with that depicted by Haswell (1907) and Gallen

(1978),

The pharynx is situated in the secand quarter of

the total length of the body from the anterior margin,

itis 3-5 mm long and possesses 10-14 pairs of shallow

lateral folds, The intestinal trunk is about as long as

the pharynx and gives off some 14-16 pairs of lateral

branches, which ramity and anastomose towards the

periphery of the body. The anterior end of the intes-

tinal trunk trifurcates, a central branch passing anter-

iorly over the cerebral organ and the outer branches

passing forwardly round the cerebral organ,

The male and female pores are well separated and

situated in the third quarter of the body. Between

them lics 4 corrugated area, probably functioning as

a genita] sucker, It is covered by a tall glandular

epithelium supported by a thickening of the muscu-

lature of the body wall,

The yasa deferentia appear on either side of the

anterior region of the pharynx, with which ihey run

parallel. Immediately posterior to ihe pharynx, ihey

converge towards the median line to open together

into the posterior end of an elongate seminal vesicle,

provided with a thin muscular wall, This vesicle lies

ventrally, is directed anteriorly, and terminates in a

narrow ejaculatory duct, The latter leads into the

anterior end of the prostatic organ through a distinet

papilla projecting into the lumen of the organ, The

latter 1s elongate and two or three times larger than

the seminal vesicle, dorsally to which it lies. [1 is

provided with a thick inner layer of circular muscle

fibres and an outer layer of longitudinal fibres and is

lined with a tall glandular epithelium thrown into

radial folds. The prostatic organ is directed pos-

teriorly, and its hinder end narraws to continue as a

much-convoluted gjaculatory duet invested with a

thick layer of muscle fibres. The ejaculatory duct

Opens into a long cirrus-sac having a narrow lumen

and lined with cuticular spines that gradually become

larger towards the exterior. The cirrus-sac is coated

with a very thick wall of radial muscles, among which

circular and longitudinal museles are scattered. There

is no antrum masculinum, becwuse the cirrus-sac Opens

directly to the exterior. The male pore or exterior

opening of Lhe cirrus-sav js situated at 6-10 ain from

the posterior end of the body.

The female genital pore lies at 2-3 mm posteriorly

to the male pore and opens into a narrow vagina

externa or antrum femininum, lined with a glandular

epithelium similar to that of the corrugated pad. The

vagina extends dorsally, but soon curves anteriorly

and immediately receives efferent duets of the invest-

378

ing ‘“‘shell”-glands to form a narrow “shell”-chamber.

From this chamber, the vagina continues anteriorly

as the vagina interna, lined with a low ciliated epi-

thelium, to near the muscular mass of the cirrus-sac,

dorsally to the male opening, where it makes a short

posteriorly-directed dorsal loop. Shortly before ter-

minating, the vagina receives a short common uterine

canal, thus leaving a very short prolongation, men-

tioned by Haswell (1907) in his generic definition, as

an “unsymmetrical diverticulum”, but apparently

comparable with the rudimentary Lang’s vesicle

found in some other polyclads. This very short narrow

vesicle is lined with a tall ciliated epithelium, similar

to that lining the uterine canals. Immediately before

and after receiving the common uterine canal, the

vagina is invested with innumerable eosinophilous

gland-cells opening into the vagina and producing a

secretion similar to that of the ‘‘shell”-glands—this

is an unusual feature among polyclads, but it is known

in the planocerid genus Disparoplana Laidlaw. From

their common entrance into the vagina, the paired

uterine canals extend anteriorly alongside the pharynx

and vasa deferentia to a level between the cerebral

organ and the pharynx, where they are confluent in

the median line. The entire vaginal system is enclosed

in a dense parenchyma packed with numerous muscle

fibres.

Remarks: In the main, the specimen described

above agrees with the description of this species given

by Haswell (1907) and exceedingly well with the

worm depicted by Galleni (1978). There are, however,

one or two differences, for Haswell found the epi-

thelium of the prostatic organ to be thrown into

longitudinal folds, whereas this epithelium in the pres-

ent specimen appears to be thrown into radial folds,

and one of his figures (2) suggests that a branch of

each vas deferens extends posteriorly beyond the male

copulatory complex. Moreover, Haswell speaks of an

“ootype”, but this appears to be that which is gen-

erally referred to by authors on polyclads as the

“shell” chamber.

In two serially sectioned specimens examined by

Haswell, there was evidence of hypodermic injection

of spermatozoa by other individuals through the

epidermis in the region of the corrugated area or

genital sucker, but in the present serially sectioned

specimen no such evidence has been found.

Along the Australian coasts, Echinoplana tener-

rima was known hitherto only from Port Jackson,

N.S.W., where it appears to have been common more

than seventy years ago. Recently, Galleni (1978) has

described and figured specimens of this species from

the Tuscan coast of Italy. Now that £. tenerrima

appears to be widely distributed along the coast of

South Australia, it seems reasonable to assume that

the occurrence of this species in the warm-temperate

waters of the Mediterranean is due to its being trans-

REC. S. AUST, MUS,, 18 (16): 361-384

June, 1982

ported in a manner similar to that mentioned above

for the North American polyclad Euplana gracilis.

Family CESTOPLANIDAE Lang, 1884

Cestoplana meridionalis Prudhoe, 1982

(Fig. 9)

Locality: South Australia—holotype (SAM/V2613-

V2620 consisting of uncut portions and serial sections)

and = paratype (BM/1980.5.1.7), Beachport,

W. Zeidler, 9.11.77.

Morphology: The slender holotype measures 20 mm

long and 3 mm wide at the broadest region near the

posterior end of the body. It has more or less parallel

sides and tapers towards both extremities. The ground

colour of both dorsal and ventral surfaces in the

preserved condition is greyish, but there is a tendency

for the body to appear brownish, due to granules of

chocolate-brown pigment in the epithelium, which

when sloughed off the brownish colouration is lost.

There is no indication of a ventral sucker or adhesive

depression in the posterior region of the body. The

bilobed cerebral organ lies at 1.25 mm from the ante-

rior end of the body. Eyes appear in the median area

of the body, at a short distance posteriorly to the

cerebral organ and spread fanwise over the anterior

region to the submarginal zones (Fig. 9A).

The mouth is situated at about 2.6 mm from the

hinder end of the body and opens into the posterior

region of a short pharyngeal chamber, which contains

a slightly “ruffled” pharynx about 1 mm in length.

The intestinal trunk is long, reaching anteriorly from

behind the pharyngeal chamber to near the cerebral

organ. It possesses numerous pairs of lateral branches,

which ramify towards the margins of the body, but

do not anastomose.

The male genital pore is situated at about 0.65 mm

posterior to the mouth and 0.4mm anterior to the

female genital pore. The testes, unlike the ovaries, do

not appear to extend into the post-pharyngeal region

of the body. A pair of vasa deferentia lie on either

side of the pharyngeal chamber and take a convoluted

course posteriorly as far as the hinder level of the

female copulatory complex, where they loop inwardly

to run towards one another and near the male antrum

they unite to form a short ejaculatory duct. This duct

extends dorsally and soon opens into a muscular,

elongate-oval, seminal vesicle, which is directed anter-

iorly and disposed ventrally to the intestinal trunk in

the region between the genital pores. The anterior

end of the seminal vesicle narrows abruptly to form

a short duct, which leads into the proximal region of

the prostatic organ. The latter is a small somewhat

pyriform structure lined with a tall glandular epi-

thelium and invested with a thick musculature,

through which pass gland-cells opening into the lumen

of the prostatic organ. This organ opens directly into

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST. 379

FIG. 9. Cestoplana meridionalis. A. Head-end of holotype (bar-scale-0.5 mm) B. Head-end of paratype (bar-scale-0.5 mm.) C. Sagittal

section of copulatory complexes of holotype. af., antrum femininum; am., antrum masculinum; po., prostatic organ; ps., penis-sheath;

5., stylet; sc., “‘shell”-chamber; sv., seminal vesicle; vi., vagina interna.

380

a small conical penis-papilla, tipped with a short spike

of scleroid material. The armed penis-papilla is dis-

posed vertically in a spacious penis-pocket which leads

into the male antrum through a distinct penis-sheath,

The male antrum is short, bulbous or somewhat pyr-

iform.

The ovaries form a broad chain on either side of

the median line and extend posteriorly from the

cephalic region to near the hinder end of the body.

The female genital opening is situated at about

1.5mm from the posterior margin of the body. It

opens into a narrow antrum femininum or vagina

externa, which leads into a wide, dorso-ventrally com-

pressed, “‘shell”-chamber. It seems that the efferent

ducts of the ‘‘shell”-glands open into the chamber

only through its dorsal wall and into a short portion

of the distal region of the vagina interna. From the

“shell”-chamber, the vagina interna, lined with a cil-

iated epithelium, makes a short posterior and down-

ward curve to receive the uterine canals at its inner

end. The uterine canals extend anteriorly from their

union with the vagina, but are not yet fully developed.

The paratype specimen is also slender, but a little

larger than the holotype, measuring 22 mm long and

3.5mm wide. Its coloration is also chocolate-brown

dorsally. The distribution of its eyes (Fig. 9B) is

different from that in the holotype, inasmuch as there

is a narrow median space, widening anteriorly, and

devoid of eyes, a feature often seen in specimens of

other species of Cestoplana. This apparent difference

in the two present specimens probably arose because

of a contraction of the cephalic region of the holotype

at fixation. Although neither set of gonads has

reached maturity, the male and female copulatory

complexes are almost fully developed and are more

or less comparable with those in the holotype.

Remarks: The diagnostic features of Cestoplana

meridionalis are the chocolate-brown colouration of

the body, the lack of a ventral adhesive pad poste-

riorly, the union of the vasa deferentia to form a

common duct entering the seminal vesicle and the

armed penis-papilla. The combination of these fea-

tures enables this species to be readily differentiated

from hitherto described species of Cestoplana.

Haswell (1907) erected the species Cestoplana aus-

tralis for an immature polyclad found between the

tide-marks at Woollahra Point, Port Jackson, N.S.W.

Apparently it possesses a light ground colour, becom-

ing reddish orange posteriorly. In addition, close to

each lateral border there is a band of “vivid vermil-

ion” and a median band of similar colour along the

length of the body. Haswell recognized the very close

resemblance of this species with the European Ces-

toplana rubrocincta (Grube, 1840,) and Kato (1944)

has considered the two forms to be synonymous.

REC. S. AUST. MUS., 18 (16): 361-384

June, 1982

Family PSEUDOCEROTIDAE Lang, 1884 emend.

Poche, 1926

Pseudoceros reticulatus Yeri and Kaburaki, 1918

Localities: South Australia—Il specimen

(BM/1980.7.14), under stones, 5m deep, Tipara

Reef, 14km west from Port Hughes, S.A. Shepherd;

1 specimen (AM/W17748), 5m deep, Penneshaw,

Kangaroo Island, N. Coleman, 12.ii1.78.

Tasmania—several specimens (BM/1980.5.1.2-4

and AM/W17751), on reef, 5m deep, Georges Bay,

St Helens, N. Coleman, 6.10.77.

Morphology: The specimen from Tipara Reef is

somewhat discoid in outline, measuring about 25 mm

both long and wide, whereas the specimen from Kan-

garoo Island is oval in outline and measures 45 mm

long and 25mm wide. The several specimens from

Tasmania are mainly a little smaller and distorted,

but after some manipulation they were found to be

oval in outline, measuring 18 mm-30 mm long and

10 mm-15 mm wide. The body is rather delicate, and

the dorsal surface of the Tasmanian specimens is

dark grey, has thick reticulation of grey or reddish

brown, with much lighter interstices; ventrally the

body is lighter than the dorsal surface, with no retic-

ulation. In these specimens, the dorsal surface also

shows many dark grey pigment spots of varying size,

as well as numerous small dark spots, due to under-

lying ovaries. In fact, these specimens agree excep-

tionally well with the coloured figure of this species

given by Yeri and Kaburaki (1918). In one or two

specimens, there are indications of a whitish margin

to the body, frequently interrupted by dull grey

patches. A dark median band exists over the main-

gut and is dappled with whitish patches. In a koda-

chrome taken by Neville Coleman, the living animal

from Tasmania is translucent, brownish, with a dif-

fusion of white over most of the dorsal surface of the

body, interspersed with white spots. The white

patches in the median band and the interruptions to

the whitich marginal band observed in the preserved

material are quite apparent in the living worm.

The specimens from South Australia are somewhat

lighter in colour. They possess the dark median band,

similarly dappled, and their marginal zones show a

reticulation of grey, but the areas between the median

band and the marginal zones are of a light yellowish

brown. The dorsal surface is likewise spotted with

dark grey pigment, but ventrally the spots are absent.

The dark dots seen in the Tasmanian specimens are

less apparent here, for the yolk material in the ovaries

is much lighter in colour. In a kodachrome, also taken

by N. Coleman, the living animal from Kangaroo

Island appears translucent, brownish, with a greyish

reticulation covering the dorsal surface.

A pair of tentacles appear as mere folds of the

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST. 381

anterior margin of the body and are tipped with

white. Each tentacle contains numerous eyes beneath

the dorsal and ventral surfaces, the number of which

increases, with the size of the body. The cerebral eyes

lie dorsally to the cerebral organ, and in the larger

specimens they form a compact rounded group,

whereas in the smaller specimens they appear to be

arranged more or less in two semicircles disposed one

behind the other.

The histological condition of the available material

is not good, nevertheless the subepithelial muscula-

ture of the Tasmania material appears to consist of

five layers of fibres, both dorsally and ventrally.

There is an outer and an inner layer of longitudinal

muscles and between them are two layers of muscles

forming a criss-cross pattern enclosing a central layer

of circular muscles.

The copulatory complexes are very similar in both

sets of specimens and agree well with those described

by Yeri and Kaburaki (1918). There is, however, one

feature of disagreement, for in the original specimens

the male copulatory complex lies closely posterior to

the pharynx, whereas in the specimens of both the

present sets the complex is situated ventrally to the

hinder region of the pharynx, a feature possibly due

to contraction at fixation.

Remarks: Pseudoceros reticulatus appears to be

common in the warm-temperate waters of the east

coast of Japan (Kato, 1944) and has been recorded

from the coast of Vietnam (Dawydoff, 1952). Its

occurrence in southern Australian waters indicates

that this species is either widely distributed in the

Indo-West Pacific region, or that it has been artifi-

cially transported by some means or other from the

northern area of this region to southern Australia.

Pseudoceros lividus sp.nov.

Locality: South Australia—holotype (AM/

W17750), on sponge, 5 m deep, Kingscote, Kangaroo

Island, N. Coleman, 13.iii.78.

Morphology: The holotype has been preserved in

formalin and is much contracted. It is oval in outline

and is 5 mm long and 3 mm wide. It does not appear

to be sexually mature. In a kodachrome taken by

N. Coleman, the living worm is bluish, dorsally and

ventrally, with a white margin. Because of the heavy

pigmentation, it has not been possible to make out

the arrangement of the eyes, even when an attempt

was made to clear the worm in methyl! salicylate. A

small ventral sucker is centrally situated in the

median line.

Remarks: Over 100 species of Pseudoceros have

been described, mainly from the Indo-Pacific region.

Hyman (1954) pointed out that the differentiation of

species in this genus has been based almost solely on

the pattern of coloration, although very little is known

of variation in this feature. Nevertheless, two or three

species have been described as showing much variety

in colour, but there are one or two instances of where

the pattern of coloration is remarkably constant.

Only two of the known species of Pseudoceros have

a bluish ground colour and these are P. concinnus

Stummer-Traunfels, 1933, nec (Collingwood, 1876)

Kaburaki, 1923, and P. tristriatus Hyman, 1959, In

both of these species, the dorsal surface is bluish with

a median and two submarginal bands of yellow, and

the present form being without any bands of colora-

tion is regarded as a new species.

Thysanozoon skottsbergi Bock, 1923

Locality: South Australia—l specimen (SAM/

V2666), Beachport, W. Zeidler, 9.ii.77; 1 specimen

(BM/1980.5.1.19), only “South Australia” known,

S. A. Shepherd.

Remarks: Both specimens are mature and closely

resemble those recorded by the present writer (1977)

from Gun Island and Fremantle, Western Australia.

Thysanozoon skottsbergi was originally recorded

from Masatierra Island, one of the Juan Fernandez

group, off the coast of Chile.

Family EURYLEPTIDAE Stimpson, 1857

Cycloporus australis Prudhoe, 1982

(Fig. 10)

Locality: South Australia—holotype (AM/

W177499) and paratype (BM 1980.5.1.22), American

River, Kangaroo Island, N. Coleman, 8.iii.78.

Morphology: The two specimens, preserved in for-

malin, are broadly oval in outline and measure 9-

10 mm long and 6,5-7 mm wide. Dorsally, they are

smooth and brownish, with widely-scattered dark spots,

due to underlying ovaries, and a lighter coloured

marginal band. In a kodachrome taken by

N. Coleman of the two specimens when alive, both

appear translucent and tinged with brown, with

whitish spots scattered over the dorsal surface and

yellowish patches distributed intermittently along the

margins of the body. Again, we have an instance of

where in the living animal the ovarian follicles are

seen through the dorsal epidermis of the body as

whitish dots, but when preserved in formalin the yolk-

material in each follicle becomes darkened and the

ovaries appear as dark spots. The ventral surface of

the body is of a uniform lighter colour, except where

the uterine canals filled with eggs appear as whitish

lines on either side of the median line. A muscular

ventral sucker lies in the middle region of the body.

The marginal tentacles, even when alive, according

to the kodachrome, are inclined to be indefinite, and

in the preserved condition they appear as slight folds

382 REC. S. AUST. MUS., 18 (16): 361-384

of the anterior margin of the body. Numerous eyes

occur in the tentacles, more especially ventrally, while

the numerous cerebral eyes lie over the cerebral organ

in two distinct rectangular clusters, about twice as

long as wide.

The dorsal epithelium of the body is taller than

the ventral epithelium and tightly packed with rhab-

dite cells, each of which contains several rhabdites.

The basement membrane is relatively thick, whereas

the subepithelial musculature appears to be rather

insignificant.

The mouth lies at about 1 mm from the anterior

margin of the body, close behind the cerebral organ,

and opens into a short pharyngeal chamber enclosing

a campanulate pharynx. From the pharyngeal cham-

ber, the median intestinal trunk extends to the pos-

terior region of the body, giving off 8 or 9 pairs of

lateral branches as it proceeds posteriorly. Anteriorly,

the intestinal trunk trifurcates, a central branch pass-

ing directly over the cerebral organ and a pair of

branches passing alongside the organ. The lateral

intestinal branches ramify and anastomose rather

loosely and terminate in numerous elongate vesicles

in the marginal regions of the body. The vesicles open

to the exterior through minute pores in the peripheral

epithelium. It seems likely that the yellowish patches

seen on the margins of the living animal are due to

undigested material lying in the terminal vesicles of

the digestive system.

The male genital pore is situated at 0.3 mm pos-

terior to the mouth and the female pore 1.3 mm

posterior to the male pore. The testes are widely

distributed ventrally to the intestinal branches and

reach to near the margins of the body. The vasa

deferentia arise on either side of the median line at

about midway between the female genital pore and

the ventral sucker. Proximally, they are narrow thin-

walled canals, but distally they are wide and lined

with a well-developed glandular epithelium, an unu-

sual feature in polyclads. On reaching anteriorly as

far as the hinder margin of the pharyngeal chamber,

they turn medially, narrow and, without uniting, open

separately into a seminal vesicle. This vesicle is bul-

bous, with a thick muscular wall, and lies ventral to

the hinder region of the pharyngeal chamber. From

June, 1982

the seminal vesicle, a short ejaculatory duct extends

anteriorly to open into a penis-papilla. The prostatic

organ is an elongate oval structure lying between the

pharyngeal chamber and the ejaculatory duct. It is

considerably larger than the seminal vesicle, and is

lined with a relatively tall glandular epithelium,

invested with a well-developed musculature. From its

anterior end, a short prostatic duct leads into the

ejaculatory duct as it opens into the penis-papilla.

This papilla is very small, but armed with a strong

stylet lying in a muscular penis-pocket, which opens

into a spacious antrum masculinum through a thick

conical penis-sheath.

The widely scattered ovaries lie dorsal to the intes-

tinal branches and extend to near the margins of the

body. They are larger than the testes. The female

copulatory complex presents the typical cotylean

structure which seldom undergoes modification. The

female genital pore opens into a widened antrum

femininum or vagina externa, lined with a densely

ciliated epithelium. The female antrum leads through

a narrow aperture in its dorsal wall into a swollen

dorsoventrally flattened “‘shell’”-chamber, into which

open the efferent ducts of innumerable investing

“shell’’-glands. From the “‘shell”-chamber, a muscular

vagina interna extends dorsally to receive the common

thin-walled uterine passage. A pair of symmetrically

disposed uterine canals extends anteriorly as far as

the male complex and then turns to run posteriorly

to a short distance beyond the vaginal complex. The

short uterine canals each bears 3 lateral uterine ves-

icles, suggesting that when the canals are fully devel-

oped and more elongate they may have about 9 pairs

of vesicles.

Remarks: According to a differential key to the

species of the genus Cycloporus Lang given by

du Bois-Reymond Marcus and Marcus (1968), the

present form bears a very strong resemblance to

C. gabriellae Marcus, 1950, from Brazil and the

Caribbean. Apart from a difference in coloration, the

greater number of eyes enables C. australis to be

readily differentiated from other species of the genus

so far described. In their key, Marcus and Marcus

did not include C. maculatus Hallez, 1893, which is

distinguished by the cerebral and tentacular eyes

merging to form an elongate cluster.

POLYCLAD TURBELLARIANS FROM THE SOUTHERN COASTS OF AUST. 383

gs” 9 SS

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S fo)

O00 Gen 0

Go Of nD 0°

0, 0008 OR

fog 090 6

°o o

0° 0? p80? 0°

000782 OO OUD O

Coe lO, o-°

OD 6 Soo Pe oboe

oO go inches

0,994 0° OG Ae 7 O

0 “ee ore 6

C00 Bao ia a

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am % po Sv af sc

FIG. 10. Cycloporus australis. A. Arrangement of eyes. ph., pharynx (bar-scale-0.5 mm) B. Sagittal section of copulatory complexes.

af, antrum femininum; am., antrum masculinum; it., intestinal trunk; ph., pharynx; po., prostatic organ; s., stylet; sc., “‘shell’-

chamber; sv., seminal vesicle; v., vagina.

384 REC. S. AUST. MUS., 18 (16): 361-384

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MARCUS, E. 1950. Turbellaria brasileiros (8). Bolm.Fac.Filos

Cienc.Univ.S.Paulo Zool.No.15: 5-191.

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no. 13: 1-115.

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RECORDS oF THE

SOUTH AUSTRALIAN

MUSEUM

VOLUME 18 NUMBERS 17 —18 April, 1983

No. 17 HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN

MUSEUM. I. NEMATODA

by LESLEY R. SMALES

No. 18 REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK

SYSTEM, NORTHERN TERRITORY

by MICHAEL J. TYLER, GRAEME A. CROOK AND MARGARET DAVIES

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

I. NEMATODA

BY LESLEY R. SMALES

Summary

This is the first of an intended series of papers listing the helminth types held by the South

Australian Museum and the Australian Helminthological Collection.

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM.

I. NEMATODA

LESLEY R. SMALES

South Australian Museum, Adelaide, South Australia S000*

ABSTRACT

SMALES. LR 1983. Helminth type specimens in the South

Australian Museum, J. Nematoda. Ree 8. tat Wis. BS (17):

385-413. Type specimens of 412 species of Neniatodia, held

in the South Australian Muscum or the Australian Helmin-

tholvgival Collection are catalugued, Species are organized

into Superfamilies or Families and bated alphabetivally

according to the original name of the species. Where a name

change hus oecurred the most revent avecpiable name os

included,

INTRODUCTION

This is the first of an intended series of papers

listing the helminth types held by the South Australian

Museum and the Australian Helmintho-

logical Collection.

The Museum's. collection of helminths was estab-

lished soon after the arnval in 1921 of T, Harvey

Johnston to take up his appointment to the newly

created Chair of Zoology at the University of Adelaide,

He became an Honorary Curator of Helminthology

in the Museum in 1922, a position he held until his

death in 1951. As well as assuming an active role in

the governing of the Museum (Board of Governors,

1927-39; Honorary Director of the Museum, 1928-

31: Chairman of the Museum Board, 1940-51), he

began to augment the Museum collections almost

immediately. In addition he was co-author in a series

of papers on the taxonomy of Australian helminths,

the type material of which was eventually deposited

in the Museum.

The first helminth registrations were three trematode

holotypes in the Invertebrate "E” register in 1927 and

five more types were registered there in the 1930's.

A register for Vermes (“V") was started in 1935 and

23 helminth types. including cight transferred from

the “E” register, were recorded in 1937 and 1938.

Both registers then fell into disuse. Type material that

had been deposited in the Museum was loaned to P.

M. Thomas in 1951 and stored at the University for

ready access and safe keeping, No further helminth

matenal was registered until 1957 when 3 lots were

recorded in the “E"' register, There was a spasmodic

flow of material from 1963 on. However it was not

until 1974, following the removal of the marine inver-

tebrates collections to the Museum’s Goldsbrough

House annex. and the appointment of W. Zeidler as

Curator of Marine Invertebrates that any systematic

allempl was made to register the now considerable

collection of helminth types. He re-established the

“V"" register:

In 1977 responsibility for helmunihs was transferred

lo D.C. Lee, Curator of Arachnids. With the help of

his assistant, A. M. Edwards, the back-log material

was registered, in the “V" register. as well as material

returned from the University. The collection, previ-

ously stored as accession lois, was arranged to reflect

the systematic lramework of the group by using a

shelf number system prefixed by “HEL. Lee excluded

annelids from the register and transferred all other

registered helminths into the “V” register so that

South Australian Museum numbers prefixed by V

indicate helminths.

When T. H. Johnston arrived in Adelaide he already

had a considerable private collection, mostly from

Qucensland and New South Wales. This was contin-

ually added to unul his death, when responsibility for

ihe now substantial collection, housed in the Univer-

sity of Adelaide, was assumed primarily by P.. M,

Thomas (nee Mawson) and L. M. Angel. By 1972

Professor J. Arundel (Veterinary Parasitology, Uni-

versity of Melbourne) had begun to express concern

over the future of this valuable and still expanding

collection. He commenced negotiations with the Aus-

tralian Society of Parasitology in 1974 and in 1975

had begun the.search for funding to provide permanent

housing and curation.

Negotiations with the South Australian Museum

resulted in an after by the Museum to holise the

collection, without accepting responsibility for its

curation. On the retirement of Miss Angel, from the

University, in 1976, part of the collection was moved

to the Museum, with Miss Angel as Honorary Asso-

ciate to tend it. At the 1977 Annual General Meeting

of the Australian Socicty of Parasitology a resolution

was adopted that the collection should be properly

housed and curated in a Museum, At the Annual

General Meeting of 1978 the following proposals were

adopted: that the Society would support an application

from the Museum to the Australian Biological Study

Resources Interim Council for funds ia support a

curator of the collection; that a committee be estab-

lished to report on the collections of parasites held

in Australia: that the committee seck an appropriate

name for the collection. At the Annual General Meet-

ing, of 1979 the constitution was. amended to include

*Present address, Gippsland Institute of Advanced Education, P:O. Box 42, Churchill, Victoria ak45,

386

among the objects of the Society

establishment and proper curation of collections of

Australian Parasites”. The committee recommended

at the 1980 Annual General Meeting in Perth that

the collection be known as The Australian Helmin-

thological Collection.

The remainder of the collection was moved to the

Museum in 1980 when Mrs Thomas (now also an

Honorary Associate of the Museum) retired from the

University of Adelaide in 1980. Through the funding

programme of the Bureau of Flora and Fauna, The

Australian Biological Resources Study, grants have

been made available for a temporary research assistant

to undertake some curatorial work. However the long-

term prospects for the safety of the collection remain

in doubt. A survey of Australian collections conducted

by D. M. Spratt for the Australian Society of Para-

sitology in 1979 indicated that the combined South

Australian Museum Collection and Australian Hel-

minthological Collection constituted considerably

more than half the total helminth collection in this

country. The index-catalogue of helminth species of

the world, arranged by genera and cross-referenced

for Australian hosts remains unique. New accessions

continue to be received so that the collection now

comprises over 11,000 bottles and some thousands of

slides, most of which, as yet, remain uncatalogued.

Free-living species are listed separately under their

family according to Filipjev, 1934. Parasitic forms,

listed alphabetically under the original generic name,

are grouped in superfamilies (also listed alphabetically)

as designated by C I H Keys to the Nematode parasites

of Vertebrates. Where a name change has occurred

the most recent acceptable name is given, together

with the relevant reference. Host species names are

those used in the original descriptions. The following

abbreviations were used in the text. BAN-

ZARE = British, Australia and New Zealand Antarc-

tic Research Expedition, 1929-1931; N.T. = Northern

Territory; N.S.W. = New South Wales;

Qld = Queensland; S.A. = South Australia;

S.A.M. = South Australian Museum;

A.H.C. = Australian Helminthological Collection,

Tas. = Tasmania; Vic. = Victoria; W.A. = Western

Australia.

Individual collectors have not been acknowledged

for each species because designations have not always

been made in the literature. However the following

people are known to have made substantial donations

of nematode material: Dr T. L. Bancroft, Dr M. J.

Mackerras, Mr A. S. LeSouef, Professor J. B. Cleland,

Dr F. H. S. Roberts, Dr S, J. Edmonds, Mr

B. Munday, Dr J. H. Arundel, Dr I. Beveridge, Dr

D. M. Spratt and Mr R. Green. The preparators of

the South Australian Museum and the Northern Ter-

ritory Administration, Animal Industry Branch have

also made substantial donations.

REC. S. AUST. MUS., 18 (17): 385-413

April, 1983

FREE LIVING NEMATODA

Family Chromadoridae

Harveyjohnstonia kartanum Mawson, 1953

Holotype ¢, allotype 2 V3075

From littoral rock scrapings Pennington Bay, Kan-

garoo Island, S.A.

Trans. R. Soc. S. Aust., 76: 39-40

Family Enoplidae

Metoncholaimus brevispiculum Mawson, 1957

Holotype ¢, allotype 2 V3076

Jetty piles, Brighton, S.A.

Trans. R. Soc. S. Aust., 80: 101-2

Pontonema hackingi Mawson, 1953

Holotype ¢ V1862, allotype 2? V1863

Off Port Hacking, N.S.W.

Trans. R. Soc. S. Aust., 76: 37-8

Symplocostomella johnstoni Mawson, 1953

Holotype ? V1861

Trawled at 7200m, 5 miles east of Point Gibbon,

N.S.W.

Trans. R. Soc. S. Aust., 76: 38

Thoracostoma australe Mawson, 1953

Holotype ¢ V1859, allotype 2 V1860

Dredging Station off Port Hacking, N.S.W.

Trans. R. Soc. S. Aust., 76: 36

Family Monhysteridae

Steineria pulchra Mawson, 1957

Holotype 4, allotype 2 V3077

Weeds on jetty pile, Outer Harbor, S.A.

Trans. R. Soc. S. Aust., 80: 103-5

PARASITIC NEMATODA

Superfamily Acuarioidea

Acuaria colluricinclae Mawson, 1972

Trans. R. Soc. S. Aust. 96 (3), p. 145

Holotype 3, allotype 2 V1511 from Colluricincla rufi-

ventris

Gizzard: collected Eyre Peninsula, S.A.

Acuaria flindersi Johnston & Mawson, 1941

Trans. R. Soc. S. Aust. 65 (1), p. 31-2

Holotype 3, allotype 2? V1420 from Hieracidea orien-

talis

Gizzard: collected Flinders Is.

Acuaria microecae, Mawson, 1972

Trans. R. Soc. S. Aust. 96 (3), p. 145

Holotype 4, allotype 2 V1512 from Microeca leuco-

phaea

Gizzard: collected Waikerie, S.A.

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM 487

Acvaria mirafrae Mawson, 1972

Trans, R, Soe, S. Aust. 96 (3), p, 147

Holotype 3, V1513 from Mira/fra javanica

Gizzard: collecied Northern Territory.

Paratypes AHC5226

Acuaria petterae Mawson, 1972

Trans. R. Soc. S. Aust. 96 (3), p. 144

Holotype 3, allotype 2 V1510 from La/age leucomela

Gizzard; collected Katherine Gorge, N.T.

Paratypes AHC5219

Acuaria streperina Johnston & Mawson, 1941

Proc, Linn. Soc. N.S. W. 64 (3-4), p. 254-5

Holotype 2, allotype 2° V1432 from Strepera melan-

oplera

Gizzard: collected Waikerie, S,A,

Cheilonematodum halcyonis Johnston & Mawson,

1941

Prac. Linn. Soc, N.S. W. 66 (3-4), p, 253-4

Holotype 4 V3045, allotype 9 V3046 trom JTaleyon

SQNclUs

Stomach: collected Milson [s., N.S.W,

Paratypes AHC900

Chevreuxia australis Johnston & Mawson, 1941

Trans. R. Soe. S. Aust. 65 (2), p. 259

Holotype 2 V1446

Stomach: collected Tailem Bend, S.A.

Cosmocephalus australiensis Johnston & Mawson,

1952

Trans. R. Soe. 8. Aust. 75, p. 34-5

now Svnhimanius australiensis (Johnston & Mawson,

1952) Beveridge & Barker 1975

Holotype ¢ V1866, allotype 2? V1867 from Aydromys

chrysogaster

Stomach: collected Tailem Bend, S.A.

Paratypes AHCI 701

Cosmocephalus jaenschi Johnston & Mawson, 1941

Trans, R: Soc. 8. Aust. 65 (2), p. 259

Holotype 2 V 1445, allotype 2 V1452 from Phalacro-

corax carbo

Stomach: collected Tailem Bend, S.A.

Dispharynx pelecani Johnston & Mawson, 1942

Rec. S. Aust. Mus. 7 (2), p. 185

now Synhimantus (Dispharynx) pelecani (Johnston. &

Mawson, 1942) Chabaud 1975

Holotype 2, allotype 2 V1453 from Pelecanus conspi-

cillatus

Under lining of gizzard: collected Tailem Bend, S.A.

Echinuria querquedulae Johnston & Mawson, |942

Trans. R, Soc, 8. Aust. 66 (1), p. 71

Holotype , allotype ? V1293 from Querquedula gib-

berifrons

Gizzard: collected Tailem Bend, S.A.

Paryseria diomedeae Johnston & Mawson, |942

Trans. R. Soc, 8. Aust. 66 (1), p. 69-70

now Sleguophorus diomedeae (Johnston & Mawson,

1942) Johnston & Mawson, 1945

Holotype 2, allotype 3 V1349 from Diomedea exulans

Stomach: collected Port Jackson, N.S_W.

Paratypes AHC309, 366

Paryseria macronectes Johnston & Mawson, 1942

Trans. R. Sac. S. Aust. 66 (1), p. 70

now Stegophorus macronectes (Johnston & Mawson,

1942) Johnston & Mawson, 1945

Holotype 2 V1348 from Macronectes giganteus

Stomach: collected Brighton, S.A.

Paryseria pachyptilae Johnston & Mawson, 1942

Trans. R. Soe. S. Aust. 66 (1), p. 70

now Sfegophorus pachyptilae (Johnston & Mawsarr,

1942) Johnston & Mawson, 1945

Holotype 2 V 1346, allotype 6 V1347 fram Pachyptilla

witata

Stomach: collected Sellicks Beach, S.A.

Seuratia marina Johnston & Mawson, 1941

Trans. R. Soe. 8. Aust. 65 (2), p. 259-60

Holotype ¢ V2824, allotype 2? V2825 from Pelago-

droma marina

Stomach: collected Flinders Is., Bass St,

Paratypes AHC995, 7798

Stamimerinema suffodiax Beveridge & Barker. 1975

J. Helminth, 49, p. 212-5.

Holotype 2 V6, allotype 2 V62 from Antechinus

stuartl

Stomach: collected Sherbrook Forest, Vic

Paratypes V63

Stegophorus heardi Mawson, 1953

Parasitology 43 (3-4), p. 295-6

Holotype $ V2898 from Oveaniies vceanicus

Stomach: collected Heard Is.

Superfamily Ankylostomatoidea

Unetnaria hydromvidis Beveridge, 1980

J. Parasitol 66 (6) p. 1207-31

Holotype ¢ V1868, allotype 2? V1869 from Midreniyr

chrysogaster

Small intestine; collected Daintree, Qld

Paratypes AHC6348 & V1870-87

Superfamily Aproctoidea

Aprocta bakeri Bain & Mawson, 1981

Ree. S. Aust. Mus. 18 (13) p. 269-71

Holotype 4 V¥2100 from Cureus orru

Nasal cavity: collected Warwick, Qld

Aprocta bouwlengeri Bain & Mawson, 1981

Rec, S. Aust. Mus. 18 (13) p. 269

Holotype 2 V2098, allotype 4 V2099 from Strenere

graculina

Head and nasal cavity: collected Qld

388

Aprocta corvicala Johnston & Mawson, 1940

Trans. R. Soc. 8. Aust. 64 (2), p. 358

Holotype 4, allotype 2 V1402 trom Corvus coronoides

Orbit: collected Musgrave Ranges, 8.A,

Paralype AHCS77

Austrofilaria vestibulata Johnston & Mawson, 1940

Trans, R. Soe. 8S. dust. 64 (2), p. 357

now proeta Vvestibulata (Johnston & Mawson, 1940)

Bain & Mawson, 1981

Holotype 4, allotype 2? V1406 from Aphelocephala

nigriciua

Orbit: collected Musgrave Ranges, S.A.

Cotypes AHC87&

Diomedenema divomedeae Johnston & Mawson, 1952

Trans. R. Soe, S. Aust. 75, p. 32

Syntypes V2827 from Diomedea chrysostoma

Body cavity: calleeted Brighton, 8.A.

Paralypes AHC310

Pseyudaprocta copemani Bain & Mawson, 198]

Rec, S. Aust. Mus, 18 (3) p., 266-7

Holotype @ V209%6, allotype 4 Y2097 from Petroica

multicolor

Collected Maggs ML, Tas,

Pseudaprocta myzarithae Johnston & Mawson, 1940

Trans. R. Sac. 8. Aust. 64, p, 358-9

Holotype 2 V1403 from Myzantha flavigula

Body cavity: collected Renmark, §,A,

Superfamily Ascaridoidea

Anisakis kogiae Johnston & Mawson, 1939

Ree, S. Ase. Mus. & (3), p, 203-6

Holotype 4, allotype @ V1337 from Kogia breviceps

Stomach: calleeted Port Victoria, S.A.

Cantracaecum hancrofli Johnston & Mawson, 194]

Trans, Ro Soe. 8. Aust. 68 (1), p. 112-3

Holotype ¢, allotype ? V1433 from Pelecanus

conspicilarus

Stomach: collected Burnett R., Old

Contracaceum clelandi Johnston & Mawson, 1941

Trans, R. Soe, S. Aust, 65 (1), p. E13

Holotype @, allotype ¢ V1434 from Pelecanus

conspicillatus

Stomach: collected Perth, W.A.

Contracaecum eudyptulae Johnston & Mawson, 1942

Proc, Linn. Sac. N.S.W. 67 (1-2), p. 93-4

Holotype 2, allotype 2 V1294 from Eudyptula miner

Digestive tract: collected Encounter Bay, S.A.

Paratypes AHC| 709

Contracaecum magnicollare Johnston & Mawson,

194)

Trans. R. Sov. 8S. Aust. 65 (1), p. 114

Holotype 4, allotype 2 V1435 from Anous stolidus

Stomach: collected N-W Islet, Capricorn Group,

Barrier Reef

REC, §. AUST. MUS., 18 (17): 385-413

April, 1983

Contracaccum murrayense Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p. 344

Holotype 2, allotype ? V139! from Mecullochella

mracquartensls

Stomach: collected Tailem Bend, S.A,

Paratypes AHC389

Contracaecum ogmorhinae Johnston & Mawson, 1941

Ree. S. Aust. Mus. 6 (4), p. 431-2

Holotype 3, allotype ? V1268 from //ydrurga leptonys

Stomach: collected Port Adelaide, S.A.

Contracaecum podicipitis Jonnston & Mawson, 1949

Trans. R. Soe. S. Aust, 73, p. 67

Holotype 4 V3047, allotype ? V3048 from Podiceps

cristatus

Stomach: collected Tailem Bend, S.A,

Paratypes AHC306

Contracaecum sintlabiarum Johnston & Mawson,

1941

Trans. R. Soc. S. Aust. 65 (1), p. 113-114

Holotype 2, allotype 2 V1436 from Plotus novaehol-

landiue

Stomach: collected Burnett R., Qld

Paratypes AHC994

Goezia fluviatilis Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p. 342-3

Holotype 4, allotype @ V1393 from Plectroplites

ambiguus

Gill mucous: collected Tailem Bend, §.A.

Ophidascaris varani Johnston & Mawson, 1947

Trans. R. Soc, S. Aust. 71 (1), p. 23-4

now Amplicaecum mackerrasxae (Johnston &

Mawson, 1947) Thomas, 1959

Holotype é V2876 from Waranus varius

Alimentary canal; collected Toowoomba, Old

Paratypes AHC31 54

Paraheterotyphium australe Johnston & Mawson, 194%

Rec, S. Aust. Mus, 9 (1), p, 102-4

Holotype 2, allotype 2 V815 from Hydrus platurus

Intestine: collected Little Bay, Sydney, N.S.W.

Paratypes AHC1438

Paranisakis australis Johnston & Mawson, 1943

Trans. R, Soe. S. Aust. 67 (2), p. 190

Holotype 4, allotype 9 V1289 from Urolophis

australis

Intestine: collected Sydney, N.S.W.

Phocasearis hyvdrurgae Johnston & Mawson, 194]

Rec. S. Aust. Mus. 6 (4), p. 432

now Contracaecum osculatum (Rud) Johnston &

Mawson, 1945

Holotype V1272 from Hydrurga leptonyx

Stomach: collected Port Adelaide, 8.A.

Note: Re-examination of P. Hydrurgae type material,

by the authors, revealed it to be a pre-adult stage

of C. oxsculatum

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM SKY

Porracaecum circinum Johnston & Mawson, 1941

Trans, R, Soe. 8. Aust, 65 (1), p. 30-1

Holotype 2 Vi4d18 from Circus assimilis

Intestine: collected Orroroo, S.A.

Porrocaecum clelandi Johnston & Mawson, 1941

Prac. Linn. Soc. N.S.W. 66 (3-4), p. 252

Holotype 2, allotype 2 V1425 from Oreocinela lunulata

Intestine: collected Bunya Mts, Qld

Porrocaecum kogiae Johnston & Mawson, 1939

Ree. S. Aust. Mus. 6 (3), p. 266

Holotype ¢, allotype 2 V1338 from Kogia breviceps

Stomach: collected Spencer's Gulf, S.A.

Paratypes AHC1606

Porrocaecuni lobibycis Mawson, 1968

Parasitology 58, p, 284-5

Holotype ¢ V1467 from Lohibyx novaehollandiae

Duodenum: collected Naracoorte, S.A.

Paralypes AHC4495

Porrocaecum streperae Johnston & Mawson, 1941

Proce. Linn. Soc. N.S.W. 66 (3-4), p. 253

Holotype 2 V1426 from Strepera versicolor

Intestine: collected Mt Kosiusko, N.S.W.

Stomachus aceanicus Johnston & Mawson, 1951

Ree. 4. Mus, 22 (4), p. 293

now Anisakis eceanicus (Johnston & Mawson,

1951) Hariwich, 1974

Stomach: collected stranded on coast of N.S.W.

Paratypes AHC]| 699

Terranova chilosevilit Johnston & Mawson. 1951

Ree, A. Adys. 22 (4), p. 291

from Chiloseyllium punctarun

Stomach; collected Halfway Is., Keppel Islands, Qld

Paratypes AHC1661

Superfamily Camallanoidea

Procamallanus murrayensis Johnston & Mawson,

1940

Trans. R. Soc. S. Aust. 64 (2), p. 347

Holotype 2, allotype 2 VL380 from Pseudaphritis

wrviller

Alimentary canal: collected Swan Reach, 8.A,

Superfamily Cosmocercoidea

Aplectana flindersi Johnston & Mawson, 1941

Rec. S. Aust. Mus. 7, p. 148

now Maxvachonia flindersi (Johnston & Mawson,

1941) Mawson, 1972

Holotype @ V1408 from //yla_jervisiensis

Rectum: collected Kangaroo Is., S.A.

Cosmocera (imnodynastes Johnston & Simpson, 1942

Trans. R. Soc, S. Aust. 66 (2), p. 174-6

Holotype 2 V1275 from Liminodynastes dorsalis

Intestine: collected Adelaide, S.A.

Paratype AHC2372

Maxyvachonia brygooi Mawson, 1972

Trans, R, Soe. 8. Aust. 96 (2), p. 102-4

Holotype 2 V1503 from Amphiholurus decressi

Rectum: collected Eyre Peninsula, 8.A,

Maxvachonia chahaudi Mawson, 1972

Trans. R. Sac. S. Aust. 96 (2), p. 102

Holotype 4, allotype 2 V1502 from Morethia

lincoucellata

Rectum: collected Eyre Peninsula, S.A.

Maxvachonia ewers! Mawson, 1972

Trans. R. Soc. 8. Aust, 96 (2), p. 105-6

Holotype 4, allotype 9 V1504 from Litoria nasuta

Rectum: collected Brown R.. New Guinea

Paratypes AHC5188

Spironoura hylae Johnston & Simpson, |942

Trans. R, Soe, S. Aust, 66 (2), p. 173

now Falcaustra simpsoni (Johnston & Simpson,

1942) Chabaud, 1978

Holotype 4 V2892, allotype V2893 from Hila aurea

Intestine: collected Sydney, N.S.W.

Paratypes AHC2313

Note: S, /i/ae was found to be a nom praeoec. by

Johnston & Mawson, 1944

Spironaura elsevae Johnston & Mawson. 1941

Rec, Aust. Mus. 21 (1). p. 15

now Falcausira elsevae (Johnston & Mawson, 1941)

Chabaud, 1978

from Elseva dentata

Intestine: collected Northern Australia

Paratypes AHC1666

Superfamily Dioctophymatoidea

Eustrongvlides phalacrocoracis Johnston & Mawson,

194]

Trans. R. Soc. .S. Aust, 65 (2), p, 255-6

Holotype 2. allotype 2? V1450 from Phalacrocoran

carbo

Sub-peritoncal tissue of stomach; collected Tailem

Bend, S.A.

Fustrongvlides plotinus Johnston & Mawson, 1941

Trans. R. Soc, S. Aust. 65 (2), p. 256

Holotype 4 V1451 from Anhinga novachallandiae

Body cavity: collected Burnett R., Qld

Superfamily Diplotriaenoidea

Diplotriaena alpha Johnston & Mawson. 1940

Trans, R. Soc, S. Aust. 64 (2), p. 359-60

Holotype @ V1394 from Strepera graculina

Peritoneum: collected Mt Irvine, S.A.

Diplotriaena beta Johnston & Mawson, 1940

Trans. R. Soe, S. Aust. 64 (2), p. 360

Holotype 2 V1395 from Corvus coronoides

Body cavity: collected Eidsvold, Qld

390

Diplorriaena beveridgei Bain & Mawson, 1981

Rec. 8, dust. Mus, 18 (3), p. 278-80

Holotype 2 V2103, allotype ¢ V2104 from Corvus

orrit

Nasal cavity; collected Warwick, Qid

Diplotriaena delta Johnston & Mawson, 1940

Trans. R. Soe. S. Aust. 64 (2), p. 360

Holotype 6 V1397 from Malurus lambert

Body caviiy: collected Ooldea, S.A.

Diplolriaena epsilon Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p. 360

Holotype 4, allotype ° V1398 from Cracticus

destructar

Body cavity: collected Brisbane, Qld

Diplotriaena gamma Johnston & Mawson, 1940

Trans. R. Soc. 8. Aust, 64 (2), p. 360

Holotype 2 V1396 from Spres superbus

Body cavity: collected Adelaide S.A.

Diplotriaena smithi Bain & Mawson, 1981

Rec, 8. dust. Mus, 18 (13), p. 280

Holotype 2 V2105, allotype ¢ V2106 from Acantho-

genys rifogularis

Collected the Bunkers, S.A.

Diplotriaena spratti Bain & Mawson, 1981

Ree. S. Aust. Mus. 18 (13), p. 277

Holotype @ V2101, allotype ¢ V2102 from Oreoica

eulteralis

Body cavity: collected Petermann Ra., N.T.

Paratypes V2669-80

Diplotriaena zeta Johnston & Mawson, 1940

Trans. R, Soc, 8. Aust, 64 (2), p. 361

Holotype @ V1399 from Acanthogenys rufigularis

Body cavity; collected Monarto, S.A,

Hamatospiculuimn chibiae Johnston & Mawson, 194)

Ree. A. Mus. 21 (1), p. 14-5

from Chihla bracteata

Skin of neck: collected Russell Is., Qld

Paratypes AHC1667

Hamatoaspiculum halcyonis Johnston & Mawson, 1941

Rec. A. Mus. 21 (1), p. 14

from Halevon pyrrhapygius

Eye balls and roof of mouth: collected Mt Lyndhurst,

S.A.

Paratypes AHC893, 1079

Hamatospiculum howense Johnston & Mawson, 1940

Trans, R. Soc. S. Aust, 64 (2), p, 355-6

Holotype 4, allotype 2 V1404 from Halcyon vagans

Sub-cutaneaus tissue; collected Lord Howe Is.

Hamatospiculum meneilli Johnston & Mawson, 1941

Rec, A. Mus. 21 (1), p. 12-14

from Ninox boobook

RECS, AUST. MUS., 18 (17): 385-413

lpril, 1983

Tissues adjacent to skull: collected Hayman Is.,

Whitsunday Group, Qld.

Paraltypes AHC862

Superfamily Dracunculoidea

Anguillicola australiensis Johnston & Mawson, 1940

Trans, R, Soc. S. Aust. 64 (2), p. 351

Holotype 2, allotype 2 V1392 from Anguilla

reinhardtii

Swim bladder: coliected Prospect Reservoir, Sydney,

N.S.W.

Paratypes AHC586

Philometra plectroplites Johnston & Mawson, 1940

Trans. R. Sac. S. Aust. 64 (2), p. 348-9

Holotype @ V1384 from Plectroplites ambiguus

Body cavity: collected Murray Bridge, S.A.

Philometra percalates Johnston & Mawson, 1940

Trans. R. Soc. §. Aust. 64 (2), p. 349

Holotype 4, allotype 9 V1385 from Percalates colon-

orum

Body cavity: collected Tailem Bend, S.A.

Superfamily Filarioidea

Carinema dubia Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p, 357-8

now Cardiofilaria duhia (Johnston & Mawson, 1940)

Sonin, 1961

Holotype 4, allotype @ V1400 from Pseudopsittacus

melennani

Abdominal cavity: collected North Qld,

Carinema graucalinum Johnston & Mawson, 1940

Trans. R. Soc. §. Aust, 64 (2), p. 358

now Cardiofilaria graucalinum (Johnston & Mawson,

1940) Sonin, 1961

Holotype 3, allotype @ VI401 from Graucalus

melanops

Abdominal cavity; collected Fraser Is,, Qld.

Dipetalonema annulipapillatum Johnston & Mawson,

1938

Trans. R. Soc. Aust, 62 (1), p, 117-8

now Breinlia (Johnstonema) annulipapillatum

(Johnston & Mawson. 1938) Chabaud & Bain,

1976

Holotype ¢ V2826, allotype ? V3] from Onychogale

fraenata

Sub-cutaneus: collected Burnett R., Qld

Note: Spratt & Varughese, 1975 found that the holo-

type of D. annulipapillatum was damaged, one

spicule missing and badly discoloured.

Dipetalonema boltoni Spratt & Varughese, 1975

Aust. J. Zool. Suppl. Ser. 35, p. 70-4

now Brejnlia (Breinlia) boltoni (Sprau &

Varughese, 1975) Chabaud & Bain 1976

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM 491

Holotype 4 V1130, allotype 9 V1I31 from Macropus

agilis

Peritoneal cavity: collected Townsville, Qld

Paratype V1132

Dipetalonema dasyuri Johnston & Mawson, 1938

Trans. R, Soc. S. Aust. 62 (1), p. 109

now 8Sreinlia (Breinlia) dasyuri (Johnston &

Mawsan, 1938) Chabaud & Bain, 1976

Holotype 4, allatype ? V1126 from Dusyurus viverrinus

Body cavity: callected Vic,

Cotypes AHC2946

Dipetalonema dentonensis Spratt & Varughese, 1975

Aust, J Zool. Suppl. Ser, 35, p. 48-52

now Breinlia (Breinlia) dentonensis (Spratt &

Varughese, 1975) Chabaud & Bain, 1976

Holotype 2 VII41, allotype 2 VI 142 from Macropus

Blganteus

Subcutaneous connective tissue: collected *Denton’

Morella, Qld

Paratypes V1143

Dipetalonema lutreali Mackerras, 1962

Aust. J. Zool. 10 (3), p. 406-9

new Dipetalonema (Chenofilaria) johnstoni

(Mackerras, 1962) Chabaud & Bain, 1976

from Rattus lutreolus

Connective tissue: collected West Burleigh, Qld

Paralypes AHC3993

Dipetalonema niundayi Spratt & Varughese, 1975

Aust. J. Zool. Suppl. Ser. 35, p. 67-70

now Breinlia (Breinlia) mundayi (Spratt &

Varughese, 1975) Chabaud & Bain, 1976

Holotype 4 V1144, allotype 2 V1145 from Afacropus

rufogriseus frulicus

Pericardium: collected Rass, Tas.

Paratypes V1146

Dipetalonema pearsoni Spratt & Varughese, 1975

Aust. J. Zool. Suppl. Ser. 35, p. 33-6

now Dinetalonema (Chenofilaria) pearsoni (Spratt

& Varughese, 1975) Chabaud & Bain, 1976

Holotype 3 V1139, allotype 2 V1140 from /soodon

macrourus

Subcutaneous tissue: collected Woolwonga, N.T.

Dipetalonema pseudocheiri Spratt & Varughese, 1975

Aust. J. Zool. Suppl. Ser. 35, p. 60-3

now Breinlia (Breinlia) pseudocheirt (Spratt &

Varughese, 1975) Chabaud & Bain, 1976

Holotype ¢ V1127, allotype @ V1128 trom Pseudo-

cheirus convalutor

Peritoneal cavity: collected Launceston, Tas.

Paratypes V1129

Dipetalonema rarum Johnston & Mawson, 1938

Trans, R. Soc. 8S. Aust. 62 (1), p. 114

now Breinlia (Breinlia) rarum (Johnston &

Mawson, 1938) Chabaud & Bain, 1976

Syntype 9? V1123 from Onvchogale fraenata

Subcutaneous: collected Vic.

Dipetalonema roberts Johnston & Mawson, 1938

Ree. S. Aust. Mus. 6 (2), p. 188-9

now Breinlia (Breinlia) roberisi (Johnston &

Mawson, 1938) Chabaud & Bain, 1976

Halotype 4, allotype 2 V1124 from Macropus robustus

Body cavity: collected Normanton, N. Qld

Dipetalonema tenue Johnston & Mawson, 1938

Trans. R. Soc. S. Aust. 62 (1), p. 118-20

now Breinlia (Breinlia) robertsi (Johnston &

Mawson, 1938 after Spratt & Varughese, 1975)

Chahaud & Bain, 1976

Holotype 2 V1125 from Macropus robustus

Body cavity: callecied Cockatoo Creek, N.T.

Dipetalonema venacavincola Spratt & Varughese, 1975

Aust. J. Zool. Suppl. Ser. 35, p. 36-40

now Spraitia venacavincola (Spratt & Varughese,

1975) Chabaud & Bain, 1976

Holotype 2 V1147, allotype 2 V1148 from Trichusurtis

Caninus

Inferior vena cava & heptic

Tamborine, Qld

Paratypes V1149

veins: collected

Johnstonema andersoni Spratt & Varughese, 1975

Aust, J, Zool. Suppl. Ser. 35, p. 14-17

now Brein/ia (Jahnstonema) andersoni (Spratt &

Varughese, 1975) Chabaud & Bain, 1976

Holotype 4 V1135, allotype 2 V1136 from Megaleia

rufa

Subcutaneous connective tissue: collecied Quilpie, Qld

Paratypes VJ 137

Johnstonema woerle] Spratt & Varughese, 1975

Aust. J. Zool. Suppl. Ser. 35, p. 17-20

now Breinlia (Johnstonema) woerlei (Spratt &

Varughese, 1975) Chabaud. & Bain, 1976

Holotype 4 V1138 from Petrogale venustula

Heart: collected Cannon Hill, N.T.

Josefilaria mackerrasse Moorhouse, Bain & Wolf, 1979

Annal. Parasit. Jlum. et comp. 54 (6), p, 645-51

from Macrederma gigay

Fascia of pectoral muscles: collected Ridgeland, near

Rockhampton, Qid

Paratypes V1913-4

Liromasa miniopteri Mackerras, 1962

Aust. J. Zool. 10 (3), p. 403-4

fram Miniopteris schretbersi

Heart: Mt Pleasant, Qld

Paratypes AHC3994

Paralemdana cleland) Johnston & Mawson, 1940

Trans, R. Sac. S. Aust. 64 (2), p. 356-7

Holotype 2 V1405 from Strepera graculina

Body cavity: collected Scone, N'S.W.

392 REC. 8. AUST. MUS., 18 (17); 385-413

Piratuba queenslandensis Mackerras, 1962

just, J. Zoal. 10 (3), p. 443

from Varanus (ristis orientalis

Lungs: collected Innisfail, Qid

Paratypes AHCO3995

Piratuha varanicola Mackerras, 1962

stust, J. Zool, 10 (3), p. 444

from Varanus tristis orientalis

Pleural covering of lungs: collected Innisfail, Qld

Paratypes AHC4000

Saurofilaria innisfailensis Mackerras, 1962

Aust J. Zool, 10 (3), p. 448

possibly how Macdonaldius innisfailensis (Mack-

erras, 1962) Chahaud & Bain, 1976

from Physignathus leseurii

Subperitoncal tissue: Innisfail, Qid

Paralypes AHC3966

Vagrifilaria ausiralis Johnston & Mawson, 1942

Proe. Linn, Soe. N.S.W. 67 (1-2), p. 93

now Aprocta australis (Johnston & Mawson,

1942) Bain & Mawson, 198]

Holotype 4, allotype 2 V1295 from Cenfropus phasi-

anus

Colon: collected West Burleigh, Qld

Superfamily Gnathostomatoidea

Tangua ophidis Johnston & Mawson, 1948

Rec, S. dust. Mus, 9 (1), p, 104-5

Holotype ¢ V2894. allotype 2 V2895 trom Nairix

mairil

Alimentary canal: collected N-W coast, Qld

Paratypes AHC 1441

Superfamily Habronematoidea

Ascarophis australis Johnston & Mawson, 1944

Trans. R. Soe. S. Aust. 68 (1), p. 62

Holotype 4 V1255, allotype 2 V1256 from Threplertus

macnlosus

Alimentary. canal: collected Cape Border, S.A.

Asearophis cooperi Johnston & Mawson, 1945

Trans. R. Soc. S. Aust. 69 (1), p. 115-6

Holotype 4 V1257, allotype @ V1258 from Platyce-

phalus bassensis

Alimentary canal: collected Rapid Bay, S.A.

tscaruphis murrayensis Johnston & Mawson, 1947

Rev. S. Aust. Mus. 8 (4), p. 553

Holotype 2 V1253, allotype 2 V1254 from Plectroplites

UHM eUMS

Alimentary canal: collected Tailem Bend, S.A.

Crassicauda magna Johnston & Mawson, 1939

Rec. S. Aust. Mus. 6 (3), p. 266-8

Holotype 2 V2833 from Kogia breviceps

Neck connecttve tissue: collected Pt Victoria, S.A.

Paratypes AHCI599, 1607

April, 1983

Cyrnea dentifera Johnston & Mawson, 1941

Proc, Linn, Sac, N.S. W. 66 (3-4), p. 255

now Excisa dentifera (Johnston & Mawson, 1941)

Chabaud, 1958

Holotype 3. allotype 2 V1429 from Eupodotis australis

Gizzard: collected Mt Leibig. N.T.

Cyrnea (Pracyrnea) dollfusi Mawson, 1968

Parasitology 58, p. 750-2

now Procyrnea dollfusi

Chabaud, 1975

Holotype 2, allotype 2 V1489 from Ninox novaesee-

lundiae

Gizzard: collected Berrimah, N.T-

Paratypes AHC4674

Cyrrnea (P.) falco Mawson, 1968

Parasitology 58, p. 752-3

now Procyrnea falco (Mawson, 1968) Chabaud,

1975

Holotype 2, allotype @ V1490 from Falco longipennis

Gizzard: collected Humpty Doo, N.T.

Paratypes AHC4679

(Mawson, 1968),

Cyrnea spiralis Mawson, 1968

Parasitology 58, p. 753-5

now Microhadjelia spiralis (Mawson, 1968) comb,

nova Mawson, pers. comm.

Holotype 4, allotype 2 V1491 from Philemon argen-

ticeps

Gizzard: collected Berrimah, N.T.

Paratypes AHC4648

Excisa hiloba Mawson, 1968

Parasitology 58, p. 755-8

Holotype 2, allotype 2 V1492 from Podargus strigoides

Gizzard: collected Brisbane, Qld

Paratypes AHC4670

Geapetitia chibiae Mawson, 1966

Parasitology 56, p. 716-7

Holotype 2 V1455 from Chihia bracteala

Wall of proventriculus: Milner’s Swamp, N,T.

Geopetitia falco Mawson, 1966

Parasitology 56, p. 717-8

Holotype & V1456 from Falco longipennis

Under lining of proventriculus: collected Humpty

Doo, N.T.

Geopetitia streperae Mawson, 1966

Parasitology 56, p. 715-6

Holotype 4, allotype 2? V1454 from Strepera melan-

oplera

Wall of proventriculus: collected Naracoorte, S.A.

Paratypes AHC4642

Hahronema aegotheles Johnston & Mawson, 1941

Trans. R. Soc. S, dust. 65 (2), p. 256-7

now Alainchabandia aegotheles (Johnston &

Mawson, 1941) Mawson, 1968

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Holotype 4, allotype 2 V1447 from Aegotheles cristata

Under gizzard lining: collected Tailem Bend, S.A.

Habronema paraleptoptera Johnston & Mawson, 1941

Trans. R. Soc. S. Aust. 65 (1), p. 32-3

now Cyrnea (Procyrnea) paraleptoptera (Johnston

& Mawson, 1941) Chabaud, 1958

Holotype 4, allotype 2 V1421 from Cerchneis cen-

chroides

Stomach: collected West Burleigh, Qld

Hedruris hylae Johnston & Mawson, 1941

Rec. S. Aust. Mus. 7 (1), p. 148

Holotype 4, allotype 2 V1261 from Hyla jervisiensis

Stomach: collected Kangaroo Is., S.A.

Hedruris longispicula Thomas, 1959

Trans. R. Soc. S. Aust. 82, p. 160-1

Holotype 2 V2829 from Lygosoma challengeri

Alimentary canal: collected Springbank, Qld

Note: The description of this species was published

in the only paper by Mrs Thomas in which acci-

dentally she did not use the name Mawson.

Microtetrameres aegotheles Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 253

Holotype ¢, allotype 2 V1540 from Aegotheles cristata

Proventriculus: collected Markanka, N.T.

Microtetrameres cacomantis Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 253

Holotype 4, allotype 2 V1534 from Cacomantis vari-

olosus

Proventriculus: collected Tobermory, N.T.

Microtetrameres cerci Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 255

Holotype 6 V1537 from Circus assimilis

Proventriculus: collected Petermann Ra., N.T.

Microtetrameres coracinae Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 253

Holotype ¢, allotype 2 V1530 from Coracina novae-

hollandiae

Proventriculus: collected Culburra, S.A.

Microtetrameres cractici Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 251

Holotype ¢ V1528 from Cracticus torquatus

Proventriculus: collected Eyre Peninsula, S.A.

Microtetrameres eopsaltriae Mawson, 1977

Rec. S. Aust. 17 (14), p. 253

Holotype ¢, allotype 2 V1536 from Eopsaltria australis

Proventriculus: collected Heatherleigh, S.A.

Microtetrameres gymnorhinae Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 251

Holotype ¢, allotype 2 V1538 from Gymnorhina tibicen

tibicen

Proventriculus: collected Canberra, A.C.T.

Microtetrameres melophagidea Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 248

393

Holotype 4, allotype 2? V1529 from Acanthogenys rufo-

gularis

Proventriculus: collected Pt Augusta, S.A.

Paratypes AHC5914

Microtetrameres mirafrae Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 249

Holotype 6 V1535 from Mirafra javanica

Proventriculus: collected N.T.

Microtetrameres ninoctis Mawson, 1977

Rec. S. Aust Mus. 17 (14), p. 256

Holotype ¢, allotype 2 V1525 from Ninox novaezee-

landiae

Proventriculus: collected Berrimah, N.T.

Microtetrameres paraccipiter Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 253

Holotype 2 V1527 from Accipiter fasciatus

Proventriculus: collected Darwin, N.T.

Microtetrameres philemon Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 249

Holotype ¢, allotype ? V1539 from Philemon argen-

ticeps

Proventriculus: collected Coomalie Ck, N.T.

Microtetrameres raptoris Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 255

Holotype ¢, allotype 2 V1526 from Falco peregrinus

Proventriculus: collected Pt Augusta, S.A.

Paratypes AHC5906

Microtetrameres sphecotheres Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 253

Holotype ¢ V1532 from Sphecotheres flaviventris

Proventriculus: collected Katherine Gorge, N.T.

Microtetrameres streperae Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 251

Holotype 4, allotype 2? V1533 from Strepera versicolor

Proventriculus: collected Waikerie, S.A.

Microtetrameres tytonis Mawson, 1977

Rec. S. Aust. Mus. 17 (14), p. 257

Holotype ¢, allotype 2 V1531 from Tyto alba

Proventriculus: collected Banka Banka, N.T.

Tetrameres anseranas Mawson, 1979

Trans. R. Soc. S. Aust. 103 (7), p. 178-80

Holotype ¢ V2681, allotype 2 V2682 from Anseranas

semipalmata

Proventriculus: collected Humpty Doo, N.T.

Paratypes AHC6941

Tetrameres australis Johnston & Mawson, 1941

Trans. R. Soc. S. Aust. 65 (2), p. 262

Holotype 4, allotype 2 V1451 from Chenopsis atrata

Proventriculus: collected Tailem Bend, S.A.

Tetrameres bizurae Johnston & Mawson, 1941

Trans. R. Soc. S. Aust. 65 (2), p. 261-2

Holotype 4, allotype 2 V1414 from Biziura lobata

Proventriculus: collected Tailem Bend, S.A.

194 RECS, AUST. MUS., 18 (17): 385-413

Tetrameres calidris Mawson, 1968

Parasitology 58, p. 300

Holotype ¢ VI410 from Calidris canutus

Proventriculus: collected Casuarina Beach, N.T,

Paratypes AHC4491

Tetrameres cladorhyncht Mawson, 1968

Parasitology 58, p. 299-300.

Holotype 2, allotype ¢ V1413 from Chladorhynchus

leucocephalus

Proventriculus: collected St Kilda, Vic.

Paratypes AHC4514

Tetrameres dacelonis Mawson, 1979

Trans. R. Soc. S. Aust. 103 (7), p. 180

Holotype ¢ V2685, allotype ? V2686 from Dacelo

novaeguinede

Proventriculus: collected Brisbane, Qld

Paratypes AHC6942

Tetrameres greeni Mawson, 1979

Trans. R. Soc. §. Aust. 103 (7), p. 180-2

Holotype ¢ V2683, allotype ? V2684 from Pelecanus

conspicillatas

Proventriculus: collected Temmink, Brisbane, Qld

Teirameres pelecani Johnston & Mawson, 1942

Trans. R. Soc. S, Aust. 66 (1), p. 72-3

Rec. S, Aust. Mus. 7 (2), p. 185-6

now Microtetrameres pelecani (Johnston & Mawson,

1942) Mawson, 1979

Holotype é V1416, allotype & (juvenile) V1417 from

Pelecanus conspicillatus

Plesiotype 2 V2719 as Microtetrameres pelecani

Trans, R. Soc. S. Aust. 103 (7), p. 178

Proventriculus: collected Brisbane, Qld and Tailem

Bend, S.A.

Tetrameres scolopacidis Mawson, 1968

Parasitology 58, p. 301-2

Holotype 4, allotype 2 V1412 from Evolia acuminata

Proventriculus: collected Woods Well, S.A.

Spinitectus bancrofit Johnston & Mawson, 1940

Trans, R. Soc, S, Aust. 64 (2), p. 345-6

Holotype 4, allotype 2 V1381 from Mogurnda adspersa

Intestine: collected upper Burnett R,, Qld

Spinitectus percalates Johnston & Mawson, 1940

Trans. R. Soc, S. Aust. 64 (2), p. 345

now Spinitectus plectroplites (Johnston & Mawson,

1940) Johnston & Mawson, 1947

Holotype 4, allotype 2 V1382 from Percalales colan-

orum

Intestine: collected Lower Murray R., 8.A,

Note: A new collection from the type host contained

males of S. plectraplites not previously found and

additional females. Examination of this material

by the authors indicated that 8. percalates falls

as a synonym of S. plectroplites.

Spinitectus plectroplites Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p. 345

elpril, 1983

Holotype @ V1383 from Plectroplites ambiguus

Gill mucus: collected Tailem Bend, S.A,

Stellocaronema charadrti Mawson, 1968

Parasitology 58, p. 294-6

Holotype 4, allotype 2 VL468 from Charadrius cucul-

latus

Under gizzard lining: collected Beachport, 8.A-

Stellocaronemad glareolae Mawson, 1968

Parasitology 58, p. 296

Holotype 2 V1469 from Glareola isabella

Under gizzard lining; collected Pt Augusta, 5.A.

Paratypes AHC4498

Viguiera chabaudi Mawson, 968

Parasiiology 58, p. 761-2

Holotype 4, allotype 2? V1494 from Podargus strigaides

Under gizzard lining: collected Blackwood, $.A.

Viguiera chibiae Mawson, 1968

Parasitology 58, p. 763

Holotype @, allotype ? V1495 from Chibia bracteata

Under gizzard lining; collected Miiner’s Swamp, N.T.

Paratypes AHC4653

Vigniera longicollis Mawson, 1968

Parasitoloxy 58, p. 759-61

Holotype 4, allotype 2 V1493 trom Collurincincla

rufiventris

Under gizzard lining: collected Eyre Peninsula, S.A.

Superfamily Heterakoidea

Heterakis catheturinae T. H. Johnston, 1912

Proc. R, Soc. Qld. 24, p. 73

from Catheturinus lathami

Caecum: collected Burnett R,, Qld

Paratypes AHC272

Superfamily Metastrongyloidea

Antechinostrongylus disgubernaculus Spratt, 1981

J. Parasit, 67 (1) p, 91

Holoyipe @ V2066, allotype 2 V2067 from Antechinus

swainsonil

Lung: collected Nadgee Nature Reserve, N.S. W,

Paratypes V2068

Filaroides (Filaroides) pilbarensis Spratt, 1979

Aust. J. Zool. Suppl. Ser. 67 (5), p, 8-11

Holotype ¢ V1570, allotype 2 V1571 from Anfechinus

rosamundae

Lungs: collected Woodstock Sth., W.A.

Paratypes ¥1572-80

Marsupostrongylus coulstoni Spratt, 1979

Aust. J. Zool. Suppl, Ser. 67, p. 16-18

Holotype 3 V1581, allotype 2 V1582 from Vombatus

Urslnus

Terminal bronchioles of lungs: collected Lucyvale,

Vic.

Paratypes V1 583-4

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Marsupostrongylus dorrigoensis Spratt, 1979

Aust. J. Zool. Suppl. Ser. 67, p. 19-2 |

Holotype ¢ V1585, allotype 2 V1586 from Thyplogale

thelis

Larger bronchioles of lung: collected Dorrigo, N.S.W.

Paratypes V1 587-94

Marsupostrongylus lancealatus Spratt, 1979

dust. J Zool. Suppl. Ser. 67, p. 25-28

Holotype 4 V1595, allotype ° V1596 from -tntechinus

Staarul

Terminal) bronchioles, alveoli, tung parenchyma: col-

lected Powelltown, Vic.

Paratypes V1597-8

Marsupostroigylus longilarvatus Spratt, 1979

Aust. J. Zool. Suppl. Ser. 67, p. 28-31

Holotype 3 V1599, allotype 2 V1600 from Wallabia

bicolor

Terminal bronchioles of lung: collected Bondi State

Forest, N.S,W.

Paratypes V1601-7

Marsupostrongylus riinesi Spratt, 1979

Aust. J. Zool. Suppl. Ser. 67, p. 31-34

Holotype 4 V1608, allotype 2 V1609 from Tricho

surus Ycaninus

Terminal bronchioles of lung: collected Mt Glorious,

Qld

Paratypes V1610-3

Parafilaroides hydrurgae Mawson, 1953

Parasitology 43 (304), p. 294

now Filaroides (Parajfilaroides) hydrurgae (Mawson,

1953) Anderson, 1978

Holotype 2 V2905 from Hydrurga leptonyx

Lung parenchyma: collected Heard Is,

Paratypes AHC3272

Superfamily Muspiceoidea

Durikainema macropi Spratt & Speare, (in press)

Holotype 2 V2849, allotype ¢ V2850 from Macropus

giganleus

Non-periferal blood: collected Durikai, Qld

Paratypes V2851-8

Riouxgolvania beveridgei Bain & Chabaud, 1979

Annales de Parasitol. 54 (2), p. 207-225

Holotype 2 V1915 from Minlopterus schreibersi

Wing membrane: collected Palluma Dam, 120k N of

Townsville, Qld

Superfamily Oxyuroidea

Aleuris brachylopi Johnston & Mawson, 1944

Trans, R. Soe. §. Aust. 68 (1), p. 61

Holotype 4, allotype @ V1271 from Brachylophus fas-

clalus

Intestine: collected Fiji

Austroxyuris finlaysoni Johnston & Mawson, 1938

Rec, 8. Aust. Mus. @ (2), p. 191-3

Holotype 2, allotype 2 V24 from Petauroides volans

var. puinor

Caecum: collected Fitzroy R., Qld

Macropoxyuris brevigularis Mawson, 1964

Parasitolagy 54, p. 257

Holotype 2 V2882, allotype 9 V2883 from Macropus

canguru

Lower intestine: collected Cunnamulla, Qld

Macropoxyuris longigularis Mawson, 1964

Parasitology 54, p, 253-6

Holotype 6 ¥2880, allotype 2 V2881 from Muacropus

canguru

Lower intestine: collected Cunnamulla, Qld

Oxyuris (8. 1.) acuticaudata Johnsten & Mawson,

1938

Rec, S. Aust. Mus. @ (2), p. 194

now Austroxyuris finlaysoni (Johnston & Mawson,

1938) Mawson, 1964

Holotype ? ¥28 from Petauroides voldans var. minor

Caecum and intestine: collected Fitzroy R., Qld

Oxvurts (8. 1.) potaroo Johnston & Mawson 1939

Trans. R. Soc, 8. Aust, 63 (2), p. 309

now Potoroxyuris poloroo (Johnston & Mawson, 1939)

Mawson, 1964

Holotype 2, allotype 4 V1375 from Potorous tridactylus

Intestine: collected Gippsland, Vic.

Note: re-examination of the onginal material by

Mawson revealed males which had not previously

been described.

Parathelandros oedurae Johnston & Mawson, 1947

Trans. R. Soc. S. Aust. 71 (1), p. 25-6.

now Skrjabinodon oedurae (Johnston & Mawson,

1947) Inglis, 1968

Holotype 4 V1245, allotype 2 V1246 from Oedura

robusta

Large intestine: collected West Burleigh, Qld

Passalurus parvus Johnston & Mawson, 1938

Ree, S. Aust. Mus. 6 (2), p. 193

now Paraustroxyuris parvus (Johnston & Mawson,

1938) Mawson, 1964

Holotype 3 ¥25 from Petauroides volans vav. minor

Intestine: collected Fitzroy R., Qld

Pharyngodon kartana Johnston & Mawson, 1941

Rec, S. Aust. Mus. 7 (1), p. 145-6

Holotype 3, allotype ? V1260 from Gymnodacivlus

multi

Intestine: collected Kangaroo Is., S.A.

Paratypes AHC1655

Pharyngodon limnodynasies Johnston & Mawson,

1942

Proc. Linn. Soc. N.S.W. 67 (2), p. 94

Holotype % V1409 from Limnedynastes dorsalis

dumerilli

Intestine: collected Tailem Bend, S.A.

4396

Skrjabinodon smythi Angel & Mawson, 1968

Trans. R, Soc. S. Aust. 92, p. 69-70

Holotype 2 V2884, allotype 2 V2885 from Phyllodac-

tvlus marmoratus

Intestine: collected Port Gawler, S.A,

Paratype AHC8663

Syphacia trichosuri Johnston & Mawson, 1938

Rec. S. Aust. Mus. 6 (2), p. 194

now Adelonema trichosuri (Johnston & Mawson, 1938)

Mawson, 1978

Holotype 2 V29 from Trichosurus vulpecula

Allotype ¢ V3049 as Adelonema trichosuri

Trans. R. Soc. S. Aust. 102 (8), p, 223

Intestine: collected West Burleigh, Qld

Thelandros kartana Johnston & Mawson, 1941

Rec, S. Aust. Mus. 7, p. 145

Holotype 3 V1248, allotype ¢ V1244 from Hemiergis

peront

Paratypes V1244

Intestine: collected Kangaroo Is., S.A.

Thelandros trachysauri Johnston & Mawson, 1947

Trans. R. Soc. S. Aust. 71 (1), p. 24-5

Holotype @ V1242, allotype 2 V1243 from Trachysau-

rus rUROSUS

Intestine: collected Adelaide, S.A,

Superfamily Physalopteroidea

Bancroftinema dentatum Johnston & Mawson, 1941

Trans. R. Soc, S, Aust. 65 (1), p. 33-4

Holotype ? V1422 from Hieracidea berigora

Stomach: collected Eidsvold, Qid

Paraleptus australis Johnston & Mawson, 1943

Trans. R, Soc, S. Aust. 67 (2), p. 188

Holotype 3, allotype 2 V1290 from Heterodontus phi|-

lipi

Alimentary canal: collected Pt. Willunga, S.A.

Physaloptera banfieldi Johnston & Mawson, 1941

Ree. Aust, Mus. 21 (1), p. 11

from Melomys banjieldi

Stomach: collected Dunk Is., Qld

Paratypes AHC1664

Physaloptera confusa Johnston & Mawson, 1942

Proc. Linn. Soc. N.S, W. 67, p. 90-1

now Abbreviata conjusa (Johnston & Mawsen, 1942)

Chabaud & Bain, 1976

Holotype 4, allotype 2 V1298 from Notechis scuratus

Stomach: collected Tailem Bend, S.A.

Physaloptera demansiae Johnston & Mawson, 1947

Ree. §. Aust. Mus. 9 (1), p. 105

now Abbreviata demansiae (Johnston & Mawson,

1947) Chabaud & Bain, 1976

Holotype 4 V2871, allotype ¢ V2872 from Demansia

psammaphis

Stomach: collected Sydney, N.S.W.

Paratypes V2903-4

REC. 5. AUST. MUS,, 18 (17); 385-413

April, [983

Physaloplera gallardi Johnston & Mawson, 1942

Ree, A, Mus, 21 (2), p. 114

now Abbreyiata gallardi (Johnston & Mawson, 1942)

Chabaud & Bain, 1976 from Amphiholurus bar-

batus

Intestine: collected Narrara, N.S.W.

Paratypes AHC2273

Physaloptera hieracideae Johnsion & Mawson, 1941

Trans, R. Soc. S. Aust. 65 (1), p. 31

Holotype 2 V1419 from Hieracidea orientalis

Gizzard: collected Flinders Is., Bass Strait,

Physaloptera papuensis Johnston & Mawson, 1940

Prac, Linn, soc. N.S. W. 65 (5-6), p, 475

Holotype 4, allotype 2 V1278 from ? bandicoot

Stomach: collected Mt Lamington, Papua New Guinea

Physaloptera parvicollaris Johnston & Mawson, 1940

Proc. Linn. Soc, N.S.W. 65 (5-6), p. 474-5

Holotype 4, allotype 9 V1280 from Perameles nasuta

Stomach; collected Sydney, N.S.W.

Physaloptera peragale Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (1), p. 100

Holotype 4, allotype ° V1578 from Peragale minor

Stomach: collected MacDonald Downs, N.T.

Paratypes AHC2664

Physalaptera sarcaphili Johnston & Mawson, 1940

Proc, Linn. Soc. N.S.W. 65 (5-6), p. 474

Holotype 3, allotype 9, V1279 from Surcophilus

harrisii

Stomach: collected Tas, Biol, Survey

Physaloptera thalacomys Johnston & Mawson, 1940

Proc. Linn, Soc. N.S, MW. 65 (5-6), p. 474

Holotype @, allotype 29 V1277 from Peragale minor

Stomach: collected MacDonald Downs, N.T.

Proleptus trygonorrhinae Johnsion & Mawson, 1943

Trans, R, Soc. 8. Aust. 67 (2), p, 187-8

Holotype 4, allotype 2? V1288 from Trigonorrhina

fasciala

Spiral valve; collected Pt. Willunga, S.A,

Proleptus urolophi Johnston & Mawson, 1951

Trans, R. Soe, S. Aust, 74 (1), p. 23

Holotype 4 V2926, allotype 2? V2927 tram Urolophus

testaceus

Spiral valve: collected Pt Jackson, N,S.W.

Skrjabinoplera — goldmanae = Mawson, 1970

Trans. R. Soc. S. Aust. 94, p, 223-4

Holotype 2, allotype 2 V2811 from Amphibolurus bar-

hatus

Stomach: collected Cobar, N.S,W,

Superfamily Rhabditoidea

Rhabdias hylae Johnston & Simpson, 1943.

Trans, R. Soe. 8S. Aust. 66, p. 176-8

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Syntypes V2808 from Afv/a aurea

Lung: collected Sydney, N,S,W,

Superfamily Rictularioidea

Rictularia carstairsi Mawson, 1971

Trans. R. Soe. Aust. 95 (2), p, 61-3

Holotype 2 V2123, allotype 4 V2124 from Rattus

villosissimius

Duodenum and stomach; collected Brunette Downs

Stn., N.T.

Paratypes AHC5032

Rictularia mackerrasue Mawson, 1971

Trans. R. Soc, S. Aust. 95 (2), p. 63-4

Holotype 2 V1267 from Rattus fuscipes assimilis

Stomach: collected Innisfail, Qld

Rictularia pearsoni Mawson, 1971

Trans. R. Soc, S. Aust. 95 (3), p. 175-6

Allotype 2 V2870 from Rattus fuscipes murrayi

Stomach: collected Pearson Is,, S.A,

Rictularina spinosa Johnston & Mawson, 1941

Proc. Linn, Soc. N.S. 66, p, 254

Holotype 2? V1424 from Myiagra rubecula

Stomach: collected Stradbroke Is., Qid

Superfamily Seuratoidea

Cucullanus heterodonti Johnston & Mawson, 1943

Trans, R, Soe, 8, Aust, 67 (2), p. 187

Holotype 4, allotype @ V1291 from Heterondontus

phillips

Intestine: collected Port Willunga, §.A,

Cucullanellus enidglanis Johnston & Mawson, 1945

Trans. R. Soe. 8. Aust, 69 (1), p, 116

now Dichelyne (Cucullanellus) cnidglanis (Johnston

& Mawson, 1945) Petter, 1974

Holotype 4 V1249, allotype @ V1250 from Cnidoglanis

megastomus

Intestine: collected Port Willunga, S.A.

Cucullanellus sheardi Johnston & Mawson, 1944

Trans, R, Soe, 8S, Aust, 68 (1), p. 64

now Dichelyne (Cucullanellus) sheardi (Johnston &

Mawson, 1944) Petter, 1974

Holotype ¢ V1251, allotype 2? V1252 from Threpterius

maculasus

Intestine: collected Cape Borda, S.A.

Echinonema edmondsi Chabaud et al. 1980

Annales de Parasit, 55 (4), p. 433-5

Holotype 3 V2071, allotype 2 V2072 from Dasyurus

hallucatus

Stomach and small intestine: collected Cannon Hill,

N.T.

Paratypes V2073-91

Echinonema meridtonalis Chabaud et al, 1980

Annales de Parasit. $5 (4), p. 436-7 from fsoodon

obesulus

397

Alimentary canal: collected Waitpinga, 8,A,

Cotypes V2932 & AHC4460

Inglisonema typos Mawson, 1968

Parasitology 58, p. 71-3

Holotype 2, allotype 2 V1697 from FPitla versicolor

Stomach; collected Swainson, Qld

Paraseuratum tandani Johnston & Mawson, 1940

Trans. R, Sac. S. Aust. 64 (2), p. 347-8

Holotype 4, allotype 2 V1389 from Tandanus tandanus

Alimentary canal: collected Tailem Bend, S.A,

Seuratinema hrevicaudatum Johnston & Mawson,

1941

Trans. R. Soc, S. Aust, 65 (1), p. 33

now Skrjabinura brevicaudatus (Johnston & Mawson,

1941)

Mawson, 1960

Holotype 6 V1685 from Ninox connivens

Alimentary canal; collected Burnett R., Qld

Seuratinema magnum Johnston & Mawson, 1941

Proc, Linn, Soc. N.S.W. 66 (3-4), p. 256

now Skrjabinura magnum (Johnston & Mawson,

1941) Mawson, 1960

Holotype 2? V1427 from Dacelo gigas

Alimentary canal: collected Pilliga Scrub, N.S.W,

Seuratinema pomatostomi Johnston & Mawson, 194|

Proc. Linn, Soc. N.S. W. 66 (3-4), p. 255-6

now Skrjabinura pomatostomi (Johnston & Mawson,

1941) Mawson, 1960

Holotype 2, allotype 9 V1428 from Poynatostomus

superciliosus

Alimentary canal; collected Baradine, N.S.W.

Seurechina chaneeti Chabaud et al, 1980

Annales de Parasit, 5§ (4), p, 429-31

Holotype 2? V2069, allotype ¢ V2070 from Dasyurus

hallucatus

Intestine: collected Koolan Is., W.A,

Superfamily Spiruroidea

Alainchabaudia alcedinis Mawson, 1968

Parasitolagy 58, p. 741-2

Holotype 2, allotype 9 V1488 from Halcyon sanctus

Gizzard: collected Brisbane, Qld

Cyathospirura dasyuridis Mawson, 1968

Parasitolagy 58, p. 77-8

Holotype 4, allotype ° V1508 from Dasyureps macu-

latus

Stomach: collected N.S.W.

Paratypes AHC5194

Gongylonema alecturae Johnston & Mawson, 1942

Prac. Linn, Soc. N.S. W’. 67 (1-2), p. 92-3

Holotype 4, allotype 2 V1296 from Alectural latham/

No host site data: collected Eidsvold, Qld

398

Gongylonema. beveridgei Mawson, 197)

Trans. R.. Soc. 8. Aust. 95 (3), p- 176-7

Holotype 4 V2822, allotype 2 V2823 from Rattus

juscipes murrayi

Stomach: collected Pearson Is., §.A.

Paratypes AHC7799

Protospirura marsupialis Baylis, 1927

Rec. §. Aust, Mus. 6 (2), p. 190-1

Allotype 4 V30 (Johnston & Mawson, 1938) from

Trichosurus vulpecula

Stomach: collected Eidsvold, Qld

Superfamily Strongyloidea

Beveridgea corneri Mawson, |980

Trans. R. Soc. 8S. Aust. 104 (4), p. 81

Holotype 4, allotype 2 VI910 from Macropus agilis

Stomach: collected Elizabeth Downs, N.T,

Paratypes AHC7801

Buccosirongylus australis Johnston & Mawson, 1939

Trans. R, Soe, S, Aust, 63 (3), p, 142

Holotype 4, allotype 2 ¥1558 from Macropus wilcoxi

Stomach: collected Eidsvold, Qld

Buccostrongylus buccalis Johnston & Mawson, 1939

Trans. R, Soe, S. Aust. 63 (3), p. 141-2

Holotype 4, allotype 2? V1557 from Macropus dorsalis

Stomach: collected Upper Burnett District, Qld

Buccostrongylus labiatus Johnston & Mawson, 1939

Proc, Linn. Soc. N.S. W. 64 (5-6), p. 527

Holotype 4, allotype $ V1371 from Macropus ruficollis

Stomach; collected Bathurst district, N.S.W.

Paratypes AHC7291

Buccostrongylus setifer Johnston & Mawson, 1939

Proc. Linn. Soc. N.S. W. 64 (5-6), p. 527-8

Holotype 2 V2815, allotype 2 V2816 from Macropus

ruficollis

Stomach: collected Bathurst, N.S.W.

Cassunema exigua Beveridge & Johnston, 1981

Systematic Parasit. 3, p. 78-80

Holotype é V1994, allotype 2 V1995 trom Thylogale

stigmatica

Stomach; collected Tolga, Qld

Paratypes V1996-05

Cloacina annulata Beveridge, 1979

J. Helminth. 53, p, 369-70

Holotype ¢ V1645, allotype 2 V1646 from W’allabia

hicolor

Stomach: collected Bellbird Ck., Vic.

Paratypes V1647-50

Cloacina australis Johnston & Mawson, 1938

Trans. R, Soe, 8. Aust, 62 (2), p. 275

now Cloacina daveyi (Johnston & Mawson, 1938)

Mawson, 1977

REC. S, AUST, MUS., 18 (17); 385-413

atpril, 1983

Holotype 4, allotype 2 V1358 from Macropus robustus

Stomach: collected Mt Licbig, N.T-

Note! Mawson examined specimens identified as

AMacropostrongylus australis Yorke &

Maplestone, 1926 by Baylis (1934), She re-

described the species from this material and

additional specimens from the same host in

Papua. From Yorke & Maplestone’s figures A/,

australis appeared to be referable to Cloacina as

were the specimens identified by Baylis. As C.

australis (Yorke & Maplestone) predated C. aus-

tralis Johnston & Mawson, 1938 Mawson pro-

posed C. daveyi as the new name,

Cloacina bancrofterum Johnston & Mawson, 1939

Trans. R. Soc. S, Aust. 63 (3), p. 133

Holotype 4, allotype 2? V155! from Macropus dorsalis

Stomach; collected upper Burnett R., Qld

Cloacina burnetiiana Johnston & Mawson, 1939

Trans. R. Soc. §. Aust. 63 (3), p. 130-1

Holotype 4, allotype 2 V1548 from Macropus dorsalis

Stomach: collected upper Burnett R., Qld

Cloacina castor Bevendge, 1979

J. Helminth, 53, p, 364-7

Holotype 3 V1651, allotype ? V1652 from Wa/labia

bicolor

Stomach: collected Teesdale, Vic,

Paratype V1653

Cloacina clarkae Mawson, 1972

Trans. R, Soe, S. Aust. 96 (2), p. 109-112

Holotype 4, allotype 2 V1506 from Macropus eugenii

Stomach: collected Kangaroo Is., S.A.

Cloacina communis Johnston & Mawson, 1938

Trans. R. Soc. S. Aust. 62 (2), p. 275-277

Holotype 4, allotype 2 V1355 fram Macropus robustus

Stomach: collected Mt Liebig, N.T,

Paratypes AHC8143

Cloacina curta Johnston & Mawson, 1938

Trans. R. Soc. §, Aust, 62 (2), p. 279-280

Holotype 4, allotype 2 V1354 from Macropus robustus

Stomach: collected Mt Liebig, N.T.

Cloacina digitata Johnston & Mawson, 1940

Proc. Linn. Soc. N.S.W’. 65 (5-6), p. 469

Holotype 4, allotype 2 V1282 from Macropus dorsalis

Stomach: collected Burnett R., Qld

Cloacina dubia Johnston & Mawson, 1938

Trans. R. Soe, 5. Aust. 62 (2), p, 280-1

Holotype 4, allotype 2 V1357 from Macropus rohustus

Stomach: collected Mt Liebig, N.T.

Cloacina edwardsi Mawsan, 1972

Trans, R. Soc. 8. Aust. 96 (2), p. 112

Holotype @, allotype ° V1507 from Wallahia bicolor

Stomach: collected Sunday I!s., Vic.

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM 499

Claacina elegans Johnston & Mawson, 1938

Trans. &. Soc. S. Aust. 62 (2), p. 271-2

Holotype 4, allotype ? V1247 from Petrogale lateralis

Stomach: collected Hermansburg, N.T.

Cloacina ernabella Johnston & Mawson, 1938

Trans. R. Soc. S. Aust. 62 (2), p, 281-2

Holotype 4, allotype 2 V1351 from Petrogale lateralis

Stomach: collected Ernabella, 8.4,

Cloacina expansa Johnston & Mawson, 1939

Proc. Linn. Soc. N.S.W. 64 (5-6), p, 529

Holotype 4, allotype 9 V1564 from Macropus major

Stomach: collected Coonamble, N.S.W,

Cloacina frequens Johnston & Mawson, 1938

Trans. R. Soe. 8. Ausi. 62 (2), p, 273-4

Holotype 4, allotype 2. V1359 from Macrapus robustus

Stomach: collected Mt Liebig, N.T.

Cloacina gallardi Johnston & Mawson, 1940

Proc, Linn. Sac. N.S. W. 65 (5-6), p, 470

Holotype ¢, allotype 9 V1283 from Macropus

ualahatus

Stomach: collected Ourimbah, N.S.W.

Cloacina hydriformis Johnston & Mawson, 1938

Trans. R. Soc. 8. Aust. 62 (2), p. 273

Holotype 2, allotype 2 V1364 from Macropus robustus

Stomach: collected Mt Liebig, N.T.

Paratypes AHC1 841

Cloacina inflata Johnston & Mawson, 1938

Trans. R. Soc. S. Aust. 62 (2), p, 285-6

Holotype 4, allotype 2 V1353 from Mucropus rufis

Stomach: collected Mt Liebig, N,T.

Cloacina kartana Mawson, 1975

Trans. R. Soe. S. Aust. 99 (1), p. 40-1

Holotype 3, allotype 2? V1266 from Macrapus eugenii

Stomach: collected Kangaroo Is., S.A.

Paratypes AHC5829

Cloacina liebigi Johnston & Mawson, 1938

Trans. R. Soc. S. Aust, 62 (2), p. 284-5

Holotype ¢, allotype 2 V1363 from Macropus rufus

“Stomach: collected Mt Liebig, N.T.

Cloacina linstowi Johnston & Mawson, 1940

Proc, Linn, Soc. N.S.W. 65 (5-6), p. 469-70

Holotype 4, allotype ? V1284 from Macropus ruficollis

Stomach: collected Burnett R,, Qld

Cloacina longispiculata Johnston & Mawson, 1939

Trans. R. Soc. S. Aust. 63 (3), p. 131-2

Holotype 4, allotype 9 V1550 from Macropus agilis

Stomach: collected Gregory R., Carpentaria District,

Qld

Cloacina macropodis Johnston & Mawson, 1938

Trans. R. Sav. S. Aust. 62 (2), p, 278-9

Holotype 4, allotype 2 V1356 from Macropus robustus

Siomach. collected Mt Liebig, N.T.

Cloacina magna Johnston & Mawson, 1938

Trans. R. Sac, 8. Aust. 62 (2), p. 277

now C. cummunis (Johnston & Mawson, 1938) Maw-

son, 196]

Holotype 4, allotype 2 V1362 from Macrapus robustus

Stomach: collected Cockatoo Ck,, N.T-

Cloacina magnipapillata Johnston & Mawson, 1939

Prac. Linn. Sac. N.S.W. 64 (5-6), p. 350

Holotype 2, allotype 2? V1566 from Macropus major

Stomach: collected Coonamble, N.S.W.

Cloacina mawsonae Beveridge, 1979

J. Helminth. 53, p. 364-7

Holotype ¢ V1619, allotype 2 V1620 from HWallabia

bicolor

Stomach: collected Bonang, Vic.

Paratypes V1621-4

Cloacina minor Johnston & Mawson, 1938

Trans, R, Soe. S, Aust, 62 (2), p. 283

now Cloacina longelabiata (Johnston & Mawson, | 938)

Johnston & Mawson, 1939

Holotype 4, allotype 2 V1360 from Macropus robusius

Stomach: collected Mt Liebig, N.T.

Note: the authors assignment of Macropostrogv/us

rilnor Davey & Wood, 1938 to Cloacina neces-

Sitated the renaming C. minor Johnston & Maw-

son

Cloacina mundayi Mawson, 1972

Trans. R. Soc. 8. Aust, 96 (2), p. 109

Holotype 3, allotype 2 V1505 fram Macrapus

rufogrisea

Stomach: collected Tarraleah, Tas,

Cloacina obtusa Johnston & Mawson, 1939

Proc. Linn. Soc, N.S.W. 64 (5-6). p. 530

Holotype ¢, allotype 2 V1565 from Macrapus major

Stomach: collected Coonamble, N.S.W.

Cloacina papillata Beveridge, 1979

J. Helminth, 53, p. 367-8

Holotype $ V1625, allotype 2 V1626 from Wallabia

bicolor

Stomach; collected Marlborough, Qld

Paratypes V1627-44

Cleacina parva Johnston & Mawson, 1938

Trans, R. Soc. 5. Aust, 95 (3), p. 171

from Petrogale pearsoni

Stomach: collected Pearson Is.

Paralypes AHC6983

Cloacina petrogale Johnston & Mawson, 1938

Trans. R. Soc, §, Aust. 62 (2), p, 277-8

Holotype é V2896, allotype 2 V2897 from Petroga/?

lateralis

Stomach: collected Mt Liebig, N-T.

Paratypes AHC1846

Cloacina pollux Beveridge, 1979

J, Helminth. 53, p. 375-6

400

Holotype ¢ V1654, allotype 2 V1655 from Wallabia

bicolor

Stomach: collected Townsville, Qld

Paratypes V 1656-70

Cloacina robertsi Johnston & Mawson, 1939

Trans, R, Soc, S, Aust. 63 (3), p. 129-30

Holotype @, allotype 2 VI547 from Petrogale

penicillata

Stomach: collected Upper Burnett R., Qld

Cloacina setonicis Mawson, 1961

Trans. R, Soc. S. Aust. 85, p. 81-2

Holotype 2 V2899, allotype 9 V2900 from Seionix

brachyurus

Stomach: collected Rottnest Is., W.A.

Paratypes AHC4532

Cloacina similis Johnston & Mawson, 1939

Trans. R. Soc. S, Aust, 63 (3), p, 131

Holotype 2, allotype 2 V1549 from Petrogale penicil-

lata

Stomach: collected Upper Burnett R., Qld

Cloacina smalesae Mawson, 1975

Trans. R, Soe, 8S, Aust. 99 (1), p. 39

Holotype 3, allotype 2 V1514 from Macropus eugenti

Stomach: collected Kangaroo Is., 8.A,

Paratypes AHC5605

Cloacina thetidis Johnston & Mawson, 1939

Proc, Linn. Soc. N.S. 4’, 64 (5-6), p, 532

Proc, Linn, Soc, N.S.W. 65 (5-6), p. 471

Holotype 2 V2877 from Macropus thetidis

Allotype é V1281 from Av. parma

Stomach: collected New England, N.S,W.

Ourimbah, N,S.W.

Cloacina vesiibulata Johnston & Mawson, 1940

Trans. R. Soe, S. Aust, 64 (1), p. 97

Holotype 2? V1379 from Macropus melanops

Stomach: collected Mundalla, S.A.

and

Cloacina wallabiae Johnston & Mawson, 1939

Proc. Linn. Soc. N.S.W’. 64 (5-6), p. 532

Holotype ¢@, allotype ¢@ VI567 from Macropus

ualahbatus

Stomach: collected Lower Hawkesbury, N.S.W.

Corallosirongylus hypsiprymnodontis Beveridge, 1978

J. Parasit. 64 (4), p. 657-9

Holotype 4 VIL83, allotype 2 V1184_ from

Hypsiprymnodon moschatus

Intestine: collected Boar Rocket Rd., via Atherton,

Qld

Paratype V1185-92

Coronostrongylus coronatus Johnston & Mawson, 1939

Trans. R, Soc. S. Aust. 63 (3), p. 145-6

Holotype 4 V1560, allotype 2 V1287 from Macropus

wilcaxt

Stomach; collected Eidsvold, Qld

REC. S. AUST. MUS., 18 (17): 385-413

April, 1Y83

Cyelostrongylus alatus Beveridge, 1982

Aust. J. Zool. Suppl. Ser. 83, p. 93-4

Holotype 2 V1768, allotype 2? V1769 from Macropus

rufogriseus banksianus

Oesophagus: collected Swanfels, Qld

Paratypes V1770-9

Cyclostrongvlus clelandi Johnston & Mawson, 1939

Proc. Linn. Sac. N.S.W. 64 (5-6), p. 518

now Alocostoma clelandi (Johnston & Mawson, 1939)

Mawson, 1979

Holotype 3, allotype ° V1696 from Afacropus major

Oesophagus: collected Coonamble, N.S.W-

Cyclostrongylus dissimilis Johnston & Mawson, 1939

Proc. Linn. Soc. N.S. W. 64 (5-6), p. 519-20

now Macropostrongyloides dissimilis (Johnston &

Mawson, 1939) Mawson, 1977

Holotype & V2817 from Macropus ualabatus

Oesophagus: collected Milson Is., Lower Hawkesbury,

N.S.W,

Note: C. dissimilis originally described from a single

damaged male specimen, Mawson assigned the

species to Macropostrologyloides after examining

fresh material.

Cyclostrongylus elegans Beveridge, 1982

Aust. J. Zool, Suppl. Ser. 83, p. 97-8

Holotype 4 V1792, allotype 2 V1793 from Macropus

parryi

Oesophagus: collected Rivertree, N.S.W.

Paratypes V1794-03

Cyclostrongylus gallardi Johnston & Mawson, 1939

Proc, Linn. Soc. N.S. MW, 64 (5-6), p. 518

Holotype 4, allotype 9 V1561 from Afacropus ruficallts

Oesophagus: collected Ourimbah, Gosford District,

N.S.W.

Cyclostrongylus perplexus Beveridge, 1982

Aust. J. Zool, Suppl. Ser. 83, p. 89-90

Holotype ¢ V1780, allotype 9 V!78) from Macropus

rufogriseus banksianus

Oesophagus: collected Grampian Ra., Vic,

Paratypes V1782-91

Cyclostrongylus wallabiae Johnston & Mawson, 1939

Proc. Linn. Soc. N.S. W. 64 (5-6), p. 517-8

Holotype 2 V2818, allotype 2 V2819 fram Macropus

ualabatus

Oesophagus: collected Lower Hawkesbury, N.S,W,

Paratypes AHC1672

Foliostoma macropodis Beveridge & Johnson, L98L

Systematic Parasit. 3, p. 84-5

Holotype 4 V2027, allotype ¢ V2028 from Thylogale

Stigmatica

Stomach: collected Tolga, Qld

Paratypes V2029-33

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Typedontis thetidis Johnston & Mawson, 1939

Proc. Linn. Soe. N.S. WW’, 64 (5-6), p, 534

now Hypodontis macropi (Johnston & Mawson, 1939)

Beveridge, 1979

Holotype 4 V1214, allotype 2 V1215 from Macropus

thetidis

Caccum; collected New England, N.S.W.

Paratypes V1220-3

Lahtostrongylus macropodis Johnston & Mawson,

1938

Trans: R. Sac. S. Aust. 62 (2), p. 266-7

Holotype 2 V2765 trom Afacropus pobustus

Stomach: collected Cockatoa Ck., N.T.

Lahiostrongylus petrogale Johnston & Mawson, 1938

Trans. R. Soc. S. Aust. 62 (2), p. 270-1

Holotype 4, allotype 2 V1367 from Petrogale lateralis

Stomach: collected Mt Licbig, N.T.

Macroponema beveridgei Mawson, 1978

Int. J. Parasitol. 8, p. 164-5

Holotype 4 V1154, allotype 9 V1155 fram Macropus

antilopinus

Stomach: collected Elizabeth Downs, N.T,

Paratypes AHC5999

Macropanema camani Mawson, 1978

Int. J. Parasitol, 8, p, 165-6

Holotype 6 V1156, allotype 2 V1157 from Macropus

giganteus

Stomach: collected Yan Yean, Vic,

Macropicola acydromi Mawson, 1978

Trans. R. Soc. S. Aust. 102 (4), p, 113-5

Holotype 4, allatype 2 V1687 from Macropus fuligi-

nasus ocyvdronus

Large intestine: collected Albany, W.A.

Macropostrongyloides

Mawson, 1978

Aust, J, Zool. 26, p. 777-9

Holotype 4 V1301, allotype 2 V1302 from Macropus

fuliginesus melanops

Caecum: collected Pine Plains Stn via Patchewollock,

Vic,

vamaguiii Beveridge &

Muacraposirangylus dissimilis Johnston & Mawson,

1939

Proc, Linn. Soc. N.S. W. 64 (5-6), p. 526

now 4rundelia dissimilis (Johnston & Mawsan, 1939)

Mawson, 1977

Holotype 4, allotype 2 V1563 from Macropus ualabatus

Stomach; collected Lower Hawkesbury, N.S.W.

Note: fresh collections of material from Wallabia

bicolor (type host) had more visible details of the

buccal capsule which allowed Mawson to amend

the original description.

Macropostrongylus irma Johnston & Mawson, 1940

Rec. Aust. Mus. 20 (5), p. 363

Holotype 3 V1671, allotype 2 V1672 from Wallalna

ira

Stomach: collected Cranbrook, W.A.

Cotypes AHCI816

Macropostrongylus pearsoni Johnston & Mawson.

1940

Trans. R. Sac. S. Aust. 64 (1), p, 98-9

now Popovastrongylus pearsoni (Johnston & Mawson,

1940) Mawson, 1977

Holotype é, allotype 2 V1377 from Petragale pearsoni

Stomach: collected Pearson Is,

Note: The original description was based on two spec-

imens, a male and a female. Additional collections

have provided material in better condition so

that Mawson was able to prepare a more detailed

description.

Macropostrongylus lesouefi Johnston & Mawson, 1939

Prac. Linn, Soc. N.S.W. 64 (5-6), p. 525

Holotype ¢ V2830, allotype 2 V2831 from Macropus

ruficollts

Stomach: collected Sydney Zool. Gardens

Macropostroneylus wallahiae Johnston & Mawsart,

1939

Proc. Linn, Soc. N.S. W. 64 (5-6), p. 525-6

now Popovastrongy his wallabiae (Johnston & Mawson,

1939) Mawson, 1977

Holotype 4, allotype ? V2832 from Maecropus riujicullis

Stomach: collected Bathurst, N.S.W.

Maplestonema typicum Johnston & Mawson, 1939

Proc. Linn, Soc. N.S.W. 64 (5-6), p. 524

Types V1562 from Mfacropus ualahatus

Stomach: collected Lower Hawkesbury, N.S.W.

Note; Subsequent finding of moulting larvae in new

collections of material showed that types. are lar-

val stages of a Cloacina species. Mawson, pers.

comm.

Monilonema lacunosa Beveridge & Johnson, 1981

Systematic Parasit, 3, p. 80-2

Holotype ¢ V2006, allotype 2 V2007 from Thylogale

suigmatica

Stomach: collected Mission Beach, Old

Paratypes V2008-26

Oesophagonastes leptos Mawson, 1965

Parasit. 55, p. 159

now Cyeclostrongylus leptos (Mawson, 1965) Mawson,

1977

Holotype é V1702, allotype 2 V1703 from Macropus

dorsalis

Oesophagus: collected Logan Village, Qld

Paratypes AHC6292

Note: Re-examination of type material by Mawson

revealed that the type species of Cyclosrrongylus

and Oesophagonastes are identical. Oesophagon:

astes thus became a synonym of Cyclostrongylus.

402 REC. S. AUST. MUS., 18 (17): 385-413 April, 1983

Oesaphagostomoides longispicularis Beveridge, 1978

Aust, J. Zoal. 26, p. 599-602

Holotype ¢ V1 340, allotype 2 V1339 from Vombatus

ursinus

Colon: collected Darirnouth, Vic.

Paratypes V1341]

Papillastrongylus labiatus Johnston & Mawson, 1939

Trans, R. Sac. S. Aust. 63 (3), p, 143-4

Holotype 4, allotype 2 V1559 from Macropus dorsalis

Stomach; collected Eidsvold, Qld

Paramacropostrongylus toraliformis Beveridge &

Mawson, 1978

Aus) J. Zool. 26, p. 783-6

Holotype ¢ V1299, allotype ¢ V1300 from Macropus

giganteus

Caecum: collected Yan Yean, Vic-

Paramacropostrongvlus typicus Johnston & Mawsan,

1940

Trans. R. Soc. §, Aust, 64 (1), p. 98

Holotype 4, allotype @.V1376 from Macropus melanops

Stomach: collected Mundulla, $.A.

Parapharyngostrongylus dentatus Beveridge, L982

Aust. J. Zool. Suppl. Ser. 83, p. 46-7

Holotype ¢ V2167, allotype 2 V2168 from Macropus

dorsalis

Stomach: collected Emu Park, Qld

Paratypes V2| 69-78

Pararugopharynx protodemnodontis Magzoub, 1964

Trans. R. Soc. S. Aust. 88, p. 50-1

Holotype 4, allotype 2 V1265 from Protonemnodon

rilfogrisea

Oesophagus and stomach: collected Logan Village,

Qld

Paratypes AHC4918

Parazoniolaimus collaris Johnston & Mawson, 1939

Proc. Linn. Soc. N.S.W. 64 (5-6), p. 522-4

Holotype 3, allotype 2 V1372 from Macropus ualabatus

Stomach; collected Lower Hawkesbury, N.S.W,

Pharyngostrongylus alpha Johnston & Mawsan, 1938

Trans. R, Soc, S. Aust. 62 (2), p. 264-6

now Rugopharynx australis (Moiinig, 1926) Mawson,

1964

Holotype 4, allotype 9 V1365 from Macropus rufus

Stomach: collected Mt Liebig, N.T.

Pharyngastrongylus beta Johnston & Mawson, 1938

Trans. R, Soc. S. Aust. 62 (2), p. 266

now Rugopharyax australis (Moiinig, 1926) Mawson,

1964

Holotype @, allotype 2 V1366 from Macropus rufus

Stomach: collected Mt Liebig, NT.

Pharyngostrongylus delta Johnston & Mawson, 1939

Trans. R. Soc. S. Aust. 63 (3), p. 136

now Rugopharynx delta (Johnston & Mawson, 1939)

Yamaguti, 1961

Holotype 4, allotype 2? V1553 from Macropus dorsalis

Stomach: collected Upper Burnett R., Qld

Pharyngostrongylus dendrolagi Beveridge, 1982

Aust. J. Zool. Suppl. Ser. 83, p. 31-2

from Dendrolagus dorianus

Stomach: collected Lumusa, P.N.G,

Paratypes V1819-20

Pharyngosirongylus diversus Beveridge, 1982

Aust. J. Zool. Suppl. Ser. 83, p. 22-4

Holotype é V1704, allotype 2 V1705 from Macropus

parma

Stomach: collected Niagara Park, N.S.W-

Paratypes V1706-13 & AHC7624

Pharyngosirongylus epsilon Johnston & Mawson, 1939

Trans. R, Soc, S. Aust. 63 (3), p. 137-8

now Rugopharynx epsilon (Johnston & Mawson, 1939)

Yamaguti, 196]

Holotype 3, allotype 2? V1554 from Macrapus dorsalis

Stomach: collected Eidsvald, Qld

Pharyngostrongylus eta Johnston & Mawson, 1939

Trans. R. Soc. S, Aust. 63 (3), p. 139-40

Holotype 4, allotype 2? V1556 from Macropus sp.

Stomach: collected Millmerran, Darling Downs. Qld

Paratypes AHC2568

Pharyngostrongylus gamma Johnston & Mawson,

L939

Trans. R. Soc. §. Aust. 63 (3), p. 134-6

Holotype 4, allotype 2 V1552 from Macropus dorsalis

Stomach: collected Upper Burnett R., Qld

Pharyngastrongylus iota Johnston & Mawson, 1939

Proc. Linn. Soc. N.S. 64 (5-6), p. 514-5

Holotype 4, allotype 2 V 1369 from Macropus ruficollis

Stomach: collected Ourimbah, Gosford District,

N.S. W.

Pharyngostrongylus kappa Mawson, 1965

Parasitology 55, p. 147-50

Holotype é V1700, allotype 2? V1701 from Macropus

canguru

Stomach: collected Inglewood, Qld

Pharyngostrongylus lambda Mawson, 1965

Parasitology 55, p, 151-2)

Holotype 3, allotype 2 V1698-9 from Macropus robus-

tus

Stomach: collected Marble Bar, W.A.

Paratypes AHC6289, Syntypes AHC7226

Pharyngostrongylus nelsoni Beveridge. 1982

Aust. J. Zool. Suppl. Ser. 83, p. 19-22

Holotype 4 V1726, allotype 2 V1727 from Petrogale

venustula

Stomach: collected Mt Borrodaile. N.T.

Paratypes V 1728-47

Pharyngostrongylus parma Johnston & Mawson, 1939

Proe. Linn. Soc. N.S G4 (5-6), p. 516

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

now C'yclostrangylus parma (Johnston & Mawson,

1939) Mawson, 1977 from Mucropus parma

Oesophagus: collected Narara, Gosford District,

NLS.W.

Paratypes AHC1827

Pharyngostrongylus papillatus Beveridge, 1982

Aust. J. Zool. Suppl. Ser. 83, p. 17-9

Holotype ¢ V1714, allotype 2 V1715 from Macropus

antilopinus

Stomach: collected 80 km S.W, of Darwin, N.T.

Paratypes V1716-25 & AHC7612

Pharyngostrongylus setosus Beveridge, 1982

Aust. J. Zool. Suppl. Ser. 83, p. 44-5

Holotype 4 V1748, allotype ¢ V1749 from Thylogale

Stigmatica

Stomach: collected El Arish, Qld

Paratypes V1750-7

Pharyngostrongvlus sharmani Beveridge, 1982

Aust. J. Zool. Suppl. Ser. 83, p. 32-4

Holotype 2 V2139, allotype 2 V2140 from Macropus

hernardus

Stomach: collected El Sharana, N.T.

Paratypes V2141-60

Pharyngostrongylus theta Johnston & Mawson, 1939

Prac, Linn, Soc. NSW. 64 (5-6), p. 514

now Rugopharynx theta (Johnston & Mawson, 1939)

Yamaguti, 1961

Holotype 4, allotype 2? V1368 from Macropus thetidis

Stomach: collected New England, N.S.W.

Pharyngostrongylus zeta Johnston & Mawson, 1939

Trans. R. Soe. S. Aust, 63 (3), p. 138-9

now Rugopharynx zeta (Johnston & Mawson, 1939)

Yamaguti, 196)

Holotype 4, allotype 9 V1555 from Petragale penicil-

lata

Stomach: collected Upper Burnett R., Qld

Phascolostrongylus stirtont Mawson, 1955

Trans, R. Soc. S. Aust. ‘78, p. 5-6

now Oesophagostomoides stirtont (Mawson, 1955)

Beveridge, 1978 from Lasiorhinus latifrons

Colon; collected Blanchetown, §.A-

Syntype 2 V2812

Popovastrongylus irma, Mawson, 1977

Trans. R. Soe. 8S. Aust. 11 (2), p, 57-8

Holotype é V1671, allotype 2 V1672 from Macropus

irma

Stomach: collected Perth, W.A.

Potorostrongylus finlaysoni Johnston & Mawson, 1939

Trans. R. Soc, 8S. Aust. 63 (2), p. 308-9

Holotype ¢, allotype 2 V1374 from Potorous tridactylus

Intestine: collected S. Gippsland, Vic.

Potorastrongylus aepyprymnus Mawson, 1974

Trans. R. Sac, 8. Aust. 98 (3), p. 137

403

Holotype 6 V1692, allotype 2 V1693 from 4epyprym-

nus rufescens

Stomach: collected Warwick, Old

Paratypes AHC5811

Rugopharynx omega Beveridge, |982

Aust. J. Zool. Suppl. Ser, 83, p, 69-71

Holotype 2 V1758, allotype 2 V1759 from Macrupus

rufogrieseus banksianus

Stomach: collected Warwick, Qld

Paratypes V1760-7

Rugopharynx rosemariae Beveridge & Presidente,

1978

Int. J. Parasitol. 8, p. 379-84

Holotype 3 V1193, allotype 2 V1194 from Macropus

giganteus

Stomach: collected Dartmouth, Vic.

Paratypes V1195-1206

Rugopharynx rufogrisea Magzoub, 1964

Trans. R, Soe. §. Aust. 88, p. 49-50

Holotype ¢, allotype 2 V1264 from Protodemmnon rufo-

grisea

Stomach: collected Logan Village, Qld

Paratypes AHC4917

Spirestrongylus kartana Mawson, |955

Trans. R. Soc. S. Aus. 78, p. 2

now Cyelostrongylus kartana (Mawson, 1955) Maw-

son, 1977

Holotype 6 V2886, allotype 2 YV2887 from Macropus

eugenil

Oesophagus: collected Kangaroo Is., $.A.

Paratypes AHC70

Spirostrongylus spirostrongylus Yorke & Maplestonc,

1926

Nematode Parasites of Vertebrates, p. 68

from Macropus agilis

Oesophagus: Qld, probably Townsville

Syntypes AN.C8660

Thylonema arundel Beveridge, 1981

J. Parasit. 67 (1), p. 103-5

Holotype @ V2042, allotype 2 V2043 from Thylogale

stigmatica

Stomach: collected E] Arish, Qld

Paratypes V2044-55

Thylanema barkeri Beveridge, 1981

J. Parasit. 67 (1), p. 105-7

Holotype 3 V1982, allotype 2 V1983 from Thy/ogule:

sugmatica

Stomach: collected Mission Beach, Qld

Paratypes V1984-93

Thylonema thylonema Beveridge. 1981

J, Parasit 67 (1), p. 101-3

Holotype 4 V2034, allotype 2 V2035 from Thrlogale

stigmatica

Stomach: collected Mission Beach, Qld

Paratypes V2036-4]

Thvlostrongvlus parvus Beveridge, 1982

etust. J. Zool. Suppl. Ser. 83, p. 53-4

Holatype ¢ Vi804, allotype ¢ V1805 from Thylogale

thetis

Stomach: collected Emu Vale, Qld

Paratypes V1806-15

Thylostrongylus tasmaniensis Beveridge, 1982

Ausi. J. Zool, Suppl. Ser. 83, p. 53-4

Holotype 4 V2161, allotype 2 V2162 from Thylogale

hillardierti

Stomach: collected Golconda, Tas.

Paratypes V2163-6

Trigonostonema longibursata Beveridge, 1981

J. Parasit 67 (1), p. 95-7

Holotype 4 V2059, allotype 2 V2060 from Thylogale

stigmatica

Siomach: collected Tolga, Qld

Paratypes V2061-5

Trigonostonema trigonostoma Beveridge, 1981

J. Parasit 67 (1), p. 94-5

Holotype 2 V2056, allotype 2 V2057 from Thy/logale

stigmatica

Stomach: collected Mission Beach, Qld

Paratypes V2058

Hvodwardostrongylus obendorfi Mawson, 1976

Trans. R. Soc. S. Aust. 100 (3), p. 121-123

Holotype 2 V1694, allotype 2 V1695 from Macropus

parry't

Ocsophagus: collected Dorrigo, N.S.W,

Zoniolaimus hancrofii Johnston & Mawson, 1939

Trans. R. Soe. §. Aust, 63 (3), p. 123-4

now Lubiastongylus bancrofti (Johnston & Mawson,

1939) Kung, 1948

Holotype 4, allotype 2 V1541 from Macropus parry!

Stomach: collected Eidsvold, Qld

Zoniolaimus hipapillosus Johnston & Mawson, |939

Trans. R. Soc, S. Aust. 63 (3), p. 125

now Labiostroneylus bipapillosus (Johnston & Maw-

son, 1939) Kung, 1948

Holotype 2, allatype 2 V1542 from Macropits major

Stomach: collected upper Dawson Valley, Qld

Zoniolaimus clelandi Johnston & Mawson, 1939

Prac, Linn. Soc. N.S. W. 64 (5-6), p. 521

now Labiostrongylus clelandi (Johnston & Mawson,

1939) Kung. 1948

Holotype 4, allotype 2 V1686

Stomach; collected Lower Hawkesbury, N.S.W.

Zoniolaimus communis Johnston & Mawson, 1939

Trans. R. Soe. 8. Aust. 63 (3), p. 125-6

now Labiostrongylus compiunts (Johnston & Mawson,

1939) Kung, 1948

Holotype 4, allotype 2 V1543 from Macropus ualabatus

Stomach: collected upper Burnett R,, Qld

REC. S, AUST, MUS., 18 (17): 385-413

innit, LIAS

Zoniolaimus insularis Johnston & Mawson, 1939

Trans, R. Soe. 8. Aust. 63 (3), p. 126-7

now Labiostrangylus insularts (Johnston & Mawson,

1939) Kung, 1948

Holotype 2, allotype 2 V1544 from Macropus welsbyi

Stomach: collected Stradbroke Is.

Zoniolaimus onychogale Johnston & Mawson, 1939

Trans. R. Soc. S. Aust. 63 (3), p. 128

now Labiostrongylus onychogale (Johnston & Maw-

son, 1939) Kung, 1948

Holotype 2, allotype 2 V1546 from Onychogale frenata

Stomach: collected Upper Burnett R., Qld

Zoniolaimus ualabatus Johnston & Mawson, 1939

Proc. Linn. Soc. N.S. W, 64 (5-6), p. 521-2

now Labiastrongylus ualabatus (Johnston & Mawson,

1939) Kung, 1948

Holotype 4, allotype 2 V1614 from Macropus ualabatus

Stomach: collected Lower Hawkesbury, N,S,W.

Zoniolaimus uncinatus Johnston & Mawson, 1939

Trans. R. Soe. S. Aust. 63 (3), p. 127-8

now Lubiostrongy/us uncinatus (Johnston & Mawson,

1939) Kung, 1948

Holotype 4, allotype 2 V1545 from Macropus dorsalis

Stomach: collected Upper Burnett R., Qld

Family Strongyloididae

Parastrongyloides australis Mawson, 1960

aust. J, Zoal. 8 (2), p. 281-2

Holotype 4, allotype 2 V1459 from Jsoodon obesulus

Intestine: collected Cherry Gardens, 5,A.

Parustrongyloides peramelis Mackerras, 1959

Aust. J. Zoal. 7 (2), p. 99

from Thylacis obesulus

Small intestine: collected Mt Glorious, Qld

Paratypes AHC3886

Parustrongyloides trichosuri Mackerras, 1959

Aust. J. Zool, 7 (2) p. 96-9

from Trichosurus vulpecula

Small intestine: collected Brisbane, Qld

Paratypes AHC3888

Strongyloides. thylacis Mackerras, 1959

Aust. J. Zool, 7 (2), p. 88-96

from Thylacis obesulus

Small intestine: collected Brisbane, Qld

Paratypes AHC3887

Superfamily Subuluroidea

Lahiobulura baylisi Mawson, 1960

Aust. J. Zool. & (2), p. 279-80

Holotype 4, allotype 2 V1464 from /seodon obesulus

Alimentary canal: collected Mt. Nebo, Qld

Paratypes AHC3285

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Labiolulura inglisi Mawson, 1960

Aust. J, Zool. 8 (2), p. 280-1

Holotype 4, allotype 2 V1462 from Isvodon obesulus

Alimentary canal: collected Cherry Gardens, S.A.

Paratypes AHC3346

Leipoanema ellisi Johnston & Mawson, 1942

Trans, R, Soe. S. Aust. 66 (1), p. 73

Holotype 4, allotype 2 V1292 from Leipoa vcellata

Intestine: collected Tailem Bend, §,A,

Paratypes AHC146, 308

Subulura clelandi Johnston & Mawson, 1941

Proc. Link. Soc. N.S. W. 66 (3-4), p, 251

Holotype 4, allotype 2? V1430 from Podargus strigoides

Intestine; collected Perth, W,A,

Subulura peragale Johnston & Mawson, 1940

Proc. Linh. Soc. N.S.W. 65 (5-6), p. 472-3

now Labiobulura peragale (Johnston & Mawson, 1940)

Mawson, 1960

Holotype 4, allotype 2 V1276 trom Peragale minor

Stomach: collected MacDonald Downs, N,T,

Superfamily Thelazoidea

Metathelazia naghiensis Spratt, 1980

J. Parasit. 66 (6), p. 1032-35

Holotype ¢ V1821, allotype 2 V1822 from Perameles

nasuta

Lung parenchyma: collected Nadgee State Forest,

N.S,W,

Paratypes V1823-30

Oxyspirura bancrofti Johnston & Mawson, 1941

Proc. Linn, Soe. N.S.W. 66 (3-4), p. 255

Holotype ? 1423 from Philemon citreogularis

Orbit: collected Eidsvold, Qld

Rhahbdochona jJaenschi Johnston & Mawson, 1940

Trans, R. Soe. 8. Aust. 64 (2), p. 348

Holotype 3, allotype @ V1390 from Psendaphritis

urvillei

Intestine: collected Tailem Bend, S.A.

Thelazia pittae Johnston & Mawson, 1941

Trans. R. Soc. S. Aust. 65 (2), p. 256

Holotype 2, allotype 2 V1449 trom Pitta macklotti

2Orbit; collected North Qld

Superfamily Trichostrongyloidea

Amidastomum biziuriae Johnston & Mawson, 1947

Ree. S. Aust. Mus. 8 (4), p. 551

Holotype 3 V1259, allotype 2 V2878 from Biziura

lohata

Trans. R. Soe. S. Aust, 82, p. 152-3

Gizzard: collected Tailem Bend and Purnong, S.A.

Amidostamum tribonyx Mawson, 1980

Trans. R. Soc. S. Aust. 104 (1), p. 10-12

Holotype 4 V1864, allotype 2? V1865 from Tribonvx

ventralis

405

Gizzard: collected Swan Reach, S.A.

Paratypes AHC6833

Amphibiophilus egerniae Johnston & Mawson, 1947

Trans. R. Soe, S. Ausl. 71 (1), p. 23 from genta

dahli

Intestine: collected ltari Rocks, Musgrave Ranges,

S.A,

Topotype 2 V2901, 2 V2902

Amphicephaloides thylogale Beveridge, 1979

Aust. J. Z00l. 27, p, 169-74

Holotype é V1470, allotype 2? V1471 from Thylovale

billardierii

Duodenumi: collected Launceston, Tas.

Paratypes V 1472-80

Asymimetracantha tasmaniensis Mawson, 1960

Aust. J. Zool. 8 (2), p. 273-6

Holotype 4, allotype 2 V1460 from Isoodon obesulus

Small intestine: collected Dunorlan. Tas.

Paratypes AHC3334

dAustroheligmonema magna Mawson, 1961

Aust, J, Zool. 9 (5), p. 815-17

now Nippostrongylus magnus (Mawson, 1961) Dur-

ette-Desset. 1970

Holotype 3, allotype 2 V1330 from Rairus assimilis

Intestine: collected Innisfail, Old

Paratypes AHC3851

Austroheligmonema typicum Mawson, 1961

Aust. J. Zool. 9 (5), p. 815-7

now Nippostrongylus typicus (Mawson, 1961) Durette-

Desset, 1970

Holotype 4, allotype 2 V1329 from Ratins assinwills

Intestine; collected Mt Nebo, Qld

Paratypes AHC3852

Austrostrongylus acinocercus Mawson, 1960

Aust. J. Zool. & (2), p. 265-7

now }Woolleya acinacercus (Mawson, |960) Mawson,

1973

Holotype 4, allotype ° V1463 from Perameles nasuta

Intestine: collected Innisfail, Qld

Paratypes AHC3350

Austrostrongylus aggregatus Johnston & Mawson, 1940

Prov. Linn, Soc. N.S.W’. 68 (5-6), p, 472

Holotype 4 V2890, allotype 2 V2891 from Macrapus

ualabatus

Intestine: collected Milson J., N.S.W,

Austrostrangylus chandleri Mawson, 1973

Trans, R. Soc. S. Aust, 97 (4), p. 262-5

Holotype 4, allotype ¢ V1516 from Macrapus rufon

griseus

Intestine: collected Bathurst, N.S.W.

Austrostrongylus hvdromyos Mawson, 1961]

Aust. J. Zool. 9 (5), p. 822-3

now Woolleya hvdromyas (Mawson, 1961) Mawson,

1973

406

Holotype 4, allotype 2 V1333 from Aydromys vhry-

sogaster

Intestine; collected Innisfail, Qld

Paratypes AHC3857

Austrostroneylus hypsiprymnodontis Mawson, 1973

Trans. R. Sac, S. Aust. 97 (4), p. 265

Holotype 2, allotype ? VIS17 from Hypsiprymnodon

mroschatus

Intestine: collected Qld

Paratypes AHCS449

dustrostrongylus minulus Johnston & Mawson, 1938

Rec, &. Aust. Mus, 6 (2), p. 195

Holatype 4, allotype 9 V26 from Macropus dorsalis

Intestine: collected Eidsvold, Qld

Atustrostrangylus paralypicus Mawson, 1973

Trans. R. Soe. 8. Aust. 97 (4), p. 262

Holotype 4, allotype ? VI515 from Macropus rufo-

Briseus

[ntestine: collected Bathurst, N.S.W,

Austrostrongylus poteroo Johnston & Mawson, 1949

Trans. R. Soe, 8. Aust, 73, p. 64-5

now Paraustostroneylus potoroo (Johnston & Mawson,

1949) Mawson, 1973

Holotype ¢ V2820, allotype 2 V2821 from Potarous

tridactylis

Intestine; collected King Is., Tas,

Paratypes AHC7796, 6040

Austrostrangvlus thylogale Joinston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (1), p. 99

Holotype 4, allotype 2 V1373 trom Thylogale eugenii

Intestine: collected Kangaroo Is,, S.A,

Austrostrongylus wallabiae Johnston & Mawson, 1939

Proc, Linn. Sac. NSW. 64 (35-6), p. 534

Holotype 4, allotype 2 V1370 from Macropus ruficollis

Intestine: collected Bathurst, N.S.W.

Beveridgiella pearsoni Humphery-Smith, 1980

Bull. Mus. nain. Hist. nat., Paris Ser 4 2 A (4),

1003-6

from Isnodon macrourus

Intestine: collected Townsville, Qld

Paratypes V2701

Copemania ohendorfi Durette-Desset & Beveridge,

L981

Annal, Parasit. (Paris) 56 (1), p. 63-6

from Dasyurus maculatus

Intestine: collected Warmambool, Vic.

Cotypes V2690, 269]

Dessetostrongvlus moorhousei Humphery-Smith, 1980

Bull. Mus. nam. Hist, nat., Paris Ser 4 2 A (4),

p. LOO7-8

from Antechinus swainsonit

Intestine: collected Nadgee, N.S.W.

Paratypes V2702-9

REC. S. AUST. MUS., 18 (17): 385-413

April, L983

Desyetostrangylus maudii Aumphery-Smith, 1980 Bull.

Mus. natn. Hist. Nat. Paris Ser 4 2 A (4) p. 1008-

trom ntechinus swainsonil

Intestine: collected Nadgee, N.S.W.

Paratypes V2710-8

Filarinema peramelis Johnston & Mawsan, 1938

Rec. S. Aust. Mus. 6 (2), p. 196-7

now Mackerrostrongylus peramelis (Johnston &

Mawson, 1938) Mawson, 1960

Holotype 4, allotype 2 V27 from [savdon obesulus

Intestine: collected West Burleigh, Qld

Globocephaloides affinis Johnston & Mawson, 1939

Trans. R. Soc. S. Aust. 63 (3), p, 147-8

now Globocephaloides macropedis Yorke &

Maplestone, 1926 after Beveridge, 1979

Holotype 2? V1216 from Macropus dorsalis

Intestine: collected Eidsvold, Qld

Paratype 2 V1219

Globocephaloides thetidis Johnston & Mawson, 1939

Proc. Linn. Soc, N.S.W. 64 (5-6), p. 533

now Globocephaloides macropodis Yorke &

Maplestone, 1926 after Beveridge, 1979

Holotype 3 V1218, allotype 2 V1285 from Thylogale

thetis

Intestine: collected New England, N.S.W.

Note: type material for Globocephaloides wallabiae

Johnston & Mawson, 1939 is missing, presumed

lost. Beveridge (1979) has also put this species

as a synonym of Globocephaloides macropodis

Yorke & Maplestone, 1926.

Heligmonaides emanuelae Mawson, 1961

Aust. J. Zool. 9 (5), p. 809-10

now QOdilia emanuelae (Mawson, 1961) Durette-

Desset, 1973

Holotype 4, allotype ? V1332 from Rattus conatus

Intestine: collected Innisfail, Qld

Heligmonoides mackerrasae Mawson, 1961

Aust. J, Zoal. 9 (5), p. 808-9

now QOdilia emanuelae (Mawson,

Desset, 1973

Holotype 2 allotype 2 V1331 from Melomys cervinipes

Stomach and intestine: collected Innisfail, Qld

Paratypes AHC3860

Hepatojarakus fasciatus Mawson, 1961

Aust. J, Zool. 9 (5), p. 822

Holotype 3, allotype 2 V1335 from Raltus conatus

Abdominal mesentery; collected Innisfail, Qld

1961) Durette-

Hepatojarakus pycnofasciatus Mawson, 1961

Aust. J. Zool. 9 (5), p. 821-2

Holotype 4, allotype 2 V1336 from Rattus assimilis

Liver and small intestine: collected Innisfail, Qld

Paratypes AHC3856

Ichthostrongylus clelandi Mawson, 1954

Trans, R. Soc. §. Aust. 77, p. 162-3

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Holotype 2 V2888, allotype 2? V2889 from Enilssola

antarclica

Spiral valve: collected Encounter Bay, S.A.

Paratypes AHC1412, 8602, 8603

Longisiriata brachybursa Mawson, 1961

Aust, J, Zool, 9 (5), p. 805-6

now Odilia brachybursa (Mawson, 1961) Durette-

Desset, 1973

Holotype 2, allotype 2° V1327 from Melomys cervinipes

Intestine; collected Innisfail, Old

Paratypes AHC3862

Longistriata melomyos Mawson, 1961

Aust. J. Zool. 9 (5), p, 801-4

now Odilia melamyos (Mawson, 1961) Durette-Desset,

1973

Holoytpe 4, allotype 2 V1328 from Melanomys cer-

VIMIpes

Intestine: collected Innisfail, Qld

Paratypes AHC3861

Longistriata polyrhahdole Mawson, 1961

Aust. J. Zoul. 9 (5), p. 807

now Qdilia polyrhabdote (Mawson, 1961) Durette-

Desset, 1973

Holotype 4, allotype 2 V1325 from Rartus assimilis

Intestine: callected. Mt Glorious, Old

Paratypes AHC3854

Longistriata uromyos Mawson, 1961

Aust. J. Zool. 9 (5), p. 804-5

now Odilia uramyos (Mawson, 1961) Durette-Desset,

1973

Holotype 4, allotype 2 V1326 from Uramys caudi-

maculatus

Intestine; collected Innisfail, Qld

Paratypes AHC3858

Mackerrastrongylus isoodon Durette-Desset &

Cassone, 1980

Bull. Mus, natn. Hist. nat. Paris Ser 42 A (4), p. 946-

Holotype 4 V2699, allotype 2 V2700 from Isoodon

macrourus

Intestine: collected Townsville, Qld

Mastonema melomyos, Mawson, 1961

Aust. J, Zool, 9 (5), p. 818-9

now AMfammanidula melomyos (Mawson,

Durette-Desset, 197]

Holotype ¢, allotype 2 V1334 from Melomys lutillus

Mammary gland: collected Whyanbeel, Qld

Paratypes AHC3859

1961)

Nicallina phascogale Mawson, 1960

Parasitology 50, p. 425-6

now Patricialina phascogale (Mawson, 1960) Inglis,

1968

Holotype 4, allotype 2? V1457 from Phascogale flavipes

Small intestine; collected Innisfail, Qld

Paratypes AHC3337

a07

Nicollina antechini Beveridge & Barker, 1975

J. Parasitolagy V61 (3), p. 489-93

Nasal cavity: collected Powelltown, Vic.

Paratypes V53

Nicollina baylisi Mawson, 1973

Trans. R. Soe, §. Aust. 97 (4), p. 270

Holotype ¢, allotype ? V1269 from Tachyglossus acu-

leatus

Small intestine: collected Tas,

Paratypes AHC5459

Nicollina calabyi (Mawson, 1973)

Trans. R, Soe, 8, Aust. 97 (4), p. 273

now Beveridgiella calabyi (Mawson, 1973) Humphery-

Smith, 1980

Holotype 4, allotype 2 V1270 from Myrmecobius fas-

claus

Small intestine: collected W.A.

Nicallina cameroni Thomas, 1959

Trans. R. Soc. 8S. Aust. 82, p. 154-5

Holotype ¢ V2873, allotype ? V2874 from Tachyglos-

sus aculeatus

Small intestine collected Kangaroo Is., S.A.

Paratypes AHC829

Note: This species was described in the only paper in

which Mrs Thomas accidentally did not retain

use of the name Mawson.

Nicollina inglisi Mawson, 1973

Trans, R, Soc. 8. Aust. 97 (4), p. 273

now Beveridgiella inglisi (Mawson, 1973) Humphery-

Smith, 1980

Holotype 4, allotype 2 V1520 from Myrmecohiuy fas-

claus

Small intestine: collected W, A,

Nicollina fota Mawson, 1960

Aust. J. Zool, & (2), p. 264-5

now Beveridgiella tora (Mawson, 1960) Humphery-

Smith, 1980

Holotype 4, allotype 2 V1465 from /soodon obesu/ts

Small intestine: collected Dunorlan, Tas.

Nicollina mundayi Mawson, 1973

Trans. R. Soc. S. Aust, 97 (4), p. 270-3

Holotype 2, allotype 2? V1519 from Yachyglossus acu-

leatus

Small intestine: collected Tas..

Paratypes AHC58

Oswaldocruzia limnodyaastes Johnston & Simpson,

1943

Trans. R. Soc. S. Aust. 66, p, 172

Holotype 2, allotype 2 V1274 from Limnodynasies

dorsalis

Intestine: collected Adelaide, S.A.

Paratypes AHC2363

408

Paraustrostrongylus bettongia Mawson, 1973

Trans. R. Soc: S, Aust. 97 (4), p. 267

Holotype 2, allotype 2 V1518 from Bettongia gaimardi

Intestine: collected Tas.

Paratypes AHC5415

Paraustrostrongvlus trichosuri Mawson, 1973

Trans. R. Soe. S. Aust, 97 (4), p. 267-8

Holotype 4, allotype 2 V1407 from Trichosurus vul-

pecula

Intestine: collected D'Aiguillar, Qld.

Paratypes AHC5443

Paraustrosirongylus ratti Obendorf, 1979

dust, J. Zool, 27 (2), p. 867-9

Holotype 3 V810, allotype V811 from Raitus fuscipes

Intestine: collected Gladysvale, Vic.

Paratypes V812-3

Patricialina birdi Humphery-Smith & Durette Desset,

1981

Bull. Mus. natn. Ilist. nat., Paris, Ser 4.3 A (1), p.

128-30,

Holotype 4 V2721, allotype 2 V2722 from Antechinus

swainsonit

Intestine: collected Nadgee, N.S.W-

Peramelistrongvlus skedastos Mawson, 1960

Aust. J. Zool. 8 (2), p. 268-71

Holotype ¢, allotype 2? V1461 from Isoadon obesulus

Stomach and intestine: collected Innisfail, Qld.

Profilarinema hemsleyi Durette-Desset & Beveridge,

1981

Ann. Purasil. (Paris) 56 (2), p. 188-91

from Trichosurus vulpecula

Stomach: collected W.A,.

Cotypes V2694-8

Sprattellus harrigani Durette-Desset & Cassone, 1980

Bull, Mfus. natn. Hist. nat,, Paris Ser 4 2 A (4),

p. 949-51

from Phascogale tapoatafa

intestine: collected Vic.

Cotypes V2694-8

Sprattellus woolleyue Durette-Desset & Cassone, 1980

Bull. Mus. natn, Hist. nat. Paris Ser 4 2 A (4),

p. 951-53

from Anlechinus stuartii

Intestine: collected Sandy Point, Western Point, Vic.

Cotypes V2692-3

Tetraboitirivostrongylus mackerrasae Mawson, 1960

Parasitology 50, p. 427-8

Holotype 2, allotype 2? V1458 from Phascogale flavipes

Intestine: collected Innisfail, Qld.

Paratypes HC3340

Trichostrongylus (s.L) incertus Johnston & Mawson,

1941

REC. 8. AUST. MUS,, 18 (17): 385-413

dprtl, 1983

Trans. R. Soc. 8. Aust. 65 (2), p. 254-5

Holotype 2 V1448 from Hydroprogne caspia strenua

Intestine: collected Tailem Bend, S.A.

Woolleya antechini Humphery-Smith & Durrette-

Desset (in press)

Holotype 4 V2723, allotype 2 V2724 from Antechinus

sruartil

Intestine: collected Powelltown, Vic.

Woalleva didelphis Humphery-Smith & Durette-Des-

set (in press)

Holotype ¢ V2728, allotype 2 V2729 from Antechinus

swainsonit

Intestine: collected Nadgee, N.S.W.

Wooalleya hickmani Mawson, 1973

Trans. R. Soc. S, Aust. 97 (4), p. 277-8

now Patricialina hickmani (Mawson, 1973), Hum-

phery-Smith & Durette-Desset, 1981

Holotype é, allotype 2? V1523 from Aniechinus stuartil

Intestine: collected Condor Ck., A,C.T-

Hoolleva martini Mawson, |973

Trans. R. Soc. 8. Aust. 97 (4), p, 275

now FPatricialina martini (Mawson, 1973) Humphery-

Smith & Durette-Desset, 1981

Holotype 3, allotype 2 V1522 from Anlechinomys

spencert

Intestine: collected Sandringham, Qld.

Paratypes AHC7295

Woalleya monodelphis Mawson, 1973

Trans. R. Soc, S. Aust. 97 (4), p. 278-9

Holotype 4, allotype 2 V1524 from Antechinus stuartil

Intestine: collected Condor Ck,, A.C.T.

Woolleya sprenti Mawson, 1973

Trans. R. Soc. S. Aust. 97 (4), p, 275

Holotype 4, allotype 2? V1521 from Dasyurus viverrinus

Intestine: collected Icena, Tas.

Paratypes AHC5447, 4940

Superfamily Trichuroidea

Anatrichosoma haycocki Spratt (in press)

Holotype 3 V2859, allotype 2 V2860 from Antechinus

sWwainsonil

Paracloacal glands: collected Nadgee State Forest,

N.S.W,

Paratypes V2861-69

Cupillaria cooperi Johnston & Mawson, 1945

Trans. R. Soc, S. Aust. 69 (2), p. 245

Holotype ¢ V1230, allotype 2 V¥1231 from Callionpnius

calaurapomus

Alimentary canal: collected St Vincent's Gulf, S.A,

Capillaria ellisi Johnston & Mawson, 1945

Trans. R. Soc. S. Ausi. 69 (2), p. 247

Holotype $ V1237, allotype 2 V1238 from Chenopsis

atrala

Alimentary canal: collected Tailem Bend, S.A.

HELMINTH TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Capillaria graucalina Johnston & Mawson, 1941

Proc. Linn. Soc. N.S.W. 66 (3-4), p. 250

Holotype 4, allotype 2 V1431 from Coracina novae-

hollandiae

Alimentary canal: collected West Burleigh, Qld

Capillaria grallinae Johnston & Mawson, 1945

Trans. R. Soc. S. Aust. 69 (2), p. 247-8

Holotype ¢ V1235, allotype 2 V1236 from Grallina

cyanoleuca

Alimentary canal: collected Tailem Bend, S.A.

Capillaria jaenschi Johnston & Mawson, 1945

Trans. R. Soc. S. Aust. 69 (2), p. 245

Holotype ¢ V1240, allotype 2 V1241 from Phalacro-

corax sulcirostris

Alimentary canal: collecied Tailem Bend, S.A.

Capillaria latridopsis Johnston & Mawson, 1945

Trans. R. Soc. S. Aust. 69 (2), p. 245

Holotype $ V1227, allotype ? V1228 from Latridopsis

forsteri

Alimentary canal: Kangaroo Is., S.A.

Capillaria lepidopodis Johnston & Mawson, 1944

Trans. R. Soc. S. Aust. 68 (1), p. 66

Holotype 2 V1229 from Lepidopus caudatus

Alimentary canal: collected St Vincents Gulf, S.A.

Capillaria miniopterae Thomas, 1959

Trans. R. Soc. S. Aust. 82, p. 151-2

Holotype 4 V2809, allotype 2 V2810 from Miniopterus

blepotis

Alimentary canal: collected Canungra, Qld

Paratypes AHC3155

Note: The description of this species was published

in the only paper in which Mrs Thomas did not

retain use of the name Mawson.

Capillaria murrayensis Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p. 342

Holotype 2 V1387 from Mccullochella macquariensis

Intestine: collected from Tailem Bend, S.A.

Capillaria orectolobi Johnston & Mawson, 1951

Trans. R. Soc. S. Aust. 74 (1), p. 24

Holotype ¢ V2879 from Orectolobus devisi

Alimentary canal: collected Port Willunga, S.A.

Paratypes AHC8662

Capillaria plectroplites Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p. 341-2

Holotype 4, allotype 2? V1386 from Plectroplites

ambiguus

Gills: collected Swan Reach, S.A.

Capillaria pomatostomi Johnston & Mawson, 1945

Trans. R. Soc. S. Aust. 69 (2), p. 247

Holotype 2 V1239 from Pomatostomus superciliosus

Alimentary canal: collected Tailem Bend, S.A.

409

Capillaria praeputialis Obendorf, 1979

Aust. J. Zool. 27, p. 872-7

Holotype ¢ V806, allotype 2 V807 from Rattus Jascipes

Praeputial glands: collected Gladysdale, Vic.

Paratypes V808-9

Capillaria recurvirostrae Mawson, 1966

Parasitology 56, p. 282

Holotype 3, allotype 2 V1466 from Recurvirostra

novaehollandiae

Duodenum: collected St Kilda, Vic.

Capillaria rhinobati Johnston & Mawson, 1945

Trans. R. Soc. S. Aust. 69 (2), p. 243-4

Holotype ¢ V1232, allotype 2 V1233 from Aptycho-

trema banksii

Alimentary canal: collected Rapid Bay, S.A.

Capillaria rickardi Beveridge & Barker, 1975

J. Helminth. 49, p. 219-220

Holotype ¢ V64, allotype 2 V65 from Antechinus

stuartii

Stomach: collected Powelltown, Vic.

Paratypes V66

Capillaria strigis Johnston & Mawson, 1944

Trans. R. Soc. S. Aust. 68 (1), p. 65-6

Holotype ¢ V1234 from Ninox novaeseelandiae

Alimentary canal: collected Invercargill, N.Z.

Capillaria tandani Johnston & Mawson, 1940

Trans. R. Soc. S. Aust. 64 (2), p. 342

Holotype ? V1388 from Tandanus tandanus

Intestine: collected Tailem Bend, S.A.

Capillaria thomascameroni Mawson, 1969

J. Fish. Res. Board Canada, 26, p. 1104-5

Holotype ¢, allotype 2 V1509 from Larus novaehol-

landiae

Oesophagus, proventiculus: collected West Is., S.A.

ACKNOWLEDGEMENTS

I would like to thank Mrs P. M. Thomas (Mawson)

and Dr I. Beveridge without whose help and advice

this list could not have been prepared. This work was

supported by an ABRS grant.

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ANGEL, L. M. and MAWSON, P. M. 1969. Helminths from

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BAIN, O. and CHABAUD, A. G. 1979. Sur les Muspiceidae

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INDEX OF GENERIC NAMES

Abbreviaia, Physalopteroidea 396

Acuaria, Acuarioidea 386, 387

Adelonema; Oxyuroidea 396

Alainchabaudia; Spiruroidea 397

Aleuris, Oxyuroidea 395

Alocostoma, Strongyloidea 400

Amidosiomum, Tnchostrongyluidea 405

Amphibiaphilus; Trichostrongyloidea 405

Amplicaecum: Ascaridaidea 388

Amphicephaloides, Trichostrongyloidea 405

Anatrichosoma; Trichuroidea 408

Angullicala; Dracunculoidea 390

Aaisakis, Ascaridoidea 388

Aniechinostrongy lus, Metasirongyloidea 394

Aplectana, Cosmocercoidea 389

Apracta, Aproctoidea 387, 388

Arundelia; Strongyloidea 40!

Ascarophis, Habranematoidea 392

Asymmetracantha, Trichostrongyloidea 405

Austrofilaria, Aproctoidea 388

Austraheligmonema, Trichostrongyloidea 405

Austrostroneylus, Trichostrangyloidea 405, 406

dustroxyuris, Oxyuroidea 395

Bancrafiinema; Physalopteroidea 396

Beveridgea; Strongyloidea 398

Beveridgiella, Trichostrongyloidea 406, 407

Sretnlia (Breinlia); Filarwoidea 390, 391

Bretnlia (Johnstonema), Filarioidea 390, 391

Buccastrongylus: Strongyloidea 398

Capillaria, Trichuroidea 408, 409

Cardiafilaria; Filariodesa 390

Carinema; Filarioidea 390

Cassunema, Strongyloidea 398

Chetlonematadunt, Acuarioidea 387

Chevreuxia; Acuarividea 387

Cloacina; Strangyloidea 398, 399, 400

Comtracaceunt, Ascaridcidea 388

Copemania, Trichostrongyloidea 406

Corollostrangylus;, Strongyloidea 400

Coronastrangylus, Strongyloidea 400

Cosmoucephalus, Acuarioidea 387

Cosmocerca, Cosmocercaidea 389

Crasstcauda; Habronematoidea 392

Cucullanellus; Seuratoidea 397

Cucullanus, Seuratoidea 397

Cyathospiruya, Spiruroidea 397

Cyelostrongylus, Strongylaidea 400

Cyrnea; Habronematoidea 392

Dessetostrongylus, Trichostrongyloidea 406

Dichelyne (Cucullanellus). Souratoidea 397

Diomedenema, Aproctoidea 388

Diplotriaena, Diplotriaenoidea 389, 390

Dipetalonema; Filanoidea 390, 391

Pimetalanema (Chenofilaria), Filarioidea 391

Dispharynx, Acuarioidea 387

Durikainema; Muspiceoidea 395

Echinonema, Seuratoidea 397

Echinuria, Acuanaidea 387

Eustrongylides, Dioctophymatoidea 389

Excisa, Habranematoidea 392

Faleaustra, Cosmocercoidea 389

Filarinema, Trichostrongyloidea 406

Filaroides (Filaroides), Metastrongyloidea 394

Filaroides (Parafilaraides), Metastrongyloidea 395

Foliastoma;, Strongyloidea 400

CGeopetitia, Habronematoidea 392

Globocephaloides; Trichostrongyloidea 406

Goezia, Ascandoidea 388

Gongylonema; Spiruroidea 397, 398

Hahronema, Habronematoidea 392, 393

Hamatospiculunt, Diplotriaenoidea 390

Harveyjohastonia; Chromadoridae 386

Hedruris, Habronematoidea 393

Heligmonoides, Trichostrongyloidea 406

Hepatojarakus, Trichostrongyloidea 406

Heterakis, Heteraloidea 394

Hypodontis, Strongyloidea 401

[chthyostrongylus, Trichostrongyloidea 406

Tnglisonema; Seuratoidea 397

Johnstonema:, Filarioidea 391

Jasefilaria; Filarioidea 391

Lahiobulura; Subuluroidea 404, 405

Labiostrongyvlus; Strongyloidea 401

Leipaanema; Subuluroidea 405

Litomosa; Filarioidea 391

Longistriata; Trichostrongyloidea 407

Macdonaldius; Filarioidea 392

Mackerrastronevlus, Trichostrongyloidea, 406, 407

Macropicola; Strangyloidea 401

Macroponema, Strongyloidea 401

Macropostrongyloides; Strongyluidea 401

Macropostrongvlus; Strongyloidea 401

Macropoxyuris; Osyuraidea 395

Mammanidula; Tnchostrongyloidea 407

Maplestonema; Strongyloidva 40\_

Marsupostrongylus; Metastrongyloidea 394, 395

Mastonema; Tnchostrongyloidea 407

Maxvachonia; Cosmocercuidea 389

Metathelazia; Thelazioidea 405

Metoncholaimus; Enoplidae 386

Microhadjelia; Habronematoidea 392

Microtetrameres; Habronematoidea 393

Montlonema; Strongyloidea 40}

Nicollina; Trichostrongyloidea 407

Nippostrongylus; Tnchostrangyloidea 405

Odilia: Trichostrongyloidea 406, 407

Ovsophagonasres; Strongyloidea 401

Oesuphagostomoides; Strangyloidea 402

Ophidascaris; Ascaridoidea 388 | ;

Oswaldocruzia; Trichostrongyloidea 4007

Oxypspirura; Thelazoidea 405

Oxyuris; Oxyuroidea 395

Papillostrongylus; Strongyloidea 402

Parajilaraides; Metastrangyloidea 395

Paraheteroiyphlum; Ascaridoidea 388

Paralemdana; Filatioidea 391

Paraleptus; Physalopteroidea 396

Paramacropastrongylus; Strongyloidea 402

Paranisakis; Ascaridoidea 388

Paraphyarvngostrongylus,; Strangyloidea 402

Pararugopharvnx; Strongyloides 402

Faraseuraluny, Seuratoidea 397

Parastrongyloides, Strongyloididae 404

Parathelandros; Oxyuroidea 394

Pardustrostrongylus; Trichostrangyloidea 40%

Paraustroxyuris; Oxyuroidea 395

Parazoniolaimus; Strongyloidea 402

Paryseria; Acuarioidea 387

Passalurus; Oxyuroidea 395

Patricialina; Trichostrongyloidea 407, 408

Peramelistrongylus; Trichostrongyloidea 408

Pharyngodon; Oxyuroidea 395

Pharyngostrongylus, Strongyloidea 402, 403

Phascolostrangylus; Strongyloidea 403

Philometra; Dracunculoidea 390

Phocascaris; Ascaridoidea 388

Physaloptera; Physalopteroidea 396

Piratuba; Filarioidea 392

Pontonema; Enoplidae 386

Popovasirongylus; Strongyloidea 403

Porracaecum; Ascaridoidea 389

Poterostrongvius; Strongyloides 403

Potaraxyuris,; Oxyuroidea 395

Provumallanus; Camallanoidea 389

Pracyrnea; Habronematoidea 392

Profilarinema; Trichostrongyloidea 408

Proleptus; Physalopteroidea 396

FProtospirura; Spiruroidea 398

Pseudaprocta; Aproctoidea 388

Rhahdias; Rhabditoidea 396

Rhabdochona: Thelazoidea 405

Rictularia, Rietularoidea 397

Rictularina; Rictularoidea 397

Riouxgolvania; Muspiceoidea 395

Rugopharvax; Strongyloidea 403

Saurofilaria; Filariaidea 392

Seuratia; Acuanoidea 387

Seuratinema; Seuratoidea 397

Seurechina; Seuratoidea 397

Skrjabinodon; Oxyuroidea 395

Skrjabinoptera; Physalopteroidea 396

Skejabinura; Seuratoidea 397

Spinilectus; Habronematoidea 394

Sptronoura; Cosmocercoidea 389

Spirostrangyvlus; Strongyloidea 403

Sprattellus; Trichostrongyloidea 408

Sprattia; Filarioidea 391

Stammerinema; Acuarioidea 387

Steineria; Monhysteridae 386

Skegophorus, Acuarioidea 387

Stellocaronema; Habronematoidea 394

Stomachus; Ascaridoidea 389

Strongyloides; Strongyloididae 404

Subulura; Subuluroidea 405

Symplocostomella; Enoplidae 386

Synhimantus; Acuarioidea 387

Synhimantus (Dispharynx); Acuarioidea 387

Syphacia; Oxyuroidea 396

Tanqua; Gnathostomatoidea 392

Terranova; Ascaridoidea 389

Tetrabothriostrongylus; Trichostrongyloidea 40%

Tetrameres; Habronematoidea 393, 394

Thelandros; Oxyuroidea 396

Thelazia; Thelazoidea 405

Theracostoma,; Enoplidae 386

Thylonema; Strongyloidea 403

Thylostrangvlus; Strongyloidea 404

Trichostrongylus; Trichostrongyloidea 408

Trigonostonenia; Strongyloidea 404

Uneinaria; Ancylostomatoidea 387

Vagrifilaria; Filarioidea 392

Vieutera; Habronematoidea 394

Woodwardostrongylus; Strongyloidea 404

Woalleya; Trichostrongyloidea 405, 408

Zoniolaimus; Strongyloidea 404

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK

SYSTEM, NORTHERN TERRITORY

BY MICHAEL H. TYLER, GRAEME A. CROOK AND MARGARET DAVIES

Summary

The Magela Creek system occupies the eastern fringe of the Northern Territory floodplain, and the

adjacent escarpment country. Twenty-four species of frogs occupy this area.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM,

NORTHERN TERRITORY

MICHAEL J. TYLER, GRAEME A, CROOK AND MARGARET DAVIES

Department of Zoology, University of Adelaide, Box 498 G.P.O., Adelaide, South Australia 5001.

ABSTRACT

TYLER, M, J,, CROOK, G, A. & DAVUES, M , 1983. Reproducuve

biology of the frogs of the Magela Creek System, Northern

Termitory. Ree. S. dust. Mus. 18 (18); 415-440,

The Magela Creck system occupies the eastern fringe

of the Nerthern Territory floodplain, and the adjacent

escarpment country. Twenty-four species of frogs

occupy this area,

We have examined facets of the reproductive biology

of the frog fauna over two entire wet seasons. Data

are recorded on the onset, duration and termination

of activity, calling and breeding, and are correlated

with temperature and relative humidity. Most species

breed ai the onset of the wet season before the flood-

plain is completely inundated.

For each species the eggs and form of the spawn

clump are described. Developmental history is derived

{rom laboratory observations on spawn transported

to Adelaide, Tadpoles are described and illustrated at

various stages of their development, and growth curves

are plotied.

Details of observations on activity are recorded.

Several species bask in the sun and attain body tem-

peratures of up to 36,4°C,

Fecundity and reproductive modes are platted. We

note limited reproductive diversity and high selective

advantage in abbreviation of the developmental span.

Introduction

Very few studies have been undertaken in Australia

on diverse anuran communities, With the exceptions

of the unpublished observations. of Straughan (1966)

in eastern Queensland, and Humphries’ (1979) study

near Canberra, A.C.T., emphasis to date has centred

upon interaction between particular congeneric com-

ponents within communities,

With the support of the Supervising Scientist for

the Alligator Rivers Region we undertook a study of

the frog fauna of a portion of the floodplains and

adjacent escarpment in the Northern Territory from

October 1978 to April 1979 and October 1979 to

March 1980. (Tyler and Crook, 1980),

The principal objectives of the study were to deter-

mine which species were likely to colonise the uranium

mining, sites there (in particular the retention ponds

and tailings dams) and, further, the nature of skeletal

abnormalities in the populations. Such data would

permit an assessment to be made of the value of frogs

as indicator organisms for monitoring the long-term

impact of uranium mining,

Atihe commencement of the study six of the species

now known to occur at the site were undescribed,

and so our early efforts concentrated upon establishing

ihe composition of the fauna, Similarly the spawn

and tadpoles of 21 of the 24 species were unknown,

In consequence we were obliged to concentrate activ-

ities upon fundamental descriptive studies of life his-

tories, and it was ocither possible nor a component

of the contract to examine community structures in

any detail,

Because the study site included floodplain and

escarpment, it embraced the two exiremes of habitat

that occur in the northern portion of the Northern

Terntory, and potentially at least appeared to permit

extrapolation of results 1o more distant areas.

The study area experiences striking climatic

extremes with marked winter dry seasons, and summer

wet seasons. Here we describe the reproductive habiis

of the frog fauna, describe the embryonic and larval

morphology and examine the reproductive regimes in

an area where most species have a limited breeding

season,

Description of Study Site

Magela Creek is a tributary of the East Alligator

River, Northern Territory, located approximately 230

km east of Darwin (Fig, 1). Magela Creek drains the

western periphery of the Armhemland Plateau, and

the adjacent, low-lying floodplain, and flows only in

the wet season. The term “Magela Creek system’ is

used 10 embrace the drainage area, which represents

an interface of ecologically divergent zones.

Our studies were centred at the Ranger uranium

mining site at Jabiru located at 24.5 m above sea

level. The majority of our observations were made

within a 30-km radius, with the extreme of that range

being north towards the East Alligator River (Fig, |),

416

MELVILLE

BATHURST

ISLAND

REC. 8. AUST. MUS., 18 (18): 415-440

elpril, LON

@ OENPELL!

CANNON HILL

FIG. 1. Study site within the Magela Creek System, Northern

Terntory.

The soils are diverse, but their significance as a

factor influencing frog distribution is associated with

porosity, and the extent to which they can be pene-

trated by the fossorial species,

There is a lack of uniformity in the nomenclature

and delineation of plant communities in the area

surrounding Magela Creek (Christian & Aldrick, 1977;

Williams, 1979). The system that we have adopted is

applicable to frog communities, and involves the fol-

lowing seven units, each of which is meaningful in

an ecological sense, and significant in the diversity

and chronological sequence of breeding sites occurring

there, Data on dominant plant species within the

units are from Christian & Aldrick (1977),

1. Sandstone scrub and woodland in escarpment

The breeding sites on the escarpment vary from

pools on the open rock face (Fig. 2A) to streams

through deep crevices in the sandstone. Tadpoles have

been found in shallow depressions on exposed rock

faces, where the water temperatures reach up to 45°C,

and where there is little aquatic algal growth. The

pools fill with rainwater early in the wet season, and

are then used as breeding sites. Further storms cause

overflowing, and water runs over the rocks and into

small, scrub-lined streams which flow only during

these storms, For the remainder of the time the stream

beds are a series of shallow, isolated pools among

sandstone rocks with spinifex, sedges and wiry grasses

along the banks. The streams extend into deeper,

sandy-floored streams (eroded from the sandstone),

which persist for much longer after the wet season

has ended. The narrow gorges created by these streams

are often very deep and have vertical faces, There is

low light intensity and the water is cold (<10°C).

The larger streams support a woodland including

Eucalyptus dichromophloia, E. phoenicea, and E.

miniata, smaller trees Owenia, Xanthostemon and

Erythrophloeum with wiry grasses and spinifex in the

understorey,

2, Sandstone rainforest remnants

These relict forests differ from the woodlands

through the domination of an undescribed myrtaceous

evergreen tree species up to 22m high. Other non-

eucalypt trees are present; the understorey is sparse,

and the leaf litter is thick and moist. Such forests

occur in the broken terrain or sandstone plateau and

escarpment, usually in narrow gorges at the base of

the escarpment, The streams usually have defined

channels with rocky bottoms, and are permanent (Fig.

2B). Frogs utilize the pools in small side streams

which flow only after storms.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

FIG, 2. Habitat types: (a) Pool on rockface in sandstone escarp-

ment, Birndu; (6) Remnant rainforest, Radon Creek: (c) Open

sclerophyll forest at foot of escarpment, Birndu; (d) Inundated

grassland.

3. Open sclerophyll forest

The open forest is dominated by Eucalyptus tetro-

donta and EF. miniata with the occasional Erythro-

phloeum. The understorey is varied including

Livistona, Xanthostemon, Planchonia, Buchanania,

Grevillea, Melaleuca and Acacia. Annual and perennial

grasses are also present (Fig. 2C). In the wet season

shallow depressions in this woodland provide breeding

sites for several frog species. Heavy storms inundate

the whole forest floor, so creating many microhabitats

in which frogs abound.

4. Inundated grasslands

In the open sclerophyll forest and along the creeks

are large areas predominated by annual and perennial

grasses (Fig. 2D), These areas have sandy sail, and

any depressions fill with water early in the wet season

providing breeding sites for several frog species. Inun-

dation of these areas occurs once the wet season has

set in (usually in early January). More frog species

then utilize the areas of shallow water outside the

deeper depressions.

5. Fringes of billabongs

The main body of water of the billabongs is not a

significant habitat for frogs. Of greater significance

are the pools and grassy areas on the fringes among

the Melaleuca and Pandanus (Fig. 3A). These areas

fill with water early in the wet season and predatory

fishes are excluded for long enough for tadpoles to

complete metamorphosis.

6. Floodplains

These are large expanses of sedgeland which are

inundated for two to six months of the year (Fig. 3B).

The principal plant genera there are Cyperus, Scirpus,

Eleocharis, Fairena and Sesbania with some prasses.

A wide variety of herbaceous plants and water lilies

occur in areas where there is inundation for six to

eight months, The frogs usually inhabit the edges of

the floodplains, and occupy small islands of rock and

vegetation when the floodplains are inundated.

7, Artificial pools, borrow pits, ponds etc,

Buffalo wallows are important breeding sites on the

floodplains and inundated grasslands in the early

months of the wet season. Man-made borrow pits

beside the roads are important throughout the wet

season, providing shallow temporary pools in areas

where such pools would not occur naturally (Tyler.

1979). The soil is gravelly and the vegetation is sparse.

418 REC. S. AUST. MUS., 18 (18): 415-440 April, 1983

FIG. 3. Habitat types: (a) Billabong fringe; (6) Floodplain from

A air (c) iis airstrip with temporary pools; (d) Retention

‘ond No. 2.

The large cleared areas around the pools provide a During the study period there was substantial alter-

larger number of potential calling sites for males than ation to the structure of the site and hence to the

the surrounding open forest. Dams, drainage ditches area in which we undertook observations. Principal

and even pools on the road surface are used as breed- changes affecting frog breeding sites were the comple-

ing sites by some species (Fig. 3C). Indeed any dis- tion of construction and the filling of the retention

turbance of the environment causing the formation ponds and the tailings dam, so increasing the area of

of pools increases the breeding potential of frogs in static water by more than | km?, and the grading of

the area, and tailings dams, retention ponds, siltation the airstrip which eliminated numerous shallow pools.

pools and other earthworks have to be considered in 36

a similar sense (Fig. 3D). MEAN MONTHLY MAXIMA

The rainfall follows a distinct seasonal pattern. There

is a wet season generally heralded in late October 354

with occasional showers in the late afternoon or eve-

ning, but the greatest volume of rain occurs from

December to March. The maximum rainfall recorded

at Jabiru Airstrip in one month since 1971 is

767.9 mm in February 1980. Annual totals there range

from 1121 mm to 2282 mm. As indicated above,

during the study the 1979-80 period was atypical

because there was no rain from June to September, 28

followed by a late onset of the wet season and then 1 My aac.

the record February falls, leading ultimately to the

second highest annual total.

30-

TEMPERATURE (°C)

MEAN MONTHLY MINIMA

Shade temperature maxima and minima were

39.9°C in October 1979 and 14.1°C in August 1979. — . ; .

Mean monthly maxima and minima at Jabiru Airstrip Qe ROE FE i Wh eR IN tad

during the study period are depicted in Figure 4.

FIG. 4. Temperature variation at Jabiru during the study period.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

Methods

Ficld observations were undertaken during the wet

seasons by G. A. Crook, He resided at Jabiru fram

October 1978 to April 1979, and from October 1979

to March 1980. M. J, Tyler visited the study area in

Navember-December 1977, November 1978, February

1979, October 1979, November-December 1979, and

February 1980. Supplementary field assistance was

provided on various occasions by M. Davies, A. A,

Martin and J. Brooks,

Initially the field programme concentrated upon

establishing the nature of the fauna, leading to the

collection of 24 specics, of Which six were hitherto

undescribed, Biological data recorded included surtace

activity, calling activity of males, spawning, the phys-

ical form of spawn clumps, and the physical appear-

ance of the tadpoles. Water temperatures at brecding

sites were recorded with a Wesco rapid-reading ther-

mometer (reading 0-50°C) and relative humidity

measured with a Zeal Whirling Psychrometer,

Spawn and tadpoles were obtained at night and

Iransported to Adelaide by air in insulated containers.

Upon arrival at the University of Adelaide the spec-

imens were transferred to 4 constant temperature lab-

oratory maintained at 30° +0.5°C. Spawn and small

quantities of tadpoles were transferred to white

‘Advance’ plastic containers of IL capacity. Denser

aggregations were maintained in 5L capacity all-glass

aquaria. The cultures were aerated sufficiently to cle-

vate oxygen levels, without reducing the water tem-

perature.

Water was changed daily. The water used was passed

through a commercial softener and filter, followed by

passage through an activated carbon dechlorinator

and a bacterial filter’ Water was allowed to stand in

the plastic containers or glass aquaria for 24 hours

prior to use to equilibrate to room temperature. The

light regime of the room was controlled and seasonally

adjusted 10 that at Jabiru.

Tadpoles were fed on washed and boiled lettuce

leaves, Tadpoles were sarnpled regularly and preserved

in Tyler’s (1962) tadpole fixative. Selection of speci-

mens for preservation followed ihe attainment of the

progressive developmental stages of Gosner (1960),

Subsequently specimens were measured with a micro-

scope eyepicce micrometer or dial callipers to an

accuracy of 0.01 mm.

A representative collection of the species studied

has been deposited in the South Australian Museum.

410

IDENTIFICATION OF SPECIES

Species Composition

Three families of frogs occur in the Magela Creek

sysiem. They represent the Hylidae (2 genera, 16

species), Leptodactylidae (5 genera, 7 species) and

Microhylidae (1 genus, 1 species): a total of 24 species.

Of that total only 1wo species: Linria personata Tyler,

Davies & Martin and Upero/eia arenicola Tyler, Dav-

ies & Martin are currently known only from that

area. Most of the remainder are widely distributed

across northern Australia,

Distribution and Abundance

In Table | we have plotted the species com position

of the fauna, together with our assessment of relative

abundance based on the frequency with which rep-

resentatives were encountered, ‘Absent’ indicates our

failure to observe or hear a species in a particular

area during the entire study; ‘Rare’ includes 1-10

observations; ‘Common’ involves at least 25 sightings,

whereas ‘Abundant’ indicates the presence of a species

in large numbers on any night within the period that

we found the species to be active.

Identification

To permit identification in the field we have pre-

pared dichotomous keys to spawn, tadpoles and adults.

To assist the identification procedures we have divided

the total fauna into two groups—the species found

on ihe escarpment, and those found elsewhere (which

we term ‘lowland’ for want of a more appropriate all-

embracing word). Within the ‘escarpment’ category

we have included spectes such as Limnodnastes

ornatus and L. conyexiusculus, which are in no way

adapted to that environment, but occasionally they

penetrate its fringe.

Excellent colour photographs of many of the species

appear in handbooks of the frogs of Australia by

Barker and Grigg (1977) and by Cogger (1979). How-

ever, because there are so many additional species

not included in those publications, and confusion

about the use of some names, we recommend that

their photographs be used to confirm identifications

reached here, rather than as primary identification

sources,

(a) KEY TO FROG EGGS FOUND LPON THE ESCARPMENT

|. Spawn terrestrial, laid in leaf litter. Sphenaphryne rabusta

Spawn laid in water... , . A Pak — * 2

2. Spawn in form of foam nest me = - 3

Spawn not in form of foam nest... — are = \

3, Eggs slightly pigmented, or unpigmented, foam nest deposited

in concealed sites; foam persists for several days _ 4

Eggs black with white vegetal pole, foam nest floating on

surface of puols; foam nest collapses within six

hours Limnodynastes ornattts

$20

REC. 8. AUST. MUS.,

18 (18); 415-440

April, 198s

TABLE 1; DISTRIBUTION AND ABUNDANCE OF THE FROG FAUNA OF THE MAGELA CREEK AREA,

2 a % z

& = Fe)

33 Og 2 i 5

as ‘ 4 = g wes

az a ee gs

0 Absent us 2 Bb S = a 2 g

| Rare 22 v4 z s ap ‘a ae

2 Common ES Bg = x 8 = 38

3 Abundant aa Se z 5 = 3 e.%

as 2 & = a = S ts

“we we = ° baa] c <6.

Litaria biedler ..... 7 0 0 2 0 3 2 0

EB. faerulea (000 6 ee ee cee et Fe ee UHH et ' 2 2 1 ! ! ] 0)

L. ooplandl 0.2 ce ne eee eee tee 3 2 0 Q 0) 0 0

L dahlit.. 0 0 i 0 2 3 |

CORRES SS nip og see eh melons 7 ee ee 0 0 | 0) 2 0) 3

Lo sp. ar. latopalmata —- te itl stemtch wales 5 Mtn eh eee 0 0 3 0 2 | 3

L. mweiriand 2... 3 3 0 0 Q) 0 0)

L. wicrobelas 2s 2 e e eb Er ete ee Q 0 i} 0 3 2 0

ey eo Ee a Pe ee 0 0 3 2 2 0 1

L. personatd oy oe eens q 0 0 0) i) 0 0

L. rothit 0 0 2 | 2 2 2

Lube: nck bee e's Fete Ss SS - 0 0 2 | 2 2 3

Ro farrtiets 6b ice ose 5} bog eters sia 0 0 3 0) 2 1 3

L. wotjulumie FAB YS — (ei rele wees PORTS Perse ye \ 2 2 l 0 0 0 2

('yclorana australis . : 0 0 2 i 2 2 3

(, IaMBIPeS - bison seas: -- ia) () 1 0 ! 1 3

Limaody’ nastes convexiusculus , 1 0 2 0 2 3 0

L ormatus...... rare & Aiewil RAPS AD 2 0 2 l 0 0 3

Notaden melanoscaphus rw Re Ala eee o- 0 0 3 3 0 0 Q

Mewistolous lignarius ...... 1 lla tclamal et 2 2 0) 0 0 0 Q

Rantdellu bilingud 2.64 0 te ee ee 0 0 3 1 2 2 0

Upernleia arenicola , TE ok pertain Se SR he 1 0 0 0 0 Q) Oo

Uperoleta inundata 9 >: wf see olla rie we totes 0 0 3 2 0 0 0

Sphenophryne robust oy cpp 2 eee ete 0 2 0 0 2 0) 0

_ Spawn floating on surface

. Dark ‘brown;

_ Foam nest under boulders in streambeds or pools, eggs creamy

yellow ...., . Megistoalotis lignartus

Foam nest in small hollows among grass tussocks; eggs

white... _ Limnodynastes convextusculus

. Large number of! browrypaiite eggs Jaid as surface

film... a Litoria caerulea

Eggs laid singly 01 or in small clumps ie pee: ee oe |

L, wot/ulumensis

Spawn below surface of water rate

. Spawn laid as small clumps or film firmly attached to sandstone

rock face _L. metriana

Spawn laid as single cpgs or in small clumps on bottom of

rock pools. ; _ L. eoplandi

The spawn of Uperoleia arenicoln and Litoria personata 1s

unknown,

(hb) KEY TO TADPOLES FOUND UPON THE ESCARPMENT

| With distinctive gold or yellow dorso-lateral stripes

. 2, pecsnnam

Lacking gold ¢ or yellow dorso-lateral stripes » : cow

. Body flattened dorso-laterally, eyes dorsal, narrowly stivarated:

Papillary border widely separated superiorly

Limnodynasles ofnatus

Body not flauened dorso-laterally, eyes dorso-lateral -. 3

. Black colour’ , Poe FNS) te, Ses siotd oF

Brown colour Pee Leo

Papillary border narrowly separated titpedionti sari dapillac;

4/3, 5/3, 6/3 labial tooth rows; strong horny

beak... ween _. Megistolotis lignarius

Papillary border widely separated superiorly; large papillae;

5/3 labial tooth rows; horny beak weakly developed

. Limnodynastes convexiusculus

conspicuous white patch near base of

abdomen , : Litoria coplandi

Lacking white patch near hase of abdomen Pe 6

. Papillary border widely separated superiorly; horny beak rather

weakly developed. Tail fin lightly pigmented; tail muscle

pigmentation in two stripes: one lateral halfway along tail,

one along dorsal edge - Litoria caerulea

Horny beak strongly developed. Upper tail fin pigmented.

Tail muscle heavily pigmented 7

7. Tadpoles small (<26 mm); papillary border siarnowly separated

superiorly, Ventral surface dark, becoming lighter. Dark

brown stripe along dorsal edge of tail muscle

1h L. meiriana

Tadpoles ‘medium-sized (<45 mm); papillary border separated

superiorly. Ventral surface light from stage 26

Mase. - : L. wotjulumensis

It is assumed that Sphenophryne robusta has no free-swimming

tadpole.

_ Green or olive; large (up to 110 mm); expanded discs on toes,

conspicuous toe webbing . Litoria caerulea

Not green or olive .

. Tympanum not visible

. Very small (up to 25 mm length)

Size within range of 25-75 mm. — .. ath 5

. Brown with dark stripe on side of head sometimes extending

along sides; cylindrical toe discs. Found in leaf litter

only - ne Sphenophryne robusta

Brown to slate grey with orange parotoid glands; no toe discs.

Metatarsal tubercle enlarged. Found in sand

streambeds .... . Uperdleta arenicola

. Inner metatarsal tubercle enlarged; usually sandy with darker

back patterns, rounded snout; <45 mm in length. Slight

web on toes ._-. _ Limnodynastes ornatis

{nner metatarsal tubercle not enlarged: grey or brown with

dark mottling on dorsum, slightly elongate snout, up to

65 mm in tenets Conspicuous cream supralabial gland, No

webbing, - : _ L. convexiusculus

. Robust; large, prominent, oval inner, but no outer metatarsal

tubercle. Toe discs absent. Hindlimbs short; males have

black nuptial spines. . .. Megistolotis fignartus

Not robust; no large, prominent, oval: inner metatarsal tubercle,

Toe discs present. Long hindlimbs -. 200 )..5)....,,7

. Brown, ovidiameter |.2

» Ovum diameter |.6-|.9 aim .

. In small clumps (of <

. Very

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

- Feet with trace of webbing. Small (<35 mm). Conspicuous

head stripe. Toe dise expanded - Lituria personata

Hindlimbs with conspicuous webbing. ..--. _., &

. Conspicuous head sinipe finishing midway along body, toe

discs small. Large se (up to 70 mm SV length)

L. wotjlumensis

Toe dings well developed. No head ‘stripe _ a oe:

. Small (<25 mm), diurnal and nocturnal. Heavy mottled dark

brown band on legs, dorsal pattern . L. meiriana

Up to 40 mm, nocturnal, Mottled dorsal pattern, large inner

metatarsal tubercle ee wi L. caplandi

fd) KEY TO LOWLAND FROG EGGS

| Terrestrial, found an leaf litter in Melaleuce swamps

ee aL be . Sphenaphryne robusta

Deposited in water-_ . SHU Sr Loko ee sot UP ene

. Deposited in foam nest . - ge re

Not deposited in foam nest _- f.- E. aB

. Barely pigmented: foam nest in cbaenled ; areas in inundated

grassland; foam persists for several days

Limurodvnastes vonvextusculus

Black with white vegetal pole; foam nest conspicuous, floaling

on surface of pools; foam nest collapses within six hours

. 1. ernatus

. In large loosely anached tiurnps on bottom of pools - 5

Not in large loosely atiached clumps. --—- poe

-1,5 mm; capsule diameter 1,9-2.5 mm

. Cvelorana longipes

Brown, ovidiameter I. 3-1, bn mm; = capsule diameter 2 2.5-2.2 mm

ee C. australis

. Laid Siriely ot orm small efutaps on bottom of pools in inundated

ermassland..,, _. Uperaleia inundata

Not taid singly or in ‘small clumps on bottom of pools in

inundated grassland - to. Fhe oO. @

. Laid as free-floating clumps. se 4:

Nat laid as free-floating clumps 10

. Capsule large (6.1 mm); brown and white egg; three jelly

envelopes Litoria (nermns

Capsule medium (4. 5-5. 4 mm); ‘dark brown and white egg 9

_ L. wothlumensis

Ovum diameter 1.3-1.4 mim = L. rothii

= 20 epps) attached lo yegelauion li

Not in small clumps fof < 20 eggs) attached wo

vepetalion 6. uy, e -- AS

small ovum (0,8-0,9 fairl, Broa and cream

f..5-.3 ie Pe ae _ L. micrabelos

Small (1.0-|.2 mm) dark brown and cream .--,..,-_,.12

- Ovum diameter 1.0 mm; capsule 2,5 mm (1.9-2,5 mm); three

jelly envelopes - L. hicular

Ovum diameter |,2 mm; capsule dianeter 1.7 mm; one jelly

envelope Ranidella hilingua

Surface film in shallow water (<5 em) in inundated grassland,

Black animal pole, white vegetal. pole; discrete jelly envelope

around ovum with loose sheet of jelly enclosing all

eges ., Notaden melanascaphuy

Surface film in ephemeral o ar temporary Bane, Animal pole

not black. Discrete jelly capsules - _- 7 14

. Dark brown foe ce B 15

Pale brown ,, 14... LL wid lt

. Surface film of 200-350 eggs, capsule 2.2-2.9 mm

..., Literta caertilea

Surface film or clump; =< 100 eggs laid: capaule 3.2-3.7 mm

416

. Ovum dhimetér 1,041 3 min; thee "jelly envelopes

J, nasuta

Ovum diameter 1.4 mm; two jelly envelopes L. forniert

- Ovum diameter 1.0-1,1 mm; capsule 1.9-2.3 mm

L. rubelle

Ovum diameter |.2~!.3 mm; capsile 3.4-3.9 mm

L. sp. nr. datepaltnata

. Black,

, Tooth raws vary 2/3 to 6/3:

. At

. Tail muscle pigmentation not in broad bands,

(2) KEY TO LOWLAND TADPOLES

to 70 mm), 5/3 labial tooth rows

a. . Limnddynastes cunverctuyculyy

Not black . og et ee 2

eyes dorsal, narrowly separated:

body dorso-ventrally flattened . L, ornelus

Tooth rows always 2/3; eyes dorso-lateral,—.. 4

large ne

i Large (up to 70 mm). Bulbous bodies, usually dense and active

in aggregations —. : fpes ts 14 4

Medium-sized (up ta §0 mm). Not in dense and acrive:

ageregalions ; 6

. Lacking nasal groove, ‘Dorsal patlern strong “mouling, tail

muscle with broad lateral stripe and along dorsal edge,

Stnped back pattern develops in stage 41. Papillary barder

widely separated superiorly Litaria dahl

Nasal groove in stages 32-41% tail THuscle mottled; papillary

border not widely separated superiorly. 5

stage 41 back pattern with well-defined dark

patches Cyclorana longipes

At stage 41 back lacking dark patches: ansaiy rows of

smal] glands present F . aayiralis

Papillary border separated inferiorly 7

Papillary border not separated infenorly 8

Papillae large in lateral area. Absent superiorly and just nwo

inferiorly, flanking short third labial tooth row; homy beak

strong, ventral surface while. Small (<25 mm)

Ranidella bilineua

Papillae large, widely separated superiorly, separated inferiorly;

horny beak weakly developed: ventral surface motiled red-

purple and white. Smal! (<34 mm)

Uperoleta inundatu

. Third lower row of labial teeth reduced to median structore

of approx, 10 teeth |, _ # 4

Third lower row of labial teeth ‘not reduced . 10

. Papillae large, border not well<leveloped and widely separated

superiorly, Horny beak very weakly developed. Tadpoles

< 38 mm in length, Dark brown dorsum, Ventral surface

dark brown becoming lighter. Very shallow water in inun-

dated grassland wee... Notaden melanoscaphus

Pagallae large, border well-developed and separated superiorly,

Horny beak well-developed. Tadpoles up to 58 mm long:

early stages pale yellow dorsum with dark brown dorsal

and dorso-lateral stripes: during later development (after

27) the stripes fade and the dorsal and lateral surfaces

become marled brawn with the pale yellow background

Dorsal surface of head becomes flattened, In temporary

pools , Liloria rothii

~ Conspicuous band of white muscle on posterior of abdomen

until stage 38. At 38 white patch develops just pusterior

of cye and develops into broad white stripe from eye to

groin. Another white stripe antedor and belaw eye passing

beneath eye and ends behind shoulder, Tadpole mol

tled brown early, uniform brawn on dorsum

later. . L. hivolar

No conspicuous band of “white muscle on posterior of

abdomen A he ' a nt, lI

<25 mm 12

Tail muscle pigmentation in broad bands, very small (=< 20

mm): horny beak weakly developed. Papillary border widely

separated. Dark dorsal pattern and dark head and lateral

stripe ee — bo ntierobelos

. Papillary border widely separated superiorly; homy beak rather

weakly developed, tail fin lightly pigmented: tail muscle

pigmentation in two stripes: one lateral halfway along tail

one along dorsal edge - L, vaeriuled

Papillae separated — superiorly; horny beak = well

developed — - wT Baote: =f. 13

- Val muscle has three stapes: one lateral, one

on dorsal and on ventral edges. Tail fin edge

becomes darker with deyelopment. Small tadpolis

(<= 35 mm), In stage 4i broad. white lateral stripe and

harrow vertebral stnpe appear White patch anterior to

and below eye at's ac Ly rubettes

422

ios)

LO.

. Inner metatarsal tubercle enlarged. ,

‘Tadpoles mottied brown. Tail muscle mottled brown. Up to

50 mm. Ground-dwelling hylids (L. wolulurmensis, L.

nasuta, tC. torniers, L. sp. nr lalopalmete, L. irlermiis),

Further specific identification is only possible from stage 41

onwards as follaws:

at stage df

(i) Toes Strongly webbed —— ii

Tow webbing not well developed - il

(ii) Stmped back pattern, long hindlimbs with Jonpitudinal

stripes _£. Hasulet

Brown back paltern, broad conspicuous dark head stipe

: L, wotpilumensis

Gil) Cankinvans dark beta stripe on posterior edge of tibia

L. tormer!

No continuous stripe i inermis and L. sp. nr fatopalmata

Specific identification of these species is only possible after

stage 46.

At stage 46

Warty dorsal surface, indistinct head stnpe, some flecking on

the dorsal surface L. inermis

Smooth dorsal skin, distinct head stripe

L. sp. ar /atepalmata

(f) KEY TO LOWLAND FROGS

Inner metatarsal tubercle not enlarged

. Medium to large size (greater than 35 mm), Outer metatareit

tubercle absent of not enlarged . . 4

Small (to 35 mm). Enlarged outer metatarsal tubercle, Stout-

bodied: short stocky limbs; conspicuous parotoid glands

and glands at comer of mouth; grey with highly vanable

dark brown to red dorsal markings. Found in inundated

grassland Uperoleja inundata

. Inner melatarsal tubercle: nor black... .., 4

Inner metatarsal tubercle black. Rounded snout. ‘Stout body,

short robust limbs. Pale brown with dark brown dorsal

markings. When seized, exudes copious quantities of yellow

sticky secretion from Jarge number of dorsal glands. Found

in inundated grassland Notaden melanascaphus

. Belly smooth, Rounded snout, Medium-sized (up 19 45 mm),

Sandy with variable dark dorsal markings. Toes one-third

webbed. 'Tympanum not visible, Conspicuous lateral fringes

on toes, Males have nuptial pads on first two fingers.

Calling occurs while free-floatmg in water

Limnodynastes ornatus

Belly granular. Stlightiy clongated snout. Large size

(50-110 mm), Toes slightly webbed) Tympanum visible,

Male nuptial pads on first finger only. — oecars on

land .... .45, . A ssi

. Medium-sized (to 55 mira “Well-defined ‘dark patches on

dorsal surface, usually with a vertebral stripe

Cyelorana. longipes

Very large (to 110 mm). Pale brown with patehes. A contrasting

dark brown head stripe from. snout to shoulder

a Tr. or C. australts

. Toe discs Conspicuous—expanded to ewice width of

digit eee? el eee ee en

Toe discs inconspicuous 10

. Small (< 30 mm) and green; thin brown stripe along flanks;

white stripe present from nose to groin Litoria biealor

Medium to large size (35-110 mm) se B

. Brown or grey, 35-60 mm snout to vent Jength ere

Very large green or olive oe an yo 110 mm). Common

around dwellings . L. caerulea

. Large (to 60 mm). Extensive 10 webbing, Yellow and black

markings in the groin and posterior of thigh. Elongate

limbs. |, Co ed rothit

Medium (10 45 mm). Webbing. halfway up toes. Broad head

stripe from nose 10 pheaider: somelimes continuing along

flanks. Short, robust limbs . *, L, nibella

Small (< 25 mm) . tino mine cei pen.

Medium to large (=> 30 mm) | 13

RECS, AUST, MUS,,

17.

18 (18); 415-440

. Toes at Icast partly webbed ,

. Toes fully webbed: large (to 85 mm),

. Striped dorsal pattern absent -

April. 1YNd

. Slender body: granular belly . -- 12

Robust body; smooth belly; mottled brown: dark head stripe

occasionally continuing along flanks, toe discs small and

cylindrical, tympanum not visible; no webbing on toes;

found in leaf liter in Melaleuca swamps

Sphenophryne robusta

Upto 25 mm, Brown with variable dark markings on back,

No Loe discs. Unwebbed toes -- _ Ranidella hilingue

Up to 20) mm; brown with darker lateral stmpes. dark eye

stripe From nose 10 midway alang flank; beneatlr this, while

stripe from nose lo shoulder. Toe discs present. Toes slightly

webbed - Litoria micrahelos

; .

‘Yoes unwebbed; large (lo 65 mm); grey or olive with discrete

dark patches on dorsal surface, conspicuous cream supral-

abial gland; stout body; tympanum not visible. Found in

inundated grassland and floodplains

Limnodynastes converiusculus

1B)

Toes less than fully webbed; medium (to 50 mm) 16

. Up to X5mm) green with brown and olive back markings.

cream patches on posterior of thigh, Belly smooth and

white. Eyes prominent and dorsal. Found on flovd plain:

mostly aquatic . . . Litoria dahlit

Up to 70 mm, brown, contrasting broad dark head stnipe

continuing along sides of body, Belly granular and cream.

Eyes dorso-lateral - 4, L, wotjulumensts

Conspicuous brown and white longitudinal stripes on back,

Broad dark head stripe, Streamlined shape with elongated

snout and extremely long legs. Black and yellow markings

on posterior of thigh. Sinipes s and dark patches on legs.

Toes half webbed. _ LL. nasuta

Continuous dark brown stripe on antenor of tibia: red-brown

dorsal surface in breeding season; pale brown la grey, dry

season LL. tarnilert

No continuous tibial snipe brown dorsal surface in breeding

season - .

. Usually indistinct head swipe! warty dorkatii some molting,

thigh markings not prominent, Males call from open areas

usually on steeply sloping banks L. inermis

Distinct head stripe; smooth dorsum; occasional moatiling;

black and yellow markings on posterior of thigh; males call

from open areas among grass lussocks adjacent to water

Th lh eee he a L. sp. nr latwpalmata

ACCOUNT OF SPECIES

Family HYLIDAE

Cyclorana australis (Gray)

The largest frog in the area, C. ausrraliy is robust,

60-100 mm snout Lo vent Jength and a drab grey-

brown. This species is fossorial, spending the day and

dry season underground, usually al the base of trees

or under logs. It is active at night from October to

April, although the number of adults. observed drops

sharply after January, when metamorphosing juveniles

predominate.

Breeding activity: Large numbers of males congregate

early in the wet season. On warm nights dunng and

following rain, they call from areas devoid of vege-

tation or at the base of grass tussocks near water.

Breeding occurs at air temperatures of 24,5-28,5°C,

78.5-96.0% R.A. AL deposition sites the water tem-

perature is 31.0-33.5°C and the water depth 5

-100

cm.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM 423

Eggs: Spawn is deposited at the edge of temporary

pools. Initially it floats on the surface, but it sinks to

the floor during early development. The clump frag-

ments when disturbed. Eggs are laid in clumps of up

to 7 000 although 100-1 000 are most common. The

animal pole is dark brown and the vegetal pole white.

Two jelly envelopes surround the ovum. Mean capsule

diameter is 2.5 mm (2.3-2.8 mm) and mean ovum

diameter 1.6mm (1.5-1.8 mm).

Developmental history: Hatching occurs at stage 17

in 1-2 days. The tadpoles become robust with a rotund

body, elongate snout and deep fins (Fig. 5) and are

pale brown or dull grey. The anus is dextral and the

spiracle sinistral. The tooth row formula is 2/3 and

there is a well-developed, elevated border of papillae

surrounding the oral disc. There is a strong horny

beak. The papillary border is separated superiorly by

a broad median gap. A naso-orbital groove is present

at stages 32-41. Stage 37 of Gosner (1960) is inappl-

icable because the metatarsal tubercle appears con-

currently with differentiation of stages 37-38. Tadpoles

attain a length of almost 70 mm at stage 41 (see Fig.

12) [mean 54.8 mm (40.9-68.2)].

Spawn of C. australis and C. longipes are often

mixed, and the tadpoles form dense and very active

aggregations. Tadpoles constantly rise to the surface,

gulp air and then submerge, so producing the impres-

sion of boiling water. The tadpoles prefer the slightly

cooler, deeper water. but low oxygen saturation levels

induce them to rise to the surface.

Metamorphosing juveniles are found from Decem-

ber to April [mean S-V 21.3 mm (19.8-25.4 mm)],

usually at the edge of pools where they feed on insects

and on other frogs. Many juveniles are emerald green.

Notes: Tyler and Martin (1975) described and illus-

trated the tadpole and tadpole mouthparts of C. aus-

tralis but had only poorly preserved series at stages

36-39. Tadpoles of C. australis cannot be distinguished

from those of C. longipes until stage 41 when the

mottled pattern on the dorsum of the latter species

appears.

Cyclorana longipes Tyler and Martin

A robust, fossorial species attaining a maximum

S-V length of 55 mm. Its habits are similar to those

of C. australis, from which it is distinguished morpho-

logically by its smaller size and densely mottled back

pattern.

Breeding activity: Males congregate in large numbers

from November to mid-December. Breeding occurs

at air temperatures of 23.0-28.5°C, 85-96% R.H. At

deposition sites water temperatures range 33.0-40.0°C

and water depth is 5-100 cm.

Eggs: Spawn is deposited on the surface of the water

at the edge of temporary pools in loosely-attached

clumps of 50-2 000 eggs. Eggs sink to the floor in

early development. The animal pole is brown and

FIG. 5. Left lateral view of tadpole of (a) Limnodynastes con-

vexiusculus, (b) L. ornatus, (c) Notaden melanoscaphus,

(d) Cyclorana longipes, (e) C. australis. Scale bar = 5 mm.

424

vegetal pole white. Two jelly envelopes surround the

ovum. Mean capsule diameter is 2.2 mm (1.9-2,5

mm) and mean ovum diameter 1-4 mm (1-2-1.5 mm).

Developmental history; Tadpoles are identical in

smaller maximum size al Stage 42; mean 42.3 (32.8-

59.5), Maximum growth occurs dunng stages 25-28

(Fig. 12), extending from a mean of 7.7 mm (6.8-&.3

mm) at stage 24 to 28.2 mm (24.6-30,5 mm) aL stage

28. The tadpoles possess numerous long gill filaments

al Stages 22-24.

At 30°C juveniles metamorphosed after 40 days.

At Jabiru juveniles were collected on 12 December

1978, in an area where pools were not formed until

after | November 1978: a maximum ot 42 days for

complete development. At metamorphosis juveniles

have a mean snout to vent length of 20 mm (14%.1-

21.1 mm).

Litoria bicolor (Gray)

A small, green tree trog bearing a white lateral stripe

an the head and body. Commonly found on vegetation

surrounding billabongs and streams. It 1s a nociurnal

species and spends the day al the base of leaves of

Pandanus and similar situations, Early in the wet

season males call during the day,

Breeding activity: From November to Apnl males

call from the stems of grass, bushes and small trees

inundated by or surrounding water. The nature ol

breeding sites appears io change during the wet scason,

Frogs are found near billabongs in the carly wel season

(October-December) buy later im temporary pools by

the roadside, and in swamps if there are sufficient

calling sites for males. Huge choruses congregate carly

in the wet season, but much fewer males call a1 the

temporary pools. Breeding occurs al air temperatures

of approximately 26°C, 92-96% RH.

Fees; Spawn is deposited in lemporary pools where

there is emergent vegetation. Eggs are small [mean

capsule diameter 2.5 (1.9-2.9 mm), mean ovum diam-

eter |. mim] with a black animal pole and a white

vegetal pole. They are laid below the surface in clumps

of 10-20 eggs, allached to submerged vegetation, Three

iclly envelopes surround the ovum,

Developmental history. The eyes are widely spaced.

the anus dextral and the spiracle simstral (Fig 6)-

Phe dorsam is mottled brown in the curly stages,

Pigmenicion mereasing progressively until, at stage

4). the tadpole is uniform brown, At stage 38 a white

patch appears just posltenor lo the eye and develops

laterally until, at stage 44, iLas a broad while siripe

from the eye to the groim. By stage 4] another white

stripe appears antero-venirally to the eye. passing

beneath it, and terminating just dorsal and posterior

lo the pectoral region. The ventral surface is white in

RC, S., AUST, MUS. 18 (18), 415-440

dpi, L989

these later stages. The tail muscle has stripes of pig-

ment laterally and on the dorsal and ventral edges.

the lail fins becoming increasingly pigmenied with

age, and the edges of the fins very dark. From stages

25-38 a conspicuous band of white muscle 1s present

at the poastenor end of the abdomen. The remainder

of the abdomen is dark brown, The tooth row formula

is 2/3. A well-developed, elevated border of small

papillae, surrounds the oral disc, except for the super

jor margin (Fig, 7), There is a strong horny beak.

Tadpoles attain a maximum length of 40.1 mm at

slage 42 [mean 33.7 (274-40.1)] with maximum

growth occuring between stages 25 and 27 [mean | 1.2

(6.4-19.6) at stage 25 to mean 21,8 (20.3-22.7) at stage

27 (Fig. &)}. At 30°C metamorphosis was completed

in 77 days: no comparative data are available for field

populations,

Litoria caerulea (White)

A large (up to 76 mm S-V), green, tree frog with

well-developed parotaid glands and greatly expanded

finger and toe discs. lt spends the day hidden in

crevices and hollows and is commonly found in

dwellings.

Breeding activity; Ts species does not form large

breeding choruses; males call individually or in small

groups trom exposed, clevated surfaces (e.g. walls,

fences, branches of trees. roadside culverts). Calling

occurs at night during or immediately after rain from

October to March. Breeding occurs fram November

(o February al air temperatures of 24-25°C, R.A.

about 96%.

/gys: Spawn is deposited m small ephemeral pools.

in whieh emergent vegelation may be present or

absent, Water temperature at deposition sites varies

28.0-38.0°C and water depths 3-50 cm, Spawn is laid

as a surface film of brown eggs with white vegetal

poles.. The nurmber of eggs in the clamp is 200-2 000,

Two jelly envelopes surround ihe ovum. Mean capsule

diameter 1.3 mm (1,l-1.4 mm).

Developmental Aistary: Eggs hatch at stage 18, and

atigin a mean lengih of & 1 mm (7.4-8.5 mm) by stage

24, A period of maximum growth follows, the embryo

reaching 28,5 mm (25,6-31.2 mm) by stage 29, Max-

imum size is reached at stage 41 (Fig. 8): mean 41.0

mm (38,6-43.6 mm). Tadpoles are of the typical,

lenuc Liloria type. Byes are widely spaced, anus dextral

und spiracle sinistral. Al stage 22 the tadpoles have

extensive external gills. Tooth row formula is 2/3 and

there is a well-developed, clevated border of large

papillae surrounding the inferior edge of the oral disc

(Fig. 7). The border does not extend superiorly to the

first upper labial tooth row, Tadpoles are mottled

brown with pigmentation increasing as develapment

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM 425

proceeds, There are twa brown, caudal stripes: a lateral

one extending halfway along the tail and a second

along the dorsal edge of the muscle for nearly the

length of the tail (Fig. 6). Pigmentation of the tail fin

increases progressively, bul a8 sparse. Ventral surface

dark, becoming progressively lighter with develop-

ment. At 30°C one juvenile metamorphosed after 38

days and measured 18 mm’'S-¥.

Litoria coplandi (Tyler)

A dark-brown frog of up to 40 mm snout to vent

length. [tis found commonly at night on sandstone

boulders in and around slreams in ihe escarpment.

The species aggregate in crevices and in caves in the

sandstone during the dry season.

Breeding activity: Males call from apen, sandstone,

rock Surfaces near water, alter rain from Sepiember

ig December. One of us (M.J.T.) has collected tadpoles

of this species in July in the Kimberley, W.A,, follow-

ing unseasonal rain,

Hees: Eggs have been found singly ar in small

clumps on the floor of shallow rock pools. We lack

data on ovum and capsule size.

Developmental history: Early Stages of development

are unknown. Al laler stages the dark brown tadpoles

can be readily identified because they have a brilliant

white patch at the posterior end of the abdomen. This

patch is.a muscle layer; it becomes less conspicuous

after stage 36 when the entire ventral surface beeomes

white. There is heavy pigmentation of the tail

myotomes over the entire surface posteriorly, but

only over the dorsal half anteriorly (Fig. 6). The tail

fin is hghtly pigmented. 4 naso-orbiial groove appears

al stage 37 but disappears by 41. Anus dextral; spiracle

sinistral. The tooth row formula is 2/3 with a complete.

well-develeped, clevated border of small papiiiae sur-

rounding the oral dise (Fig. 7). The horny beak is

well-developed. Tadpoles attain a length of more than

45 mm. Juveniles metamorphose at a mean of

15.8 mm (13.6-18.8 mm). The duration of develop-

ment is: unknown, but metamorphosing specimens

found on 2) February 1980 were collected in an area

where sufficient amounts of rain had fallen only since

the first days of January: a maximum of 52 days.

Litoria dahlii (Boulenger)

An aquatic species of the floodplain. L. dahlti is a

large frog of up to 70 mm S-V. Itis green with brown

and olive back markings, or so heavily suffused with

brown that the green is restricted 10 a mid-vertebral

stripe. The dry season and early wet Stason are passed

in eracks and crevices in the soil, When the plains

are flooded, it emerges and can be seen in shallower,

vegetated parts of the flooded areas, or basking at the

perimeter af buffalo wallows.

Breeding activity. Little is known of its breeding

habits. Males have been observed calling from a float-

ing position in the water when tain was falling during

periods of rising floods on the floodplains |l. Marris

pers, camm.].

ges, The nature of the spawn is unknown. Ova

dissected from gravid females are pigmented and haye

a mean diameter of 1,2 mm (range 1.1-1,3 mm).

Developmental history: The iadpoles have a bulbous

but elongate body. The eyes are dorso-lateral, the anus

median and the spiracle sinistral. The dorsal pattern

comprises dense mottling ona dark background, until

the longitudinal orientation of the stripe pattern of

the adult is detectable at stage 41. Ventral surface

dark, lightening to white by stage 42. Tail ryotomes

pigmented with a broad lateral stripe and another

along the dorsal edge (Fig. 6). Slight pigmentation of

the tail fin, The tooth row formula is 2/3. There is 4

well-developed, elevated border of small papillae

around the aral disc, widely separated superiorly by

a broad median gap (Fig, 7), The horny beak is well-

developed. The tadpoles reach 35 mm by stage 26.

and 52.6 mm (44.4-56.7) at stage 38. At stage 43 the

metamorphosing juveniles measured

28.2 mm (23.2-32.5 mm),

Litoria inermis (Peters)

A pale grey to brawn, medium sized (up ia 40 mm

S-V) ground-dwelling frog with slightly expanded ter-

minal digital discs. One of three closely related ‘ground

hylid’ species found in the area. Distinguished by its

warty dorsum, poorly developed head stripe. relative

limb lengih and reproductive strategy,

Breeding activity; Males. call from open areas (usu-

ally steeply sloping banks) fram Novetnber to March

The largest choruses occur early in ihe wet season-

Breeding occurs at 23.5-28.0°C, 92-96% RH. Depo.

silion sites are temporary pools with little or no emer-

gent vegetation. surrounded by gravel or sandy soils

and vegetation caver within Im of the water. Breeding

occurs al a water temperature of 33-35°C in water of

5-60 cm depth.

Eggs: Spawn is laid in free-floating surface clumps

of 96-330 eggs. Capsule diameter is large: 6.1 mm

(5.8-6.8 mm), Ovum 1.3 mm (1.3-1.4 mm), brown

with a white vegetal pole. There are three jelly envel-

opes,

Pevelopmental history; Tadpoles are of the vypical

lentic Literia form, with an elongate, oval-shaped

body (Fig. 6). Anus dextral, spiracle sinistral. Tadpoles

mottled brown on dorsum; abdomen dark, becoming

lighter during development. Tail myotomes mottled:

tail fin edges become yery dark in late developmental

stages. The tooth row formula is 2/3 and a well-

developed, elevated border of small papillae surround

REC. S. AUST. MUS., 18 (18): 415-440

April, 1983

roe]

SS ee

* nes ia

FIG. 6. Left lateral view of tadpole of (a) Litorta nasuta, (b) L.

bicolor, (c) L. tornieri, (d) L. dahlii, (e) L. inermis, (f) L.

caerulea, (g) L. sp. nr latopalmata, (h) L. wotjulumensis,

(i) L. rothii, (j) L. coplandt, (k) L. rubella. Scale bar = 5 mm.

the oral disc. There is a well-developed horny beak;

the papillary border is separated superiorly by a

median gap. A small patch of brown pigment is present

on the tip of the tail at stage 25, but this disappears

by stage 26. Maximum growth is from stages 25-27,

with the total length increasing from a mean of 6.3

mm (6.0-6.6 mm) at stage 24, to 21.2 (16.7-23.3 mm)

at stage 27. There is then a gradual increase to the

maximum mean length of 40.3 mm (38.3-43.5 mm)

(see Fig. 8). At 30°C juveniles metamorphosed after

74 days.

Litoria sp, near latopalmata

A slender pale grey to brown species; often mottled

with brown on the dorsum. Maximum length 37 mm

S-V. Distinguished from other ground-dwelling hylids

by its well-defined head stripe and smooth dorsum.

Breeding activity; Males call in large choruses from

open areas within | m of water from October to April.

Breeding has been observed from November to March,

at 24-28°C, R.H. 89-96%. At deposition sites the

water temperature was 33.6-41.2°C, and water depth

up to 100 cm.

Eggs: Spawn is deposited in temporary pools; eggs

are brown with a white vegetal pole and are laid as

a surface layer, becoming clumped when even slightly

disturbed. 50-350 eggs are laid with a mean capsule

diameter of 3.6 mm(3.4-3.9 mm), and ovum diameter

of 1.3 mm (1.2-1.3 mm). The ovum has three jelly

envelopes.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

Developmenral history Tadpoles hatch at stage L8.

They are mottled brown on the dorsum and have a

dark abdomen which lightens to white by stage 41.

The mottled pattern overlies the dark flanks, The tail

myotomes ar¢ heavily pigmented and mottled, and

the tail fin pigmented. The tadpole is of the typical

Litoria form with a dextral anus and sinistral spiracle

(Fig. 4). The tooth row formula is 2/3 and an elevated

and well-developed papillary border surrounds the

oral disc, The small papillae are separated superiorly.

Maximum growth ovcurs at stages 25-27. the total

length increasing trom 6.2 mm (6.0-6.3 mm) at stage

24 10 24.4 mm (20,]-28.5 mm) at stage 27. Maximum

length 1s reached at stage 38 with a mean total length

of 44.8 mm (42,1-48.9 mm) (see Fig. 8). This length

is Maintained until absarpiion of the tail begins at

stage 42, Two metamorphosing juveniles at stage 44

measured 14.8 and 16.1 mm respectively. AL 30°C

juveniles metamorphosed afier 87 days. In the field

juveniles were collected on 28 February 1980 in an

area where suitable pools had been available only

from January 5: a maximum of 54 days for devel-

opment.

Litoria meiriana (Tyler)

A small brown or mottled slate-coloured frog reach-

ing a maximum snout to vent length of 25 mm. It is

diurnal and occurs In leatHitter alongside sireams in

the sandstone escarpment. The dry season [s spent in

crevices and caves in the sandstone.

Breeding activity: Early in the wet season males call

trom Jeaflitter and open rock faces at the edge of

streams and pools in the escarpment. Breeding occurs

from October to December. Amplectant pairs have

been found at 26,5-27,0°C, R.H. 85.0-88.5%.

Eggs: Spawn is deposited beneath the water suface

ina film of small clumps attached to rocks or to the

floor of small, lemporary pools in sandy escarpment

Streambeds, Eggs are small and light brown. One

clutch contained 39 eggs.

Developmental history: Tadpoles are brown and have

4 dextral anus and sinistral spiracle. Tail myotomes

are mottled brown and havea dark brawn stripe along

the dorsal edge (Fig. 9). The upper tail fin is pigmented

The ventral surface lightens during development unt]

it is white at stage 42, Tadpoles have the typical

Literia tooth row formula and a well-developed, ele-

yated border of small papillae surrounding the oral

disc. The papillary border is narrawly separated supe-

norly (Fig. 9), There is a well-developed horny beak.

Maximum growth occurs at stages 25-27 increasing

from a mean of 4.05 (3.9-4.2 mm) ai stage 22 to 6.2

mm (3.7-12.6 mm) al stage 25, and up to 21.3 mm

(15.9-25.1 mm) at stage 27, The growth is then very

slight, increasing to only a mean of 25-4 mrn (25.3-

256 mm) at stage 41 (Fig. 10). At 30°C tadpoles

$27

reached stage 42 in 45 days. At Ngarradj warde djob-

keng (Birndu) juveniles were found in large numbers

on 30 November 1978, The first appreciable rains in

the area and observations of calling and spawning

were in the first days of November [G. Miles

pers, comm. ]: a maximum ol! 30 days for developmen,

Litoria microbelos (Cogger)

A small, slender, brown species attaining a maxi-

mum S-V length of 14 mm. It occurs among long

grass at the edges of pools and streams. The day is

spent under logs, stones and vegetation near

streambeds or flooded grassland.

Breeding activity: Males call from the base of grass

tussacks or en grass stems during or immediately

fallowing rain, from October to February. Early in

the wet season this species and 1. bicolor form large

breeding choruses in pools al the edge of billabangs,

Where L. bicolor is present, L. microbelos males call

trom the base of the grass tussocks; however, if Z.

hicalor is absent, £. microbelos calls from the grass

stems normally occupied by L. hicelor.

Eggs: Eggs are deposited in small clumps among

grass stems. Mean ovum diameter is 0.8 mim

(0,8-0.9 mm); the animal pole is brown and the vegetal

cream. No complete jelly capsules were found,

Developmental history: (Derived from specimens

iaken at Mitchell Plateau, W.A.) Embryos hatch at

stage 20 at a total length of 3.0 mm (2.8-3.1 mm). At

stage 24 total length is 4.1 mm (4.0-4.2 mm) increasing

to 11.4 mm (10.8-12.0 mm) at stage 27. No data are

available for later stages, There is dark pigmentation

on the dorsal surface, and a dark head and lateral

stripe. There are bands of pigment on the tail muscle.

The tail is lightly pigmented. The tooth row formula

is 2/3. A border of relatively large papillae surround

the aral disc. This is widely separated superiorly. The

horny beak 1s rather weakly developed.

Litoria nasuta Tschudi

An exceptionally long-legged, elongate-bodied spe-

cies attaining a maximum S-V length of 50 mm, It is

brown with dark brown and white longitudinally

orientated back markings, and a conspicuous dark

head stripe. It is abundant, eacept upon the escarp-

ment

Breeding activity: Males call from open areas

between grass tussocks within | m of water from

Navermnber to March, Breeding occurs from November

ta February. Amplectant pairs have been found at

25.0-26.5°C, R.H. 96%, At spawn deposition sites

water temperatures were 30.5-41.2°C, and water

depths of 15-30 cm.

428

Eggs; Spawn is deposited in small, temporary pools

on inundated grassland, usually where there 1s sparse

emergent vegetation. Eggs are dark brawn with cream

vegetal hemispheres, Spawn is laid as a surface film

which becomes a clump when disturbed. There are

50-100 eggs/clump, with a mean capsule diameter of

3.4 mm (3.2-3.7 mm) and ovum diameter of 1,1 mm

(1,0-1.3 mm), There are three jelly envelopes.

Developmental history: The tadpole is of typical

lentic Litoria form. The anus is dextral and spiracle

sinistral. The tooth row formula is 2/3 with a well-

developed, elevated papillary border surrounding the

oral disc and separated superiorly (Fig, 7), Tadpoles

are motiled brown with the Ventral surface whitening

during development. The tail myotomes are mottled,

and the tail fin pigmented (Fig. 6). The characteristic,

striped, back patiern of the adults becomes evident

at stage 42. The extreme leg length, striped leg pattern

and reduced toe-webbing distinguish this specics at

silage 41 from closely-related species. Eggs hatch at

stage 18 at a mean length of 1.9 mm (1.8-1.9 mm).

They attain 6.0 mm (5.2-6.3 mm) by stage 24, where-

upon there is maximum growth until stage 28,

33.3 mm (29.0-38.3 mm) (Fig. 12). The maximum

mean length of 48.8 mm (44.8-55.7 mm) 1s reached

al slage 39, Tadpoles remain at this size until absorp-

tion of the tail commences at stage 43. Ar 30°C

juveniles metamorphose afler 31 days at a mean length

of 18.7 mm (17.3-21.6 mm),

Litoria personata Tyler, Davies & Martin

A small, rock-dwelling species, attaining a maximum

S-V length of 35 mm. It is brown or grey with a

conspicuous dark head stripe. The dry season is spent

in cracks and crevices in the escarpment.

Breeding activity: Males call from open rock faces

and elevated positions at the edge of streams in the

escarpment from November to late January, Tadpoles

and transforming juveniles were collected in late April

1978 by M. J. Tyler.

Egas: The eggs are unknown.

Developmental history; Tadpoles have been found

in temporary pools upon the escarpment and in stream

beds at its base. The tadpoles are highly distinctive,

being dark brown dorsally with gold to yellow dorso-

lateral stripes. The ventral surface is creamy white.

There is a dark prey band between the eyes, and an

arrowhead-shaped dark prey paich anteriorly. The

body shape is typical of other Lyroria fram louie hab-

i1ats, The body is flattened, the tail muscle is well

developed and the fins narrow. The tooth row formula

is 2/3, with a complete, elevated papillary border

surrounding the oral disc. The horny beak is weakly

developed. Anus dextral. and the spiracle sinistral.

Tadpoles measured 41_8-55.1 mm at stages 41-45

REC. S. AUST. MUS., [8 (18). 415-440

dprit, MSZ

(Tyler et at,, 1979). They form aggregations and are

often seen miainiaining a static. horizontal position

§-10 em beneath the surface.

Litoria rothii (de Vis)

A large tree frog attaming 60 mm snout to vent

Jength. It is brown and has bright yellow and black

markings in the groin and upon ihe posterior surface

of the thigh, 11 is commonly seen on buildings in the

area.

Breeding activily: Males call between November

and March following rain from elevated positions

neat water, Breeding occurs from November to March

at 27.5-28.5°C, R.A. approximately 96%.

Eggs: Spawn is deposited in small clumps in tem-

porary pools in which emergent vegetation may be

present or absent. Eggs are dark brown with a cream

vegetal pole and have a mean diameter of 1.4 mm

(1.3-1.4 mm). The capsule is 4,6-5.4 mm. It ts fragile

and easily ruptured. There are three jelly envelopes.

Developmental history: Tadpoles are pale yellow on

ihe dorsal surface and white yentrally when they

hatch at stage 18. There are dark brown stripes, one

branches forward antenur to the eyes and another on

each side of the body fram the nostri! to ihe end of

the tail. A bright yellow spot is visible on the dorsal

surface just anterior 10 the eyes. At stage 23 external

gill filaments are present. Anus dextral, spiracle sin-

istral. The body is of the typical “irorja leniie waier

form, but the tail fins are much deeper than in other

species in the area (Fig, 6), The stripes fade during

development and the dorsal and lateral surfaces

become mottled brown. The dorsal surface of the

head becomes very flat. The tooth row formula is

2/3. The third lower row of labial teeth is absent or

reduced io a small, median structure bearing less than

ten teeth. A border of well-developed large papulae

is elevated and sourrounds the oral disc except supe-

riorly (Fig. 7). There is a well-developed horny beak,

The tadpoles undergo a maximum growth period in

stage 25 inereasing in total length from 7.0 mm (6.8-

74mm) at stage 24 to 19.9 mm (7.2-38.2mm) at

stage 25 increasing to 31.4. mim (25.3-43.0 mm) in

stage 26. A maximum of 58,2 mm was measured in

a stage 40 iadpole (Fig. 8). Ai 30°C a juvenile meta-

morphosed after 146 days: in the field, huwever, a

large juvenile was collected on 11 March 1980 in an

area where water was not present until after 5 January

1980; a maximum of 65 days for development.

Litwria rubella (Gray)

A small brown tree frog of up 10 40 mm S-V, it is

abundant throughout the area except in the escarp-

meni. The day ts spent in hollows, under logs and

rocks. in dry streambeds and on banks. in dwellings

and cisterns.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM 424

Te ite,

WHnt¥e on WELW), sal yz yy ae ;

ELAN

swine

is f

eit r

—

| wate ied

ni Pa ( ie

4 RAWeoe

vad ipa rel

syit

wl!

ju telltaniaeay

iy, Ne ss

ii Hey

belle juan

FIG, 7. Oral discs of fa) Literia dahlii, (b) L. caerulea, (c) L.

rothi, (d) L. wotjulumensis, (e) L. bicelar, () L. nasuta, (g) L.

caplandi. (h) L. rubella, Seale bar = | mm.

(mm)

LENGTH

TADPOLE

430

REC. S. AUST. MUS.,

18 (18): 415-440 April, 1983

Litoria bicolor Total Length Litorta rubella Total Length

40+ 4

304 } | | 4 | t | if

204 Ta J |

i tyf

104 4

AIR TEMP. 30° - 32°C AIR TEMP 30° - 32°C

WATER TEMP 25°- 30°C = ft WATER TEMP 25° - 30°C

z * ae

10) T T T T T T T + T “¥ T

éo Litoria caerulea Total Length | iF t } Litoria inermis Total Length f i

304 it 1 | } } i

204 |

15 Lt AIR TEMP 30°- 32°C 7 AIR TEMP 30° - 32°C

z WATER TEMP 25° - 30°C zit WATER TEMP 25° - 30°C

zy = zt =F

10) + T r T << T T T T T T

607 4

Litoria rothi Total Length I] - Litoria sp.nr. /atopalmata Total Length

50+ - 4 |

40 J t

1 ts

= =

30- 4 |

20-4 4

ol AIR TEMP 30° - 32°C | AIR TEMP 30° - 32°C

= } t WATER TEMP 25° - 30°C eet = = WATER TEMP 25° - 30°C

fe} L 1) T T T T T T T T T T T

20 25 30 35 40 45 20 25 30 35 40 45

TADPOLE STAGE

FIG. 8. Growth gradients in tadpoles of various species of Litoria.

Breeding activity: Males call from a variety of areas

near water ranging from open grassy areas to elevated

sandy hillocks, usually on the ground. Silent males

and amplectant pairs are often found within 0.3 m of

a calling male. Calling and breeding occur from

November to April, although both activities appear

concentrated early and late in this period. Amplexus

occurs at 25.0-26.5°C, R.H. 92-96%.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

Eggs: Spawn is deposited in the form of small,

floating surface films in temporary pools with or with-

out emergent vegetation. Water temperatures of up

ta 40°C have been recorded at the deposition site,

water depth is 30-LO0 cm. Eggs have a brown animal

pole and cream vegetal pole. There are 40-300 eggs

in a clump. The capsule diameter ts 2.1 mm

(1.9-2.3 mm), and ovum diameter 1.0 mm

(1,.0-L.1 mm). There are three jelly envelopes.

Developmental history: At 30°C the eggs hatch

within three days at stage 21, Tadpoles attain maxi-

mum growth duririg stages 25-27, increasing in total

length from 5.9 mm (4.5-6.9 mm) al stage 23 to

22.2 mm (17-9-30,9 mm) at stage 27. Maximum length

js reached at stage 42 (Fig, 8), 31.2 mm

(25.3-34.8 mm), Tadpoles have a 2/3 tooth formula,

and small papillae in a well-developed, clevated border

surraund the oral dise except superiorly (Fig. 7), The

horny beak is well developed, The spiracle is left

ventro-lateral and the anus right median. The body

is rotund, mottled brown on the dorsum, and white

on the ventral surface (Fig. 6). The mottling becomes

darker during development. There is dark pigmenta-

tion on the tail muscle in three stripes: a laleral ane

flanked by others along the dorsal and ventral extrem-

ilies, This pigmentation also occurs on the outer edge

of the tail tin becoming increasingly darker during

development, A shallow nasal groove from eye to

nase 1s present in later Stages. At stage 4], a broad,

white, lateral stripe and a vertebral stripe appears and

is retained until after metamorphosis. There is also a

white patch anteriorly to and below the eye. The

remainder of the dorsum is uniform brown. At 30°C

the juveniles metamorphose afler 38 days. A collection

of tadpoles with metamorphosing juveniles present

was made on 10 February 1980 from a pool that

could noi have been filled before 5 January 1980: a

maximum of 37 days for complete development.

Litoria tornieri (Nieden)

A moderate-sized, ground-dwelling hylid (40 mm)

distinguished by a narraw continuous brown. stripe

on the anterior margin of the tibia. In the breeding

season the species ts reddish with a conspicuous, dark

brown head stripe.

Breeding activity: Males call from cover, either under

leaves or at ihe base of grass tussocks within 3 m of

water. Calling occurs from October to April; breeding

has been observed from November until March.

Breeding occurs at 25.0-26,5°C, R.H,

92-96%,

Eges: Eggs are laid in temporary pools in inundated

grassland where the water depth is Jess than 100 cm,

The dark brown eggs have a white vegetal pole and

are laid as a surface layer which becomes a clump

441

SC ae a

re oot

D AN ew.

r Se oe / a

: Fe \ alt

ae ee te ; =

ada eS 7 a a Oe ie _

SES ‘ at

rn ee)

oe ee

FIG. 9. (a) Left lateral view of tadpole, and /) oral dise af

Literia meiriana, Scale bar = 1 mm.

when disturbed. Capsule diameter is 3.4 mm

(3.2-3.7 mm) and ovum diameter 1.4 mm. There are

iwo jelly envelopes,

Develapmental history: Tadpoles ace mottled brown

on the dorsum and have a dark abdomen which

lightens to white during development. The tail

myotomes are mottled, and the tail fin pigmented,

The tadpole is of typical lentic Litoria form (Fig. 6).

The anus opens right of median and the spiracle left

lateral, The tooth row formula is 2/3, and a well-

developed, elevated border of small papillae incom-

pletely surrounds the oral disc. The horny beak is

strongly developed. Tadpoles hatch at stage 18 and

reach a mean total length of 6.5 mm (6.1-7.1 mm) at

stage 24 (Fig. 12). A period of accelerated growth

occurs, increasing the total length to 26.4. mm (23,2-

29,9 mm) at stage 27, Growth increases slowly until

it reaches a maximum of 38.8 mm (34,6-44,0 mm)

at stage 38. One juvenile was 15.1 mm at metamor-

phosis. At 30°C tadpoles reached stage 40 in 44 days.

Litoria wotjulumensis (Copland)

A large ground-dwelling hylid (reaching 70 mm

5-V), with an elongate body, extensive webbing of the

toes and a broad dark conspicuous head stripe, Its

call has a repertoire of rattles and chuckles.

Breeding activity: Males call trom open areas at the

edge of water and face the water. Calling occurs from

October to March, and the species breeds in the carly

months of the wel season. Breeding has been observe:l

at 25-27°C, R.H.. 75-92%,

LENGTH (mm)

TADPOLE

432

REC. §. AUST, MUS,, 18 (18): 415-440

tprl, 1983

Nofaden mel/anoscaphus Toia| Length if Litaria meiriana ~~ Total Length

30 I } _ = i

i T fE-*F_

F eet

20 |

a | AIR TEMP 30° - 32°C | AIR TEMP 30° - 32°C

[= zt WATER TEMP 25° - 30°C WATER TEMP 25° - 30°C

t z=

= =

= | ——

(9) SS ee Tr a T 1 T es La —_——_— — we

20 25 30 35 40

TADPOLE STAGE

30 Uperojeia jnundata Total Length {-

/ | if]

10 AIR TEMP 30° - 32°C

WATER TEMP 25° - 30°C

30 35

TADPOLE STAGE

FIG. 10. Growth gradients in tadpoles of various species of frogs.

Eggs: Spawn is laid in temporary pools in sandy or

gravelly soil. Eggs were found at water temperatures

of 32-33°C in water 10-50 mm deep. The dark brown

eggs have a cream vegetal pole and deposited as a

floating clump of 30-200 eggs, occasionally attached

to surface vegetation. Mean capsule diameter 4.6 mm

(4,2-4.9 mm), ovum diameter 1.75 (1.6-1.9 mm), There

are three jelly envelopes.

Developmental history: Tadpoles have well-devel-

oped external gills at stages 20-24, The body is typical

of lentic Litoria (Fig. 6). The tadpole is mottled dark

brown and the ventral surface is white. Anus dextral,

spiracle sinistral. The tail myotomes are heavily mot-

tled, and the upper tail fin pigmented. The tooth row

formula is 2/3 and a well-developed, elevated border

of small papillae incompletely surrounds the oral disc

(Fig. 7). The horny beak is strongly developed. At

stage 25 the total length is 12.6 mm (8.2-21.2 mm)

increasing to 40.5 mm (39.3-42.3 mm) at stage 41

(Fig. 12). Juveniles metamorphosed at a mean length

of 17.0 mm (15.5-18.0 mm). At 30°C tadpoles reached

stage 42 after 53 days.

Limnodynastes convexiusculus (Macleay)

A large, broad-bodied frog with a mottled tubercular

dorsum, common in swampy areas and abundant on

the floodplains. It is found among long grass usually

in small hollows or burrows, It is cryptic in its habits.

Breeding activity: Males call from October until

April from small hollows among long grass near water.

Eggs: There is little known of the eggs except that

they are laid in the form of a foam nest.

Developmental history: The tadpoles are a distinc-

tive, intense black with long, deep tail fins (Fig. 5).

Anus median, spiracle lateral and sinistral. The tooth

row formula is 5/3 and there is an elevated border of

large papillae which is widely separated superiorly

around the oral disc. The horny beak 1s rather weakly

developed. The body is robust and the dorsal surface

slightly flattened. Tadpoles do not occur in large num-

bers. They attain a maximum length of 70 mm.

Limnodynastes ornatus (Gray)

A rotund, fossorial frog up to 45 mm S-V with a

highly variable back pattern. This species has no pre-

ferred habitat (although it is uncommon in the larger

billabongs) and is abundant in the area.

Breeding activity: Males call whilst free-floating in

water. Calling occurs from October to March, and

breeding from November to February. Breeding has

been observed at air temperatures of approximately

26°C, R.H. 88.5-96.0%. Any pool deeper than 2 cm

is used for breeding purposes, including ditches newly

dug for sewer pipes and shallow, water-filled depres-

sions on the road surface. At deposition sites water

temperatures were 27-38°C.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

433

FIG. WI. Oral dises

(bh) Limnodynastes ornatus

har = Imm.

ges: Eges are deposited in a foam nest floating on

the surface of the water. The foam rapidly collapses

to form a thin surface film. The number of eggs per

clump is 155-1 630 with a mean of 645. Ovum black

with a white vegetal pole and diameter of 1.2 mm

(1.0-1.3 mm), The eggs have one jelly en velope; capsule

diarmeter 1.6 mm (1.45-1,8 mim).

Developmental history: The embryos hatch after 18

hours at stage 20 at a length of 4.4 mm (4,0-4.7 mm).

Long filamentous gills are present from slages 22-24,

Growth is steady throughout development, The tad-

poles are brown on the dorsal surface with a darker

patch between the eyes, and on the tail myotomes.

Eyes dorsal and close together giving the tadpole a

distinctive appearance, Anus median, spiracle sinistral.

Bady flattened dorso-ventrally (Fig. 5), Tooth rows

vary from 2/3 to 5/3. A well-developed, elevated

border of large papillae, separated superiorly, sur-

rounds the oral disc (Fig. 11). The horny beak js

weakly developed, The tadpoles form dense and acti ve

aggregations, and cannibalism 1s prevalent, particularly

in ephemeral pools. A maximum length of 31.0 mm

(29,5-32.8 mm) is reached at stage 39 (Fig. 12) and

juveniles metamorphosed at 10.3 mm (6.5-11.6 mm),

At 30°C metamorphosis was completed after 26 days.

In the field, spawn laid on 28 November 1977 devel-

oped to stage 46 on 23 December 1977: 25 days.

Megistolotis lignarius Tyler, Martin and Davies

A moderate-sized (60 mm S-V) robust species with

short, Very muscular limbs and a very large tym-

panum. Inhabits scree slopes and escarpmenis and

hides beneath rocks and boulders, and in caves during

the dry season.

Breeding activity; Males call from beneath boulders

in or near sireambeds in the escarpment, Breeding

commences jn late November,

Eggs: The creamy’ yellow eggs are deposited in a

foam nest under rocks in static water.

of (a) Notaden melanascaphus,

» (c) Cyeloranu longipes. Seale

Developmental history: Hatching occurs at stage 21,

The body and fins are intense black, Anus median,

spiracle sinistral. The tadpoles have highly efficient

suctorial mouthparts. The tooth row formula varies

from 4/3 to 6/3. A border of small papillae is well

developed, elevated and narrowly separated superiorly.

The horny beak is strongly developed, Cannibalism

has been noted in the laboratory. Tadpoles reach a

length of over 50 mm by Stage 42. A juvenile meta-

morphosed after 65 days and measured 23.3 mm

(Tyler, Martin & Davies, 1979),

Notaden melanoscaphus Hosrner

A methum-sized, extremely robust, fossorial trog

with short limbs, It is pale brown with large dark

blotches on its back, and when seized exudes a copious

amount of sticky, yellow secretion from large dorsal

glands. It is found in grasslands throughout the area

and passes the day buried in sandy soil.

Breeding activity: Males call during or immediately

afier rain trom shallow, water-filled depressions. Call-

ing occurs from December to March. and breeding

from early January to March. Air temperatures range

23.0-26,5°C, R.H, 96-100%.

Eggs: The spawn is deposited as a surface film in

shallow, ephemeral pools in inundated grassland.

Water temperatures range 27-36°C and the depth

1-30 cm. Eggs have a black animal pole and white

vegetal pole. The number of eggs is 500-1 000 with a

mean of 833. The eggs have individual capsules jained

by a sticky loose layer of jelly. The mean capsule

diameter is 2.7 mm (2.6-2.9 mm) and avum diameter

1.4 mm (1.3-1,45 mm). There is one Jelly envelope.

Developmental history: The spawn sinks as the eggs

develop, The larvae hatch at stage 20 at 5.8 mm (5.0-

6.8 mm). External gills are present at stages 22-24.

Anus dextral, spiracle sinistral and lateral (visible

from a dorsal view). Dorsal pigmentation dark brown,

with the adwt back pattern becoming apparent in

stage 41. Ventral surface dark brown becoming lighter

as development proceeds. Tail myotomes dark brown

434

(mm)

LENGTH

TADPOLE

REC. S. AUST. MUS., 18 (18): 415-440 April, 1983

705 ]

Litoria tornier’ Total Length Cyclorana australis Total Length

604 |

da! 4 | t

‘i

40 | J | |? 4 a

305 ] } if i “I

204 |

10") AIR TEMP 30° - 32°C 7 AIR TEMP 30° - 32°C

‘ zt WATER TEMP 25° - 30°C WATER TEMP 25° - 30°C

t ‘ a

ol ad 2 — —T— —T- — ~~ eee, ao ——T —= — —T

607 5

Litoria nasuta Total Length Cyclorana fongipes Total Length

40-5 4

304 | ft

20+ 1 1 HH

104 AIR TEMP 30° - 32°C | AIR TEMP 30° - 32°C

WATER TEMP 25° - 30°C f WATER TEMP 25° - 30°C

zit rt xt

ft ft zt

ot — + — T i — a: —— —- a a ——

505 4

Litoria wotjulumensis Total Length Limnodynastes ornatus Total Length

40 T } 4

30+ | it a t i

20 4 i i ea

| | tt |

104 4 I

AIR TEMP 30° - 32°C } AIR TEMP 30° - 32°C

i = WATER TEMP 25° - 30°C +] zt t WATER TEMP 25° - 30°C

L =

[0] —T =" 4d 3 —T— —— —T —T ee | as | —~T —“

20 25 30 35 40 45 20 25 30 35 40 45

TADPOLE STAGE

FIG. 12. Growth gradients in tadpoles of various species of frogs.

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

and tail fins pigmented. Upper tail fin much deeper

than lower (Fig. 5). Tooth row formula is 2/3 with

one complete upper and two complete lower rows.

The third lower tooth row is reduced to a median

structure of 10 teeth (Fig. 11). A border of large

papillae is not well developed and is widely separated

superiorly. The horny beak is very weakly developed.

The tadpoles increase in total length from 7.25 mm

(6.1-8.1 mm) at stage 24, to 17.2 mm (13.7-19.2 mm).

At stage 24 a tadpole measured 25.4 mm (Fig. 10).

The tadpoles inhabit very shallow water (1-5 cm) deep

in inundated grassland. Water temperatures exceeding

41°C have been recorded there. There is heavy floc-

culent material on the floor, and the tadpoles spend

most of the time lying in this layer. In the field a

juvenile was found on 26 February 1979 in an area

where adult Notaden were not present before 4 J anuary

1979: a maximum of 53 days for complete develop-

ment.

Ranidella bilingua Martin, Tyler and Davies

A small, brown species with variable darker and

lighter markings on its back, and measuring a maxi-

mum of 25 mm. It inhabits inundated grasslands in

the wet season; the dry season is spent on the soil

under logs and rocks.

Breeding activity: An opportunistic breeder; males

call and mating occurs whenever sufficient rain falls,

even in dry season showers. Males call from the base

of grass tussocks in or around water.

Eggs: Eggs are laid in small clumps attached to

vegetation. The animal pole is dark brown and the

vegetal pole creamy white. There is a single jelly

envelope. The capsule diameter is 1.74 (+ 0.16 mm)

and ovum diamter 1.17 (+ 0.04 mm) (Martin, Tyler

& Davies, 1980).

Developmental history: Derived from Martin ef al.

(1980). Eggs hatch at stage 20 and have a total length

4.0-4.5 mm. There are no external gills present. At

Stage 25 the total length is about 6.5 mm, increasing

to 24.5 mm by stages 40-41. The tadpoles are dark

brown with the myotomes and ventral surface white.

Upper and lower fins are pigmented. The tooth row

formula is 2/3 with the third lower row very short.

Few large papillae border the lateral area of the mouth,

but are completely absent superiorly. Posteriorly there

are just two papillae flanking the third short tooth

row. The horny beak is strongly developed. Juveniles

collected at Mitchell Plateau, W.A. metamorphosed

at 7.5-8.4 mm S-V after 13-14 days.

Uperoleia arenicola Tyler, Davies and Martin

A small, squat species only found in a sandy

streambed in the escarpment of Birndu (Ngarradj

warde djobkeng) near Ja-Ja. Its colour is variable (pale

435

grey to dark slate) and there are bronze parotoid

glands. The specimens were found on steep-sided

sandy banks of the stream.

Breeding activity: Data are confined to observations

on calling males. We noted that they called without

moving from their burrows and were still covered

with fine sand.

Uperoleia inundata Tyler, Davies and Martin

A moderate-sized species of up to 30 mm S-V

length. It is grey with highly variable dark brown to

ted dorsal markings. In the wet season it inhabits

inundated grasslands.

Breeding activity: From January to April the males

call from shallow water at the base of grass tussocks,

under leaves and logs, and from grass stems in water.

Large breeding choruses are formed. Breeding occurs

during January and February, at 24.5-29.0°C, R.H.

89-100%.

Eggs: Spawn is deposited in shallow (2-5 cm), inun-

dated grasslands, water temperatures 27,2-28.6°C. Eggs

are laid in small clumps (<5) or individually, and

immediately sink to the bottom. They have a sticky,

tough outer capsule which accumulates a covering of

particles. The ovum has a dark brown animal pole

with a light brown vegetal pole. There is one jelly

envelope, and the capsule has a diameter of 1.8 mm

(1.5-2.0 mm). The ovum diameter is 1.25 mm (1.1-

1.4 mm).

fo figura x

ule STO

uot

gol " Qe Ay,

Fao lg

FIG. 13. (a) Left lateral view of tadpole, and (b) oral disc of

Uperoleia inundata. Scale bar = 1 mm.

Developmental history: Embryos hatch at stage 20

at a total length of 2.7 mm (2.5-2.8 mm). External

gills are absent. At stage 25 the total length is

10.4 mm (6.6-17.1 mm). Tadpoles are dark brown on

the dorsal and lateral surfaces, and mottled with red-

purple and white on the ventral surfaces. Anus dextral,

spiracle sinistral and visible from above. The tail

myotomes are mottled with brown and the upper tail

fin is pigmented (Fig. 13). The tooth row formula is

2/3 with the upper row divided. A poorly-developed

436

border of large papillae, widely separated superiorly

and inferiorly. surrounds the oral disc (Fig. 13). The

horny beak is weakly developed. Tadpoles grow to 4

length of 31.0 mn (23.25-33.9 mm), At stage 44 the

tadpoles had a size range of 16,0-17.7 mm (Fig, 10).

Juveniles were collected on 2 March 1979 in an area

where there was not water prior to 10 January 1979:

a maximum of 41 days for metamorphosis.

Sphenophryne robusta (Fry)

A very small, brown cryptic. terrestrial species found

in “*paperbark swamps’, rainforest remnants and in

ihe escarpment. It inhabiis leaf bitter.

Breeding activity: Little is known of us biology.

Other sphenophrynine microhylids lay eggs out of

water and exhibit direct development within the egg

capsules, It may be assumed that this species behaves

similarly.

BEHAVIOUR

Activity Patierns

Adults of most species are active only at night but

there are a few exceptions. We have observed spas-

modic diurnal calling activity by Liroria rothiv and

L. rubella, whilst in the wet season Cyclorana australis

and L. dahlii commonly bask in the sun a1 the edge

of pools. On the morning of 29 November 1977 M.D,

and M.J.T. captured two basking C. australis at

Ja Ja. The air temperature there was 33°C, Dorsal

skin temperatures were 33.0°C and 34.4°C respec-

tively, cloacal temperatures 33.4°C and 34,2°C and

ground beneath the frogs 32.0°C and 34.0°C. Our

experience with C. australis at the study site and

elsewhere in the Northern Territory indicate that the

basking frogs are not alert; they press their bodies

close to the ground and if approached quietly they

can be captured by hand, We are uncertain of the

upper limit of temperature at which basking is. dis-

continued. Beyond the study area at a site adjacent

to the Stuart Highway 99 km south of Elliot on

17.12.80, two of us (Davies, Tyler) with A. A. Martin

collected specimens beside a temporary pool where

the air temperature was 37.0°C; soil beneath one frog

was 36,2°C and the adjacent water temperature

30.0°C. The body temperature was 36.4°C. During

experiments at Adelaide designed to induce burrowing,

C ausiralis temained on the surface tor two hours

when exposed to a heat lamp elevating the surface

temperature of the adjacent soil to 40°C (J. Sanders,

pers, comm.).

Litoria dahlii is less readily approached whilst bask-

ing than is C. australis, and we have fot captured

basking specimens.

REC. S, AUST. MUS,, 18 (18); 415-440

ipril, HAs

Litovia meiriana is habitually diurnal. This small

species shelters in the shade at the perimeter of rock

pools and is unique amongst Australian anurans in

having the ability to travel across the surface of static

water in a series of rapid bounces to avoid enemies

or Lo capture small. flying insects.

Seasonality of Reproductive Behaviour

From May to September any rainfall in the Magela

Creek system isa rare event (see Jabiru rainfall records

plotted by Tyler and Crook 1980). The area is hot

and dry and activity by frogs 1s minimal,

The nature of dry season refuges influences the

onset of reproductive activity only slightly. The low-

land species in general require the formation of shallow

bodies of water, and 11 is evident that the fossorial

species ate amongst the first to breed there (Fig. 14).

We have observed the effect of rainfall upon the

escarpment and upon the floodplain and, despite the

absence of comparative rainfall records, 11 1s evident

that even slight falls upon the escarpment are sufficient

1o induce breeding. This is because the impermeable

surfaces result in rainfall flowing across the rock and

accumulating in every depression, so providing

numerous breeding sites. These breeding sites may be

produced by localised showers in the wel season, and

our observations in the Northern Territory and the

Kimberley Division of Western Australia indicate that

some species breed during these unseasonal light or

localised rains. Irrespective of how localised these

rains tay be. they are of greatest significance upon

the escarpment, benefitting species such as Liforta

personata, L. coplandi and Megistalotis lignarius.

Fecundily

Clutch size is commonly at variance with the total

egg complement because some individuals deposit

several clutches on a single mght. This habit probably

explains the wide disparity in clutch sizes recorded

in Table 2. Other species such as Litaria coplandi

and Ranidella bilingua scatter single eggs and the

complement cannot be calculated from field obser-

vations.

Salthe and Duellman (1973) demonstrate that within

any reproductive mode there is a positive correlation

between egg complement and adult size. Thus large

species tend to lay more eggs than small ones. This

is bore out by our data (Table 2) and there is very

slight interspecific variation in egg diameters. Thus

1.8mm and in L. microbelos 0.9 mm (100% variation).

This is despite the fact thal there is a 500% difference

20 mm in L, wticrohelos.

a, ACTIVE

WA CALLING ACTIVITY

ria BREEDING

FIG. 14. Wet season activity of frogs in the Magela Creek System.

For each species, the upper trace represents data derived

from 1978-79 season, and the lower trace 1979-80 season.

Capsule diameter is the sum of the ovidiameter

surrounded by the expanded (hydrated) oviducal

secretions deposited upon the ova. In the species

examined the outer capsule diameter was approxi-

mately 1.5-2.5 times the ovidiameter (Table 2). Figures

for L. inermis, L. rothii and L. wotjulumensis are

exceptional, the maximum capsule diameter of L.

inermis being 6.8 mm and maximum ovidiameter

1.4 mm (Table 2).

Reproductive Modes

In the 24 species there are only three distinct repro-

ductive modes, one of which is represented by only

one species (Table 3), and in fact 20 species share the

generalised anuran pattern of laying eggs in water and

having aquatic larvae. In general the Australian hylids

are remarkably conservative in reproductive modes,

and presumably the high proportion of hylids in the

study fauna contributed to the limited overall diver-

sity. Elsewhere in sub-tropical faunas, diverse repro-

ductive regimes have been recorded (Salthe &

Duellman, 1973; Crump, 1974). Certainly in temperate

southern Australia, diversity of reproductive modes

is associated with leptodactylid rather than hylid spe-

cies (Tyler, Watson & Martin, 1981, Table 2). Never-

theless the seasonal aridity and limited forestation of

the Magela Creek system environment probably are

inimical to many reproductive modes. For example,

there is very limited leaf litter or other low layers of

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM 437

OCT. | NOV. | DEC. | JAN. | FEB. | MAR. APR.

Litoria bicol COLL LILELIEE. LEE TTEEEEZEE

7 Yoria bicolor LLL tit ty LSS LLLILLLLLLLL LLL LE LE

° ELL

= L. caerulea LEE: tit LLM yyy yyy" LLL

o : USULLLLLLLLLLL LULL LLL ZZ

fe | 4. roth CLL, C5 WLLILULU LILIA U ttt

L. rubella LLL LA

. EE LLLALLLL ALLELE ELLE LEE EEEEEEEEEEELE, LLLLLEPLLE LLL LL

L. inermis a 7 LLL Sv ee

. LLL =

= OE§G LILLE WMI LULA Le Kiko

Sy L. sp. nr. /atopalmata LL—____ ZZIIIED

acl

vu =

1 > tapi [eZ yyyygglrPYmn"/""" dada,

ge L. tornieri LI LEE LZ A

°o

SMELL, LP 1

oO L. nasuta _— COLL LLL CLLLEELELEL EE EEEEEEEED

‘ ‘ ZZ LLL: tj La ULL LEE a)

L. wotjulumensis EE

. LLL, Yih srr iggy

L. microbelos we EEE HHH

? EZ SLL ]

Cyclorana australis i KLE ZZ

‘ zzz LLLLILLLLLLIL ITA LILLE,

C. longipes EE:

Limnodynastes MM MLL LL LELLLLLLTILL HL

convexiusculus VALLI, LLLLLLLLLLLLLA

BZ LLL

L. ornatus LZ LLLLA OLLI)

LLL YUL

Notaden melanoscaphus ‘— = LLL

‘ yj MTThibhooyy M_DAM-—A>AA/>DPU>-Jqpz-: Lp JILL DLE

Ranidella bilingua MEE J™

Uperoleia inundata c MM I fPY"»-ILLELLLL Li

Litoria dahlit l 7

L. copland ZA =)

2 ‘ ma

a. L. personata LLL LLL LLL LE LLL LLL LLL EEE

=| 4. meiriana sae om ama =

& Megisto/otis eB;

5 lignarius

wi | | I | | |

OCT. NOV. DEC. JAN. FEB. MAR. APR.

438

REC. 8. AUST. MUS., 18 (18): 415-440

April, 1983

TABLE 2: CHARACTERISTICS OF SPAWN

Clutch Clutch No, of Capsule wa:

number size jelly diam, a mre Cl

envelopes (mm) :

median range median range

ee RY

) trali oo asd ITanare z° | 7.000 2 2,5 (2.3-2.8) 1.65 (1.5-1.8)

ete Sonsings 2 { 000-1 615 2 2.2 (1.9-2.5) 1.35 (1,2-1.5)

Litoria bicolor. - 2 50-195 3 2.4 (1.9-2.9) 1,00

Litoria caerulea 4 103-328 2 2.5 (2.2-2.9) 1,25 (1.1.4)

Litoria inermis . 23 7 96-330 3 6.3 (5.8-6.8) 1,35 (1.3-L.4)

Litoria sp. nr- fatopalmata 10 52-921 3 3.6 (3,4-3.9) 1.25 (1.2-1,3)

Litoria metriana - ; ges 5 - - ° F & ; 0 ba by

itera ren he 2 50-100 3 3.4 (3.253.7) «15 (1.01.3)

Litoria rothit. -- ! 504 3 5.0 (4.6-5.4) 1.35 (1.3-1.4)

Litaria rubella . l 715 3 2.1 (1.9-2.3) 1.05 (1.Q-1.1)

Litoria torniert 2 69-190 2 3.4 (3,2-3.7) 1.4 .

Litoaria woljulumensis 4 31-50 3 4.5 (4.2-4.9) 1.75 (1.6-1.9)

Limnodynastes ornalus - 19 155-1 632 | 1.6 (1,45-1.8) 1.15 (1,0-1,3)

Megistolotis lignarius’ ! 352 - - Pry 1.85 (1,8-1.9)

Notaden melanoseaphus 4 561-1 028 | 22 (2.6-2,9) 14° (1.3-1,55)

Ranidella bilingua - - 1 3.1 (2.5-3.7) 1.12 (1.05-1.2)

Uperoleia inundata 1 400 l 1.8 (1.5-2.0) 1.25 (1.1-1.4)

> From Tyler et al (1979)

TABLE. 3; REPRODUCTIVE MODES

AMONGST THE SPECIES AND GENERA OF THE FROG FAUNA OF

THE MAGELA CREEK SYSTEM

g 2

g Ka =

= é, 3 PA 3 S =

g 3 3 iS & = 3 5

S = = a 3 3 S = =

= 5 E & = = & = a

= = to =

oC I 3 = 2 4 5 a -

(a) Eggs in water, aquatic larvae. , 2 14 - 1 i 2 20

(b) Eggs in foam nest on water, aquatic

larvae . pews} of , pu - ~ 2 . = :

ceed

‘

high humidity necessary to create the conditions under

which direct development, or the delayed emergence

from egg capsules is physically possible. Evidently

similar factors operate in South America where the

greatest diversity of reproductive modes is associated

with environmental conditions rather than systematic

factors: the majority of modes occurring in species

occupying forest habitats (Crump, 1974; Lynch, 1979).

Developmental Spans

Water temperature and aggregation density both

affect the developmental span of tadpoles, although

there is substantial variation in developmental rates

within the progeny hatching from a single clump. Our

observations in the laboratory and field suggest that

this variation is a normal phenomenon, but that in

the wild the ‘slow’ developers die. In Table 4 we have

recorded the means and ranges of the shortest span

in each species. (The shortest span is the period taken

for the first individual of a clump to complete met-

amorphosis.) Recording the shortest span is realistic

because of the ephemeral nature of many of the pools.

Thus abbreviation of the developmental period ensures

survival, whereas those individuals with the longest

developmental span are most prone to perish as the

pool evaporates.

We reared in the laboratory 24 clumps of L. nasuta

spawn, and observed a distinct tendency for the spawn

laid late in the season to complete development in a

shorter span than that laid early in the season (Fig.

15).

At Jabiru water temperatures tend to be highest at

the early part of the wet season; we recorded tadpoles

in water at 42.7°C in early January. This is 3.5°C

higher than the maximum for any species of frog

known to Main (1968) but, as he suspected, is quite

typical of summer diel temperatures experienced by

eggs and tadpoles at localities in northern Australia.

(The maximum noted by Tyler, Davies and Martin

(unpublished) is 45.0°C in a rock pool occupied by

spawn of L. meiriana at Mitchell River Falls, W.A.

in February, 1979.)

REPRODUCTIVE BIOLOGY OF THE FROGS OF THE MAGELA CREEK SYSTEM

439

TABLE 4: DEVELOPMENTAL SPANS IN DAYS UNDER FIELD AND LABORATORY CONDITIONS

Field Laboratory

(ca, 30-40°C) (30°C)

n < n x +8.D.

Cyclorana australis 2-2... cece eee I 29 1 37 —

Cyclorana longipes ...0 0000000 cece ee 2 35 (33-37) 7 48 (31-66) 12

LUOrGBICOlg EA lta Seabee ag ft dbs ceed — — 2 53 (25-82) 40

Litoria Caeruled- <2 < ech cn We ene pom eee en _ —_ 10 66 (28-94) 29

Literia coplandt .....60 6000 00 occ cee ee eee 1 §2 1 = =

Litoria sp. nr latopalmata...... 000.0... 0. eee 1 54 7 96 (61-122) 23

Litoria meiriana.. 2... ee es ! 30 1 45 —

LOPE HASLEG... 3. cor eitgak Stewed oe OR bas oe —_ — 23 71.(31-120) 22

ETUOTER VOU Seg ty tare at 3 ft ot ee Sv nelle ns 4 1 65 2 144 (142-146) 3

TCO PI ATUB GHG | erie ates aos bile oa nomi lenesta te wislacis ] 37 5 45 (28-84) 22

Litoria tornieri . 0.00.6 ce ee eee ee — _ 11 97 (62-129) 24

Litoria wotjulumensis .....0000 0000600252000 — _ 5 71 (39-120) 37

Limnodynastes ornatus -. 2... 2-000 ee ee ee | 25 5 38 (21-63) {7

Megistolotis lignarius .......0000000 00 cece esa ee _ — _— 65! —

Ranidella bilingua ...........00..0.22.2-2-4,. 1 142 — = 7

Uperoleia inundata ... 0.0.0.0 022 ee ee ee _ — 1 99 =

'From Tyler et al. (1979)

2From Martin et al. (1980)

LARVAL SPAN

re) i) TS

8 3 3 8 5 S 3

ee ee ee ee | Ce en Ae) es Re fee) Gee |

iw)

g . *

P= .

oO ee .

(e}

Oo e Seee one

ba ees . . bed

FIG, 15. Larval life span of Litoria nasuta at differing egg

is]

deposition times.

Acknowledgements

We are very deeply indebted to Mr C. Miller who

bore the greatest responsibility in the rearing of the

tadpoles transported to Adelaide, and to the University

of Adelaide for providing the excellent controlled

temperature laboratory in which the work was con-

ducted. We are most grateful to Mr G. Mewett and

Mr B, Pennington of Ansett Airlines of Australia for

considerable assistance in the transportation of live-

stock,

Various persons assisted the field studies. In Dar-

ticular we thank Messrs G. Miles and I. Morris of

the Australian National Parks and Wildlife Service,

and our colleague Dr R. Marchant.

We thank Mrs Jean Russell-Price for typing the

manuscript and Mrs Ruth Hughes for expertly drafting

many of the figures.

Finally we express our gratitude to the Supervising

Scientist for the Alligator Rivers Region who funded

the study, and permitted the publication of this com-

ponent of the results.

440

REFERENCES

BARKER, J. and GRIGG, G. (1977)

Australian frogs.” (Rigby, Sydney.)

CHRISTIAN, C. S, and ALDRICK, J. M. (1977) “Alligator Rivers

Study. A review of the Alligator Rivers Region environmental

fact-finding study.” (Australian Government Publishing Serv-

ice, Canberra.)

COGGER, H. G. (1979) “Reptiles and Amphibians of Australia,”

2nd Edition. (Reed, Sydney.)

CRUMP, M. L. (1974) Reproductive strategies in a tropical anuran

community. Univ. Kansas Mus. Nat. Hist. Misc. Publ. (61):

1-68,

GOSNER, K. L. (1960) A simplified table for staging anuran

embryos and larvae with notes on identification. Herpetologica

16: 183-190,

HUMPHRIES, R. B. (1979) Dynamics of a breeding frog com-

munity. Ph.D. thesis, Dept of Environmental Biology, Aus-

tralian National University, Canberra. (Unpublished)

“A field gutde to

LYNCH, J. D. (1979) The amphibians of lowland tropical forests.

In W. E. Duellman (Ed.) “The South American Herpetofauna:

Its origin, evolution, and dispersal”. Mus. Nat. Hist. Univ.

Kansas, Monogr. (7): 189-215.

MAIN, A. R. (1968) Ecology, systematics and evolution of Aus-

tralian frogs. Adv. Ecol. Res. 5: 37-86.

MARTIN, A. A., TYLER, M. J. and DAVIES, M. (1980) A new

species of Ranidella (Anura: Leptodactylidae) from north-

western Australia. Copeia 1980(1): 93-99.

SALTHE, S. N. and DUELLMAN, W. E. (1973) Quantitative

constraints associated with reproductive mode in anurans.

In J. L. Vial (Ed,) “Evolutionary Biology of the Anurans”.

(University of Missouri Press, Columbia.)

April, 1983

SLATER, P. and MAIN, A. R. (1963) Notes on the biology of

Notaden nichollsi Parker (Anura: Leptodactylidae). W. Aust.

Nat. 8: 163-166.

STRAUGHAN, I. R. (1966) An analysis of species recognition

and species isolation in certain Queensland frogs. Ph.D. thesis,

University of Queensland (unpublished),

TYLER, M. J. (1962) On the preservation of anuran tadpoles.

Aust. J. Sci. 25: 222.

TYLER, M. J. (1979) The impact of European man upon

Australasian amphibians. 7n M. J. Tyler (Ed.) “The Status

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TYLER, M. J. and CROOK, G. A. (1980) Frogs of the Magela

Creek System. Department of Zoology, University of Adelaide

(mimeographed).

TYLER, M. J., DAVIES, M. and MARTIN, A. A. (1978) A new

specics of hylid frog from the Northern Territory. Trans. R.

Soc, S. Aust. 102: 151-157.

TYLER, M. J, and MARTIN, A. A. (1975) Taxonomic studies

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todactylidae) from north-western Australia, Aust. J. Zool.

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TYLER, M. J., WATSON, G. F. and MARTIN, A. A. (1981) The

Amphibia: diversity and distribution. pp. 1277-1301. In A.

Keast (Ed.) “Ecological Biogeography of Australia”. W. Junk,

The Hague.

WILLIAMS, A. R. (1979) Vegetation and stream pattern as indi-

cators of water movement on the Magela floodplain, Northern

Territory. Aust. J. Ecol. 4: 239-247.

RECORDS oF THE

SOUTH AUSTRALIAN

MUSEUM

VOLUME 18 NUMBERS 19-23 MAY 1984

No.

19 THE TAXONOMY OF THE ATRAX ADELAIDENSIS SPECIES —

GROUP (MACROTHELINAE: MYGALOMORPHAE) WITH NOTES ON

BURROWING BEHAVIOUR

by M. R. GRAY

20 THE PSOCOPTERA (INSECTA) OF SOUTH AUSTRALIA

—_"

by C. N. SMITHERS

HELMINTH TYPE — SPECIMENS IN THE SOUTH AUSTRALIAN

MUSEUM.

Il ACANTHOCEPHALA AND CESTODA

by LESLEY R. SMALES

HELMINTH TYPE — SPECIMENS IN THE SOUTH AUSTRALIAN

MUSEUM.

Il. TREMATODA

by LESLEY R. SMALES

23 AREVISION OF THE GENUS APLEROTUS DALLAS (HETEROPTERA:

PENTATOMIDAE: PENTATOMINAE) WITH DESCRIPTION OF A

NEW SPECIES FROM SOUTH AUSTRALIA

by IMTIAZ AHMAD, NASEER AHMAD KHAN and SYED KAMALUDDIN

SOUTH AUSTRALIAN MUSEUM

North Terrace, Adelaide

South Australia 5000

THE TAXONOMY OF THE ATRAX ADELAIDENSIS SPECIES-GROUP

(MACROTHELINAE: MYGALOMORPHAE) WITH NOTES ON

BURROWING BEHAVIOUR

BY M. R. GRAY

Summary

The funnel web spiders, genus Atrax, making up the adelaidensis species-group from South

Australia are described: Atrax adelaidensis sp. n., A. eyrei sp. n. and A. flindersi sp. n. Their

distribution in relation to that of the genus is outlined. The absence of a serrula in all species and the

distinctive modification of the first leg in males of A. adelaidensis are recorded. The unique burrow

structure comprising shaft plus side-chamber with trapdoor is described and the presence of a

surface door in early juvenile burrows is noted.

THE TAXONOMY OF THE ATRAX ADELAIDENSIS SPECIES-GROUP

(MACROTHELINAE: MYGALOMORPHAE) WITH NOTES ON BURROWING BEHAVIOUR

M. R, GRAY

The Australian Museum, Sydney

ABSTRACT

GRAY, M. R., 1982. The taxonomy of the 4trax ade-

laidensix species-group (Macrothelinae: Mygalomor-

phae) with notes on burrowing behaviour. Rec. S.

Aust, Mus. 18 (19): 441-452.

The funnel web spiders, genus Arrax, making up

the adelaidensis species-group from South Australia

are described: Atrax adelaidensis sp. n,., A. eyrel sp.

n. and A, flindersi sp. n. Their distribution in reta-

tion to that of the genus is outlined. The absence of

a serrula in all species and the distinctive modifica-

tion of the first leg in males of A. adelaidensis are

recorded. The unique burrow structure comprising

shaft plus side-chamber with trapdoor is described

and the presence of a surface door in carly juvenile

burrows. is noted,

INTRODUCTION

Funnel web spiders (genus Atrax O. P. Cam-

bridge, 1877) have a distribution that is essentially

gondwanian. They are commonly represented in

coastal and highland habitats from southern Queens-

land to Tasmania, North of the Gladstone area in

southern Queensland records are available for three

specimens only (all females), from Queensland,

Papua New Guinea and the Solomon Islands respec-

tively. These records have not been confirmed or

added ta since the orginal collections of 60 to 80

years ago and one of them definitely represents a

locality data errors Anacpsiada (syn, Afrax) ven-

tricosa Rainbow and Pulleine 1918. recorded from

Cloncurry, northwest Queensland in fact was col-

lected from Tamborine Mountain, southeast Queens-

land and is probably a juvenile A. validus Rainbow

and Pulleine, In the drier conditions to the west of

the Great Dividing Range their numbers fall off

rapidly, To the west of the Grampian Range area

in Victoria there appears to be a gap in the distri-

bution of the genus until it reappears again in the

highlands of the Mt. Lofty-southern Flinders Ranges

and the southern Eyre Peninsula of South Australia

(Fig, 32). These westernmost, isolated represen-

tatives of the genus Afrax form the adelaidensis-

group,

NOTATION FOR SPINES

Leg surface abbreviations for spine counts: pd

prodorsal; rd = retrodorsal; py = praventral; rv

May, 1984

South, New South Wales 2000

retroventral. The leg spination data given in the

holotype and paratype descriptions refers to the

range of variation encountered (total spine counts

per surface) both in these specimens and the remain-

ing type series; left and right side leg spine counts

are included,

Atrax adelaidensis group

Diagnosis: Characteristic burrow with — side

chamber closed by trapdoor. Cephalic area

moderately raised, serrula absent. Males lack apo-

physis on leg 2, sometimes with tibia and metatarsus

of leg 1 swoilen.

Comments: The presence of the adclaidensis-

group was first reported by Main (1964) who thea

referred them to the genus Hadronyche L. Koch,

1872. Main (1967, 1976) also noted the burrow

structure, including the presence of a side chamber

closed by a trapdoor. Gray and Sutherland (1978)

indicated that at least three species could be recog-

nised and referred them to the genus Alrax, noting

that their burrow structure set them apart from all

other members. of the genus. Their placement within

Atrax is clearly indicated by the presence of three

tooth rows in the fang grooye, spinose tarsi and

many cuspules om both the labium and maxillae,

The adelaidensis group shows two morphological

features of particular interest. These are the absence

of a serrula and the modification of the first legs

in males of A, adelaidensis, In a survey of mygalo-

morph genera Platnick (1977) recorded the pre-

sence of a serrula in an unidentified species of funnel

web spider, Examination of many eastern funnel web

specics by the present author indicates thut serrula

development in Atrax is highly yariable—in some

species the serrula is large and obvious, in others

reduced or absent (Plates 5-7), The modification of

the first leg in males of A. adelaidensis (Fig, 2) 1s

most unusual for this genus. In other funnel web

species it is the second leg that may show signifi-

cant modification in the form of tibial and metatarsal

apophyses. Not only the position but also the type of

modification differs in A, ddelaidensis, where the

entire first metatarsus and tibia are swollen. A slight

swelling of the second metatarsus is alsa evident.

This modification of the first leg may be of signi-

ficance in the context of the generic relationships of

dd2

Ajtrax, The systematic position of the genus as a

memiber of the family Dipluridac has Jong been

problematical, Diplurid genera possessing male

second leg modifications such as Evagrus Ausserer

and Allothele Tucker are not closely related to

Atrax, Simon (1892) used the presence of numerous.

cuspules on the labinm and maxillac as key charace

ters in his recognition of the genera Atrax, Porrothele

Simon and Macrothele Ausserer in his Macrotheleae

and Raven (1980) considers that these. geneta com-

prise a natural grouping within his Hexathelidae.

Zoogeographically, as well as morphologically and

behaviourally, the New Zealand genus Porrothele ts a

logical candidate for relationship with Avray.

Thickening of Jeg segments in these genera may be

independent derivations but could be additional

indicators of relationship.

The distribution limits of the Adelaidensis species

are not clear as yet. However, the general areas of

Species distribution can be approximately defined us

follows: 4. adelaidensis—Mt. Lofty Range and

adjacent coastal plain; A. eyrei—southern Eyre

Peninsula; A, flindersi—southern Flinders Range

(Fig, 32). These areas correspond with open forest

habitats of the type best developed in the Mt. Lofty

Rangs. In drier areas such as the Flinders Range

the spiders tend to be concentrated in tiparian

habitats where open forest (though with a reduced

understory) persists.

KEY TO FEMALES

1, Prodorsal surface of patella CE with 18 to 26 spines.

Pigmented lawer lateral surface of abdomen marked

with severa] paler bars .. ..., Atrax adelaidensis

—Prodorsal surface of patella ILL with 6 to 9 spines.

Lateral] abdominal surface without pale bars... . 2

2. Abdominal pigmentation almost uniformly distributed,

only slightly reduced laterally and ventrally M.O.Q,

length to posterior width ratio more than 1:2

Leno , Atras. ayrel

—Abdoniinal pipticniatiost porniusntritsd ‘dorsally and

posteriorly, lateral and ventral surfaces much paler,

M.O,Q, Jength to posterior width ratio less than 1;2

Puicxcctkcoarhias cevworevyr end ctasee 00a ys:net CELESTE, Fee Alrax flindersi

KEY TO MALES

1. Vibia and metatarsus of lag 1 swollen. Tibia 1 with

2 ventral spines distally; metatarsus I with 25-30

ventral spines .,...., Atrax adelaidensis

—Leg; | unmodified, ‘Tibia 1 with 6-10 ventral spines;

metatarsus L with 14-16 ventral spines |

A tite flindersi

Atrax adelaidensis sp. n.

(Figs. 1-11, pl. 5)

Diagnosis: Prodorsal surface of patella HL with

18 to 26 spines, 8 to 11 on patella ITV, Female

abdomen pigmented dorsally and laterally but three

REC, §, AUST. MUS. 1& (19): 441-452.

May, 1084

to five pale bars on lower lateral surface of abdo-

men; ventral abdomen pale. Tibia I and. metatarsus

T and II of males swollen, Male metatarsus I with

25-30 spines ventrally. Middle third of ventral fang

graave with few or no teeth in male.

Pemale—HOLOTYPE N1979146

Measurements (nim) and markings: Total Jength

(including cheliccrae) 18.98, Carapace length 6.78;

width 5.88; height 3.35, Abdomen length 9.30;

width 6.40, Colour pattern—Carapace dark glossy

brown-black, sparsely haired, Black pigment around

eyes, Dorsal and Jateral surfaces of abdomen dark

maroon-broWn. Ventrally and above the spinnerets

pigmentation greatly reduced. Three to five pale

bars extend from yentral surface up anto darkly

pigmented lateral surface, Above spinnerets, small

sector of pigmented cuticle cut off by narrow semi-

circular band of non-pigmented cuticle. Abdominal

cuticle finely rugose, depressed intervals between

rugosities formed by numerous, small, non-pig-

mented, sub-circular to elongate depressions. Along

dorso-lateral surfaces three pairs of pale: markings,

anterior pair roughly circular, posterior pairs thin,

laterally elongated lines.

Cephalathorax: Eyes—Size (mm): AME. 0,19,

ALE 0.32, PLE 0.26, PME 0,17. Interdistances

(mm); AME-AME 0,34, AME-ALE 0.24, ALE-

PLE 0.23, PLE-PME 0,13, PME-PME 0,73,

M.O.Q, length 0.51 mm; width, anterior 0.69 mm,

posterior 0.91 mm. M.O.0O. length/posterjor width

ratio 121.78, Anterior cye row straight, width 1.67

mm. Posterior eye row recurved, width 1.68 mm.

Carapace—longer than wide in ratio 1:00.87; raised

in head region, height/length ratio 1:2.02. Fovea

deep and strongly procurved, anterior margin

smoothly curved. Chelicerae—inner margins of fang

groove with 11 teeth (9 large) outer margin with

11 large teeth; centre of groove with 23 small teeth

in single row, less regular apically and basally.

Labium—wider than long in ratio 1.45;1, cuspules

numerous. Maxillae—divergent with shart, conical

lobe apically, Cuspules mumerous, few apically;

serrula absent. Sternum—subcircular; length 3,81

mm, width 3.62 mm, Surface gently convex with

thin cover of long and short hairs. Three pairs of

sigilla, posterior pair large and oval, anterior pair

very small, ciccolar.

Legs: 4123. Length (mm); I 15.45; 12 14.39;

Hi 11.64; IV 15.57. Spination: I—rmetatarsus vy

5-6, tarsus py 3-8, rv 3-4, v 0-1; 1l—melatarsus p

1, v 5-7, tarsus pv 4-11, rv 3-5; I1l—patella pd

18-26, tihia p 7-8, ¢ 2-3, metatarsus p 5-8, pa 2-4,

rd 2-3, v 5-7, tarsus py 10-13, ry 2-5; ['¥V—patella

pd 8-11, tibia p 2, metatarsus p 6-7, rd 0-2, v 6-7,

tarsus py 10-12, Trichobothria in single raw on

443

ATRAX ADELAIDENSIS SPECIES-GROUP

FIGS. 1-11: Atrax adelaidensis. 1, Carapace, dorsal (female), x6; 2, Leg 1, retrolateral (male), x11; 3, Leg 3, prolateral

(female): patella, tibia and metatarsus, x11; 4, Male copwatory organ, x20; 5, Female internal genitalia, dorsal,

x20; 6, Eyes (female), x20; 7, Body, dorsal (male) x6; 8, Palp, prolateral (male), x6; 9, Tooth pattern in fang

groove, left hand side (female), x20; 10, Tooth pattern in fang groove, right hand side (male), x20; 11, Body, side

view (female), x6.

444

larsus, bothria collariform, Superior tarsal claws

with three to six pectinations, inferior claw smooth.

Tarsal scopulae absent. Tarsal organ distal to tri-

chabothria, a low circulac mound with faintly

sculpted, concentric grooves,

Abdomen; Spinnerets—Posterior lateral spin-

merets: length 2.35-2.58 mms; width 0,79-0,82; ratio

of length of apical, middle and basal segment 2.2:

1:1.6; basal separation 1.10 mm; terminal segment

conical, longer than wide in ratio 1,95;1, Posterior

median spinnerets: Jength 1.02; width 0.52 mm,

Genitalia—a pair of spermathecae open ventrally

into common copulatory bursa, Middle part of each

spermatheca swollen.

Male—PARATYPE N1979145

Measurements (mm) and markings: Total length

(including chelicerae) 16.1. Carapace length 5.92,

width 5.58, height 2,71, Abdomen length 6.8, width

4.8, Colour pattern—similar to female except that

the dorso-lateral abdominal markings more evident;

lateral abdomen uniformly pigmented, no pale bars

present.

Cepholathorax: Eyes—Size (mm): AME 0.23,

ALE 0,32; PLE 0.23; PME 0.14. Interdistances

(mm): AME-AME 0,17; AME-ALE 0.16; ALE-

PLE 0.20, PLE-PME 0.11, PME-PME 0.69:

M.O,Q. length 0.47 mm, anterior width 0,61 mm,

posterior width 0.97 mm. M.O,Q. length/posterior

width ratio 1;2,06, Anterior eye row slightly pro-

clirved, width 1.55 mm. Posterior eye row recurved,

width 1.60 mm, Carapace—longer than wide in

ratio 1;0.94, Raised in head region, height/length

ratio 1:2.18. Chelicerae—inner margin of fang

groove with 10-11 teeth (9 large): outer margin

with 13 teeth (11-13 large); central groave with a

Taw of 2-4 small teeth in apical third, none or

reduced in central third, a row of 7-10 small teeth

in basal third. Palp—distal prodorsal femur with

4-6 sinuous bristles; patella with 4-7 dorsal apical

bristles, Embolic process of palpal bulb short

and broad with well developed apical flange. Labium

—wider than long in ratio 1:1.61, cuspules

numerous, Sternum—tength 3.63 mm, width 3.03

mm.

Legs; 1423 (or 4123), Length (nim): 1 18.41

(19,30); 11 17.55 (18.93); If 14.39 (15,35); IV

17.85 (19.60). Anterior legs modified, tibia I and

metatarsus I swollen; metatarsus IT weakly swollen.

Tarsi weakly scopulate. Spination: I—tibia v 1-2,

metatarsus p 0-2, y 25-30, tarsus py 10-12, rv 7-11,

v 0-2; Il—femur d 0-1, patella r 0-1, bia p 0-1,

v 1-2, metatarsus p 0-1, r 0-2, y 12-16, tarsus py

10-16, rv 6-21, v Q-1; [1l—patella pd 18-23, tibia

pd 9-11, ¢ 3, ¥ 1-7, metatarsus p 8-13, pd 0-4, r

REC, S. AUST. MUS, 18. (19); 441-452,

May, 1984

1-6, rd 2-5, v 8-14, tarsus p 12-16, r 0-2, rv 6-9,

v 0-1; IV—femur d 4, patella pd 7-8, tibia p 2-3,

r 2-4, d 1-2, v 2-4, metatarsuis p 8, rd 1-2,.v 8-12,

tarsus p 14-17, rv 6-11, v O-1_ Single row of 3 to 5

weak bristles on dorsal surfaces of femora I, If, IV.

Abdomen; Spinnerets—postcrior lateral spin-

nerets; length 2.60 mm; maximum width 0.56 mm}

ratio of length of apical, middle and basal segments

1.55:121.79; basal separation 0,95 mm; apical sep-

ment conical, length/width ratio 2/82:1. Posterior

median spinverets; length 0,77 mm, width 0.31 mm.

Material examined: Holotype female (N1979146

S.A. Mus. coll), Hackuey, Adelaide, S.A. 16.11,

1973, J. Batt; burrow in sloping, littered ground on

banks of Torrens River. Paratype male (N1979145

S.A. Mus. coll), St. Peters C.G.S., Adelaide, S.A,

3,6,1971, D, Edwards Paratype female (KS 4511

Aust, Mus. call), Beefacres Recreation Reserve,

S.A., 16.11.1973, N. Miles; found in damp sandy

soil under tree stump. Paratype male (KS 4512

Aust, Mus. coll.), Belair, Mt Lotty Range, S.A.

24.5.1972, Two paratype males (N1979147.

N1979148 S.A, Mus, coll.), Clare, S.A, 5.8,1979.

Paratype female (N1979149 S_A. Mus coll.), Ros-

trevor, Nr. Adelaide, $.A., May 1974, B. Brewer; in

burrow, shaded alluvial soil by creek, Juvenile,

Hahndorf, Mt. Lofty Range, S.A.

Atrax eyrei sp. v1.

(Figs. 12-19)

Diagnosis; Prodorsal surface of patella ITT with

8 to 9 spines; O to 1 on patella TV, Abdominal

pigmentation almost uniform, only slightly paler

laterally and ventrally. Posterior median cyes widely

separated; M,0.Q, Jength/posterior width ratio

much greater than 1:2,

Female—HOLOTYPE KS 4509

Measurements (mm) and markings: Total length

(including chelicerae) 16.14, Carapace length 6.62;

width 5.93; height 3.38, Abdomen length 8.1; width

5.9. Colour pattern—Carapace dark, glossy brown-

black, sparsely haired. Black pigment around eyes.

Abdomen dark maroon-brown dorsolaterally, ventral

surface only slightly paler, Abdominal cuticle finely

rugose with small, subcircular to elongate nonpig-

mented depressions between rugosilies. Dorsoiateral

abdominal surfaces with four pairs of non-pigmented

Markings, anterior pair sub-circular, remainder

laterally elongate.

Cephalothorax; Eyes—Size (mm): AME 0.23,

ALE 0.30, PLE 0,19, PME 0.13, Tnterdistances:

AME-AME 0.28, AME-ALE 0.20, ALE-PLE 0.18,

PLE-PME 0.06, PME-PME 0.91, M.O.Q. length

0.47, anterior width 0,71, posterior width 1.21.

M.0.Q. fength/posterior width ratio 1;3,37.

ATRAX ADELAIDENSIS SPECIES-GROUP dd5

TARR AAA

: the

FIGS, 12-19; Atrax evrei. 12, Body, dorsal (female), x6; 13, Abdomen, side view (female), x6; 14, Tooth patlern in

fung groove, left hand side (female), x20; 15, Spinnerets, ventral (female), x20, 16, Femule internal genitalia,

dorsal, X20; 17, Eyes, dorsul (female), x20; 18, Leg 3, prolateral (female); patella, tibia and metatarsus, x Jl, 19,

Maxillae, labium and sternum, ventral (female), x6.

Anterior eye row straight, width 1.62 mm) posterior

eye row recurved, width 1.61 mm. Carapace—

longer than wide in ratio 1:0,90; raised in head

region, height/length ratio 1:1.96. Fovea procurved,

anterior margin slightly indented on each side of

midline. Chelicerag—inner margin of fang grooye

with 13-14 teeth (J0 Jarge), outer margin with

10 large teeth, central groove with 20-23 small teeth

in irregular cow. Labium—wider than long in ratio

1.45:1, numerous. cuspules. Maxillae—divergent

with short conical Jobe apically, serrula absent;

cuspules numerous, few apically, Sternum—subcir-

cular; length 3.90 mm, width 3.65 mm. Surface

gently convex with thin covering of long and short

hairs. Three pairs of sigillae, posterior pair large

and oval, anterior pair very small, circular.

Legs; 4123. Length (mm): I 16.02; Il 14.40;

THE 12.41; IV 16,03. Spination: I—metatarsus v 8,

tarsus pv 3, rv 5, v 1; [[—metatarsus p 0-1, v 9,

446

abe

PLATES 1-4, Atrax flindersi. 1, open forest habitat, Mt. Remarkable, S.A.; 2, juvenile in fee

REC. 8S. AUST. MUS. 18 (19): 441-452.

May, 1984

ding position, door stand-

ing open, silk trip lines radiating from burrow rim; 3-4, surface door of juvenile, 3, closed; 4, open.

tarsus pv 4-5, rv 4; I[I—patella pd 8-9, tibia p 4,

r 1-2, metatarsus p 4-5, pd 2, rd 1-2, v 9, tarsus

pv 8-9, rv 4; 1V—patella pd 0-1, tibia p 1, meta-

tarsus p 2, v 5-8, tarsus pv 7-9, rv 2. Trichobothria

in single row on tarsus, bothria collariform. Superior

tarsal claws with 5-6 pectinations, inferior claws

smooth or with 1 pectination. Tarsal scopulae

absent.

Abdomen: Spinnerets—posterior lateral spin-

nerets: length 2.67-2.72 mm; width 0.90-0.94 mm;

ratio of length of apical, middle and basal segments

1.08:1:1.60; basal separation 1.30 mm; terminal

segment conical, longer than wide in ratio 1.86:1.

Posterior median spinnerets: length 0.94 mm; width

0.47 mm. Genitalia—pair of short, broad sperma-

thecae opening ventrally into a common copulatory

bursa.

Material examined: Holotype female (KS 4509

Aust, Mus. coll.), 6.5 km south of Coulta, Eyre

Peninsula, S.A. 18.12.1952. B. Y. Main (cat. no.

52/600). Paratype female (KS 4510 Aust. Mus.

coll.), between Coulta and Wangary, Eyre Penin-

sula, S.A., 18.12.1952. B. Y. Main (cat. no. 52/

611). Juvenile male, Nr. Port Lincoln, Eyre Penin-

sula, S.A., 16.12.1952, B. Y. Main (cat. no. 52/

540).

Atrax flindersi sp. n.

(Figs. 20-30, pls 1-4, 6)

Diagnosis: Prodorsal surface of patella ILI with

5 to 10 spines; 0 to 3 on patella TV. Abdominal

pigmentation mainly dorsal; lateral and ventral

surfaces paler. Femur I of male with 7-9 spines

dorsally, femur If of male with 5-7. Tibia I of male

with 6-10 ventral spines, tibia Il of male with 8-10.

ATRAX ADELAIDENSIS SPECIES-GROUP

447

PLATES 5-8. 5, Atrax adelaidensis, tarsal organ, x1250. 6-8, apical lobe of maxilla showing serrula structure variation

in three species of funnel web spider; 6, A. flindersi (serrula absent), x165; 7, A. robustus, x90; 8, A. versutus, x60.

Female—HOLOTYPE KS 983

Measurements (mm) and markings: Total length

(including chelicerae) 20.02. Carapace length 6.90;

width 6.12; height 3.37. Abdomen length 9.1; width

7.3. Colour pattern—carapace dark glossy brown-

black, sparsely haired. Black pigment around eyes.

Abdominal pigmentation dark maroon-brown dor-

sally, pigmented area narrow anteriorly, broader

posteriorly. Lateral and ventral abdomen very lightly

pigmented and tinged with pink. Abdomen cuticle

finely rugose with small subcircular to elongate

non-pigmented depressions between rugosities.

Dorso-lateral abdominal surfaces with four pairs of

non-pigmented markings, anterior pair subcircular,

remainder elongated and rather indistinct.

448

Cephalothorax; Eyes—Size (mm): AME 6.23,

ALE. 0.34, PLE 0,31. PME 0.18. Interdistances:

AME-AME 0.27, AME-ALE 0.25, ALE-PLE 0.26,

PLE-PME 0,11, PME-PME 0.78. M.O.Q, length

0,65, anterior width 0.67, posterior width 1.12.

Anterior eye row straight, width 1,65 mm; posterior

eye row recurved, width 1.80 mm. Carapace—longer

than wide in ratio 1:0.89; raised im head region,

height/length ratio 1:2.01. Fovea deep and strongly

procurved, anterior margin with broad shallow in-

dentation centrally, Chelicerse—inner margin of

fang groove with 14-16 teeth (12-15 large); outer

margin with 15-18 teeth (12 large); central groove

with 23-27 small teeth in irregular row, divided into

two rows apically, Labium—wider than long in

ratio 1.4071, numerous cuspules. Maxillac—diver-

gent, ameally with short conical lobes, serrula

absent; cuspules numerous, few apically, Sterrum—

subcircular to oval; length 4.04 mm, width 3,66 mm.

Surface gently convex with thin covering of long

and short hairs, Three pairs of sigilla, posterior pair

large and oval. Anterior pair yery small, circular.

Legs; 4123. Length (mm): T 16,33; IT 14.93;

IIT 13,36; TV 16,79. Spination: I—metatarsus v 7-9,

tarsus. pv 4-5, rv 5-6; [[—mietatarsus p 0-1, v 8-10,

tarsus pv 4-5, rv 4-8; I[1]—patella pd 5-10, tibia

p 4, r 1-2, v 6-1, metatarsus p 2-5, pd 2-3, rd 2,

v 7-10, tarsus py 10-14, rv 4, y 0-1; [V—patella pd

0-2, tibia p O-1, y 0-1, metatarsus p I-4, rd 0-1, v

4-8, tarsus pv 6-15, rv 0-3. Superior tarsal claws

5-7 pectinations, inferior claws smooth, Tarsal sco-

pulae absent. Tarsal organ distal to trichobothria,

low circular mound with faint concentric grooves.

Abdomen: Spinuerets—posterior lateral spin-

nerets; length 3,53 mm; width 1.02 mm; ratio of

length of apical, middle and basal segments 1.78: 1:

1,62; basal separation 1.48 mm; terminal segment

conical, longer than wide in ratio 2,56:1, Posterior

median spinnerets: length 0.94 mm, width 0.47 mm,

Genitalia—a pair of rather slender spermathecae,

terminally swollen, opening ventrally into common

copulatory bursa.

Male—PARATYPE KS 980

Measurements (mini) and markings; Total length

(including chelicerae) 16.80. Carapace length 6,23;

width 6.29; height 3.06, Abdomen length 7.50;

width 5,70, Colour pattern—as for female; paired

abdominal markings slightly more emphatic,

Cephalotherax: Eyes—Size (mm): AME 0.26,

ALE 0,35, PLE 0.23, PME 0,15, Interdistances

(mm): AME-AME 0.26, AME-ALE 0,12, ALE-

PLE 0.24, PLE-PME 0.12, PME-PME 0,82,

M,O.Q. length 0.58, antetior width 0.74, posterior

REC. 5, AUST, MUS, 18 (19): 441-452,

Muay, 1¥d¢

width 1.18. M.O.Q, length/posterior width ratio

1:2.03, Anterior eye row straighi, width 1.59 mm.

Posterior eye row recurved, width 1.64 mm, Cars-

pace—slightly wider than long in ratio 1:0.99.

Raised in head region, height/icngth ratio 1:2.04.

Foyea decp, procurved, anterior margin smoothly

curved, Chelicerae—inner margin of fang groove

with 12 teeth (10-12 Jarge), outer margin with 14-

15 teeth (12-13 large), central groove with 18-19

small teeth in single row, Palp—temur with 7 pro-

dorsal, apical spines; patella with 14 dorsal, apical

spines. Embolic process of palpal bulb straight and

slender with well developed apical Hange. Labium—

Wider than long in ratio 1.54;1, Sternum—subcir-

cular; length 3.46 mm; width 3.26 mm. Sigillae

as in female but placed closer to lateral margins

and anterior sigilla much more distinct.

Legs: 4123. Length (mm): 1 19,46; I 18,24;

TH 17,82; TV 21.39, First and second legs unmodi-

fied, Tarsi of legs IIT and IV mildly swollen. Tarsal

Scopulae present, strongly developed on legs II,

TIT and SY. Spination: I—femur p 0-2, d 7-9, tibia

p 0-1, v 6-10, metatarsus p 0-2, v 12-15, tarsus

py 8-9, rv 9-12, v 1-3; I—femur d 5-7, tibia v

8-10, metatarsus y 12-13, tarsus py 9-10, rv 8-10,

v O-1; Til—femur d 0-1, patella pd 5-8, tibia p

4-7, r 1-4, d 1-4, d 1, v 4-6, metatarsus p 2-8; r

2-3, rd 2-3, y 11-15, tarsus p 8-12 r 0-2, rv 8-14;

IV—femur d 5-6, patella p 0-3, tibia p 1-4, r 3-5,

d 0-3, y 4-6, metatarsus p 3-5, re 0-2, v 11-15,

tarsus p 9-15, rv 8-11,

Abdomen: Spinnerets—Posterior lateral spin-

nerets: length 3.30 mm; width 0.71 mm; ratio af

length of apical, middle and basal segments 1.12:1:

1.24; basal separation 1.40 mm; terminal segment a

slender cone, longer than wide in ratio 3.49: 1. Pos-

terior median spinnerets cylindrical; length 0.91 mm,

width 0.36 mm,

Material examined: Holotype female (KS 983

Aust. Mus. coll.), Mt, Remarkable (lower slopes),

3 km, north of Melrose, Flinders Range, S,A,,

23.4.1973, M, and G. Gray. Dry open and eucalypt

forest, burrow with side passage in littered riparian

slope; juvernles in side passage, Paratype male (KS

980 Aust. Mus, coll.), Mt, Remarkable (lower

slopes), 3 km north ot Melrose, Flinders Range,

S.A... 25,4,1973, M. and G. Gray; pitfall trap in

dry open forest on southern eastern slope, Paratype

males (KS 979 and KS 985 Aust. Mus. coli.) and

female (KS 981 Aust. Mus. coll.), Mt. Remarkable

(lower slopes), 3 km north of Melrose, Flinders

Range, $.A,. March-April 1973, M. and G. Gray;

dry open euclaypt forest, sloping ground with

medium-heavy litter cover,

ATRAX ADELAIDENSIS SPECIES-GROUP 449

“9

)

Fal

)

}

ceeP9 9 OF on 50

rey

pal

0°

-c

2 0

ia §

e%9 0°250

wewt0U

ofa ok OF

er* geoo

FIGS, 20-30: Atrax flindersi, 20, Leg 3, prolateral (female): patella, tibia and metatarsus, x11; 21, Abdomen, side view

(female), x6; 22, Carapace, dorsal (female), x6; 23, Palp, prolateral (male), x11; 24, Male copulatory organ, x20;

25, Tooth pattern in fang groove, right hand side (female), x20; 26, Tooth pattern in fang groove, right hand

side (male), x20; 27, Eyes, dorsal (female), x20; 28, Body, dorsal (male), x6; 29, Female internal genitalia, dorsal,

x20; 30, Leg 1, retrolateral (male), x11.

450

BEHAVIOURAL OBSERVATIONS ON

ATRAX FLINDERSI

Field observations

Summer collections of A. flindersi were made on

riparian slopes in the summer-dry, open eucalypt

forest habitats of the Mt. Remarkable area (Plate

1). The retreat is a ground-burrow consisting of a

main shaft plus a side chamber closed by a door

(Fig. 31). The burrow has a single entrance which

is hidden amongst the well-developed litter layer of

the forest floor. The burrow may be sited in an

open situation or in the shelter of a grass tussack,

log or stump. This retreat structure contrasts

markedly with that typical of many easterm members

of the genus occupying wetter forest habitats (Gray

1978),

The entrance to the adult burrow consists of a

silk collar which in 4. flindersi is produced around

one side to form a short cowl-like structure which

lies limply across the entrance partially occluding

it, The silk of both collar and cowl is well disguised

hy attached litter and fine soil particles.

The silk lining of the main shaft is entire and

closely adherent to the soil walls. Total burrow

depth varied from 21 to 25 mm; burrow diameter

varied from 1-1.5 cm near the entrance to 2-2,5

cm in the end chamber. The general inclination of

the burrow is at 90° to the ground slope, though the

end chamber below the lJevel of the side chamber

may be less steeply inclined,

BIG. 31. Burrow structure of Alrax flinderst

REC_S. AUST, MUS, 18 (19): 441-452

May, 1984

The silk-lined, flask-shaped side chamber (length

3-4.5 em; widih 2-2.5 em in adult burrows) was

placed between three-quarters and four-fifths of the

way down the burrow of adult specimens, In the

burrows of juveniles the side chamber may be only.

half-way or even less down the burrow. The side

chamber is closed by a strong, externally concave,

s0il trapdoor, hinged vertically (diameter 0.85-1 5 cm

in adult burrows), The usual orientation of the side

chamber was slightly upslope, but this was not in-

variable, A shallow excavation in the wall of the

main shaft at the leyel of the trapdoor probably

ptovides space for manocuvring during door open-

ing operations.

The functions of the side chamber seem mainly

concerned with the provision of a secure refuge

against predators, particularly when moulting, and

a protective brood chamber for the eggs and spider-

lings. Several Afrax burrows were found to contain

occupants which could represent predators—a cen-

tipede, a large carabid beetle and a spider—the latter

belonged to the genus 4nane and was found with

the remains of the presumed original occupant. One

burrow contained a brood of 21 spiderlings in the side

chamber, The need to function as a brood chamber

during the hot summer months could account for the

trend suggesting that adult side chambers are found at

greater depths relative to total burrow depth than

those of fuvenjles; brood chamber function would

require long term constancy of high humidity—

moderate temperature conditions which would not

be so eritical for the temporary predator/moulting

refuge function which must be the primary role of

the shallower side chamber of juvenile burrows.

Morphological adaptations to aridity are not

readily upparent in any of the adelaidensis species.

Consequently, in swnmer-drought regions such as

the Flinders Range the construction of a relatively

deep, vertical ground burrow has obvious advan-

tages in avoiding desiccation and overheating. Soil

characteristics may also be significant, particularly

in relation to their water-holding ability which

directly influences soil and burrow air-water humi-

dity, The burrows of A. /lindersi were excayated

in a yellow-grey, gritty loam. Evidence from trap-

door spider burrow studies indicates that such soil

would require severe surface drying conditions to

render the soil air significantly undcrsaturated helow

10 cm from the surface (Gray 1969). The surface

litter cover would provide a further cushioning

effect against environmental drying.

Laboratory observations

Fifteen juveniles (probably second instar) taken

from the side chamber of a burrow in the ML.

Remarkable area were set up individually in glass

jars with 10 cm depth of lightly littered soil. All

ATRAX ADELAIDENSIS SPECIES-GROUP

eS TTL TIOIN Ge PURE WER

SPIDERY GLhals aFnay

451

4 adelaidensis

m eyrei

e flindersi

FIG, 32, Distribution records for the Atrax adelaidensis species-group in South Australia. Inset—overall distribution of

the genus Atrax.

juveniles excavated small burrows and twelve of

them constructed flap-like, but quite rigid, soil

trapdoors closing the surface burrow entrances

(Plate 2, 3). The remainder constructed an inde-

finite soil-silk entrance mound. When hunting, the

juveniles would prop the door wide open at an

angle of 70-100° and sit in the burrow mouth with

the tarsi of the palps and first two pairs of legs

resting on the burrow rim (Plate 4). Some shect-

like silk was placed on the ground around the burrow

entrance, usually radiating outwards as prey trip-

lines. In catching prey the spiderlings were seen to

fully emerge from the burrow up to a distance of

2 cm, Examination of these burrows revealed that

no side chamber was present, the burrow consisting

of a simple, vertical shaft. The discovery of surface

door building behaviour in A. flindersi spiderlings

is interesting in view of its absence in the adults

and larger juveniles. However, in the context of the

environmental conditions that the juveniles face,

adaptive advantages can be seen. The major advan-

tages would seem to be that the microclimate within

a shallow burrow closed by a surface door is likely

to be more stable than that found within an open

or partially closed burrow; and in the absence of a

side chamber refuge, the surface door may serve to

decrease predation on spiderlings. The burrows of

larger juvenile spiders kept in captivity showed no

sign of a surface door, a mixture of soil and silk

usually partially closing the entrance leaving only a

small opening to one side or none at all. A side

chamber with door was always present in these

burrows.

The initial stages of burrowing activity by some

of these juveniles were observed. The litter adjacent

to the burrow site was silked together and drawn in-

wards. The soil surface to be excavated was thinly

silked over, a process repeated with each phase of

digging, and the chelicerae used to loosen the soil.

Both chelicerae and palps combined to scrape up

and carry away the soil as silk-impregnated bundles

which were initially deposited (and tamped down

by the palps) just outside the burrow lip. Frequent

360° traverses were made with the spinnerets about

the lip area of the burrow so building up a soil-

impregnated silk collar attached to the surrounding

452

litter. This collar was periodically raised slightly

by the palps and first legs and the whole jerked in

centrally.

At this stage a shallow inclined burrow approxi-

mately 2 cm deep and 1.5 cm wide had been ex-

cayated. A second opening or “window”, about

half the size of the original entrance, was then

made in the thin roof of the burrow; this “window”

apparently proyided room for manoeuvring while

excavating the bottom of the burrow, the spider's

abdomen being protruded out of it during turning

and digging operations. Subsequent activity took

place entirely within the burrow. Soil was pro-

gressively added to the internal margins of both

entrance and window until the latter was com-

pletely closed over. The entrance was eventually

reduced to a small irregular hole 2-3 mm wide

REC, 8. AUST, MUS. 18 (19): 441-452

May, 1984

located near the lower edge of the original entrance.

This location arose because the direction of deposi-

tion of soil proceeded from the upper toward the

lower margin of the entrance. The soil-impregnated

silk curtain so formed over the burrow entrance

can be seen to have affinities both with the cowl-like

entrance structure noted in the field situation and

the soil door built by captive second instar juveniles.

ACKNOWLEDGEMENTS

My thanks to Dr. B. Y. Main for the loan of speci-

mens from the Eyre Peninsula and locality data for

the Melrose area, I am grateful to Mr. D. C. Lee

of the South Australian Museum for his co-operation

in the provision of facilities and the loan of material.

Drawing assistance was provided by G. E. J. Gray.

REFERENCES

GRAY, M. R. 1968. “Adaptations to Aridity in Trapdoor

Spiders”. M.Sc. Thesis, University of Western Australia,

Library.

GRAY, M, R., and SUTHERLAND, S. K. 1978. Venoms

of Dipluridae: 121-148—/n “Systematics and Distri-

bution in Arthropod Yenoms” (S. Bettini, ed.) Hand-

book Exper. Pharmac., 48: 121-148.

MAIN, B. Y. 1964. “Spiders of Australia”

Jacaranda Press, Brisbane.

(2nd ed.).

MAIN, B. Y. 1976. “Spiders.” Australian Naturalist Library

—wWilliam Collins, Sydney.

PLATNICK, N, I. 1977. The Hypochiloid Spiders: a

Cladistic Analysis, with notes on the Atypoidea (Arach-

nida, Araneae). Amer. Mus. Novit, 2627: 1-23.

RAVEN, R, 1980. The evolution and biogeography of the

mygalomorph spider family Hexathelidae (Araneae,

Chelicerata). J. Arachnol. 8: 251-266.

SIMON, E. 1892, “Histoire Naturelle des Araignees”, Vol.

1, 2nd ed., Libraire Encyclopedique de Roret, Paris.

THE PSOCOPTERA (INSECTA) OF SOUTH AUSTRALIA

BY C. N. SMITHERS

Summary

The number of Psocoptera known from South Australia is increased from 8 to 45, including 16 new

species and 21 new records for the State. The fauna appears to be predominantly associated with

bark and dried leaves as opposed to green foliage. The relationships of the fauna are discussed

briefly within the context of present records of the Australian fauna.

THE PSOCOPTERA (INSECTA) OF SOUTH AUSTRALIA

Cc. N. SMITHERS

The Australian Museum, 6-8 College Street, Sydney, N.S.W. 2000

ABSTRACT

SMITHERS, C. N., 1982. The Psocoptera (Insecta) of

South Ausiralia. Rec. S. Aust, Mis. 18 (20): 453-491,

The oumber of Psocoptera known from South

Australia is increased from 8 to 45, including 16

new species and 21 new records for the State. The

fauna appears to be predominantly associated with

bark and dried leaves as opposed to green foliage.

The relationships of the fauna are discussed briefly

within the context of present records of the Aus-

tralian fauna.

INTRODUCTION

Two significant recent collections of Psocoptera

from South Australia have prompted a detailed

study of all material available from this area the

results of which are presented here.

The first Psocoptera from South Australia were

described by McLachlan (1866). He described

Psocus pallipes McLachlan from Adelaide, trans-

ferring it to Prepsocus in a postscript to the same

paper. It has since been recorded from Western

Australia, Queensland and New South Wales. He

also described Psocus striatifrons McLachlan from

“Australia, meridionali”, transferring it immediately

to Stenopsocus Hagen, This species has not been

recorded again and it is possible that it may not

be South Australian even if it docs occur elsewhere

in southern Australia, The type locality is not pre-

cisely known, Stenvpsocus is a Palaearctic and

Oriental genus with one species in New Guinea and

eastern Australia which has been found as far south

as northern New South Wales. §. striatifrons is cer-

tainly not the same species. Detailed study of the

type of S. striatifrans is necessary to clarify its status

and confirm its generic position. Banks (1939)

described Zelandopsocus sinuosus Banks from “Mt.

Lofty Range”. This species is now placed in Aus-

tropsocus Smithers and has been recorded from

Tasmania, Victoria, Australian Capital Territory,

New South Wales and Queensland, Smithers (1964a)

recorded the widespread Lepinotus reticulatus Endet-

lein and Psyllipsocus ramburii Selys-Longchamps

from caves in South Austraha and Phlotodes aus-

tralis (Brauer) from several localities (Smithers

1964b), ‘Thornton and New (1977) recorded

Haplephallus bundoorensis New and H. medialis

Thornton and New from South Australia. The

former is known from Victoria, Queensland and

New South Wales and the latter fram New South

Wales, Australian Capital Territory and Victoria.

So. by 1977 eight species had been recorded from

South Australia,

This paper is based on collections in the Aus-

tralian Museum and the South Australian Museum.

These include 41 species, of which four have already

been recorded fram South Australia. Four of the

recorded species are not represented in ihe present

material, 21 are new records for the State (marked

with ao asterisk in the Jist belaw) and 16 are new

species, In all, 45 species (including $. striatifrons)

are now known from South Australia,

The Psocoplera so far recorded from South Aus-

tralia are nearly all inhabitants of bark or dead

leaves; there seem to be very few species on green

foliage, Precise habitat information is, however,

available for only a small proportion of the material.

The species which may be associated with green

foliage are Caceilius seenifuscatus (Tillyard), Stenep-

socus siriatifrons (if it is a Stenapsocus) and Cladio-

neura punctata sp. n. although even these cannot be

confirmed as inhabitants of fresh leaves, It seems

unlikely that the main method of collecting, by

beating, would give samples biassed toward bark

dwellers as opposed to inhabitants of green Icaves.

The fauna as so far known seems, therefore, to be

poor in inhabitants of green leaves,

Echinopsocus grayi sp. 0., Pachyiractes rugesus

sp. mn. and Sphaeropsecepsis recens (Hickman) are

leaf litter insects. chmepteryx (Loexophelia)

brunnea Smithers is an inhabitant of twigs and

smaller branches. The two species of Tragiidae,

Cerobasis guestfalica (Kolbe) and Lepinotus reti-

culatus and also Psyllipsocus ramburii are very

widespread domestic species which have all been

taken in outdoor habitats, the last two being known

also from caves. Lachesilla pedicularia (L.) has

been recorded from several parts of the world, In

Europe it is an inhabitant of trees and shrubs and is

often found in homes, The only South Australian

record is fram packing straw from England. The

two species of Ectopsocidae, Ectopsecus californicus

(Banks) and E, cetrdtus Smithers, are inhabitants of

dried leaves cither on plants of as leaf litter as are

Pentacladus eucalypti Enderlcin and Propsocus pal-

lipes. Mesopsocus reticulatus sp. n. is the first species

of the family to be recorded from Australia, Several

454

members of this family in Africa occur in fairly dry

habitats on the twigs of shrubs and the European

species are found on bark of branches and twigs.

The most important tamily of bark dwelling psocids

is the Psocidae. The family is well represented in

South Australia, comprising about 30% of the

psocoptcran fauna so far known. The two myop-

socids, Phlotodes australis and Ph. hickmani

(Smithers), feed on algae and fungi on bark; the

former is very common on damp suburban paling

fences,

Until the fauna of the whole of the continent is

hetter known only tentative comments can be made

ou the relationships of the South Australian fauna

as a whole although a few elements can be clearly

discerned. There is a worldwide clement, repre-

sented by such species as Cerobasiy guestfalica,

Lepinolus reticulatus and Psyllipsecus ramburii

which have probably established themselves inde-

pendently of human influence, although they are also

found in domestic situations, Ecropsocus californicus

and Lachesilla pedicularia are similarly very wide-

spread species weli able to establish themselves in

new areas. There is an element composed of Aus-

(ralasian species, such as Caecilius semifuscalus,

Peripsocus maoricus (Tillyard), Propsocus pallipes,

Pentacladus eucalypti, Phlotodes australis and the

philotarsids, which are widespread in coastal regions

of Australia and which, in some cases, have ranges

which include New Zealand and Norfolk Island. To

what extent there has been human influence in the

overseas distributions is not known but such species

as Ph, australis could easily be carried on timber,

Together with these species can be included those

which have a more restricted southern and eastern

Australian distribution such as Echmepieryx brunnea

Smithers, Peripsocus edwardsi New, P. hickmani

New, Spilopsocus ruidis Smithers and Tanystigma

tardipes (Edwards). There appear to be two other

elements occurring in South Australia one of which

has affinities with Western Australia, including

species such as. Lasiopsocus michaelseni Enderlein

and Ectopsecus cetratus, While the other has aflini-

ties in an easterly direction with Victoria and Tas-

mania, represented by such species as Phlotodes

hickmani, Spilopsocus masseyi New and members of

the Blasle macrops species group, Finally, there is

the interesting species Sphaeropsocopsis recens

which js known from Tasmania and South Aus-

iralia and Which appears to have close relatives only

in South America, Angola and in Ballic amber

Mesepsocus reticulatus is an anomalous specics in

that the genus is known from many parts of the

world but. has not yet beea mentioned in records

for other parts of Australia. Its occurrence in South

Australia places it in an unusual, isolated position

zoogeographically,

REC. S. AUST. MUS. 18 (20); 453-491.

Muy, 1084

TAXONOMIC ACCOUNT OF SOUTH

AUSTRALIAN PSOCOPTERA

Smithers (1970, p, 372 er seq.) has given a key

to the families of Australian Psocoptera. In that

key the Mesopsocidae (not previously recorded

from Australia) would run to couplet 6. As it has

glabrous wings Mesopseeus will run to the Psilopso-

cidae, a family not yet recorded for South Australia,

Mesopsocus differs from psilopsocids in having 4 tall

areola postica and clear, hyaline wings; the psilop-

socids have a shallow areola postica and darkly

patterned wings.

LIST OF SPECIES OF PSOCOPTERA KNOWN FROM

SOUTH AUSTRALIA

(* New records for South Australia, ** South Australian

species not represented in present material),

LEPIDOPSOCIDAE

Echtnopsocus grayi sp. 1.

“Eehmepteryx (Loxopholia) brunnea Smithers

TROGHDAB

*Cerobasis yuestfulica (Kolbe)

**Lepinoias reticuluias Bnderlein

PSYLLIPSOCIDAE

**Psyllipsocus ramburii Selys-Longchamps

PACHYTROCTIDAE

Pachytroctes rugosus sp. 1,

SPHAEROPSOCIDAE

*Sphucropsocopsis recens (Hickman)

CAECILITDAE

*Caveilius semifascatus (Tillyard)

STENOPSOCIDAE

*"Stenapsocus. siriatifrans (McLachlan)

LACHESILLIDAE

*Lachesilla pedicntaria (1...)

ECTOPSOCIDAER

*Ecrepsocus californicus (Banks)

*Eeltopsocus cetratus Smithers

PERIPSOCIDAE

* Peripsocus edwardsi New

*Peripsocus maoricius (Tillyard)

*Peripsacus hickmani New

Peripsocus norialis sp. ti.

Peripsacus hollowayi sp, 1

PSEUDOCAECILIIDAE

Cladionewra punctata sp. n.

ELIPSOCIDAE

*Pentacladus cucalypri Enderlein

Propsocus pallipes (McLachlan)

*Spilopsocus masseyt New

*Spilopsocas reidis Smithers

PHILOTARSIDAE

#4 d4ustropsecks situasus (Banks)

*Haplophallus guitaluy (Tillyard)

Haplophallus hundaorensis New

Huplophallus medialis Thornton and New

*Haplophallus sinus Thornton and New

*“Aaronicla rawlings! Smithers

PSCOPTERA OF SOUTH AUSTRALIA

MESOPSOCIDAE

Mesopsocus reticulatus sp. n,

PSOCIDAE

*Lasiapsocus michaelsenit Enderlein

Lasiopsocus dicellis sp, 1.

Blaste macrops sp. n.

laste magnifica sp. a.

Blaste angusta sp. n.

*Prycla umbrara New

‘Pryeta glossepicra New

Prycta longipenniy sp, 0

Prycta hollowayae sp. ni.

*Yanystigma. tardipes (Edwards)

Tanystigma elongata sp, n,

Tanystigma bifureata sp. nt.

Psocidus mouldsi sp. n,

Psacidus parilla sp. n.

MYOPSOCIDAE

Phiotodes australis (Brauer)

*Phlotades hickmani (Smithers)

KEY TO ADULT PSOCOPTERA FROM SOUTH

2(1)

3 (1)

5 (2)

6 (2)

AUSTRALIA

Antennae with more than 20 segments, never

secondarily annulated. Tarsi 3-segmented. Plera-

stigma not thickened, or absent. Paraprocts with

strong posterior spine ..,...... 2a

Antennae usually with 13 Begmenits, if 15- to “47-

segmented then some segments are secondarily

annulated. Tarsi 2- or 3-segmented, Pterostigma

thickened or not. Paraprocts without posterior

BOM G: Siti. — copes res rrerimden stain america ee

Antennae 12- to 17-segmented, some secondarily

annulated. Tarsi 3-segmented, Pieromlems not

thickened _.....,.,, a]

Antennie usually [3-seemented. ‘Parsi 2 or 3-

segmented, if latter then flagellar segments not

secondarily annulated, Pterostigma thickened ,, 6

Head long and yertical, Maxillary palp without

sensillumm on inner side of second segment. Cuz

and IA end together at wing margin (i.e,

nodulus present) (Psyllipsocidae) bf see

a . Psyllipsocus perma biirie

Head short. Maxillary palp with sensillum on inner

side of second segment. Cus and IA end

separately at wing margin (Le, no nodulus),, 4

Claws with preapical tooth. Body and wings bear-

ing scales (Lepidopsocidae) _.. . . .,,,,..., 18

Claws without preapical tooth. Body and wings

not scaly (Trogiidae) 2.0. 19

Wings, when present, flat, with complete venation,

In apterous and alate forms eyes situated near

vertex (Pachytroctidae) .... Pachytroctes rugasus

Wings, when present, with incomplete venation,

curved, elytriform. Fyes situated well below

vertex a alia aaa :

Sphaeraptacagsis ¢ recens

Labial palps broadly triangular, laterally diverg-

ing. Lacinia narrow towards apex, Female gona-

pophyses reduced to a pair of inconspicuous,

acuminate valves, with a basal seta... __ 7

Labial palps short and appressed, somewhuat

circolar. Lacinin not usually narrowed towards

7 (4)

8 (6)

9 (8)

10 (9)

11 (10)

12 (9)

13 (12)

14 (13)

15 (8)

16 (15)

17 (12)

18 (4)

19 (4)

455

apex. Female gonapophyses usually of three

valves, the external Valve setose; if reduced then

not in form of two acuminate valves ......... 8

Cui, fused with M, or joined to it by a cross-

vein (Stenopsocidae) — Stenapsacus striatifrons

Cen not fused gor joined to M (Caeciliidac) , -

nna Caecilius semifuseatus

Areola postica free or absent. Females sometimes

brachypterous or apterous but withoul glandular

setae on head , rit 9

Cu, fused wits M in : fully “winged ‘fortns.

Females occasionally brachypterous, but then

glandular setae are present — -.,,,.,.,,. 15

Tarsi 3-segmented 0004002022. .---, 10

Tarai 2-segmented viccecccccccccrercreercrercecres ve TD

Fore and hind wings withott setae (Mesopsocidae )

restgst Mesopsocus. reticulatus

Fore. and ‘hind wines with. at least a few marginal

setae, even in brachypterous forms; usually with

DBVIPUG MOLE 1) ears T BIE LUN seoauh sere BY

Hind wing with margin entirely setose, Male hyp-

andrium strongly sclerotized. Female subgenital

plate with median lobe (Philotarsidae) .... 20

Hind wing with at most setae on margin between

Reis and Ry45, Brachyptery common, Male

hypandrium lightly sclerotized. Female sub-

genital plate usually bilobed (Elipsocidae) __ 24

Areola postica absent

Areola postica present 0.0. jecceecereee EB

Wings without setae (Lachesillidae) a

ne Henne Lachesilla pedicularla

Wings STORES ec ps a iettiidenselinaesatanienseapanceenscittice LG

Distal parts of veins in fore wing wilh more than

one row of setae (Pseudocaeciliidae) .

WES Alea fda le tdabld a lan ol Cladioneura punctata

Distal ‘parts of veins in fore wing with one row

of setae (Elipsocidae) . 24

Tarsi 2-segmented (Psocidae) ... ..__.,, 27

Tarsi S-segmemted 2... ccc cscs ceererenrens 16

Fore wings without setae. Wing pattern of

numerous, confluent, irregular dark areas giving

wing a densely mottled appearance (Myo-

psocidac) . 7 coronene

Wing pattern. bold, made up of large ‘hyaline and

coloured areas or without pattern (Elipsocidae)

. 24

Hind wing with Rs and M fused for a length,

Ry) meets margin al acute angle pF pelpepeetne

: . 44

Hind v wing with Rs aid M foined by a crossvein.

R; meets margin at right angle or almost so

(Ectopsocidae) cece Oe AB

Fore wing with Rs branched. Hind wing developed

vad , Echmepteryx hrunnea

Fore wing Swe Rs Sirtipte:. ‘Hind wing reduced to

small, pomted, flap ............ Echinapsecus grayi

Fore wings reduced but present as broad flaps

Tyan: Lepinotus reticulatus

Fore. inet r victaalty absent, represented by a

small tubercle . . Cerabasis guestfalica

aS6

20 (11)

21 (20)

22 (20)

23 (22)

24 (16)

25 (24)

26 (24)

28 (27)

29 (28)

40 (29)

31 (30)

32 (31)

#3 (30)

REC. 8. AUST, MUS. 18 (20):

Setae of fore wing veins sited on distinct dark

brown spots, at least in basal half of wing;

flagellar segments 6-8 with light apices ,...,.. 21

Setae of fore wing veins not sited on dark brown

spats, sind segments 6-8 without light apices

In fore wing Cuy bare, setae of upical veins sited

on distinct brown spots —- Aaroniella rawlingsi

In fore wing Cup setose, setae of apical veins not

sited on brown spots Haplaphalius medialis

In tore wing Cus selose or wing very reduced.

Antennal apex aitenuated, with a single, long

sel sites fa: 23

In fore wing Cus ‘Hare. Avitennal apex ‘not ‘allen-

uated, not with a single long seta

_ Haplophallus sinus

Femora and tibiae largely dark chocolate brown:

female brachypterous ,..........

Haplophullus. bandaorantis

Femora, and tibiae light brown; female macro-

plérous or brachypterous Haplophallus guttatus

Areola postica fused with M or joined to it by a

crossvein . |i conrenneranrvecoecoeyen@nrecionneen

Areola postica tres 26

M with more than 3 branches

i a . Pentacladus Nacktuer

M with 3 branches _ Prapsacus pallipes

Areola postica with Cy, curved to give a convex

distal margin to cell. Hypandrium simple, with-

out postero-lateral Johes , Spilopsecus muasseyi

Arcola postica with Cuy, sinuous to give a concave

distal margin to cell. Hypandrium with rounded

postero-lateral lohes - . Spilopsocus ruidis

Fore Wing veins obviously setose .......... .

Lasiopsocus michaelseni

Fore. wing veins without or apparenily without

SCTE coy amernrtinr meen ses, eycieris co. 2B

Fore wing with overall sheng pattern...

: Blaste magnifica

Fore wing hyaline | or + patterned, in Which case

patiern is not speckled ay ae. eae)

Cell My narrow. Mg and Cony parallel -

Blaste angusta

Cell. My 1 not markedly narrow. My and C09 not

parallel a“ :

Pterostigma with spurvein ..., . — ... — 3h

Pterostigma without spurvein ........000.-... 33

Cells Ry and R5 strongly and extensively pig-

mented . . . Tanystigma elongata

Cells Rg and Rg not strongly and extensively pig

FORTE” Wi acwer siliuiluddt delle suet cene aa

Distinct pigmented area near margin only in cell

Ri invcnnenee. LTanystigma bifurcata 2

No distinct inthented area near margin in cell

Ri . Tanystigma lardipes

Fore wing hyaline except for pigmented area at

margin in cell Ry or also over whole of cell Ry

basad of hind angle of plerostigma renee) 34

Wing markings otherwise ....00...000.0, 0... 35

34 (33)

35 (33)

36 (35)

37 (36)

38 (36)

39 (38)

40 (39)

41 (40)

42 (40)

43 (42)

44 (17)

45 (44)

46 (45)

47 (16)

48 (17)

453-491, Muy, 1984

Pigmented area near margin and over whole of

eell Ry basad of hind angle of pterostigma ....-

. Prycta hollowayae &

Pismerited 2 area ve: ‘cell Ri only near margin

Tunystiema bifurcata §

DUCE COORD DOR OUe Brit erie

Psocidus moulds?

Median cells strongly pigmented

Median ‘ells nol - strontly plemented . .. 36

Area around nodulus not pigmented ,,,..,,,,, 37

Acea around nodulus pigmented .....,.. ., 38

Phullosome cGpen posteriorly

. Lastionvavins idicetles ri

Phallosome closed posteriorly

. Ptyeta hollawayde 3

Cell Ry strongly pigmented beyond hind angle

of pterostigma —. . Psocidus parilla

Cell R, pot so pigmented . 39

Cell Rs;

postica .. |... .

Cell R5 without sucty § a Spot

with pigment spot anterior to areola

Prycta glossoptera

No spot at separation of M and Cuz .... ! 41

Spot of colour at separation of M and Cuz 42

No dark areas in basal cells ....

Some dark areas in basal cells ...

Piyeia langipennis

Ptycta umbrata

Spot at separation of M and Cuy almost snr

Cuy Hh. _ 43

Spot at separation of M and Cu ‘apnall,,

$a trodmxooweocka ¢posocotta cackpott Blaste rmderops

Area around Rs and M junction pigmented ..,

Apa rsfostekacertcctentrimiac tite Lasiapsocus dicellus &

Area around Rs and M junction hyaline ...,,..,

Piycta hallawayae 2

Fore wings grey With hyaline areas, visible even in

brachypterous females Peripsacus edwardsi

Fore wing without such markings 45

Fore Wing with Rs, M basad of fusion with Rs

and nodulus narrowly bordered with brown

. Peripsocus hiekmant

Fore. wines without such markings 46

Epicranial plates pale, bordered with pale brown

. Peripsocus maaricus

Epicranial ‘plates “pale. with a few brown marks

adjacent to compound eyes, across vertex and

adjacent to median epicranial suture .......,

Pia etisiecoroccottchtehnc aro to Peripsocus notialis

Epicranial plates very dark brown, a narrow pale

stripe from epistomial suture towards back of

head on each siilorttat L plate Tats es

_ Peripsocus hoflewayt

Larger apres fore wing Jength 5.0-5.5 mm

va . Phlotodes australis

Smaller species, fort wing 2 Tensth 3.4-3.6 mm __

_ Phlotodes hiekmani

Vertex pale brown, postclypeus much darker ,,,,,..,

. Ectapsocus californicus

Vertex pale with “darker spots, postclypeal ground

colour not much darker than vertex, postclyp-

eus with dark stripes .,..... Ectopsoens cetratus

PSCOPTERA OF SOUTH AUSTRALIA

Family LEPIDOPSOCIDAE

Echinopsocus grayi sp. n.

Female

Coloration (in alcohol): Head, body, legs,

antennae and maxillary palps golden brown. A faint

suggestion of a darker, curved narrow line mesad of

each compound eyc, a darker median epicranial

suture, a median line running length of pronotum

and mesoscutum and with margins of mesoscutum

dark. Apex of abdomen darker than head and

thorax. Ocelli black, Eyes black, Fore wing trans-

parent, tinged with golden brown; ves 4 little

darker than rest of wing surface; when vicwed with

transmitted light a somewhat mottled appearance is

evident with very faint indication of broad, trans-

verse, irregular banding. Hind wing transparent,

colourless.

Morphology; Brachyptcrous. Most of scales lost

in available material, those present long and very

narrow, Somewhat thickened fore wings not reaching

apex of abdomen, Abdominal terga membranous

under fore wings (i.e. as far as tetgite 7); eighth and

ninth tergites and terminal structures well sclero-

tized. Length of body: 2.5 mm. Median epicranial

suture and anterior arms very distinct. Vertex sharp.

Head with long, dense pubescence, Postclypeus

bulbous, Antennae incomplete in all specimens;

scape and pedicel broad, remaining segments short,

about twice as long as wide. Eyes fairly large,

Pao oes rt oace 2 “7

+57

reaching level of vertex, with an occasional seta

between facets. IO/D:2.0; PO:;0.7, Ocelli well-

developed, anterior ocellus only a little smaller than

lateral ocelli, Lacinia narrow, parallel-sided with an

emarginate apex so that the end is divided into a

smaller inner tooth and a larger outer tooth, Maxil-

lary palp with elongate second segment with small

sensillum; fourth segment a little broadened distally.

Prothorax sharp dorsally, strongly pubescent,

Measurements of hind leg: F: 0.72 mm; T: 1.16

mm: ty: 0.43 mm; te: 0.11 mm; ty: 0.1L mmy rt:

4:1:1. Hind femur short and broad, dorsal edge

convex with large setae, Tibia Jong and narrow,

apex with two stout and one small spur, Claws with

preapical tooth but without evident denticles basad

of tooth. Fore wing length: 1,96 mm; width: 0.72

mm, Fore wing (Fig, 1) not reaching apex of

abdomen, narrowing towards apex; membrane some-

what thickened, Anterior margin somewhat

thickened, Distal section of Sc (ie. from stigma-

pophysis to marginal thickening) distinct. R, runs

parallel to wing margin for a long length, reaching

margin not far short of wing apex. Rs fused with

M for a short length just distad of stigamapophysis,

simple, not branched, M 2-branched, the anterior

branch, apparently M, is, in fact, Rs. Cu, forks at.

level of stigmapophysis so that the areola postica

is long. Veins not well developed but evident, indi-

cated by position of rows of long, erect setae, repre-

sented by alveolae in figure. Membrane densely

FIGS, 1-6. Echinopsocus grayi sp. n, 1, 2 Fore wing. 2, 9 Hind wing (same scale as fore wing). 3. 9 Hind wing (enlarged)

4. 9 Epiproet and paraproct. 5. Gonapophyses. 6. ¢? Paraproct.

$34

covered with scales and scale-like setac, most of

which ate lost in available specimens, Hind wing

length; 0.44 mm, width: 0.12 mm. Hind wing (Figs,

2, 3) reduced to a membranous, acuminate flap

with faint suggestion of one vein near anterior

border and another, even less distinct, running more

or less parallel with the hind margin. A. single

small seta occurs about half-way along hind margin,

Epiproct (Fig, 4). Paraproct (Fig, 4), Subgenital

plate simple, rounded behind, Gonapophyses (Fig,

5). Ninth tergite and median part of eighth tergite

more heavily sclerotized than other abdominal terga,

being the areas exposed through wing reduction.

Male

Coloration (in alcohol): As female.

Morphology, General morphology as female,

similarly brachypterous, Byes as in female, Length al

hind leg: F: 0.64 mm; T: 1.04 mm; t: 0.39 mm;

ty: 0.08 mm}; ty: 0.08 mm; rt) 4.9:1:1. Fore wing

length: 1,68 mm; width: 0.68 mm, Form and vena-

tion as in female, Hind wing as in feniale, Hypan-

drium well sclerotized, with gently rounded hind

margin, a group of strong setae in middle of hind

margin, Phallosome with external parameres pos-

teriorly strongly acuminate, their apices strongly

incurved. Epiproct simple, rounded behind, Para-

proct (Fig, 6).

Material Exantined‘ SOUTH AUSTRALIA. ? (holo-

type), & (allotype), in litter, Mambray Creek, 19 iv.

1973, M. Gray. Paratypes: 2 %, as holotype,

Holotype, allotype and paratypes in the Australian

Museum.

Discussion; Echinopsecus Enderlein was erected

on ihe basis of a single specimen, in poor condition,

of E, erinaceus Enderlein from New Guinea (Ender-

Icin 1903), No additional material of this genus has

become available until now, E. gravi has most of the

generic characters of E. erinacezs but differs in the

fore wings not haying a long apical extension

although they are somewhat pointed. Enderlein

(1903, p. 331) indicated that he was unable to

find ocelli in E. erinaceus but they are present in

E. grayi. The venation of Echinopsocys is quite

distinctive, however, and E. vrayi agrees with the

type species. R, is long; Rs is not branched, uppear-

ing to arise from M owing to the evanescence of the

basal section of that vein busad of its fusion with M;

M is 2-branched, the anterior branch reaching the

margin near the bluntly pointed wing apex. Although

most of the setae and scales have been lost from the

four available specimens, those which remain and

the arrangement of alveolac indicate that FE. perayi

is clothed in scales through which protrude a fairly

dense covering of strong, erect setae, as described for

REC, S. AUST. MUS. 18 (20): 453-491.

Alay, 198d

&. crinaveus. The immediately apparent dillerence

in wing shape. the apical extension being consider-

able in £, erinaceus whereas the apex is. bluntly

pointed in £. gray).

Although &. grayi does not. conform to the charac-

ters of the generic definition so tar as occlli and fore

wing shape are concerned it ig not considered

necessary fo erect a separale genus for it. EL suffices

to enlarge the limits of Lehinepsecus to include

species which do have ocelli and in which the wing

Shape is more nearly normal,

Echmepteryx (Loxophalia) brannea Smithers

Echmepleryx (Laxopholia) hrunnea Smithers 1965.

J. ent, Sve, Od. 4: 75, Figs. 11-16,

Material Examined: SOUTH AUSTRALIA. 1 3,

Yalata, 131°45°E, 31°30'S, 29,i%.1978, M.S. and

B. J, Moulds. 1 9, 40 km. W. Nullarbor, 130°29°E,

31°28'S, 29,i%,1978, M.S. and B, J. Moulds.

This species has been recorded [rom New South

Wales and Queensland,

Family TROGITIDAE

Cerobasis guestfalica (Kolbe)

Hyperetes euestfalicus Kalbe, 1880, Jber. wesi/.

Prover, Wiss, Kunst &: 132; pl, IV, fig. 22.

Hyperetes pinicala Kolbe, 1881, Ent, Nachr, 7:

Tichohia aliernans Kolbe, 1882. Ent, Nachr, &: 21:

Cerobasis muraria Kolbe, 1882, Ent, Nachr, 8:

Ayperetes tessulatus Hagen, 1883. Stetitn. eat. Zig.

44; 216,

Alhardia alternans (Kolbe). Jacobson and Bianchi,

1904, Neyropt. Russ. Enip. p. 496.

Cerébasts guesifalica (Kolbe),

Sretiin. ent. Ztg. (04: 13.

Material Examined: SOUTH AUSTRALIA, 5 2,

Port Elliot, 13.v.1980, G. and J. Holloway,

C, guesialice is a cosmopolitan species which is

found in domestic habitats as well as in the wild.

Roesler, 1943.

Lepinotus reticulatus Enderlein

!Clothilla inguilina (Heyden). Hagen, 1882. Siettin.

ent, Zre. 43: 526, Pl. I, Fig. 6.

!Atrapos inguilina (Heyden), Kolbe, 1888. Jb) Fer.

Naturk. Zwickau 1887: 190, 191.

Lepinetus reticulatus Enderlein, 1905, Res, Swed.

Exp. Egypt 18: 31, Fig. 9; Pl. 1, Figs 1, 2; Pl. 2,

Figs, 12, 19, 23,

Not represented in the present material, this

species has been recorded from caves in South

Australia (Smithers, 1972).

PSCOPTERA OF SOUTH AUSTRALIA

Family PSYLLIPSOCIDAE

Psyllipsocus ramburii Selys—Longchamps

Psecus pédicularius Rambur, 1842. Historie natu-

relle des Insectes, p. 323.

Psyllipsocus ramburii Selys-Longchamps, 1872. Ent.

mon. Mag. 9: 145.

Nymphopsocus destructor Enderlein, 1903. Zool.

Anz. 27: 76.

Ocelloria gravinympha Weber, 1906, N.Y. Med, J.

$4: 885, Fig. 1.

Nymphepsocus traglodyta Enderlein, 1909. Arch.

Zool. exp. gen. 5 (1): 536, Pl. 18, Figs 9-11,

13, 14.

Fita vestigator Nayas, 1913, Rev. Acad. Madrid 12:

333, Fig. 4.

Fabrella convexa Lacroix, 1915, Bull. Soc. ent. Fr.

1915: 194,

Not represented in the present material, this

species has been recorded from South Australian

caves (Smithers, 1972).

459

Family PACHYTROCTIDAE

Pachytroctes rugosus sp. n.

Female

Coloration (in alcohol): Head, thoracic nota,

legs, first two abdominal terga, a lateral patch on

each side of third abdominal tergum and eighth

tergum dark brown. Ninth tergum and epiproct pale

brown; paraprocts dark brown. Other abdominal

terga very lightly sclerotized but segments are indi-

cated by broken bands of subcutaneous brown pig-

ment, Antennae brown, a little paler than head. Eyes

black.

Morphology: Apterous. Median epicranial suture

extending about one third distance towards epi-

stomial suture; anterior arms represented by a line

of tiny papillae visible only in cleared specimen.

Hind border of head straight between eyes. Sculp-

turation of vertex consisting of short, raised bars in

haphazard arrangement, that of the postclypeus

similar but finer, Lengths of antennal segments: f,:

0.12 mm; f.: 0.12 mm. First three flagellar segments

without annulations; antennae annulated from fourth

FIGS. 7-10. Pachytroctes rugosuy sp. n. 7. Gonapophyses. 8. Subgenital plate. 9, 9 Paraproct. 10. 2 Lacinia.

460

segment. Eyes fairly small, their upper margin lying

Jevel with vertex. Small tubercles present between

facets. Ocelli absent, Lacinia (Fig, 10) apparently

with five apical teeth, the fifth not clearly separated.

Fourth segment of maxillary palp elongate, four

times as long as wide. Thoracic terga of approxi-

mately equal width and length, clearly delineated,

finely sculptured with small, pointed spicules

arranged in irregular transverse rows, Dorsally, the

first, second, eighth and ninth ierga heavily sclero-

tized, remaining terga lightly sclerotized, almost

membranous, except for a small irregular area near

the lateral margin on segment three, Second segment

laterally sclerotized. Eighth segment much shorter

than ninth, the latter being more than twice as long

as eighth and much narrower opposite epiproct than

adjacent to eighth tergite. Sculpturation as in

thoracic terga but absent froni median area in

anterior half of second abdominal segment; spicules

jess densely arranged on ninth tergite, especially in

middle of plate, Femora of pro- and mesothoracic

legs strongly broadened with long setae. Hind

femora only slightly broadened, Measurements of

hind leg: F; 0,38 mm; T: 0.49 mm; t,: 0.24 mm;

ty: 0,08 mm; ty: 0.07 mms rt: 4.8:1:1.4. Epiproct

(Fig, 9) finely papillate except for a narrow pos-

terior area; setose, Paraproct (Fig. 9), setose, papil-

fae in dorsal half only, Subgenital plate (Fig, 8)

very large, setose and papillate, the papillae arranged

roughly in transverse rows. Gonapophyses (Fig, 7).

Material Examined) SOUTH AUSTRALIA. ¢

(holotype), ex dry sclerophyll Ewucalyptas litter,

Melrose, Flinders Ranges, 16.iv.1973. M. R. Gray.

Paratypes; 2 9, as holotypes. Holotypes and para-

types in the Australian Museum.

Discussion: Pachytroctes rugosus differs from

P. achrosta Thornton and Woo (from the Galapa-

g0s) in lacking postclypeal striations and pale

thoracic terga. The gonapophyses are similar in

Shape but the external valve of P. rugosus lacks a

sensillum. It differs from P. tapinelloides Badonnel

(from Africa) in having the fourth and fifth antennal

segments annulated, P. australis Ribaga, P, dichram-

moscelis Badoonel and P, granulosus Badonnel (all

African) differ in having the sculpturation of the

vertex in the form of fine granulations, not bar-

shaped ridges. In P. bicoloripes Badonnel (African),

P. insularis Thornton and Woo (Marianas) and

P. nivecinctus Badonnel (African) the metathorax is

white, In P, aglypha Badonnel (African) the sculp-

turation of the head is indistinct and, as in P. auran-

tiacus Badonne] (African) and P. ambiguus Badon-

nel (African) there are no granulations between the

ommatidia.

The only previous record of this genus from Aus-

tralia is that of an unidentified species taken from

bags of peanuts in shell at Kingaroy, Queensland,

REC, §. AUST. MUS, 18 (20); 453-491.

May, TUs4

Family SPHAEROPSOCIDAE

Sphaeropsocopsis recens (Hickman)

Sphaeropsocus recens Hickman, 1934. Oce. Pap: R.

See, Tusm. 1933: 83, Figs. 4A-4F,

Sphaeropsocopsis recens (Hickman). Badonnel,

1963. Biol. l'Amerique australe 2: 291, 323.

Material Examined: SOUTH AUSTRALIA, 1 2,

ex Eucalyptus litter, dry sclerophyli, Melrose, Flin-

ders Ranges, 16.iv-1973, M. R. Gray. 1 2, Mt. Lofty,

26.iv.1943, H. Womersley,

This small interesting species was described from

dry grass tussocks in Tasmania (Hickman 1934),

The genus is known also from Chile and Angola.

The closely related genus SpAaeropsocus Hagen is

known from one species in Baltic amber.

Family CAECILITDAE

Caecilius semifuscatus (Tillyard)

Maoripsocus semifuscatus Tillyard, 1923. Trans.

NZ, Inst, 54; 191, Fig. 16; Pl. 18, Fig. 11.

Caecilius semifuscatus (Tillyard). Senithers, 1969.

Rec. Canterbury Mus, 8: 280, Figs, 44-48.

Material Examined; SOUTH AUSTRALIA. 20 ¢,

23 9, Wirrulla, ESE Ceduna, 28-ix.1978, M. S. and

B, I. Moulds. 1 c', 1 °, 50 km WNW Ceduna, 28.ix.

1978, M, S. and B. J. Moulds, 12 2, Pooginook

Park, 13-16.vi.1979, G. A, Holloway. 1 2, 20 km

SE Pt. Augusta, Horrock’s Pass, Flinders Ranges,

17,vi.1979, G. A. Holloway. 2 %, 15 km N Port

Broughton, 7,v.1980, G. and J, Holloway. 1 %, 10

km N Goolwa, 13.v,1980, G, and J, Holloway. 1 3,

18 km N Ardrossan, 8.v.1980, G. and J. Holloway.

] 3, Telowie Gorge, 10 km E Port Germein, 20.v.

1981, G,, J. and A. Holloway.

This species, originally described from New

Zealand, has also been recorded from Curtis Island,

Bass Strait. These records are the first from the

Australian mainland.

Family STENOPSOCIDAE

Steropsocus striatifrons (McLachlan)

Psecus striatifrons McLachlan, 1866, Trans. ent,

Sec, Lond, (3) 5: 351.

Slenepsecus striatifrans (McLachlan). McLachlan,

1866, Trans, ent, Soc, Lond. (3) S: 352.

This species was described from “Australia meri-

dionali” and is not represented in the present

material, The original locality may not have been in

South Australia and nothing referable to this species

has since been reported in the literature,

PSCOPTERA OF SOUTH AUSTRALIA 461

Family LACHESILLIDAE

Lachesilla pedicularia {L,)}

Hemervbius pedicularius Linnaeus, 1758. Systema

Naturae p, 551,

Lachesilla pedicularia (L.). Enderlein, 1919. Cat.

Coll, Selys Longchamps 3 (2); 16,

For complete synonymy see Smithers (1967)-

Material Examined; SOUTH AUSTRALIA, 1 8,

in packing straw from England, Adelaide, y.1937,

L, pédicularia is a very widespread species being

known from the Palearctic Region, Comoros, Argen-

tina and South Marianas. There ig one previous

Australian record, from Victoria.

Family ECTOPSOCIDAE

Ectopsocus californicus (Banks)

Peripsocus californicus Banks, 1903. J. N.¥. ent,

Soc, 11: 237,

Ectopsecus californicus (Banks) Badonnel, 1955.

Pub, culi. Camp. Diam, Angola 26: 185.

Ectopsocus congener Tillyard, Smithers, 1969. Rec.

Canterbury Mus. & (4): 289, Figs. 71-75.

Material Examined: SOUTH AUSTRALIA. 1 8,

9 km S$ Edithburgh, 7.v.1980, G, and J, Holloway,

1 os, 2 9, Mt, Alma, 12 km SW Victor Harbor,

12.v.1980, G, and J. Holloway, 1 3, 4 km N

Murray Bridge, 22.v.1981, G. and A. Holloway.

This species is known from North America, New

Zealand and Antipodes Islands; previous Australian

records are from Tasmania and New South Wales,

Ectopsocus cetratus Smithers

Ectopsecus. cetratus Smithers, 1972, Aust, Zool, 17

(1): 15, Figs, 1-8.

Material Examined: SOUTH AUSTRALIA. 7 34,

36 9, 3 km EK Nundroo, 132°30'E, 31°50'S, 29.1x.

1978, M.S. and B. J. Moulds, 5 ?, Wilmington,

Flinders Ranges, 6.v,1980, G, A, Holloway. 2 &,

50 km WNW Ceduna, 28.ix.1978, M. S. and B. J.

Moulds, 3 3, 10 9, 40 km E Nullarbor, 131°15B,

31°25'S, 29.ix.1978, M. S. and B. J. Moulds, 2 &,

Yalata, 131°45°E, 31°30'S, 29,ix,1978, M. 8. and

B. J. Moulds,

This species was described from Western Aus-

tralia; these are the only subsequent records for the

species.

Family PERIFSOCIDAE

Peripsocus edwardst New

Peripsocus edwardst New, 1973, J. Aust. ent, Soe.

12: 40, Figs. 1-6.

Material Examined: SOUTH AUSTRALIA, 3 4,

1 2, 20 km. SE Port Augusta, Horrock’s Pass, Flin-

ders Ranges, 17.vi1979, G. A. Holloway. 3 4,

Germein Gorge, Flinders Ranges, 11,5 km E Pt,

Germein, 7.vi,1979, G. A. Holloway. 2 ¢&, 2 &,

Germein Gorge, 19.v.1981, G. and J. Holloway.

This species, in Which the females are brachyp-

terous, has previously been recorded only fram

Victoria.

Peripsocus maoricus (Tillyard)

Peripsocopsis maoricus Tillyard, 1923, Trans, N.Z,

Inst. 54: 194, Fig. 18; Pl. 18, Fig, 12.

Peripsocus macropterus. Edwards, 1950, Pap, R, See,

Tasm. 1949; 124, Figs. 89-94.

Perjpsacus maoricus (Tillyard). Thornton and

Wong, 1968. Pacific Ins, Monoer, 19; 10, 135.

Material Examined: SOUTH AUSTRALIA, 30 4,

23 ?, 6 km W Kapunda, 18,vi,1979, G. A. Hollo-

way. 20, 6 ?, 5 n, 9 km § Edithburgh, 7-v.1980,

G. and J. Holloway. 7 3, 17 9, 2 n, Spring Gully

Park, 9 km SSW Clare. 18.vi.1979, G. A. Hollo-

way, 1 9, Pooginook Park, 15.vi.1979, G. A. Hollo-

way. 3 3, Germein Pass, Flinders Ranges, 6,v,1980,

G, and J. Holloway, 17 3, 17 9, Germein Gorge,

Flinders Range, 11,5 km E Pt, Germein, 7.vi,1979,

G. A. Holloway. 1 ¢, Wirrulla, ESE Ceduna, 28.ix.

1978, M. S. and B. J. Moulds, 21 o, 55 %, 23 n,

25 km E Peake, 23.vi,1979, G. A. Holloway, 19

3, 21 2, 1 n, 23 km E Tailem Bend, 13.v-1980,

G. and A, Holloway, 12 ¢, 24 %, 15 km W Tailem

Bend, 13.v.1980, G. and A. Holloway. 9 dy, 19 2

2 km E Parilla, 23.vi,1979, G. A, Holloway. 11 ¢,

72,3 n, 4 km E Pinnaroo, 14,v.1980, G. and J,

Holloway, 2 4, 2 9, 4 km S, Moonta, 7.v.1980, G,

and J, Holloway, 2 d, 1 8, 18 km N Ardrossan,

8.v.1980, G. and J. Holloway. 1 3, 6%, 9 n, 10 km

N Goolwa, 13.y.1980, G. and J. Holloway, 6 ¥, 20 9,

41 n, 16 km 8 Minlaton, 7.v.1980, G, and J, Hollo-

way, 2d, 13 %, Adelaide, 224x.1965, H. Womersley.

8 3,102, 7 4, 12 km SE Port Wakefield, 8.v,1980,

G. and J. Holloway, 4 6, 2 ?, 19 n, 2 km W Wil-

liamstown, 8,v.1980, G. and J, Holloway. 3 ot, cx

Cypress Pine, Alligator Gorge Rd., near Mt, Re

markable, Flinders Ranges, 17.iv.1979, G. Hollo-

way. 3 2, W end of Horrack's Pass, 19.v,1981, G,

and J. Holloway. 10 3, 17 &, ex Casuarina, Mt,

Gambier, 23,v.1981, G,, J. and A. Holloway. 30 3,

55 2, Germein Gorge, 19-20,v.1981, G. and J, Hol-

loway. 6 4, 10 %, Burra Gorge, 20 km SSE Burra,

21.v.1981, G., J. and A. Holloway, 1 ¢, 2 km SE

Williamstown, 22.v.1981, G. and A, Holloway, 1 &,

Telawie Gorge, 10 km E Pt. Germein, 20.v.1981,

G,. and J. Holloway-

462 REC. S$. AUST, MUS. 18 (20); 453-491, May, 1984

Peripsocus maoricus appears to be widely dis- Peripsocus notialis sp. n.

tributed, at least through the southern part of the Male

continent and Tasmania, It has been recorded from

Tasmania, Victoria, Western Australia and now Coloration (in alcohol): Head pale brownish

from South Australia. Tt was originally described cream with pale brown markings. A few brown

from New Zealand. marks close against upper margin of compound eyes

FIGS. 11-16, Peripsocus notialis sp. n. 11. ¢ Fore wing. 12. Phallosome. 13. Subgenital plate, 14, 2 Fore wing. 15, 2

Paraproct. 16. Gonapophyses,

PSCOPTERA OF SOUTH AUSTRALIA

and across vertex with broken patch on either side

of median epictanial suture near back of head, Post-

clypeus with anteriorly converging, pale browu

stripes nol reaching anteclypeus, Genae and labrum

as head. Antennae and muxillary palps brownish.

Eyes black. Ocella¢t tubercle brownish, Lobes of

mesothoracic notum pale brown; parapsidal sutures

paler than lobes. Legs pale brownish cream with

slightly darker tarsi, Fore wing (Fig. 11) hyaline,

uniformly very lightly tinged with pale brown; ptcro-

stigma opaque but noi darker than rest of membrane,

Hind wing, membrane us fore wing. Veins brown, all

well developed, none evaneseent, Abdomen pale;

phallusome sclerifications indistinctly visible through

hypandrium.

Morphology: Median epicranial suture distinct;

anterior arms evanescent. Antennae much thicker

than in female. Lengths of flagellar segments: f;-

0,68 mins fu: 0.48 mm, Eyes large; reaching well

above level of vertex, Faccts exceptionally large: eye

emurginate where almost in contact with antenna

base. 10/D: 0.80; PO: 1.0. Ocelli large, on well

developed tubervle. Lacinia yery narrow near distal

end, hardly divided apically. Fourth segment of

maxillary palp long, parallel sided with rounded

apex, four times long as wide. Measurements of

hind leg: F: 0.60 mm; T: 1.16 mm, ty: 0.32 mm;

ty: O16 mm; rt: 2:1; et: 22, 0. Legs long and

narrow. Fore wing (Fig. 11) with fairly broad costal

cell, Cu, evanescent just before margin, Fore wing

length: 4,0 mm} width: 1.6 nim, Hind wing length:

3.0 mm; width: 1.1 mm. Epiproct triangular with

angles rounded, sctose, sparsely spiculate in median

area near base; spicules very small. No clumal camb,

Paraproct simple, ovoid, with very large circular

trichobothrial field, the setae long and fine. Hypan-

drium simple, with a broad sclerotized band parallel

with hind margin; margin interrupted. in middle:

svlose. Phallosome (Fig. 12) closed anteriorly with

distinctly sclerotized margin interrupted posterjarly-

Sclerification of penial bulb heavy. of two, pos-

teriorly outwardly curving sclerites subtended by a

complex sect of symmetrically arranged selerites.

Female

Coluration (in alcohol): As male but marks on

head 4 little more extensive, Fore wings (Fig. 14)

as in male but costal cell, anterior half of cell R

and anterior pact of cell Ry as far as apex of ptero-

stigma hyaline, contrasting with the very pale

browish linge in rest of membrane. Hind wing

hyaline, faintly tinged with pale brawn along veins.

Terminal abdominal structures brown,

Morphology, Length of body: 2.5 mm, Lengths

of flagellar segments: f,: 0.40 mm; f: 0.26 mm,

Antennae nyuch finer than in males. Eyes fairly small,

much smaller than in males. 1O/D; 1-7; PO: 0,75.

Measurements of hind leg: F: 0.52 mm; T: 0.96

mm; ty: 0.20 mms t.: 0.16 mits rts 1.3:15 cts 8, 0,

Fore Wing length; 2.8 mm; width: 1.1 mm, Fore

wing (Fig. 14). Hind winy Jenyth: 2.2 mm) with:

0.8 mm. Epiproct (Fig. 15), Paraproct (Fig. 15)

simple, with small trichobothrial field, the setae long

und fine, Subgenital plate (Fig. 13). Gonapophyses

(Fig. 16).

Material Examined; SOUTH AUSTRALIA, 2

(holotype), ? (allotype), ex cypress pine, Alligator

Gorge Rd., near Mt. Remarkable, Flinders Range,

17.vi.1979, G. A, Holloway. Paratypes: 5 ¢, 19 9,

as holotype (one $ on slide). Other material: 5

nymphs, as holotype.

Holotype, allotype and paratypes in the Australian

Museum.

Discussion: Peripsoeus netialis is a species which

is somewhat sexually dimorphic, The female has the

anterior part of the wing hyaline in contrast to the

uniformly coloured wing of the male, The female

wings are considerably shorter than in the large nvale

and the legs are stouter, with fewer ctenidia not

regular in arrangement. In general arrangement the

sclerifications of the penial bulb resemble these in P-

norfolkensiy Smithers. & Thornton and P. suaoricts

(Tillyard) but differ in proportion and shape. The

female differs in 1O/D ratio and in wing colour.

Proportions of the gonapophyses also differ but they

resemble evch other in having a broad ventral valve.

Peripseeus. notialis is similar in size to P. edwards?

New but differs in having much paler wings, Ti P.

edwardsi the wing membrane is faintly tinged with

grey but there are discrete hyaline patches im most

cells, In P, notialis the male fare wing membratie is

very faintly but uniformly tinged with pale brown.

In the female the costal cell and a narrow strip

behind R and R, is hyaline. The female is some-

what brachypterous. There are slight differences in

proportions of the ponapaphyses and, although

simulac jn general structure, the male phallosome has

differently proportioned sclerificatiang of the penial

bulb, In P. hamiltonae Smithers, a slightly smaller

species, there isa distinet darkening across. the. wing

from stigmapophysis to nodulus, which is not present

in P. nofialiy. P. hickman’ New is a much smaller

species. than P. netialis with a fore wing Jength of

less than 3 mm as Opposed ta that of 4 mm. for

P, notiglis, In P. hyekmanj{ there is a little pigmen-

tation adjacent to the basal section of Rs and M

basad of fusion with Rs which is absent from P-.

notielis, The phullosome in P. votialis hits a trans-

verse anterior boarder with complex seclerification of

the penial bulb and narrow, outwardly curved

external purdmeres, In P, hiekmani the phallosame

464

tapers to a broken, narrow anterior end and the

external parameres are broad and short; the scleri-

fications of the penial bulb are simpler, being in the

form of a few longitudinal rods and a rngosely

sclerotized bulbous area, P. paoricus has a distinc-

tive three lobed apex to the arch formed by the dis-

tally fused inner paracneres and the wing is uni-

formly more darkly tinted than in P. notialis. P.

notialis is much larger than P, melaleucae New. The

female of P, melaleucae has tapering lateral exten-

sions of the sclerotized area of the subgenital

plate; these areas are broad in P, nojialis. The

phallosome of P. smelaleucae has short, broad,

external parameres, In P, muller’ (Tillyard), P.

morulops (Tillyard) and P. tillyardi New the wings

are much darker in general with cyen darker areas

along some of the wing veins, In P. roseus Smithers

the pterostigma is distinctively reddish in the distal

half in life (this color fades in alcohol). The main

veins have dark and light sections which give a

distinctive appearance of being broken and discon-

linuous.

Peripsocus hollowayi sp, n.

Male

Coloration (in alcohol); Head very dark brown

except for a paler area between ocellar tubercle and

antenna base, narrowing laterally to proximity of

REC, 8. AUST. MUS. 18 (20): 453-491,

May, 1984

compound eye and epistomial suture and a similar

pale narrow stripe from this area towards back of

head on each epicranial plate. Postelypeal stripes

hardly discernible (might be more easily seen on

paler specimens?). Labrum coloured as head.

Antennae dark brown, Maxillary palps dark brawn.

Eyes black. Ocellar tubercle very dark brown, Meso-

thoracic: notum very dark brown, lacking pale

median stripe: lateral lobes each with a narrow paler

mark running from scutellum to lateral margin near

wing base: a pale spot where. parapsidal sutures con-

verge, Coxae dark brown. Legs pale brown, Fore

and hind wings hyaline with dark brown veins.

Abdomen pale, terminal structures very dark brown,

the Jarge, broad phallosome clearly visible through

paler integument of abdomen.

Morphology; Median epicranial suture very dis-

tinct, as. is epistomjal suture, Length of flagellar

segments: fy: 0.56 mm; f.: 0.40 mm. Eyes only

moderately large, small for a male. IO/D: 1.7; PO:

0.88. Lateral ocelli large, anterior ocellus much

smaller. Measurements of hind leg: F; 0.62 mm) T;

1.28 mm; t: 0.28 mm; t.: 0.14 mm; rt: 2:1; ct:

15. 0. Fore wing length: 3.3 mm; width: 1.2 mm.

Hind wing length: 2.5 mm; width: 0.8 mm- Ptero-

stigma fairly shallow, hind angle not pronounced,

shallowly rounded. Epiproct (Fig. 19) heavily

BIGS, 17-20, Peripsocus hollowayi sp. o. 17, Padlloout., 18. Hypandrium, 19, 3° Epiproct. 20. Peripsacus hickmant

allosome.

PSOCOPTERA OF SOUTH AUSTRALIA

sclerotized, with semicircular hind margin very

strongly sclerotized and a small posterior median

membranous extension with rounded hind margin.

Paraproct well sclerotized with large, circular tri-

chobothrial field. Hypandrium (Fig. 18) (damaged

in preparation) with median emargination, lighily

sclerotized except for a broad, well-sclerotized

medially interrupted band paralJcl with hind margin.

Phallosome (Fig. 17) well sclerotized, broad,

rounded anteriorly; external parameres broad; inter-

nal parameres distally fused, ending in a pair of

blunt processes, Sclerifications of penial bulb heavy,

symmetrically arranged.

Material Examined: SOUTH AUSTRALIA, 1 ¢

(holotype), ex cypress pinc, Alligator Gorge Rd.,

near Mt. Remarkable, Flinders Ranges, 17.v1.1979,

G, A. Holloway, Paratype; 1 3, Germein Gorge,

Flinders Ranges, 11.5 km B Pt. Germein, 7.vi.1979,

G. A. Holloway.

Holotype and paratype in the Australian Museum.

Discussion; Peripsocus fellowayi is a large

species with dark head and thorax and hyaline wiags.

The broad phallosome, with broad, external para-

meres and two short posterior processes with

heavily sclerotized penial bulb sclerifications is

characteristic and distinguishes it from all ather

species.

Peripsocus hickmani New

Peripsoous hickmani New, 1973. J. Aust. ent. Sac.

12: 341, Figs 5, 7-10.

When this species was described from Victoria

(New 1973. p, 341. Figs 5, 7-10) male material

was not available, The males in the present material

permit description of that sex here.

Male

Coloration (in alcohol): As female (New 1973,

p. 341).

Morphology: Length of body: 2.2 mm. Median

epicrania] suture very clearly defined, Lengths of

flagellar segments: f;: 0.40 mm; £: 0.30 mm.

Antennae shorter than in female. Eyes fairly large,

larger than in female but anly just reaching level of

vertex. IO/D: 1.2; PO: 0,81. Measurements of hind

leg: F: 0.44 mm; T: 0.92 mm; t,: 0.20 mm; te:

0.12 mm; rt; 1.7;1; ct: 16, 0, Fore wings as in

female (New 1973, Fig. 7). Fore wing length: 2.8

mm, width: 1.1 mm. Hypandrium simply rounded

behind, not medially emarginate, a broad, lightly

sclerotized band parallel with margin, shghtly inter-

rupted in midline. Phallosome (Fig. 20) similar ta

that of P. tillyardi New (1973, Fig, 12) but broader;

sclerificatians of penial bulb symmetrical.

A465

Material Exainined: SOUTH AUSTRALIA, 1 3,

1%, Walata, 131°45°E, 31°30'S, 20.ix.1978, M. S-

and B. J. Moulds, 3 2,6 9,18 km WN Ardrossan,

8,v.1980, G. and J. Holloway.

Discussion; The male phallosome of P. hickmani

resembles that of P. sillyardi in general form but

differs in being wider in relation to length and in

having the external parameres almost as wide as

long, whereas in P. tillyardi they are more elongate,

distinctly longer than wide,

Family PSEUDOCAECILIIDAE

Cladioneura punctata sp. n.

Female

Coloration (in alcohol): Head ivory, with very

faint suggestion of median brown band from vertex

to labrum and very pale greyish patch mesad of each

compound eye, Median epicranial suture not

obviously coloured. Antenmac pale brown, a small

section at the distal end of each flagellar segment

paler, Eyes black, Ocellar tubercle very dark brown,

conspicuous against otherwise pale head, Labrum

brown, Maxillary palps pale, tip of fourth segment

very dark brown. Mesonotum with dark brown ante-

dorsam und lateral lobes; parapsidal sutures pale,

Metanotum with Jateral lobes brown, Pleura mainly

brawn. Coxae brown basally, pale distally. Femora

pale, without dark bands. Tibijae pale. Tarsi brown.

Claws black. Fore wings (Fig, 21) hyaline with

dark brown markings. Hind wings hyaline with a

faint infuscation adjacent to basal section of Rs and

M between separation from Cu, and fusion with Rs.

Abdomen pale with a few spots on each segment;

the dorsum of the ninth tergitc, in particular, has

an irregular row of darker, larger spots which are

conspicuous. Epiproct and paraprocts are pale.

Morphology; Length of body: 2.5 mm. Median

epicranial suture distinct, anterior arms less con-

spicuous, Head, except anteclypeus and genac, with

strongly developed erect setae, those on postclypeus

shorter and finer than most of those on vertex,

Lacinia (Fig, 25). Length of flagellar segments: f;:

0.40 mm; f.; 0.28 mm, Eyes fairly small, not reach-

ing level of vertex, [1O/D: 2.9; PO: 0.7. Ocelli large,

anterior ocellus about as large as lateral ocelli.

Measurements of hind leg: F; 0.52 mm; T: 0.96

mm; ty: 0.28 mm; tg; 0.14 mm; rt; 2:1. Cteni-

diobothria absent. Fore wing length; 2.3 mm} width:

0.9 mm, Fore wing (Fig. 21) with veins evanescent

in many places; strong setae arise in two rows mostly

alongside rather than on veins, Cus without setae,

Hind wing length: 1.8 mm; width: 0.7 mm. Veins

sparsely setose only in distal part of wing (Fig. 22).

Epiproct almost semicircular with small setae on

posterior half and a series of strong setae along

REC. $, AUST. MUS. 18 (20): 453-491,

~~ ™~ %

Ea Sw eg phat we Ey

<a Se _ PEW AR, 5

May, Las

bee

te Ne

FIGS, 21-25, Cladioneura punctata sp. n. 21, % Fore sai te ? Hind wing, 23, Gonapophyses. 24, Subgenital plate.

. acinia.

posterior part of margin. Paraproct simply rounded

behind, setose in posterior half with strong posterior

marginal setae. Trichobothrial field of about 9

trichobotliria, poorly defined. Subgenital plate (Fig.

24) posteriorly bilobed, each lobe with a few strong

marginal setae, variable in number (holotype with

four on right lobe and two on left). Gonapophyses

(Fig. 23).

Male

Unknown,

Material Examined! SOUTH AUSTRALIA, &

(holotype), 15 km N Ardrossan, 8.v.1980, G. and

J, Holloway. Paratypes; 2 9, as holotype. 1 ?, 16 km

§ Minlaton, 7.v.1980, G, and J. Holloway.

Holotype and paratypes in the Australian

Museum,

Discussion: Cladioneura punctata differs from

C, pulchripennis Enderlein in details of wing pat-

tern, fore wing development and shape of the tip of

the lacinia. In C. punetata the most conspicuous

wing colour difference is the absence of pigment over

most of cell My adjacent to the areola postica and

the presence of a spot in cell Rs at the basal

quarter. The wings in C. punctata are relatively

short and broad being only 2.3 mm long for a

body size of 2,5 mm, whereas in C. pulchripennis

the fore wings approach 3.0 mm for the same body

size. The lacinia is broader at the apex in C. punc-

fata, The gonapophyses are similar but differ a

little in proportions. In C. puxetata the marginal

setae of the subgenital plate are in two groups

whereas in C. pulchripennis the four setae are more

evenly spaced across the hind margin, C, pulchri-

pennis is the only other species in the genus.

Family ELIPSOCIDAEK

Pentacladus eucalypti Enderlein

Pentacladus eucalypti Enderlein, 1906, Zool. Jh, 23:

408; Pl. 23, Fig. 7.

Material Examined; SOUTH AUSTRALIA, 2 ¢,

Ravine des Casoars, Kangaroo Is., 28.x.1951, G. F,

Gross,

This species has been recorded from New South

Wales, Tasmania, Victoria and Queensland.

PSCOPTERA OF SOUTH AUSTRALIA

Propsocus. pallipes (McLachlan)

Psocus pallipes McLachlan, 1866. Trans. ent. Soc.

3 ser. 5: 349.

Propsocus pallipes (Mclachlan).

1866. Trans. ent. Sec, 3 ser, 5; 352,

Tricladuy froggatti Enderlein, 1906. Zool. Jb. 23:

410; Pl. 23, Fig. 6,

Tricladellus freggatti (Enderlein), Enderlein, 1909.

Srertin. ent. Zte. 70: 273,

MeLachlan,

Material Examined: SOUTH AUSTRALIA, 1 3,

15 km SW Tailem Bend, 13.v.1980. G. and A.

Holloway,

Originally described from Adelaide, this specics

is known from New South Wales, Tasmania, Queens-

Jand and Western Australia. It has not yet been

recorded from Victoria but it seems likely that it

will be found there,

Spilopsocus masseyi New

Spilopsocus masseyi New 1971. J. Aust. ent. Soc.

10: 226, Figs, 7-13,

Material Examined: SOUTH AUSTRALIA, 1 32,

Port Elliot, 13.v.1980, G, and J, Holloway, t 3,

Seal Bay, Kangaroo Island, 2-4.xii.1977, D. K.

McAlpine and M, A, Schneider, 1 2, 40 km W,

Nullarbor, 130°29'E, 31°28'S, 29.ix.1978, M. S. and

B. J. Moulds.

This species is recorded from Tasmania, New

South Wales and Victoria,

Spilopsocus ruidis Smithers

Spilopsocus ruidis Smithers, 1963, Pacific Ins. 5 (4):

894, Figs. 19-25,

Material Examined: SOUTH AUSTRALIA. 1 32,

Yalata, 131°45'E, 31°30'S, 29.ix.1978, M. S. and

B. J. Moulds.

Spilopsocus ruidis has previously been recorded

only from New South Wales.

Family PHILOTARSIDAE

Austropsocus sinuosus (Banks)

Zelandapsocus sinuosus Banks, 1939. Bull. Mus.

comp. Zool. Hary. 85: 441, Pig. 12,

Austropsocus sinuosus (Banks). Thornton and New,

1977. Aust. J. Zool. Suppl. ser, 54: 28, Figs. 99-

104.

This species was originally described from South

Australia (Mt. Lofty Range) but is not represented

in the present material.

467

Haplophallus guttatus (Tillyard)

Philotarsus guttatus Tillyard, 1923. Trans. N.Z.

Inst, 54: 181, Fig. 8.

Philotarsopsis deleaius Tillyard, 1923. Trans. N.Z.

Inst. 54: 182, Fig. 9.

Philotarsus greyi Edwards, 1950, Pap. R. Soe. Tasm.

1949: 116, Figs. 68-75,

Haplophallus greyi (Edwards).

J. ent. Sac, Od, 2: 60.

Smithers, 1963.

Haplophallus guttatus (Tillyard). Smithers, 1969.

Rec. Canterbury Mus, 8: 322, Figs. 158-162.

Material Examined: SOUTH AUSTRALIA, 1 ¢,

15 9, 40 km E Nullarbor, 131°15°E, 31°25'S, 29,

ix.1978, M, S. and B. J. Moulds. 1 ¢, 7 ?, 3 km

E Nundroo, 132°30'E, 31°50'S, 29.ix.1978, M, S.

and B. J. Moulds. 1 ¢, 1 2, Myponga, H, M, Hale.

Haplophallus gutiatus has been recorded from

many localities in New South Wales, Victoria, Wes-

tern Australia and Tasmania. It is also known from

Queensland and was originally described from New

Zealand,

Haplophallus bundoorensis New

Faplophallus bundoorensis New, 1971, J, Aust, ent.

Soe. 10 (1): 25, Figs. 1-10.

FHlaplophallus capitulatus Smithers, 1972, Aust. Zool.

17 (1): 19, Figs, 12-17,

Material Examined: SOUTH AUSTRALIA, 9 32,

5 3, 6 km W Kapunda, 18.vi.1979, J, Holloway.

2 3, 50 km WNW Ceduna, 28.iv.1978, M. 8. and

B. J. Moulds,

H, bundoorensis is known from Victoria, Queens-

land and South Australia.

Haplophallus medialis Thornton and New

Haplophallus medialis Thornton and New, 1977.

Aust. J, Zool. Suppl. ser., 54: 13, Figs, 26-32,

36.

Material Examined: SOUTH AUSTRALIA, 7

specimens ex Eucalyptus obliqua dry sclerophyll

forest, Naracoorte Cave Reserve, 25.x.1958, G. F.

Gross,

This specics was previously recorded from New

South Wales, A.C.T., Victoria and South Australia.

Haplophallus sinus Thornton and New

Haplophallus sinus Thornton and New, 1977. Aust.

J. Zool. Suppl. ser. 54: 20, Figs. 60-68.

Material Examined: SOUTH AUSTRALTA, 1 @,

Mt. Alma, 12 km SW Victor Harbor, 12.v.1980,

468

G. and J. Holloway. 1 3, 10 km N Goolwa, 13.y.

1980, G. and J. Holloway. 3 &, 1 9, ex Casuarina,

Mt. Gambier, 23.v.1981, G., J. and A. Holloway.

1 3, Yourambulla Caves, 6 km SW Hawker, 18.v.

1981, G. and J. Holloway.

H. sinus is known previously from only one New

South Wales locality.

Aaroniella rawlingsi Smithers

Aaroniella rawlingsi Smithers, 1969. Rec. Canter-

bury Mus, 8: 324, Figs. 163-168.

Aaroniella pallida New, 1971. J. Aust. ent. Soc. 10

(1): 29, Figs 11-21.

Material Examined: SOUTH AUSTRALIA. 1 4,

Mt. Gambier. 23.v.1981, G., J. and A. Holloway.

Originally described from New Zealand _ this

species has been recorded from Victoria, New South

Wales, Australian Capital Territory and Western

Australia.

REC. S. AUST. MUS. 18 (20): 453-491,

May, 1984

Family MESOPSOCIDAE

Mesopsocus reticulatus sp. n.

Male

Coloration (in alcohol): Head brown, very

slightly darker on either side of median epicranial

suture and dorsad of compound eyes, lacking the

discrete spotting usual in the genus. Median epi-

cranial suture dark brown. Postclypeus brown, ante-

clypeus pale. Labrum and genae brown. Antennae

brown. Eyes black, Ocellar tubercle a little darker

than surrounding area. Maxillary palps with all four

segments brown. Mesonotum brown, parapsidal

sutures pale brown. Pleura brown, a reddish mark

laterally on hind part of mesopostnotum. Legs

brown. Fore wings (Fig. 26) hyaline, veins and

pterostigma brown. Abdomen pale, terminal struc-

tures brown,

Morphology: Length of body: 2.2 mm. Median

epicranial suture distinct, anterior arms absent.

FIGS. 26-29. Mesopsocus reticulatus sp. n. 26. ¢ fore wing, 27. 6 Epiproct and paraproct. 28, Phallosome. 29. Hypan-

rium.

PSCOPTERA OF SGUTH AUSTRALIA

Head small, finely pubescent: vertex arched, Seulp-

turation of head on vertex and frons consistitig of a

well defined polygonal pattern of raised ridges, that

on pastclypeus of a serics of irregular, fine, trans-

verse ridges, Postclypeus not exceptionally bulbous

but reflexed anteriorly so that the anteclypeus is

somewhat set back, Posterodistal row of sensilla on

anterior margin of Jabrum made wp of three placoid

sensilla and two trichoid sensilla almost in a straight

line, Anterodistal row of four sensilla, Lengths of

flagellar segments (in mm,.)> f,: 0.39; f.: 0.23;

fy> O21; fy: 0.19; fp: 0.14; far 0.14; fy: 0.12;

Fg: 0,10; fy: 0.09; fy 9! 0.08; fia: 0.08, Antennae

fairly short, tip mucronate. Total tength: 1.81 mm.

Eyes fairly Jarge, somewhat protruding but not

reaching level of vertex, [10/D: 2.0; PO, 0.86, Ocelli

large. Pedice] with two conical sensilla; first flagellar

segment with one sensillum with minute point at

basal quarter, Fourth and tenth flagellar segments

each with a similar sensillum at distal end. Sixth

flagellar segment apparently lacks a sensillum. Legs

slender, Measurements of hind Jeg: F: 0.47 mm;

T: 1.00 m; + 1-19 mm; te: 0.07 mm; tg: 0.08 mm;

rt: 2,7:151.1; ct: 10, 0, 0. Fore wing length: 3.5

tm; width; 1,25 mm. Fore wings (Fig. 26) with

long, narrow pterostigma, Ry geatly curving, Rs

before junction with M slightly curved; fusion with

M fairly short. Radial fork about as Jong as stem.

Cu,, slightly curved before rounded apex of areola

postica so that basal side of areola postica is slightly

concave. Hind wing length: 2.6 mm; width: 0.

mm, Cuy and TA not strongly curved near wing

margin, Four minute setae on margin between

Ro+, and Ry, 5, readily visible only at magnifica-

tion greater than about 50%. Epiproct (Fig. 27).

Paraproct (Fig: 27). Hypandrium (Fig. 29). Phallo-

some (Fig, 28) narrowing anteriorly and posteriorly

with bluntly pointed anterior end to phallic frame.

Penial bulb with strong, symmetrical sclerifications.

Acdeagal arch narrow, slightly angled distally.

External parameres broad apically, with conspicuous

extension but obliqucly truncate distally,

Material Examined: SOUTH AUSTRALIA, ¢

(holotype), Germein Gorge, Flinders Range, 11.5

km E. Pt. Germein, 17,vi,1979, G. A. Holloway.

Holotype in the Australian Museum,

Discussions Mesapsecus reticulatus is. the first

species of the family to be reported from Australia;

the genus has previously been recorded fram Africa,

Europe, Asia and North and South America,

Most species have a distinctive and striking head

pattern made up of spots and stripes on the vertex

and have the more or less clearly defined postcly-

peal stripes so characteristic of many psocopterans.

Mesopsocus reliculatus obviously differs at first sight

in lacking. this patterning, The head is almost uni-

449

formly brown, The antennae are unusually short

and the truncate form of the apex of the external

parameres and the clearly defined, symmetrical

sclerificattons af the peniwl bulb differ in those

species in Which these structures have been described,

Unfortunately only one specimen, a male, is

available; it is possible that the female, as in many

other species of Mesepsocus, is apterous.

Family PSOCIDAE

Lasiopsocus michaelseni Enderiein

Lasjapsocus michaelsent Enderlein, 1907. Fauna

SW, Aust. (1) 3: 234, Figs 1-5.

Blaste (Lasiopsocys) michaelseni (Enderlein).

Roesler, 1943. Srettin, ent, Zig, 104! 3,

Material Examined: SOUTH AUSTRALIA. 2 @,

3 nymphs, Pooginook Cons. Park, 15.vi,1979, G. A.

Holloway, 1 ¢, 2 km & Parilla, 23.vi.1979, G. A

Holloway. 4 d, 5 9, 5 nymphs, 4 km E Pinnaroo,

14.y,1980, G. A, and J. Holloway, 2 ¢, 5 $, Pan-

dappa Res,, 20 km E Terowie, 16.v.1979, G. A.

Hollaway-

L. michaelseni is one of the largest Australian

psocids with a wing length of up to about 7 mm.

It was. originally described from Western Australia.

The record from New South Wales (Muogamarra

Nature Reserve) (Smithers 1977) is in errar and

the specimen referred to there belangs to L, simu-

latus Smithers,

Three of the females. from Pandappa Reserve are

brachypterous, the wings reaching only to the end of

the abdomen, Enderlein (1907) gives wing lengths

of 7.0 mm (2) and 4.7 mm (?), With the shorten-

ing of the wings venational aberration has occurred

to the extent that the fusion of Rs and M in the fore

wing is shortened so that these veins meet in a point

or are even joined by a crossvem, A similar reduc-

tion in the length of fusion occurs of Cu,, and M to

the point that these veins do not meet at all so the

aerola postica ts free and the discoidal cell open.

Lasiopsocus dicellus sp. n,

Male

Coloration (in alcohol); Head pale, but brown

as follows: a double row of irregular confluent spots

adjacent ta each compound cye, across back of

vertex and on either side of median epicranial suture;

a broad spot on frons ynterior to ocelli; a line in

position of anterior arms of epicranial suture from

ocelli ta antenna base; a ming around antenna base;

a mark below compound eye on gena; postclypeal

stripes and the labrum, Median postclypeal stripes

closer and darker than lateral stripes. Ocellar tubercle

black, Scape and pedicel brown; flagellum very

470 REC, 8. AUST. MUS. 18 (20): 453-491 May, 1984

FIGS, 30-38, Lasiopsocus dicellus sp. 1. 30, d Fore wing. 31. 2 Fore wing. 32, Phallosome. 33. Hypandrium. 34. Sub-

genital plate. 35. Ninth tergite, poslero ventral apophysis, 36, ¢ Epiproct, 37. Gonapophyses, 38. @ Ninth sternite.

PSCOPTERA OF SOUTH AUSTRALIA

dack brown, Eyes black, First and second maxillary

paip segments pale, third brown, fourth dark brown,

Dorsum of mesolborax dark, shiny brown, a little

paler where parapsidal sutures meet. Fore Iegs pale

brown except for darker apex of tibia and tarsal seg-

ments. Meso- and meta-thoracic legs similar bot coxae

dark brown, Fore wings (Fig. 30) hyaline, without

extensive pattern: pterostigma brown; veins dark

brown, Hind wings hyaline; veins brown. Abdomen

pale, terminal structures very dark brown.

Morphology: Length of body; 4 mm. Median epi-

cranial suture distinct; anterior arms not evident but

usual position indicated by brown line. Antennae

fine, bearing Jong setae, those of first flagellar seg-

ment up to more than five times longer than width of

segment. Eyes fairly large, not reaching Jevel of

vertex, 1O/D: 1,9; PO: 0,83, Ocelli large. Apex of

lacinia divided into two slightly denticulate Jobes,

the outer strongly divergent; Jacinia narrowed just

basad of apical division. Measurements of hind leg:

F; 1.04 mm} T; 2,04 mm; t,; 0,60 mm; tu> 0,20

mmm; rt: 3:1; ets 22, 2, Fore wing length: 5-0 mm;

width; 1.6 mm. Pterostigma long, narrow, Stigma-

pophysis shallow, dome-shaped. M before fusion

with Cu;, curved to give a cancave outer margin, to

discoidal cell; curved section of vein somewhat

evanescent as are Reys and R4}5 beyond separa-

tion. TA fairly thick. An occasional tiny seta present

on Veins. Hind wing length: 3.7 mm; width: 1.1.

Hind wing margin with a few fine, shert setae

between Roja and Ry, 5 near wing apex. Epiproct

(Fig. 36) sclerotized with a sinuous hind margin.

A complex basal structure arises from epiproct

where it is attached to ninth tergite. This consists of

a posteriorly directed plate which lics above epiproct,

the plate is posteriorly bilobed and strengthened

along each side by a sclerotized strip, the strips

curving away somewhat from each other behind,

From the base of the plate arise two erect, elongate,

apically 3-lobed strap-like apophyses (twisted in

illustration) which are more heavily sclerotized

dorsally than ventrally, Paraprocts wilh very

strong, median sclerotized strengthening bar, a large,

almost circular trichobothrial field distad of which

the paraproct ts extended inta 2 media-dorsally

directed tapering bar subtended by a ventrally

placed, lightly sclerotized lobe. These lobes are

curved inwards and lie behind the distal part of ihe

hypandrium. The eighth sternite sclerotized in distal

part Only, adjacent to base of hypandrium. Hypan-

drium (Fig, 36) distally upturned to end in a broad

lobe with transverse posterior margin. Ninth tergite

extended posteroventrally on each side into a distally

broadened lobe with a heavily sclerotized margin.

The broadened end is acute dorsally but broadly

rounded ventrally (Fig. 35). Phallosome (Fig. 32)

ATL

with external parameres joined anteriorly, narrow,

rodlike, diverging posteriorly, each with a pair ol

strongly developed, outwardly curved distal teeth,

Near the end of each paramere arises a small pro-

jection, probably representing the temnants of the

internal patameres,

Female

Coloration (in alecghol)- As in male but with

parapsidal sutures pale, legs darker and with dis-

linct wing pattern in various shades of brown (Fig.

31), This ts in strong contrast to the hyaline male

fore wing,

Morphology: Length of body: 5.2 mm. Eyes small.

10/D: 2.7; PO: 0.84. Lacinia as in male, Measure-

ments of hind leg: F> 1.2 mm; T: 2.1 mm; t,: 0.5

mm; ty: 0.25 mmy rt: 2:1; ct: 21, 2. Fore wing

length: 5.0 nm; width; 1.6 mm, R_ strongly

developed. Rs and M fused for a shart length. Dis-

coidal cell as in male, Hind wing lensth: 3.8 mm;

width: 1.2 mm, Marginal setae as in male but a little

more strongly developed. Paraproct broadbased,

narrower posteriorly, well sclerotized with simall

circular trichobothrial ficld. Subgenital plate (Fig.

34). Gonapophyses (Fig: 37) with finely pointed

ventral valve, dorsal valye blunt-ended, spiculate

near end. Sclerification of ninth sternite (Fig, 38)

simple, ring-like.

Material Examined: SQUTH AUSTRALIA. 1 ¢

(holotype), | 2? (allotype), Pandappa Res., 20 km,

BE. Terowie, 16,vi.1979. G, A. Holloway. Paratypes:

12 &. 6 %, data as holotype; 2 3, 1 2%, 4 km NW

Murray Bridge, 22.v.1981, G., J. and A. Holloway.

Holotype, allotype snd paratypes in the Aus-

tralian Museum,

Discussion; Lasiopsocus dicellus differs from L.

michaelseni in being much smaller and darker, with

a fore wing length of only 5.0 mm compared with

7.0 mm for L. michaelseni. The wing sctae are re-

duced to an occasional small seta, not easily seen.

The proportions of the complex structures associated

with the male epiproct are different although the dis-

tinctive structures are similar m general form and

arrangement. The phallosome differs in proportions

of the parts. The eighth sternite of the male is

sclerotized only in a marrow band adjacent to the

base of the hypandrium and the sternite is thus much

Jess heavily and extensively sclerotized than in other

genera of the Amphigerontiinae, to which sub-

family Lasiopsocus. belongs,

The female of L. michaelseni, as well as being

much Jarger than that of DL. dicellus, does not have

the extensive dark wing markings characteristic of

L, dicellus,

Males of L. dicellus are very similar to those of

L. simulatus Smithers but differ slightly in propar-

472 REC. S, AUST. MUS. 18 (20): 453-491, May, 1984

FIGS. 39-45, Blaste macrops sp. n, 39. ¢ Fore wing. 40. Phallosome. 41, Hypandrium. 42. 9 Fore wing. 43. Subgenital

plate. 44, 2 Epiproct. 45. Gonapophyses.

PSCOPTERA OF SOUTH AUSTRALIA

tions of the phallosome and the structures associated

wilh the epiproct. The pasterior lobe of the hypan-

drium hag a transverse hind margin in L. dicellis

but is more rounded in L. simularus, The female of

L. simulatus is not known.

Blaste macrops sp. n.

Male

Coloration Gn alcohol): Head pale with brown

Toarkings as follows: a wide band consisting of irre-

Bular, sometimes confluent spots adjacent to com-

pound eyes, across yertex and on either side of

median epicranial suture (these arcas leave only a

small part of epicranium clear); a broad band from

eye through antenna hase and across anterior part

of postclypeus to meet band from other side; from

the broad band, another runs from between eye and

antenna base towards the ocellar triangle, Posicly-

peus pale with rows of faint spots in positions usually

occupied by postclypeal stripes; spots darker in

anterior third. Labrum dark brown. Genae pale

with dark mark below eye. Antennae brown. Eyes.

black. Ocelli circled with black, Maxillary palps

progressively darker from pale base to dack brown

fourth segment. Meso-thoracic antedorsum dark

brown except for a small spot on each anterolateral

angle and one at junction of parapsidal furrows;

dorsal lobes dark brown, broadly pale adjacent to

parapsidal furrows and fore wing base; scutellum

pale. Metathorax with pale antedearsum and brown

dorsal lobes. Femora pale, itregularly marked. with

pale brown; tibiac pale proximally, becoming dark

brown distally; tarsi dark brown, Fore wings (Fig.

39) hyaline, marked in shades of brown. Hind wings

hyaline with a pale brown areca in distal angle of

Cu,, Abdomen pale, with irregular brown, seg-

mentally arranged markings; terminal structures

dark brown,

Morphology: Length of body; 2.8 mm. Median

epicranial suture distinct, anterior arms evanescent.

Upper region of head somewhat expanded to form

lobes on which the eyes are carried so that top of

head is broadened; vertex medially depressed.

Length of flagellar segments: f,: 0,85 mm; f.: 0.90

mm. Antennae slender with fine pubescence. Eyes

fairly large, very prominent, carried on upper lateral

head extensions, inner margins diverging strongly

behind when seen [rom above. TO/D: 2-1; PO:

0.90. Ocelh large, but ocellar tubercle not very

prominent, Measurements of hind leg: F: 0.83 mr;

T; 1.9 mm; ty: 0.6 mm; ty: O15 mm, rt: 4:15 et:

26, 4. Legs long and slender. Fore wing length: 4.5

mun; width: 1,7 mm. Se curves distally to meet R.

Rs and M fused for a length. Discoidal cell concave,

ic, M curved. M, curves, arched towards wing

margin, reaching margin just anterior to wing apex.

473

Hind wing length: 3.4 mim; width: 1.1 mm. Micro-

trichia of wing membrane a little larger in cell Cuo

than elsewhere. Epiproct rectangular with rounded

hind angles, Paraproct heavily sclerotized in basal

half, lightly so in distal half; trichobothrial field

large and circular, posterior projection small and

pointed. Hypandrium (Fig. 41). Phallosome (Fig.

40) with parameres apically separate and tapering,

divergent,

Female

Coloration (in alcohol): As. male but with a pale

median line on mesothoracic antedorsum and more

extensive fore wing markings (Fig, 42),

Morphology; Length of body: 3-7 tam, Head

much larger than i male, shaped as in male, eyes

unusually prominent for a female. Antennae fine.

10/D: 2.5; PO: 0.82. Measuremenis of hind leg:

F; 0.82 mm; T: 1,8 mm; t; 0.55 mm; te: 0.15

mm; rt: 3,6:1, Fore Wing length, 3.7 mm: width:

1.3 mm. Venation as in male but My, reaches wing

margin at wing apex. Hind wing length: 2.8 mm;

width: 0.9 mm. Epiproct (Fig. 44) heavily sclero-

tized laterally, less so medially with hind margin

slightly emarginate medially, Subgenital plate (Fig.

43) well sclerotized. Gonapaphyses (Fig, 45).

Material Examined: SQUTH AUSTRALIA, 1 ¢

(holotype), 1 % (allotype), 25 km E Peake, 23,vi.

1979, G, A. Holloway, Paratypes: 3 %, 1 ¢, Germein

Gorge, Flinders Ra., 11.5 km FE Pt. Germein, 17.vi.

1979, G. A. Holloway. 1 2, 2 km E Parilla, 23-1.

1979, G. A. Hollaway. 1 ¢, Pandappa Res.,. 20 kim

E Terowie, 6.¥i.1979, G, A, Holloway,

Holotype, allotype and paratypes in the Australian

Museum.

Discusston; Blaste macrops belongs to a group

of species in which the male hypandrium js rounded,

with a pair of lateral, incuryed, posteriorly project-

ing processes and a phallosame in which both inner

and outer parameres are elongate, joined anteriorly,

with the inner parameres free behind, tapering to-

wards end, The eyes if both sexes are prominently

placed on short dorsolateral extensions of the head

capsule and there is usually sexual dimorphism in

wing pattern with females having somewhat more

extensive and often darker markings than males,

The group includes &. fulcifer Smithers (Tasmania),

B_ furcilla New (Western Australia), B, luaulata

New (Western Australia) and B. tillyardi Smithers

(New Zealand, New South Wales and South Aus-

tralia). &. panaps Smithers (Tasmania) may alsa

be considered as belonging to this group although

wing pattern dimorphism is not pronounced and the

474

male hypandrium lacks the curved processes; the

phallosome, however, is very similar to that of B.

falcifer, as is the shape of the epiproct.

Only the most obvious differences between the

species are given here. Perusal of the descriptions

and illustrations will provide an indication of the

characteristic differences in proportions and shapes

of genitalic structures as well as wing pattern dif-

ferences by which the species can be distinguished.

REC. S. AUST. MUS, 18 (20): 453-491.

May, (984

Both B. falcifer and B. panops have a distinct

mark between the ends of Ry, and M, i.e. near

the wing apex which is lacking in B. macreps. In B.

lunulata males (females not known) there are exten-

sive wing markings in cell Cus; the band of colour

across the wing from pterostigma to areola postica

is broad and continuous; M, Rs -++- M and the edges

of the discoidal cell are bordered with brown and

the median cells and cell R; cach apically have

FIGS, 46-50, Blaste magnifica sp, n, 46, 3 Fore wing. 47. Hypandrium. 48, ¢ Epiproct, 49. ¢ Paraproct, 50. Phallo-

some.

PSCOPTERA OF SOUTH AUSTRALIA

extensive brown markings, These markings are far

more extensive than in B. maecreps (Fig. 39). In B.

tillyardi, probably the nearest known relative to 2.

maecrops, the hypandrial processes are shorter and

stouler and the inner parameres posteriorly incurved.

In female B. macrops there are irregular marks in

cell R adjacent ta M which are absent from female

B. ftillyardi and B, fureilla, Males of B- fureilla

differ from males of B. muacrops in having a tri-

angular flap arising from the base of the epiproct

(New 1974, Fig. 17).

The generic Jimits of Blaste and its nearest rela-

tives are in meed of revision on a world basis and

when this is dane it seems likely that B. macrops

und similar species will be defined as a distinct

generic group,

Blaste magnifica sp, n.

Male

Coloration (in alcohol): Head brown with dark

brown markings. A double row of irregular, con-

fluent spots adjacent to compound eyes, across back

of vertex and adjacent to median epicranial suture.

Frons with triangular patch with pale centre anterior

to ovellar tubercle, Epistomial suture very dark

brown. Aréas of top of head not occupied by the

spotting described above are mostly occupied by

brown Which does not quite reach the spots, A very

dark bund runs from compound eye to antenna

base. Genae with an irregular mark below compound

eye and two small marks above base of mandible.

Postelypeal stripes dark and clearly defined. Antecly-

peus dark in basal half, pale in distal half. Labrum pale

brown with dark brown proximal border, Scape, pedi-

cel pale brown and basal half of first flagellar segment

pale brown, remainder of flagellum brown, Eyes black.

Ocellar tubercle black. Maxillary palp with third and

fourth segments dark brown, otherwise pale. Thorax

distinctively marked, being very dark chocolate brown

with only parapsidal furrows and a narraw median

line between dorsal lobes pale, the line between the

dorsal lobes extending for a short distance onto

hind part of anledorsum. Pleura mostly dark brown,

Femora irregularly marked in various shades of

brown, darkest at distal end. Tibiae brown, darker

at each end, Tarsi brown. Fore wings (Fig. 46)

hyaline, densely speckled brown, Hind wings hya-

line, faintly tinged with grey, Abdomen pale with a

few brown marks; terminal structures dark brown,

Morphology: Length of body: 3.8 mm, Upper

angles. of head produced a little sa that the eyes

are very prominently held on short, thick “stalks”.

Median epicranial suture very distinct, anterior arms

absent, Vertex slightly lower in middle than laterally,

thus accentuating eye prominence, Postclypeus

almast square when viewed [ram front of head; not

475

very prominent. Lengths of flagellar segments: f):

1,16 mm, fa: 1,04 mm, Antennae very fine, with

long setae, some as long ws seven times flagellar

diameter. Eyes large, prominent on cephalic exten-

sions, upper margins high above level of veriex-

TO/D; 1.4; PO: 0.94. Ocelli large on prominent

tubercle; epicranial plates concave an either side of

oeellar tubercle, thus accentuating tubercle pro-

minence, Measurements of hind leg: F: 1.12 mim;

Ts 2.64 mm) t;: 0.68 tam; ta: O14 mm; rt: Sil;

ci; 29, 2. Legs lang, thin, with very spiny tibiac,

Femora narrow, parallel sided. Fore wing lengihy

6.5 mm; width: 2.1 mm, Se meets R distally, Rs

and M fused for a length, Discoidal cell incurved on

distal margin, Cu,, and M fused for a length: basal

and second sections of Cu,,, at a slight angle to one

another. Hind wing length; 4.8 mim width; J.6 mm,

Rs and M fused for a short length. Margin glabrous.

Epiproct (Fig, 48), Paraproct (Pig. 49), Hypan-

drium (Fig. 47). Phallosome (Fig. 50).

Female

Unknown.

Material Examined> SOUTH AUSTRALIA, ¢

(holotype), Pooginook Park, 15.vi.1979, G. A.

Holloway. Paratype: 3, as holotype.

Holotype and paratype in the Australian Museum.

Blaste magnifica is a very distinctive, large and

easily recognized species, It is the only species in

the genus in Which the wing is mottled with an

overall pattern of small, irregularly contluent spots,

The hypandrium is distinctive, with an unusual hind

border which is medially sclerotized and carries two

small projections, one on each side of the midline.

The phallosome is also of unusual form for the

genus. As im the case of the graup of Species asso-

ciated with B, macrops, mentioned above, it. muy

be necessary to recognize this species in a separate

genus. when the large genus Blaste is revised,

Blaste angusta sp. n.

Male

Coloration (in alcohol): Head pale brownish with

dark brown, irregular, confluent spots adjacent to

compound eyes, across vertex and adjacent to

median epicranial suture, A narrow brown band

surrounds antenna base. Postclypeal striations dark

brown, A brown ovoid patch with pale centre lics

anterior to ocellar triangle just posterior to episto-

mial suture. Genae pale with two small spots below

eye. Afitennae brown, a little paler in basal four

segments than more distally. Maxillary palps pale

basally, fourth segment yery dark brown. Meso-

thoracic antedorsum dark brown, parapsidal furrows

broadly pale; scutellum pale; a posterior lateral pale

line on each dorsal lobe. Femora pale brown, dark

476

FIGS, 51-55, Blaste angusta sp, n. 51, ot Fore wing. 52. H

REC. 8. AUST. MUS. 18 (20): 453-49L.

May, (O84

ypandrium, 53. Phallogome. 54. gd Paraproet. 55, Bpiproct and

ninth tergite,

along dorsal and lateral surfaces. Tibiae pale brown,

a little darker at cach end, 'Tarsi dark brown. Fore

wing (Fig. 51) hyaline with markings in various

shades of brown, Hind wings hyaline, a brown spot

at wing base. Abdomen pale with slightly darker,

irregular, segmentally arranged markings in basal

two thirds; distal third occupied by the almost black,

extensively sclerotized terminal structures; eighth

sternite extensively and very heavily sclerotized.

Morphology: Length of body: 3.0 mm. Median

cpicranial suture very distinct but anterior arms

absent, Median part of epistomial suture sinuous,

Length of flagellar segments: f): 0.80 mm; f.: 0.64

mm, Flagellar setae about three times as Jong as

flagellar width, Eyes large, inner margins divergent

behind when seen from above and just reaching level

of vertex. IO/D: 1.9; PO: 1.0. Measurements. of

hind leg; F: 0.50 mm; T; 1.80 mm; ty: 0,56 mm;

ty: 0,20 mm, rt; 2,8;1; ct: 27, 1. Fore wing length;

4.0 mm; width: 1.4 mm, Fore wing (Fig. 51) with

broad pterostigma, with round hind angle and Ry,

curved to give a concave pterostigma basad of hind

angle, Rs and M fused for a very short length, Sc

ends free in costal cell, runs very close to R. Third

median cell very narrow, My and distal sections

of Cu,, almost parallel but relationship varies some-

what in the four fore wings available for study. Apex

of areola postica fairly broad, Hind wing length:

3.0 mm; width; 1.1 mm. Rs and M fused for a

fairly long length, Ninth tergite (Fig. 55) very

heavily sclerotized, extended posteriorly in the

middle to which extension is attached the small

lobed epiproct. Paraproct (Fig. 54). Eighth sternite

very heavily sclerotized. Hypandrium (Fig. 52).

Phallosome (Fig. 53).

Female

Unknown,

PSCOPTERA OF SOUTH AUSTRALIA

Material Examined: South Australia ¢ (holotype),

10 km SW Renmark, L5.vi.1979, G, A. Holloway.

Paratypes: 1 2, as holotype. 1 4, Telowie Gorge,

10 km SE PL, Germoin, 20.v.1981, G, and J, Hollo-

way. 1 4d, W end of Horrock’s Pass, 19.v.1981, G.

and J. Holloway.

Holotype and paratypes in

Museum.

the Australian

Discussion: Blaste angusta is an easily recognized

species. It ig so far the only known Australian

species of Blaste in which cell My in the fore wing

is. Narrow as. well as being well pigmented. The

paramerces, distally broadened into a rough knob, are

characteristic.

Ptycta umbrata New

Prycta umbrata New, 1974. J. Aust. ent. Soc. 13:

297, Figs. 38-44.

Material Examined: SOUTH AUSTRALIA, 2 4,

1 9, 2 km W Williamstown, 8,v,1980, G. and J.

Holloway. 3 co, 11 n., Wilmington, Flinders Range,

6,v,1980, G. and J. Holloway. 1 9%, Alligator Gorge

Rd., near Mt, Remarkable, Flinders Range, 17.vi.

1979, G. Holloway. 1 ?, 6 km W Kapunda, 18-v1.

1979, G. Holloway. 1 #, 25 km E Peake, 23.vi,1979,

G. Holloway. 3 ¢, 1 ¥, Telowie Gorge, 10 km SE

Pt. Germein, 20,v981, G., J, and A. Holloway.

3 2, Germein Gorge, 19,v,1981, G. and J. Holloway.

This species has previously been recorded only

from Victoria.

Prycta glossoptera New

Ptycta glassoptera New, 1974. J. Aust. ent. Soc.

13; 302, Figs, 58-64

Material Exaniined> SQUVH AUSTRALIA, 6 3,

24 n, Pandappa Res., 20 km E Terowie, 16-vi,

1979, G. A. Holloway. 1 9 (macropterous), 2 ?

(brachypterous), I n, 2 km E Parilla, 23.vi,1979,

G. A. Holloway. 1 2 (brachypterous), 1 n, 10 km

SW Renmark, 15.vi,1979, G. A, Holloway. 1 3,

Pooginook Park, 15.vi.1979, G. A. Holloway.

This species has previously been recorded from

Victoria,

Pryeta longipennis sp. n.

Male

Coloration (in alcohol): Head greyish with brawn

markings, A double row of spots adjacent to cam-

pound cycs, ucross back of head and adjacent to

median epicranial suture, a mark along anterior arms

of suture which broadens toward lateral ends; an

avoid spot anterior to ocellar tubercle. Base of

antenna surrounded by a narrow brown band. Post-

clypeal stripes narrow. Antenna dark brown. Maxil-

lary palps pale, fourth segment very dark brown,

ay7

almost black, Mesonotum dark brown, sutures,

median antedorsal stripe and posterolateral edges of

lateral lobes pale. Legs pale, tarsi brown, Fore wing

(Fic. 56) hyaline, with a faint general tinge of

brown and slightly darker brown areas as in tigure:

Abdomen pale, irregularly marked with brown;

terminal structures dark brown.

Morphology: Length of body: 2.3 mm, Median

epicranial suture distinct, anterioc arms evanescent

but position marked by dark stripe, Lengths of

flagellar segments: fy: 0.76 mm; t,: 0.74 mm. Eyes

large, reaching above loyel of vertex, TO/D: 1.3;

PO: 0.91. Measurements of hind legs PF: 0.68 mim

T: (.60 mm; t,) 0.50 mm; to: 0.12 mm; rt: 42:7;

et! 30, 3, Fore wing length; 4.0 mm; width: 1.3 mm.

Fore wings relatively long and narrow. Se ending

free in costal cell. Pterostigma fairly narrow but hind

angle clearly evident. Rs and M fused for a short

length. Discoidal cell broad, distal section of M and

Cu, bordering cell both curved distally towards wing

apex, Basal section of Cu,, slightly sinuous almost

in a line with second section. Hind wing length:

3.2 mm; width: 0.9 mm. Rs and M fused for a

long length. A few marginal setae between Ro).

and R4,.. Epiproct (Fig. 59) lightly sclerotized,

bordered with slightly more heavily scleralized band

of variable width. The posterior medjan part ts

folded back in figure. Hypandrium (Fig, 58) slightly

asymmetrical with a lateral, sclerotized extension of

the dorsally curved, median, straplike band near the

distal end, Phallosome (Fig. 57) with a long pas-

terior median extension.

Female

Coloratian (in alcohol): As ia male. Wings (Fig.

60) slightly darker.

Morphology: Length of body: 2.6 mm, Lengths

of flagellar segments: f;- 0.60 mm; fo: 0.60 mr.

Eyes smaller than in male, not quite reaching level af

vertex. 1O/D: 1.7; PO: 0.82, Measurements of hind

leg: F; 0.56 mm; T: 1.36 mm; t,: 0.40 mm; ty:

0.13 mms tt: 3:1; et: 25, 1, Fore wing length:

3,6 mm; width: 1.0 mm. Venation (Fig. 60)

as in male. Hind wing length: 2.7 mm; width: 0.8

mm. Epiproct (Fig. 61). Subgenital plate (Fig, 64).

Gonapophyses (Fig, 62) with short ventral valve;

dorsal valve tapering to point, distally slightly curved

upwards; spicuwlate in distal third. Sclemte of ninth

sternite (Fig. 63) simple, lightly sclerotized,

Material Examined: SOUTH AUSTRALIA, ¢

(holotype), ? (allotype), Germein Gorge, 19.v.

1981, G. and J. Holloway, Paratypes: 1d. LY

Telowie Gorge, 10 km SE Pt. Germein, 20.v.1981,

G. and J. Holloway. Holotype, allotype und paratype

in the Australian Museum.

478

REC. S$. AUST. MUS, 18 (20); 453-491.

May, 1984

FIGS. 56-64, Ptyew longipennis sp, n. 56, 3 Fore wing. 57. Phallosome, 58. Hypandrium. 59. ¢ Epiproct. 60, ? Fore

wing. 61. 2 Epiproct. 62. Gonapophyses, 63, ¢ Ninth sternite. 64, Subgenital plate.

Discussion: Ptycta longipennis is very similar to P.

muogamarra Smithers, from New South Wales, It

is, however, larger and there are distinct differences

in several anatomical features, In the male the distal

median extension of the phallosome is longer and

the median strap-like upcurved part of the hypan-

drium is more nearly symmetrical, In the female the

external valve of P, longipennis is broader and,

although reduced in a similar way, the ventral valve

is more robust. In P. glossoptera the hypandrium is

strongly asymmetrical and in the other Australian

species the hypandrium bears a variety of spines. and

projections,

Females of P. emarginata New, P, glossoptera

New, P, improcera New, P. picta New and P. umbrata

all have some darker brownish marks on the fore

PSCOPTERA OF SOUTH AUSTRALIA 479

FIGS. 65-69. Piveta hollowayae sp. n, 65. 2 Fore wing. 66. 2 Epiproct, 67. Subgenital plate. 68, 9 Ninth sternite,

69. Gonapophyses.

480)

wings, The female of P. cornigera is not known but

the male does not have obvious dark marks. P.

longipennis differs from P. hallowayae sp. n.

(described below) in lacking the rugose areas along

the distal border of the epiproct and in not having

an apical thickening to the posterior median process

of the phallosome; phallosome shape also differs in

that the phallic frame is much narrower in P.

hollowayae and has an anteriorly projecting apo-

physis on each side at the posterior third of the

frame, The fore wing of the female of P. hellowayae

has an interrupted, irregular brown band from

stigmapophysis to nodulus. In P. hollowayae the

dorsal valve of the gonapophyses is broad and the

ventral valve not shortened to the extent that it is

in P. longipennis.

REC. 8. AUST. MUS. 18 (20): 453-491,

May, 1984

Ptycta hollowayae sp. n.

Female

Coloration (in alcohol); Head and appendages

very similar to Piycta emarginata New but with post-

clypeal striae not obsolete in midline and frons with

a small dark circle in midline anterior to ocelli.

Thoracic lobes dark brown but each broadly

bordered with pale areas. Fore wings (Fig. 65)

hyaline with very faint overall brown tinge and

marked in shades of brown. Hind wings hyaline

with very faint suggestion of brownish area behind

end of Cus.

Morphology: Length of body: 3.8 mm. Median

epicranial suture distinct, anterior arms absent. Head

broad across vertex, eyes large, just reaching level

FIGS. 70-73, Prycta hollowayae sp. n. 70, & Fore wing. 71. 3 Epiproct. 72. ¢ Paraproct. 73. Phuallosome.

PSCOPTERA OF SOUTH AUSTRALIA

of vertex. LO/D: 1.8; PO: £.0. Ocelli large, anterior

oeellus only little smaller than lateral oeelli. Length

of flagellar segments: f,: 0.96 mm; f,: 0.76 mm.

Measurements of hind leg: F: 0,92 mm; T; 2.0

mm; t,: 0.56 mm; ty: 0.20 mm; rt: 2.8:1; et: 22,

4, Front femora noticeably broader than femora of

middle and hind legs. Fore wing length; 4.4 mm;

width: 1.4 mm, Fore wing with Rs and M meeting

in a point, joined by a crossvein or fused tor a very

short length. Pterostigmal spurvein minute, hardly

discernible, Fore wing glabrous. Hind wing length:

3.3 mm; width: tt mm. About ten marginal setae

between Re;g sod Rass. Epiproct (Pig. 66).

Gonapophyses (Fig, 69). Dorsal valve broad with

fairly long apical, pointed extension, External valve

with distinct lobe lying adjacent to dorsal margin of

dorsal yalye, Subgenital plate (Pig. 67) with short,

truncate, posterior lube on the upper side of which

near the posterior margin is a small, irregularly shaped

thickening of the internal membrane, Y-shaped pig-

mented urea with shoct, broad, “stem”, the ends of

the “arms” broad but somewhat apically divided.

Selerificuiion of ninth sternile (Fig. 68) in form of

an ovoid, very heavily sclerotized plate,

Male

Colorahem Cn alcohol); Head and appendages

as In female but head markings reduced in uccor-

dance with greater eye size. Pore wings (Fig. 70).

Hind wings hyaline.

Morphology: Length of flagellar segments; f;:

1,04 mm; fu: 0.84 mm. Antennae thicker than in

femule, finely pubescent, 1O/D: 1,1: PO: 1.0, Eyes

mich larger than in female, reaching well above

level of vertex with inner margins diverging pos-

teriorly when viewed from above; emarginate

posleriorly-medially above. Measurements of hind

leg! Fi 1.0 mm; T: 2.24 mm; ty: 0.60 mm); t*}

0.18 mm; rt: 3.3:1; ct: 27. 4, Fore wing length;

5.0 mm; width; 1-7 mm. Pterostigmal spurvein

absent, Rs and M joined by a very short crossyein,

in length only a little greater than vein thickness.

Hind wing length: 3.7 mmy; width: 1,3 mm, A few

fine setae on margin between Ros, and Ry)s -

Epiproet (Fig. 71) lightly sclerotized with broad,

thickened, marginal band of varying width; hind

margin transverse; a basal, upstanding, broad-

margined lobe partly overlies ninth tergite, the lobe

medially slightly ¢marginate. Paraproct (Fig. 72).

Hypandrium similar ta that of Pivela glossaptera

(New 1974, Fig. 62), Phallosome (Fig, 73).

Material Exanined: SOUTH AUSTRALIA,

(holotype), & Callotype), 15 km W Tailem Bend,

13.v.1980, G. and J. Holloway. Paratypes: 1 2%, as

holotype. | 2°, 18 km N Ardrossan, 8,v,1980, G, and

48)

J. Holloway. 1 ?, | km E Edithburgh, 7.v. 1980, G.

and J. Holloway.

Holotype, allotype and paratypes in the Aus-

tralian Museum.

Discussion: Piyeta hellowayae resembles P. emiar-

ginara New (? only known) and P. glosseptera New

(both sexes know), Jt differs from bath in being

larger und the female has more extensive wing

markings. The male of P. Aollowayvae differs from

that of P, glosseprera in the form of the paraprocts

and in proportions of the phallosome (although

general shape is similar). The Jateral projections are

more pronounced in P. hollowayae, The epiproct and

the hypandrium are similar in the two species. The

anterior lobe of the epiprect appears ta be folded

back in the illustration given by New (1974, Fig.

64), In P, cornigera, PL impracera and P. wnbrata

the hyandrium bears spines and various processes

which are not present in P. hellowayae. From P.

hiuegaiara P. hollowayae dilers in the form of

proportions of the phallosame and in not having

the ventral valve of the yonapophyses reduced. Com-

parison with P. longipennis has been made above.

Tanystigma tardipes (Edwards)

Clematostigina tardipes Edwards, 1950. Pap. &. Soe.

Tasm. 1949: 95, Figs. 1-17.

Copastigma (Clematostigna) tardipes (Edwards).

Smithers, 1967, Aust, Zoal, 14 (Ll): 103,

Tanystigina rardipes (Edwards), Smithers, in press.

Aust, ent. Mag.

Material Examined) SOUTH AUSTRALIA. 1 1

9, 12 km SE Port Wakefield, 8.v.1980, G. and J.

Holloway. I 4, 3 2, 1 n, 4 km S Moonta, 7.v, 1980,

G. and J, Holloway, 1 2, Port Elljot, 13.v.1980, G.

and J, Holloway, 2%, Wilmington, Flinders Ranges,

6.v.1980, G, and J. Holloway. 1 ¢, 2 4%, 18 km N

Ardrossan, 8.vy.1980, G. and J. Holloway. 2 4, 2 2,

23 km E Tailem Bend, 13.v.1980, G. and J. Hailo-

way. 1 g, 1 2, Telowie Gorge, 10 km E Pt. Ger-

mein, 20,v,1981, G, und J, Holloway, 4 2,5 9,

Farina, 1 7.v.198L, G. and J. Holloway.

T. tardipes was described from Tasmania and has

been recorded from Victoria.

Tanystigma elongata sp. n.

Male

Coloration (in alcohol): Head pale grey-brown

with dark brown markings; A double row of con-

fluent spots adjacent to median epicranial suture,

werass back of vertex and adjacent to compound

eyes; a spot between ocellar tubercle and epistomial

suture; an irregular ring around antenna base; dis-

linct postclypeal striations. Epicramal suture dark

brown Labrun) pale brown, Genae grey-hrown.

Antennae dark brown. Eyes purplish. Ocellar

482 REC. 8, AUST. MUS, 18 (20); 453-491,

May, 1984

FIGS. 74-78, Tanystigina clongata sp. un, 74. d Fore nite. 75, Phallosome. 76, Hypandrium, 77, ¢ Puraproct, 78. @

piproct,

tubercle dark brown, First and second maxillary palp

segments very pale brown, third and fourth segments

dark brown, Mesothoracic notum dark brown

except for pale brown parapsidal sutures and pos-

terior borders of dorsal lobes. Femora brown with

darker apical band; tibiae pale brown, darker at

each end; tarsi dark brown. Fore wings hyaline with

brown pattern (Fig. 74). Hind wings hyaline, veins

brown, Abdomen pale with irregular lateral seg-

mentally areanged marks; terminal structures very

dark brown.

Morphology: Length of body: 3.1 mim. Median

epicranial suture distinct, anterior arms not evident.

Epistamial suture sinuous, curved forwards in

middle, interior to anteocellar spot. Length of

flagcllar segments: f;; 0.90 mm; f.: 0.84 mm.

Antennae fine, flagellar setae up to three times as

long as Hagellar diameter. Eyes large, reaching a

little above level of vertex. IO/D: 1.2: PO; 1,0,

Eyes slightly emarginate opposite antenna base,

Measurements of hind leg; F: 0.76 mm; T: 1.80

mm; ty! 0.46 mm; t.: 0.20 mm; rt: 2.3:1; cl: 21,

4. Tarsal segments long, ctendia large. Fore wing

length: 4.9 mm; width: 1.6 mm. Fore wing. Fore

wing (Fig. 74) with Sc ending in R. R, almost

straight basad of hind angle of pterostigma; R,

almost straight between hind angle and wing margin,

Pterostigmal spurvein obvious. Rs and M fused for

a short length, Veins somewhat evanescent at forking

of Rs, second half of Cu,), basal section of Cu,,

and the third quarter of outer margin of discoidal cell,

Cuz, and M fused for a very short Iength. Hind

margin of wing near base thin, rugose. A few small,

fine selae on wing margin from proximal end of

pterostigma fo wing apex, Hind wing length: 4.0

mm; width; 1,2 mm. Hind wing with Rs and M

fused for a length. A few small marginal setae

between Ro+s and wing apex, Epiproct (Fig, 78)

well sclerotized, less so laterally, with a pair of small,

crect basal lobes, Paraprocts (Fig. 77) with large

field of trichobothria and a lightly sclerotized, setose

dome between trichobothria and the apical spur.

Apical spur broad-based with sharply pointed,

PSCOPTERA OF SOUTH AUSTRALIA 483

FIGS, 79-84, Tanystigma clongata sp. n. 79, 2 Fore wing, 80, 9 Hind wing, 81. Subgenitai plate. 82. @ Ninth sternite.

83, 2 Epiproct, 84, Gonapophyses,

dS4

curved apex, Posterior margin of ninth tersite

strongly sclerotized im middle section, the sclera-

lizution tupering laterally, Hypandrium (Fig. 76)

yery heavily sclerotized wilh two inwardly curved,

posterior projections which are laterally serrate

between the bases of which the hind margin of the

hypandrium is medially emarginate and bers a

small median, rounded ventral sclerite, Phallosome

(Fiz 75, ulted in preparation) consistiny of two

elongate, narrow, basally fused, distally divergent

external parameres each with pointed, apicul sclerite.

Fentale

Colorqtion (in aleohol): Body ¢oloration as in

mile but with a medion pale hrawn line on ante-

dorsum of mesothorax and abdomen distmetly

banded with brown, Antennae os in male but scape,

pedicel and first two flagellar segments yery pale.

Third maxillary pal scgment pale in distal hall.

Labrnm pale, darker medially. Legs as in male,

Fore wings (Fig. 79) hyaline with pattern more

extensive than in male. Hind wings (Fig. 80) with

some very pale brown colour.

Morphology: Length of body: 3.5 mm. Length af

flagellar segments: Iy° 0.80 mm; fur O80 mm.

Antennae fine, flagellar setae about as long as

flagellar diameter, that is, setae are relatively much

shorter than in male, Eyes much smaller than in

ale. not reaching level of vertex. TIO/D: 2.2; PO:

Q.8. Anterior oeellus much smaller than lateral

ocelli. Epistomial suture sinuous anterior ta ocellar

tubercle, Measurements of hind leg: F; 0.72 mm;

T; 1.56 mm; ty): 0.38 mm; te: 0.20 mm) rts 1,9) 1)

ct: 19, 0. Ctenidia small. Fore wing length: 4.1

mm; width: 1.4 mim, Fore wing (Fig. 79) similar

lo that of tale but pterostigma a little broader,

with murgin both basad and distal of hind angle

very slightly sinuous. Spurvein obvious. Areola

pustica. joined to media by a short crossvein. Mar-

ginal Selae as in male, between base of plerostigma

and wing apex. Epiproct (Fig. 83). Subgenital plate

(Fig. 81) with long, rounded posterior lobe with

broad, irregular median band without pigment.

Gonapophyses (Fig, 84), Selerifications of ninth

sternite (Fig, 82),

Material Examined: SOUTH AUSTRALTA. ¢

(holotype). ¥ Callotype), 20 km SE Port Augusta,

Horrocks Puss, 17.vi1.1979, G. A. Hollaway. Para-

types: 1 9, Mt. Ohlissen Bragge, Wilpena Pound,

18.v.198l, G. and J. Holloway, 4 2, Germein

Gorge, 20.v.1981, G. and J. Holloway,

Holotype, allotype and paratypes in the Austra-

lian Museum.

Discussion: Tanya elongata is the only species

of the genus in which there ate extensive wing

REC. S, AUST. MLS, 18 (20): 453-491.

May, (98d

‘markings in eetls Ry, Ry and Rs, Cu, Cus and the

discoidal cell. Lt is easily recognized on this feature.

Tatystigma bifurcata sp. n.

Fomale

Coloration (in alcohol): Head pale buff with

dark brown spotting on either side of mediun epi-

cruml suture, across back of vertex and adjacent

lo compound cyes. Area between epistomial suture

and ocellar tubercle wilh ovoid brown mark en-

closing a pale central spot. Postelypeus pale bulf

with fine parallel brown lines, very distinct, running

forward ftom epistomial suture but mot reaching

unteclypeus, Labrum dark brown with median,

amerior, semicircular pale area. Genae. pale, not

marked. Scupe. pedicel and basal part of first fla-

wellar segment brown, rest of antenna dark, almost

black, Fyes black. Ocelli on very dark brown

tubercle, First and second maxillary palp sczments

pale; third segment dark in’ basal half, dark dis-

tally; fourth segment dark brown, Mesothoracic

notum dark brown with aumedian pale stripe; dorsal

Jobes. with a small pale area near postero-lateral

corner, Metanotum similar to mesonotum, Pleura

dack brown with same puler areas. Coxae dark

brown. Femora pale with some ifegular brown

marks und slight suggestion of preapical dark bund.

Vibiae pale brown, Tarsi brown. Fore wings (Pig.

85) hyaline with brown markings. Hind wings hya-

line, with faint brownish patch behind distal end

af Cus.

Morphology: Length of body: 3.5 mm. Head

with well rounded vertex. Median epicranial suture

very distinct, Postelypeus fairly bulbous. Lengths of

flagellar segments: #4: 0.64 mmy fo: 0.52 mm. Eyes

fairly large. TO/D: 1,1; PO: 0,80. Lateral ocelli

very large, anterior ocellus small, Measurements of

hind leg: F: 0.66 min; T; 1.44 mm; ty: 0.36 mm;

tos O.16 mmy rts 2.321; et: 18, 1. Fore wing length:

4.) mm; width: 1.40 mm, Subcosta well developed

basally, straight, becoming evanescent in costal cell.

Stigmopophysis well developed, dome shaped. Ry

beyond sligmapophysis fine, ie, hind margin of

pterostizma fine, Spurvein very small. Postptero-

stisinal mark indistinct as it is of same colour as

pigniented area but can be recognized by difference

in texture [rom base to hind angle of pterostigma.

Rs and M fused for a length, Rs and branches of

Rs evanescent neur bifurcation, M between Rs and

Cu,, Strongly curved to give a concave outer margin

to discaidal cell. Hind wing length: 3.2 mm; width:

1,0 mm, Se evanescent in costal cell, Rs. and M

fused for a length, Wing margin between Ross

and Rass with twelve well developed but short

setuc. Epiproct (Fig. 86), Paraproct (Fig, $9). Sub-

genital plate (Pig, 87). Gonapophyses (Fig. 90).

Sclerificationy of ninth sternite (Fig. 88).

PSCOPTERA OF SOUTH AUSTRALIA 4f5

FIGS, 85-90. Tanystigma bifurcata sp. n. 85. 2 Fore wing. 86. 9 Epiproct. 87, Subgenital plate. 88. 9 Ninth

sternite. 89, & Paraproct. 90. Gonapophyses.

186 REC. S. AUST, MUS, 18 (20): 453-491,

May, 198¢

PIGS, 91-94, Sanyitigina bifurcata 3p. 1. 91. & Bore wing. 92, Phallosome. 93, 3 Epiproct, 94, Hypundrium.

Male

Coleraia (in alcohol): Body coloration as

female. Fore wiugs (Fig. 91) hyaline. Hind wings

hyaline without brownish area behind Cus.

Morpholegy: Postclypeus not as bulbous as in

female. Length of flagellar seoments; f;: 0,80 nim;

fu; 0.64 mm. Byes very large, reaching just above

level of vertex, LO/D: 1.0; PO: 0.94, Ocellar tubercle

very well developed. Measurements of hind leg: FP:

0.72 mm; T- 1-72 mm; t)! 044mm; ta: 0.18 mm) rt:

2.4: 15 ct: 18,4, Legs long and thin. Pore wing length:

4.2 mm; width: 1.5 mm, Fore wing with indistinct

plerostigmal spurvein but postpterostigmal mark

Well developed. M distad of separation from Rs

strongly curved to give concave discoidal cell, Hind

Wing length; 3,1 mm; width; 1.l mm, A few fine

marginal setae between Roys and R445. Epiproct

(Fig, 93), Hind margin of ninth tergite well sclero-

tized with two small projections against which the

epiproct articulates. Latero-ventral margin of ninth

tergite on each side ends in a rounded apophysis,

Hypandrium (Fig. 94). Phallosome (Fig. 92),

Material Examined: SOUTH AUSTRALIA. ? (holo-

type), & (allotype), 18 km S Port Pirie, 7,v.1980,

G. and J. Holloway. Paratypes: 7 2, 7 c', as holotype.

1d 1 2%, 15 km N Port Broughton, 7.y.1980, G.

and J, Holloway, 1 2, 4 km E Pinnaroo, 14.v.1980,

G. and J, Holloway, 1 %, Alligator Gorge Rd., near

Mt. Remarkable, Flinders Range, 17.vi.1979, G. A,

Holloway, 1 @, Qverland Corner, 15.vi.1979, G. A-

Holloway,

Holotypes, allotype and paratypes in Australian

Museum.

Discussion: Tanystignia bifurcata is yery similar to

Psocidus nolialiy Smithers, described from Western

Australia, In both species the pterostigmal spur-

vein is very small, the hypandrium is symmetrical

and somewhat bilobed and (he phallosome has

apically divided external parameres, In all other

species of Tanysiigma the external parameres are not

so divided. The female yenitalia differ in details

Irom those of other species of Tanystigina and the

extent ol wing marking is considerably greater than

in females of Ps, norialis, T. pantuin (Smithers) and

T. tardipes (Edwards). [t is similar ta that of T.

dubium (New) but that species lacks the dark mark

at the wing margin in cell Ry. Ps. notialiy was not

placed in Tunystigia when the latter genus was

erecied because of the bifurcation of the external

parameres and the very small pterostigmal spurvein

in both sexes, The discovery of T. bifurcata, haw-

ever, indicates the inclusion of both in Tanystigma.

T. netialis (Smithers) comb, nov, and T. hifiureata

do stand apart somewhat from the other species

of the genus in having apically divided external para-

meres. They differ from cach other in that T. bifur-

cata has a longer phallasome and the wing mark-

ings are less extensive in female 7. notialis than

they are in T, bifureata. The statement made by me

PSCOPTERA OF SOUTH AUSTRALIA 487

FIGS 95-100. Psecidus mouldsi sp. n. 95. 2 Fore wing. 96. Subgenital plate. 97. 2 Epiproct. 98. @ Ninth sternite. 99.

@ Paraproct. 100, Gonapaphyses,

+08

(Smithers, 1972) that “.., Ps. netiulis will probably

he found to be cangenerie with ,..” Clematostigma

tardipes Edwards and C, meculiceps Enderlein has

not been supported by subsequent study of material

of &. maculiceps (Smithers, 1983).

Psocidus mouldsi sp. n.

Female

Coloration (in alcahol): The colour pattern of

this species is very similar to that of the males of

Blayre qigusta, described above. The spotting on the

head is fiver and the spots quite discrete and the

ovoid mark anterior to the ocellar triangle is very

conspicuous, The terminal structiices of the abdo-

men are dark, the Subgenilal plate being very con-

spicuous owing to the heavily sclerotized T-shaped

area, Fore wing (Fig, 95) hyaline, marked in various

shades of brawn,

Morphology; Length of body: 4,0 mm. Brachy-

ptcrous, fore Wings not reaching end of abdomen.

Lengths af flagellar segments: fy: 0.64 mm; fos 0,52

moi, Eyes nat reaching vertex level. IO/D: 3.4)

PO; 0.83. Measurements of hind leg: F: 0.64 mm;

T: 1.84 mm ty): O48 mm; te: 0.20; rt: 2.4:1;

et: 21, 2. Fore Wing length: 3.4 mm; width: 1.3 mm,

Rs and M fused for a yery short Jength, Third

median cell narrow and almost parallel sided. Hind

wing length: 2.9 mm; width: 1.0 mm, Rs and M

fused for a short length, Epiproct (Fig, 97) lightly

sclerotized with two, small, curved sclerotized ares

about half way along epiproct and midway between

middle and Jateral edge of epipract. Paraproct (Fig.

99) with two internal accessory sclerites attached

10 paraproct by membrane (displaced in ilustra-

lion). Subgenital plate (Fig, 96) with heavily

sclerotized transverse bar basad of posterior Jobe;

posterior lobe long with unusual pattern of sclero-

tization and pigment, having an ovoid, less sclero-

tized area near base of lobe. Gonapophyses (Fig.

100) with long, aeuminate ventral valve; dorsal

viilve broad, rounded behind, constricted at the

middle und supported by longitudinal sclerifica-

(ions; external valve with dorsal, well sclerotized,

posteriorly directed lobe. Sclerifications of ninth

sternite (Pig, 98) more complex than usual in the

penus,

Male

Unknown,

Material Examined: SOUTH AUSTRALTA, 2

(holotype), 40 km E Nullarbor, 131°15'E, 31°25'S,

29.1x.1978, M. S. and B. J. Moulds. Paratype: 1 3,

as holotype, Holotype and paratype in the Austealian

Museum.

The genus Psovieiys was erected by Pearman

(1934) to hold a large assemblage of species which

had been described in the genus Pocus Latreille

REC. $, AUST, MUS. 18 (20): 453-491,

Vay, 1984

but which could not be retained in his restricted,

redefinition of that genus. Psecidus, therefore, con-

tains many unrelated species. With time it iy hoped

that they will be redistributed amongst present

genera or logically placed in new genera. For the

present Psocidus remains a “holding” genus.

The relationship of Ps. mouldsi and Ps. parilla

sp. n. (described below) are not known, They both

stand apart trom other species of the family and it is

hoped that when further material, including males,

is forthcoming. their position will he clarified.

Psocidis parilla sp. n.

Feniale

Celeration (in alcohol): Head pale grey-brown

with a double row of brown, irregular, confluent

spots adjacent 19 compound eyes. across back of

vertex and on either side of median epicrantal

suture; an oval brown mark anterior to acellar

tubercle; tubercle brown; postclypeal striations

brown; genae not marked. Labrum dark brown.

Antennae almost black. Eyes black. Maxillary palp

with dark third and fourth segments; first and second

segments pale, Mesothoracie notum shiny dark

brown with pale parapsidal furrows, pale line along

postero-lateral parts of lateral lobes and hind half

of mesoascutellum pale. Pleura dark shiny brown.

Coxae dark brown, Femora brown, darker along

anterior and posterior sides with some irregular

darker dorsal marks in distal quarter. Tibiae pale

brown, darker ut each end. Tarsi dark brown. Fore

Wings (Fig, 101) hyaline with pattern in various

shades of brown, Hind wings hyaline with pale

brown patch in distal) quarter of cell Cus adjacent

to vein Cu, but not in distal corner of cell, Abdo-

men pale with dorsal and lateral irregular, seg-

mentally arranged marks. Terminal structures very

dark brawn,

Morphology: Median epicranial suture distinct.

anterior arms evanescent, Epistomial suture sinuous

in middle, curving away from ocellar triangle in

middle section. Length of flagellar segments: f,:

0.92 mm; fy: 0.68 mm. Eyes fairly large. TO/D:

2,5; PO: 0.9, Ocelli large, anterior ocellus smaller

than Jateral ocelli, Measurements of hind leg: F:

0.84 mm: T: 1,92 mm; t,: 0.44 mm; ty: 0,14 mm,

cit 3:12 et: 19, 3. Fore wing length: 4.7 mm; width:

1.4 mm. Fore wing (Fig. 101) with Se ending in R

about hal! way belween wing base and base of

pterostigma, Ry basad of hind angle of pterostigma

yery slightly curved to give a slightly cancave ptero-

Stigma, beyond apex slightly convex. Rs and M fused

for 4 length, M and Rs and Cuy, curved to give

fairly strongly concave discoidal cell. Cu,, fused

with M for a length, Cu,, sinuous basad of fusion:

basal section of Cuy, and apex of arcola postica at

PSCOTERA OF SOUTH AUSTRALIA 489

FIGS. 101-105. Psocidus parilla sp, n. 101. 2 Fore wing, 102, Subgenital plate. 103. 2 Ninth sternite. 104. 2 Epiproct

105. Gonapophyses.

40 REC. 8. AUST. MUS, 18 (20); 453-49).

Slight angle to one another, Fore wing glabrous.

Hind wing length: 3.4 mm; width: 1.1 mm. Hind

wing with Rs and M fused for a Jength, Costal cell

near base broadened, the anterior margin slightly

thickened and finely rugose in broad region, ‘This

part of costal margin would lie adjacent to a slight

thickening of the hind margin of the fore wing in

{light and these two structures probably assist in wing

coupling. A few fine marginal sctae between Rey 3

and wing apex, Epiproct (Fig, 104), Subgenital

plate (Fig. 102). Gonapophyses (Fig, 105).

Sclerifications of ninth sternite (Fig. 103).

Male

Unknown,

Material Examined: SOUTH AUSTRALIA,

(holotype), Pooginook, 5.vi.1979, A. Holloway,

Paratypes: 2 7, as holotype. 2 2, 2 km E Parilla,

23.vi.1979, G, A, Holloway. 1%, 4 km NW Murray

Bridge, 22.v.1981, G,, J, and A. Holloway,

Holotype and paratypes in the

Museum.

Australian

Discussion: See under Psocidus mouledsi-

Family MYOPSOCIDAE

Phlotodes australis (Brauer)

Psocus australis Braucr, 1865. Fer. zool. -hot. Ges.

Wien 15: 908,

Psocus griseipennis McLachlan, 1866. Trans, ent.

Soc, Lond. (3) 35: 348.

Myopsocus griseipennis (McL.). McLachlan, 1866.

Trans, ent, Soc, Lond, (3) 5: 352,

Myopsocus novaezealandiae Kalbe, 1883. But.

Nachr, 9: 145,

Psecus zelandicus Hudson, 1892, Manual of New

Zealand Entomology, p. 107; Pl. XVI, Figs. 2,

2a.

Phlotedes griseipennis (McL,). Enderlein, 1910.

5.8, Ges, naturf. Fr, Berl, 1910 (2): 67,

Atay, 1984

!Myopsocus griseipennis (McL.) Edwards, 1950.

Pap. Proe, R. Soe. Tasm. 1949; 104, Figs, 26-32,

Phlotodes australis (Braver), Smithers, 1975. Ause.

ent. Mag. 2 (4): 76.

Material Examined; SOUTH AUSTRALIA, | ¢

Belair, 28,ix.1935, H. Womersley.

This species has been recorded from all states

except the Northern Territory.

Phlotodes hickmani (Smithers)

Myopsocus australis (Brauer), Hickman, 1934. Pap.

Proc. R. Soe. Tasm. 1933: 85.

Myopsocus griseipenniy (Mcl,), Edwards, 1950.

Pap, Proc, R, Soe, Tasm, 1949) 104, Figs, 26-32.

Myopsecus hickmani Smithers, 1964. Proc. R. ent,

Suc, Lond, (B) 33: 135,

Phlaiodes hickmant (Sm.). Smithers, 1971. J. Aust.

ent. Soc, 10 (1): 24,

Material Examined: SOUTH AUSTRALIA. 3 2,

Morialta, 14.v.1940, H, Womersley. 6 ?, Magill,

6.11.1884, Tepper.

P. hickmani is known from Tasmania and Vic-

toria.

ACKNOWLEDGEMENTS

I would like to thank Professor VY. Y. Hickman

for material of Tasmanian species of Psocoptera for

comparison wilh South Australian matenal; the

several collectors who provided other material,

especially Geoffrey, Janct and Andrew Holloway

and Max and Barbara Moulds; Gordon Grass for

arranging the loan of South Australian Museum

material; the Director of the South Australian

National Parks and Wildlife Service for permission

for work to be done in areas under his jurisdic-

tion; the Australian Research Grants Committee for

providing funds in support of studies of the Pso-

coptera of the Melanesian arcs and possible source

areas; and Martyn Robinson for preparing the illus-

trations to this paper.

REFERENCES

BANKS, N, 1939. New genera and species of Neuropteroid

Insects, Bull, Mus, camp. Zool. Harv, 85 (7): 439-

S04,

CHAMP, B. R. and SMITHERS, C. N, 1966, The insects

and miles associated with slored products in Qucens-

land. 1. Psocoplera. Qd J. Agric. Anim, Sei 22. (3):

259-262,

ENDERLEIN, G. 1903. Die Copeognathen der indo-aus-

tralischen Paunengebietes, Ann, Hist. -nar. Mus. hung.

1: 179-344,

ENDERLEIN, G 1907, Copeognatha. Jn Michaelsen., W-

and Hartineyer, R. Die Fauna Siidwest-Australiens. 1:

233-240), ;

HICKMAN, Y, ¥, 1934. A contribution to the study of

earn Copeognatha, Pup. ray. Soc. Tas, (933;

77-84.

McLACHLAN, RB. 1866. New genera and species of Pso-

cidae, Trans, ent. Soe. Land. 5 (4): 345-352.

NEW, T, R. 1971. The Psocoptera of Curlis Island, Tus-

mania, J. Aust. ent. Sac, 10 (4): 223-229,

NEW, T.. R, 1973, New species and records of Peripsacus

Hagen (Psocoptera, Peripsouldac) from south east

Australia. J. Aust. ent. Soc. 12: 340-346,

NEW, T. R. 1974. Psocidae (Psocopteru) from southern

Australia. J. Aust. ent. Sac, 13: 285-304.

NEW, T. R. 1975. Aerial dispersal of some Victorian

Psocoptera as indicated by suction trap catches, J.

Aust. ent, Soe. b4: 139-184.

PEARMAN, J, V. 1934. New and little known Africun

Psocoptera. S/ylops 3 (6): 121-132.

SMITHERS, C. N, 19640. New records of Cave and Mine-

dwelling Psocoptera in Australia. J ent. See Qi. 3:

85,

SMITHERS, C. N, 1964b, The Myopsocidae (Psocopleral

of Australia. Pree. R. ens. Sec. Lund. (B) 33: 133-

138.

PSCOTERA OF SOUTH AUSTRALIA 491

SMITHERS, C. N. 1967. A catalogue of the Psocoptera

of the world. Aust. Zool. 14 (1): 1-145.

SMITHERS. C. N. 1970. Psocoptera. In “Insects of Aus-

tralia”. CSIRO, Melbourne.

SMITHERS, C. N. 1972. A collection of Psocoptera (In-

secta) from Western Australia, including four new

species. Aust. Zool. 17 (1): 15-23.

SMITHERS, C. N. 1977. The Psocoptera of Muogamarra

Nature Reserve. Rec. Aust. Mus. 31 (7): 251-306.

SMITHERS, C. N. 1983. A reappraisal of Clematostigma

Enderlein with notes on related genera (Psocoptera,

Psocidae). Aust. ent. Mag. 9 (5): 71-79.

THORNTON, I. W. B., and NEW, T. R. 1977. The Philo-

tarsidae (Insecta, Psocoptera) of Australia. Aust. J.

Zool., suppl. ser. 54: 1-62.

TILLYARD, R. J. 1923. A monograph of the Psocoptera,

or Copeognatha, of New Zealand. Trans. N.Z. Inst.

54: 170-196.

HELMINTH TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

Il. ACANTHOCEPHALA AND CESTODA

BY LESLEY R. SMALES

Summary

Type-specimens of 16 species of Acanthocephala and 80 of Cestoda are catalogued. All except two

are from Australian or Antarctic hosts. Species are listing the helminth types held by the South Aus-

of the genus. Wher a name change has occurred the most recent accepteable name is included.

HELMINTH TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

1, ACANTHOCEPHALA AND CESTODA

LESLEY R. SMALES*

South Australian Museum, Adelaide, South Australia 5000

ABSTRACT

SMALES, L. R. 1983, Helminth type-specimens jij the

South Australian Museum, TL Acanthocephala and

Cestoda. Ree. 8. Aust, Mus, 18(21)) 493-501,

Type-specimens of 16 species of Acanthocephala

and 80 of Cestoda are catalogued, All except two

are from Australian or Antarctic hosts, Species are

listing the helminth types held by the South Aus-

of the genus, Where a name change has occurred

the most recent acecptable name is included.

INTRODUCTION

This is the second of an intended series of papers

listing the helminth types held by the South Aus-

tralian Museum and the Australian Helminthological

Collection, currently housed in the Museum,

Species are arranged alphabetically according to

the original name of the genus, Full synonymies are

not given, but where a name change has occurred,

the most recent acceptable name is given together

with the relevant reference. Host-species names are

those used in the original descriptions. Types held

in the South Australian Museum have registration

numbers prefixed by V. Types held in the Australian

Helminthological Collection have registration num-

bers prefixed by AHC or AHC S.

The following abbreviations are used in the text.

A.A.E. = Australian Antarctic Expedition, 1911-

1914; BANZARE — British Australia and New

Zealand Antarctic Research Expedition, 1929-1931;

SAM = South Australian Museum; AHC = Aus-

tralian Helminthological Collection; N.T, =

Northern Territory; N.S.W, = New South Wales;

Old = Oneensland; $.A, = South Australia; Tas. =

Tasmania; Vic. — Victoria; W.A. — Western Aus-

tralia.

PHYLUM ACANTHOCEPHALA

The earliest descriptions of type material held at

the South Australian Museum are those of T. H.

Johnston, Professor of Zoology, the University of

Adelaide, Honorary Director of the Museum 1928-

1931, and Chairman of the Museum Board 1940-

1951, Later deseriptions are by S$. J, Edmonds, Hon-

* Present address: Gippsland Institute of Adyanced Educa-

tion, P.O, Box 42, Churchill, Victoria 3845.

orary Associate of the Museum, who is the foremost

Australian authority on Acanthocephala and con-

tinues active research into the taxonomy of the

group.

Acanthocephalans are parasitic, wormlike pseudo-

coelomates that lack a digestive: system, A diagnos-

tic fealure of the group is the presence of an an-

teriorly placed, invaginable proboscis or introvert,

always armed with hooks,

The adult parasites infest the digestive tract of

vertebrates. Unless otherwise stated specimens listed

below Were dissected from the small intestine.

Arhythmorhynchus limosae Edmonds, 1971

Trans. R. Soe, §, Aust, 95 (2); 58-60,

Paratypes V2185-9, 3 4, 2 9, one in fragments

from Limeosa lapponica, Townsville, Qld,, collector

A. J, Bearup, 2871/59,

Arhythmorhynchus frassoni Molin, 1858

Trans. Roy. Soc. §. Aust. 74 (1): 3 (Johnston &

Edmonds, 1951), now 4, Johnston’ (Molin, 1858

sensu Johnston & Edmonds, 1951), Golvan, 1960.

Paratypes V2183-4, both ¢, one in fragments, from

Numenius cyanopus, Gladstone, Old,, collector T, H.

Johnston, 1910.

Note: Golvan, 1960 made the Australian specimens,

described by Johnston & Edmonds, 1951, a new spe-

cies but gave no reasons for his action.

Corynosoma australe Johnston, 1937

Prac. Linn. Sec. N.S.W. 62 (1/2): 13-16.

Syntypes V2227-9, 4 & and 2 @, from Arcrocepha-

lus forsteri, Pearson Is., S.A., collector J. B. Cleland,

January 1923.

Note: Johnston, 1937, recorded the type as being

deposited in SAM but this does not appear to have

been done. Johnston & Best, 1942 mention that the

type host for C. australe was incorrectly identified

and should have been listed as Neophoca cinerea.

Corynosomg cetaceum Johnston & Best, 1942

Trans, R. Soc. §, Aust. 66 (2); 250-2,

Holotype 2 V964, allotype @ V963, from Del

phinus delphis, Gult St Vincent, S.A., collector pro-

bably 'T. H. Johnston, date unknown,

Echinarhynchus hylae Johnston, 1914

Proc. R. Soc. Old. 36: 83.

494

Now Porrorchis hylae (Johnston, 1914) Schmidt &

Kuntz, 1967.

Type V2540, cotype V2539, from Hylae caerulea

encysted in the liver. Collector probably T. H.

Johnston, date unknown.

Locality: near Brisbane, Qld.

Echinorhynchus menurae Johnston, 1912

Proc. R. Soc. Qld. 24: 83.

Now Plagiorhynchus menurae (Johnston,

Golvan, 1956.

Type V2438, from Menura superba, near Gosford,

N.S.W., collector probably T. H. Johnston, date un-

known.

Note: Johnston, 1914 recorded that the types of E.

hylae and E. menurae had been deposited in the

Queensland Museum, Brisbane. However, on exa-

mination, specimens registered in SAM were found

to be Johnston’s original material.

1912)

Echinorhynchus zanchlorhynchi Johnston & Best,

1937

A.A.E. Rep. Ser. C 10 (2): 12-3.

Holotype ° V2200, from Zanchlorhynchus spinifer,

Macquarie Island, collected during A.A.E. 1911-

1914,

E'mpodius alecturae Johnston & Edmonds, 1947

Rec. S. Aust, Mus. 8 (4): 557-61.

Now Mediorhynchus alecturae (Johnston & Ed-

monds, 1947) Schmidt & Kuntz 1977.

Syntypes V2470-8, from Alectura lathami, Eidsvold,

Burnett River, Qld., collector T. L. Bancroft, date

unknown.

Gordiorhynchus bancrofti Johnston & Best, 1943

Trans, R. Soc. S. Aust, 67 (2): 226-9.

Now Centrorhynchus bancrofti (Johnston & Best,

1943) Golvan, 1960.

Holotype ¢ V2210, cotype @ V2211, from Ninox

strenua, Eidsvold, Burnett River, Qld., collector

T. L. Bancroft, date unknown.

Gordiorhynchus falconis Johnston & Best, 1943

Trans. R. Soc. S. Aust. 67 (2): 229-30.

Now Centrorhynchus falconis (Johnston & Best,

1943) Golvan, 1960.

Holotype ¢ V2212, from Falco berigora, Her-

mannsburg, N.T., collector T. H. Johnston, date un-

known.

Illiosentis furcatus Van Cleave & Linicome, 1939

BANZARE Rep. Ser. B 6 (5): 91-8. (Edmonds,

1957).

Now Illiosentis edmondsi (Van Cleave & Linicome,

1939 sensu Edmonds, 1957) Golvan, 1960.

Lectotype ? V2336, from Upenichthys porosus.

REC. S. AUST. MUS. 18 (21): 493-501.

May, 1984

Station 76, continental shelf east of Albany, W.A.,

collector T. H. Johnston, 1929-1931.

Note: The lectotype here designated was selected

from the original material by Edmonds.

Longicollum pagrosomi Yamaguti, 1935

Trans. R. Soc. S, Aust. 74 (1): 3-5. (Johnston &

Edmonds, 1951.)

Now Longicollum edmondsi (Yamaguti, 1935 sensu

Johnston & Edmonds, 1951) Golvan, 1960.

Holotype ¢, V2349, from Mylio australis, Brisbane

River, Qld., probably collected by T. H. Johnston

in 1918.

Mediorhynchus corcoracis Johnston & Edmonds,

1951

Trans. R. Soc. Aust, 74 (1): 1-3.

Holotype g V2502, allotype @ V2501, from Cor-

corax melanorhampus, Upper Burnett River, Qld.,

collector T. L. Bancroft, date unknown.

Neoechinorhynchus aldrichettae Edmonds, 1971

Trans. R. Soc. S. Aust. 95 (2): 55-8.

From Aldrichetta forsteri (posterior gut), Port Pirie,

S.A., collector L. M. Angel, Port Pirie, S.A., date

unknown.

Paratypes ¢ V2125, 9 V2126.

Pararhadinorhynchus coorongensis Edmonds, 1973

Trans. R. Soc. S. Aust. 97 (1): 19-21.

Holotype $ V2392, allotype ? V2393, from Aldri-

chetta forsteri, Coorong, S.A., collector J. Harris,

1962.

Paratypes V2381-84, 3 ¢ andl 9°.

Note: Edmonds, 1973 recorded the type specimens

(3 and ¢) as being deposited in the Australian

Museum but examination of specimens revealed that

both primary and secondary types had been regis-

tered at SAM.

Pararhadinorhynchus mugilis Johnston & Edmonds,

1947

Trans. R. Soc. §. Aust. 71 (1): 15-7.

Holotype ¢ V2399, from Mugil cephalus, Port Wil-

lunga, S.A., collector T. H. Johnston, 1939.

Note: Paratypes 2400-05, 5 @ showing introvert

well extended, genital complex and eggs, 1 ¢ show-

ing cement glands and genital ganglion.

Polymorphus bizurae Johnston & Edmonds, 1948

Trans. R. Soc. S. Aust. 72 (1): 71-4.

Holotype ¢ V2444, allotype @ V2447, from

Biziura lobata, Tailem Bend, S.A., collectors G. G.

& B. Jaensch and L. Ellis, date unknown.

Pseudoporrorchis hydromuris Edmonds, 1957

Trans. R. Soc. S. Aust. 80: 77-8.

From Hydromys chrysogaster, Innisfail, Qld., col-

lector N. G. Elliot, 10/10/55.

HELMINTH TYPE-SPECIMENS If 495

Nole: Type specimens do not appear to have been

deposited in SAM. There 1s addifional material

V2573-9, collector B. Ewers from Port Moresby,

Papua New Guinea,

PHYLUM PLATYHELMINTHES

CLASS CESTODA

Specimens described by G. Krefft, 1873, are the

oldest material in the type collection. Krelft’s work

was subsequently revised by 'T. H. Johnston. When

Johnston arrived in Adelaide he brought with him

much of the material which had been collected and

deseribed by him between 1909 and 1916 as well as

some of Krefft’s collection. Both T. H. and S.J,

Johnston worked on Australian cestode material dur-

ing the sume period, All the species designated John-

ston were described by T. H, Johnston, Continued

interest has been shown in the Australian cestoda by

a small number of workers since then, Some 80 type

specimens from hosts collected throughout Australia

and from Antarctic waters have been registered at

the SAM or in the AHC.

Unless otherwise stated specimens listed below

were dissected from the intestine of the vertebrate

host,

Acatuhotaenia tidswelli Johnston, 1909

Proc. R. Soc. N.S.W., 43: 103-16.

Paratypes AHC 2051, from Faranus varius, near

Bathurst, N.S.W,, collector T, H. Johnston, date un-

known,

Anomotachia asymmetrica Johnston, 1913

Rep, Aust. Inst. Trop, Med, 1911; 81-2.

Paratype AHC $95, from /erodias timorensis,

Townsville, Qld., from the collection of A, Breinl,

date unknown.

Anomotaenia rhinocheti Johnston, 1911

Proc, Linit, Soc, N.S.W, 36 (1): 70-4.

Cotype AHC $82, from Rhinecheius jubaius, col-

lected in New Caledonia, held in captivity in Sydney

for six months, collector H, E, Finck, date unknown,

Anthabothriunr sexorchidtim Williams, 1964

Parasitology 54: 741-3.

Holotype V1061, trom Taeniyra lvmna, Heron Is.,

oll Rockhampton, Qld,, collector J. Pearson, June

1956.

Balanoraenia hancrofii Johnston, 1924

Proc. Linn. Soc. N.S.W. 49 (3); 339-47.

Paratypes V1437-43, including serial sections, from

Tandanus tandanus, Eidsvold, Burnett Rd,, Qld.,

collectors TV, L, & M, J. Bancroft, date unknown.

Balanorcenia newguinensis Mackiewicz & Blair, 1978

J. Helminihal. 32: 201-3.

Paratypes V1158-59 from Tandanus brevidersalis,

Brown R., near Port Moresby, Papua New Guinea,

collector D, Blair, date unknown.

Bancroftiella ardea Johuston, 1913

Rep. Aust, Inst. Trap, Med. 1911: 85-6,

Now Parvitaenia ardea (Johnston, 1913) Baer &

Bona, 1960.

Lectotype V1673, paralectotypes AHC §309-11,

from Nyeticorax caledanicus, Townsville, Qld,, from

the collection of A. Breinl, date unknown.

Nate: The lectotype, selected by Bona (1975), was

labelled in T. H. Johnston's handwriting Nycticorax

caledoanicus, Glasshouse Mts., S, Qld., collector B. B.

Taylor,

Bancroftiella tenuis Johnston, 191la

Prac. Linn. Svc. NSW. 36 (1): SO-7,

Cotype AHC $92, from Macrepus ualabaius, cal-

lector A, S. Le Souéft, date unknown.

Note; Other species in this genus known only from

birds, Johnston (1913) observed that there was no

reason to doubt the accuracy of the collection data.

Bertiella foederata Beveridge, 1976

Aust. J, Zool, Suppl, Ser. 44: 14-16.

Holotype V797, paratypes V798, 799, holotype sec-

tions AHC S968, fram Mseudocheirus peregrinus,

Inverleigh, Vic., collector I, Beveridge, 1/6/75.

Berteiella mawsonae Beveridge, 1976

Aust. J, Zool, Suppl. Ser. 44: 16-18,

Holotype W794, paratype V795, from Schainobates

volans, Dartmouth, YVie,, 20/1/74, collector L.

Beveridge,

Bertiella petavrina Beveridge, 1976

Aust, J, Zool, Suppl. Ser. 44: 21-3,

Holotype V796, from Schoinohates volans, Tumut,

N.S.W., collector R, Smith, 1964-

Riacetabulum tandani Johnston & Muirhead, 1950

Rec, §, Aust, Mus. 9 (3): 346-8.

Holotype V847, from Tandanus tandanus, Tailem

Bend and Murray Bridge, $.A., collectors G. G_

Jaensch, L, Ellis and L, M, Angel, 1947-48,

Bothridium pyihonis yar, parvum Johnston, 1913

Rep, Aust. Inst. Trop. Med, 1911: 91-3,

Type material AHC 2054, from Varanus varius,

Townsville, Old., collector A. Breinl, date unknown.

Bothriocephalus kerguelensis Prudhoe, 1969

BANZARE rep, Ser. B 8 (9): 173-4,

Cotypes V115-27, 567-9, from Notethenia cyano-

hrancha, Royal Sound, Kerguelen, 20/10/29,

10/2/30; Netothena rossi Station 64, Kerguelen,

collector T, H, Johnston.

496

Nate; Prudhoe did not designate type host, or loca-

lity of lype in any of the cestodes he described from

the BANZARE Expedition for 1929-1931. All the

material sent to SAM was designated cotype material

by the author,

Calostaurus mundayi Beveridge, 1975

J. Helminthol. 49: 133-35,

Paratype V55, from Potorous apicalis, Launceston,

Tas,, collector B. L, Munday, date unknown,

Calostaurus thylogale Beveridge, 1975

J. Helminthol. 49+ 129-33,

Paratype V54, from Thylogale hilliardieri, Lautces-

ton, Tas., collector B. L, Munday, date unknown,

Cuapiuterilepis australiensiy Schmidt, 1972

J. Parasitol. 58 (6): 1089-91,

Holotype V L090, paratype V1091-02, from Antho-

choera carunculata, Nicholl’s Reserve, §.A., cal-

lected by SAM, July 1971.

Capiuterilepis meliphagicola Schmidt, 1972

J, Parasitol, 58 (6) + 1091-2.

Holotype V1108, paratype V1109-10, from Phile-

mon corniculatus, Caloundra, Old,, collector pro-

bably 'T. H. Johnston, 1920.

Caryoaustralis sprenii Mackiewicz & Blair, 1980

Proc. Helm, Soc, Wash. 47 (2); 169-72.

Paratype V1899, from Tandanus ater, Annan R.,

Helemvale, Qld., probable collector D. Blair, date

unknown.

Choanotaenia meliphagidarum Johaston, 191 1¢

Prac. Linn. Soe. N.S,\W, 36 (1): 58-70.

Now Pseudochoanotaenia meliphagidarum (Jotin-

ston, 1911) Schmidt, 1972,

Cotypes AHC $87, 8&8, 89, 389, fram Ptilotis

leucets, Sydney and Hawkesbury District, N.S.W.,

collectors T, H, Johnsion & J. B. Cleland, 1909,

Note; Although designated cotype by T. H. Johnston

some of the material in the AHC was collected from

Meliornis sericea and Meliornis novaehellandiae as

well as Prilatis leucotiy,

Choanotaenia taylori Johaston, 19124

Mem, Old. Mus. 1: 213.

Coiype AHC $90, from Malurus cyanoclamys, near

Adelaide, §.A., collector J. B. Cleland, 1910.

Cittatdenia bancrofti Johnston, 1912¢

Prac, R, Soe, Qld. 24: 68-9.

Now Progamotaenia hancrofti (Johnston,

Nybelin, 1917 after Beveridge, 1980.

Holotype (sectioned material) V2121, from Quay-

chogale frenata, Burnett R,, Qld,, collector T. L.

Bancroft, date unknown,

1912)

REC, S, AUST, MUS. 18 (21): 493-501.

Mi ay, 198d

Cittotuenia lagerchestis Lewis, 1914

Prac, Zool, Soc., Lond. 1914; 420,

Now Pregametaenia lagorchestis (Lewis, 1914)

Nybelin 1917 after Beveridge & Thompson, 1979.

Neotypes V1322, 1323, Barrow Is., W.A., collector

L. Hemsley,

Note: The original type material [rom Lagorchestes

conspicillatus collected on Hermit Is,, Montebello

Group, W.A. by P. D, Montague 21914 has been

reported as lost (Beveridge, 1976; Beveridge &

Thompson, 1979), Two similar yet distinct species

have been confused under the name P. lagorchestis

(Lewis, 1914). The original species is redescribed

by Beveridge & Thompson (1979) as above, The

specimens described by Beveridge in 1976 represent

a new species designated P. thylegale by Beveridge

& Thompson (1979),

Cittolaenia tachyglosyi Johnston, 1913

Rep. Aust, Inst. Trop. Med. 1911: 77-8.

Now Echidnoiaenia tachyglossi (Johnston, 1913)

Beveridge. 1980a. Syntypes V1303-14, including

sectioned material, from Tachyglossus aculeatus,

Townsville, Old., from the collection of A, Breinl,

date unknown.

Cyrelorchida omulanchristota Wedl, 1855

Rep. Aust, Inst. Trop. Med, 1911: 86-8. (Johnston,

1913,)

Now C. o. australiensis (Wedl, 1855 sensu Johnston,

1913), Bona, 1975,

Syntypes $312, from Platalea regia, Townsville, Old.,

from the collection of A. Breinl, date unknown.

Davalnea cacafuina Johnston, 1913

Rep. Aust. Inst. Trop, Med. 1911: 79-80.

Now Raillietina cacaiuina (Johnston, 1913) Yama-

suili, 1959,

Paratypes V1081-4, from Cacatua galerita, Towns-

ville, Old., from the collection of A. Breitil, date un-

known.

Davaineéa himantopodis Johnston, 1911¢

Proc. Linn, Soc, N.S.W, 36 (1)) 75-9,

Cotypes AHC $83, from Himantopus leucocephalus,

Tailem Bend, S.A., collector J. B. Cleland, 1910.

Dendrouterina ausiraliensis Baer & Bona, 1960

Boll. Instit. Mus. Zool. Univ, Torino 6 (4); 12-3.

Holotype ¥1676 including sectional material, from

“grey heron”, Swan Reach, §.A., collector T, H.

Johnston, 15/12/37.

Note: Bacr & Bona (1960) record this from Austra-

lian material labelled “Grey Heron” Ardea pacifica

Latham? At the time of the collection “Grey Heron”

was olten used in §.A. for Ardea nevaehallandiae

and in T. H, Johnston's dissection notes there is

recorded “Grey Heron” Nolephoyx novaehollandiae.

N, pavifica was referred to by Johnston as the Pacific

Heron which is a much rarer bird at Swan Reach,

HELMINTH TYPE-SPECIMENS IL

Dilepis bancrelti Johnston, 1912a

Mem. Qld. Mus. 1: 21-2.

Now Hemiparonia bancroftl

Schmidt, 1972.

Type and cotype ANC 891 including sectioned

tnaterial, feom Platycercus exinuus, collector T, H,

Johnston, date unknown,

(Johnston, 1912)

Diphylloborhriunt arctocephalinum Johnston, 1937

Proc, Linn. Sov. NSW, 62 (1): 9-13.

Holotype V13, from Arclocephalus forsteri, Pearson

Is., collector J, B, Cleland, 1923.

Note: Host collection data are identical with thal for

the host of Corynesoma australe. Johnson and Best

(1942) record that the type host was incorreetly

identified and should be regarded as Neophoca

cinerea.

Echinobethrium heronensis Williams, 1964

Parasitology 54: 737-40.

Paratype V1066, holotype V1060, from Taenuira

lyma, Heron Ts,, off Rockhampton, Qld,,. collector

J. Pearson, June 1956.

Lutetrarhynchus australis Prudhoe, 1969

BANZARE Rep. Ser. B 8 (9): 176-7.

Catypes V143, 144, 605, from Mustelus antarcticus,

Hobart, ‘Tas., collector T, H. Johnston, November

1930.

Hymenolepis aklei Beveridge & Barker, 1975

J. Helminthal. 49, 220-4.

Paratypes V68-70, holotypes V67, from Antechinus

steariii, Pawelltown, Vie., collected by the authors

17/8/74.

Hymenolepis bradleyi Beveridge & Barker, 1975

J, Helminthol, 49: 224-6,

Paratypes V72-4, holotype V71, from Anrechinus

sinartii, Powelltown, Vic., collected by the authors

23/3/74.

fivmencolepis ellisi Johnston & Clark, 1948a

Trans. R. Soe. 8. Aust. 72 (1): 81-2.

Holotype ¥2803, from Pelecanus conspicillatus, col-

Ivectors G. G., Fred & B, Jaensch and L. Ellis, date

unknown. Locality: Tailem Bend, 5.A.

Hymenolepis jaenschi Johnston. & Clark, 194828

Trans, R. Soe. §, Aust. 72 (1): 79-81,

Holotype V2804, from Pelecanus canspicillatus,

Tailem Bend, S.A., collectors G. G., Fred and B.

Jaensch and L. Ellis, date unknown.

Hymenolepis murrayensis Johnston & Clark, 1948a

Trans, R, Soc. S. Aust. 72 (1): 77-9,

Holotype V2802, from Pelecanus conspicillatus,

Tailem Bend, §,A,, collectors G, G,, Fred and B.

Jaensch and L, Ellis, date unknown,

Lapwingia ddelaidae Schmidt, 1972

J Parasitol 58 (6): 1085-7.

497

Paratypes V1105-7, holotype V1104, from Lebibyx

novaehollandiae, Townsville, Old., collector and date

unknown (probably from A, Breinl’s collection 1910

to 1913),

Lateriporus maw soit Prudhoe, 1969

BANZARE Rep, Ser, B 8 (9); 190-1.

Cotypes V362-71, from Chionis minor, Kerguelen

Is., collector T, H. Johnston, 17/11/29.

Noie; Several other collection. sites are listed, Also

recorded from Chionis minor crozetlensis, Possession

Island, Crozets, 3/11/29.

Megalonchas dubius Prudhoe, 1969

BANZARE Rep, Ser, B 8 (9): 180-2.

Cotypes V156-8, from Mustelus antarcticus, Hobart,

Tas., collector T, H. Johnston, November 1930,

Megalonchos musteli Prudhoe, 1969

BANZARE Rep. Ser. B 8 (9): 179-80.

Cotypes V152-5, from Mustelus antarcticus, Hobart,

Tas., collector T. H, Johnston, Navember 1930,

Montezia diaphana Zschokke, 1907

Zentralbl, Backterial, Parasitenk, 44: 261-64,

Now Progamotaenia diaphana (Zschokke, 1907)

Baer, 1927 after Beveridge, 1976.

Syntypes V2122, from Lasierhinus latifrons bile

ducts, probably Adelaide, $.A., collector E, Angus

Johnston, date unknown,

Nore: The material registered in SAM is thought by

Beveridge (1976 and 1980b) to be some of the ori-

ginal material described by Zschokke (1907). The

material was mounted by T, H. Johnston and

labelled, in his handwriting, “part of material exa-

mined by Zschokke and Angus Johnston’. There

were no collection data given by Zschokke in the

original description, The host name was recorded as

Phascolomys vombat now a synonym of Vomrbatus

ursinus, Beveridge (1980) believes this identification

to be in error with the type host being Lasiorhinus

latifrons.

Notylocestus majer Johnston & Muirhead, 1950

Rec. S. Aust. Mus, 9 (3): 340-4.

Holotype V854, paratype V855, from Tandanus tan-

danus, Murray Bridge, S.A,, collector L, M, Angel,

December 1947.

Notylocestus minor Johnston & Muirhead, 1950

Ree. §, Aust. Mus. 9 (3): 344-6,

Holotype V856, from Tandanus tandanus, Murray

Bridge, S.A., collector L. M. Angel, December 1947.

Oochoristica antechini Beveridge, 1977

J, Helminthol. 51: 32-5,

Holotype V818, from Antechinys macdonnellensis,

27 km west of Refrigerator Well, Tanami Rd., N,T.,

collector P. Woolley, July 1975.

498

Oochoristica eremophila Beveridge, 1977

J. Helminchel, 51; 35-8,

Holotype V828, paratypes V829-35, from Ante-

chinus rosamundae, Woodstack Stn., via Marble

Bar, W.A., collector P, Woolley, November, 1975.

Ophivtaenia hylae Johnston, 1912¢

Proc. R, Soc, Old. 24: 63-4.

Colypes AHC $86, trom Myla aurea, Sydney, NS.W.,

collector S$, J. Johnston, date unknown,

Ophietaenia longmani Johnston, 1916

Mem. Old. Mus, 5: 186-94,

Cotypes AHC S85, $523, from -lypidiotes ramsayi,

Yeulba, Western QOld., collector H, A, Longman,

date unknown.

Paramoneszia johnston’ Beveridge, 1976

Aust, J. Zool. Suppl, Ser, 44: 31-4,

Holotype V760, paratypes V761-5, from Vombatus

uwrsinus, Forester R., Tas., collector B, L. Munday,

28/4/72,

Parabathriocephalus johnstoni Prudhoe, 1969

BANZARE Rep. BB (9): 174-5,

Cotvpes V128-36, 570 (rom Coryphaenides whitseni,

330 fathoms, 66° 245, 59° 09°F, callector T. H.

Johnston, 7°1/30,

Parvifaenia clavipera Baer & Bona, 1960

Boll. Inst, Mus. Zool, Univ, Torine 6 (4); 3,

Holotype V1675, from Ardea novaehollaniiae,

Townsville, Old., collector not known, possibly A.

Breinl 1910-1913,

Note: This slide labelled Bancroftiella glandularis

(Fuhrmann), in T. A, Johnson's handwriting.

Parvitaéenia paracyclorchidu Baer & Bona, 1960

Boll, Inst, Mus, Univ. Torino @ (4); 1-3 & 44,

Holotype V1674, from Ardea novaelallanliac,

Wamberal. N.S.W., collector S. J. Johnston, date un-

known,

Note: This slide labelled Baneroftiella glaneilaris in

T. H. Johnston's handwriting.

Phaycolotaenia comani Beveridge, 1976

Angst, J, Zool, Suppl. Ser. 44; 35-7,

Holotype V766, paratypes V767-8, from Fombatus

ursinus, Tageerty. Vic., collector B. J, Coman,

19/7/71.

Palyonchobothriin scleropagts Blair, 1978

J. Helinathol, 52 (2); 147-53,

Holotype V1160, paratypes VI16I-5, trom Sclero-

pages Ieichardti, Wenlock R,, Cape York Pen., Qld.,

collector D, Blair, 8/7/75.

Progamotuenia cepyprviuy Beveridge, L976

Anst, J. Zool, Suppl. Ser. 44; 40-1.

REC. 8. AUST. MUS. 18 (21): 493-501.

Muy, (984

Holotype V772, fram Aepyprymnus rufscens, Bur-

nett R., Qld., collector T. L. Bancroft, February

1912,

Progainotaenia effigia Beveridge, 1976

Aust. J, Zoal, Suppl. Ser. 44; 45-7,

Holotype V773, paratypes V774-5, from Macropus

fuliginasus (bile duct), Pine Plains Stn,, via Patche-

wollock, Vic., collector I, Beveridge, 28/6/73.

Progametuenia — gynandroalinearis

Thompson, [979

J. Hlelminthal. 53; 156-7.

Holotype V1315, paratypes VI3L6-9, sectioned

material V1320. from Lagorchestes conspicillatus,

Barrow Is., W.A., collector L. Hemsley, 1976,

Beveridve &

Progamotaenia jolinsoni Beveridge, 1980b

Trany. R. Sac. 8, Aust. Wa (4)2 69-72.

Holotype VY1I918 including sections from Lagor-

chestes conspicillatus, paratype V1919, Mingela,

Old., collector T, Beveridge, 10/5/79.

Progamotaenia lagorchestiy Beveridge, 1976

Aust. J. Zool, Suppl. Ser, 44: 55-8.

Now Prevaniantnenia thylagale Beveridee (1976)

Beveridge & Thomson, 1979.

Holotype V783, paratypes V784, from Thylogale

billardicrii, Launceston, Tus., collector B. L,

Munday,

Progamolaenia macropodis Beveridge, 1976

Aust. I. Zool. Suppl. Ser. 44; 58-60,

Holotype V776, paratypes V777-8, from Macropus

eivantens, Yan Yean, Vic., collector J. H. Arundel,

12/9/72.

Progamotaenia ruficola Beveridge, 1978

J. Parasitol, 64 (2); 273-6,

Holotype V1207-L0, paratypes V1I212-13, AHC

8650, 8651, from Macrepus rufus, Menindec,

N.S.W., collector T. Beveridge, date unknown.

Progamolaenia xspearei Beveridge, 1980b

Trans. R. Soc. 8. Aust. 104 (4): 67-9.

Holotype V1920, from Thylogale stigmatica, Tolga,

QOld., collector I, Beveridge, 16/10/78.

Proteocephalus gallardi Johnston, 19 Lb

Ann. Old. Mus, 10: 1-8.

Now Acanthotaenia vallardi Johnston (1911)

1912¢, Cotypes AHC $80, 81, from Pseudechis por-

phiyriacus, Gostord, Richmond, Hunter R, and Syd-

ney, N.S.W., Gippsland, Vie., collectors L. Gallard,

Cc. T, Musson, F. H. Taylor, W. Hall, A. S, Souell

& T. H. Johnston, 1909-1910,

Nove; Johnston exhibited same specimens of A. gal-

lardi as Tehthyetaenia sp. ata meeting of The Royal

Suciety ol N.S.W. in 1910. However, a full descrip-

tion of the species was not published (as P. gallardi)

HELMINTH TYPE-SPECIMENS If

until T9171. In his paper Johnston did not designate a

type locality, but the slides registered as cotypes with

AHC as Jedihyotaenia gallardi have Narrara, N.S.W.

and Victoria (A. §. Le Souéff collector) as localities,

Tt is known that T. A. Johnston always listed the

type locality first.

Raillietina leipode Johnston & Clark, 1948b

Ree. §. Aust. Mus, 9 (10): 87-90,

Holotype V3044, paratypes AHC 1381, from

Leipoa eeellata, Vailem Bend, S.A,, collector L.

Ellis,

Rhinebothriun himanturi Williams, 1964

Parasitology 54: 740-1,

Holotype V1063, paratype V1064, from Himanrura

grunulata, Heron Is. off Rockhampton, Old., col-

lector J, Pearson, June 1956.

Sphaeruterina punctata Johnston, 1914

Proe. R, Soc. Qld. 26: 76-80,

Cotypes AHC §369, 879, from Pachycephala rufi-

ventyix, Caloundra, N.S.W., collector T. H. Johnston,

August 1914,

TVaenia bairdii Krettt. 1873

Trans. bent. Soc. NSW, 2: 224-7,

Now Microsomacanthus collaris Batsch, 1786, afler

Yumaguti, 1959, Cotypes AHC $77, from Anas

superciliosa, N.S.W,, collector G, Krefft, date

unknown.

Nele: The specimens collected by Krefft constituted

about half the then known cestodes (Johnston,

1912b), However, his descriptions and figures were

not detailed enough for species recognition, For-

tunately most of the type material was deposited in

The Australian Museum and available for re-exa-

mination by Johnston, Although some specimens

were poorly preserved Johnston was able to find bet-

ter preserved material which he identified as the

same species as the types in question. Some of

Kreflts material as redeseribed by Johnston has

been deposited in the AHC,

Taenia chlamydoderae Kreltt, 1873

Trans, Ent, Soe. N.S.W. 2: 224.

Now Paricterolaenia chlamydodere (Keefit, 1873)

Yamaguui, 1959, Cotype AHC $71, fram Chlamy-

dera maculara, collector G. Krefit, date unknown.

Tadenia coronata Kreltt, 1873

Trans, Ent. Soe, NSW. 2: 220-1.

Now Gyrocoelia aiysrraliensis (Krelfi, 1873) John-

ston, 1921h, Cotypes AHC $72, 8344-5, N.S.W,,

collector G. Kreflt, date unknown,

Taenia flacescens Keetit, 1873

Trans. Ent, Soc. NSW. 2: 219.

Now Dioarchis flavescens (Krett, 1873) Johnston,

1912b, Cotypes $73-5, S441, from Anas super-

cilioxa, S_S.W., collector G. Kreflt, date unknawn.

499

Taenia forsteri Krefit, 1873

Trans, Ent, Soc, NSW, 2: 218.

Now Tetrabothrius forsteri (Krettt, 1873) Johnston,

Cotype AHC 578, 1912h, from Delphinus fearsteri,

Port Jackson, N.S-W., collector and date unknown.

Taenia mastersii Kreitt

Trans. Ent. Soe. N.S.W, 2; 217-8.

Cotype AHC $70, from Macropus sp., Qld., col-

lector G. Masters, date unknown,

Note: Beveridge (1976) considered that this species

possibly belonged to Carestaurus,

Taenia parudoxa Kretit, 1873

Trans. Ent. Soe. N.S.W. 2: 217-8,

Cotypes AHC 876, $386, from Podiceps aitstralis,

N.S.W.,, collector G. Kreift, date unknown,

Note: T. paradoaxa is preoccupied by Rudolphi,

1801, Johnston was unable to establish the status of

the species which awaits redescription. Krettt named

the host Little Grebe, Podicepy auviralis, This is now

a synonym of the Crested Grebe. Obviously he

meant what is now Tachybatus novaehollandiae. The

Little Grebe would not be mistaken for the Crested

Grebe or vice versa.

Taenia rigosa Krettt, 1873

Trans, Ent. Sac. N.S.W. 2; 223-4.

Now Acoelus hedlyi (Kreflt, 1873) Johnston,

1912b, Neotype AHC §339-40, from Himantupuy

leucocephalus, Hunter River, N.S.W., collector G.

Krefft, date unknown,

Note: Johnston tound the original material so dam-

aged as to be useless, but was able to compare it

with fresh material from the same host in South Aus-

tralia by J. B, Cleland. The armatuye of the cirrus

sac of both lots of material assisted in establishing

their identity.

Tholophyllacus johnston’ Mackiewicz & Blair, 1980

Proc. Helm. Soc, Wash. 47 (2): 172-4.

Paratypes V1893-4, from Yundanus glencoensis,

Ross R, near Townsville, Old., collectors probably

the authors, date unknown.

Triplotaena fimbriata Beveridge, 1976

Aust. J, Zool, Suppl, Ser. 44; 69-71.

Holotype V800, paratype V801, from Macropiis

gigdnteus, Nebo, Qld., collector A, Glassop,

17/8/73.

Triplotaenia undosa Beveridge, 1976

Aust. J. Zool. Suppl. Ser. 44: 73-5,

Holotype V802, paratype V803, AHC 8962, trom

Wallabia hicoler, Dattmouth, Vic., collector 1.

Beveridge, 22/1/74,

Valipera parvitaeniunca Baer & Bona, 1960

Boll. Inst, Mus. Zool. Torino & (4): 9.

Now Baerbania p. parvitaeniunea (Bacr & Boni,

400

1960), 1975, Syntype AHC $308, from Egrerta

sacra, Masthead Is., Qld,, collector T, H,. Johnston,

1917.

Zasteropicola clelandi Johnston, 1912a

Mem, Qid. Mus. 1: 214-5.

Cotypes AHC S84, S464, fram Zosleraps ceeeru-

lescens, neighbourhood of Sydney, N.S.W., collectors

J. B. Cleland, J, O. Heimrich and T. H. Johnson,

dates unknown,

REC. 8. AUST. MUS. i8 (21): 493-501,

Atay, 1984

ACKNOWLEDGEMENTS

T would Jike to thank Mrs. P. M, Thomas (Maw-

son), Miss L. M. Angel and Dr. 5. J. Edmonds

without. whose adyice and assistance these lists of

helminth types could nol have been prepared. This

wark was funded by an Australian Biological Re-

sources Study grant to the South Australian Museum.

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three new species. Trans. R. Sac. S. Aust, 95 (2):

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HELMINTH TYPE-SPECIMENS II 501

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HELMINTH TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

HI. TREMATODA

BY LESLEY R. SMALES

Summary

Type-specimens of 13 species of Monogenea and 70 species of Digenea are catalogued. All are

from Australian or Antarctic hosts. Trematode species are listed alphabetically, according to the

original name of the genus, under either Monogenea or Digenea. Where a name change has

occurred the most recent name is included. The hosts are given under the name recorded by the

author.

HELMINTH TYPE-SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM

il. TREMATODA

LESLEY R. SMALES*

South Australian Muscum, Adelaide, South Australia 5000

ABSTRACT

SMALES, L. R.. 1983. Helminth type-specimens in the

South Australian Museum, UI. Trematoda, Ree, S,

Ausi, Muy. 1B (22): 503-511,

‘Type-specimens of 13 species of Monogenea and

70 species of Digenea are catalogued. All are from

Australian or Antarctic hosts, Trematode species are

listed alphabetically, according to the original name

of the genus, under either Monogenea or Digenea.

Where a name change has occurred the most recent

name is included. The hosts are given under the

name recorded by the author.

INTRODUCTION

This is the third and final paper of a series listing

the Helminth Types held by the South Australian

Museum and the Australian Helminthological Col-

lection, currently housed in the Museum. There is

also a substantial collection of larval material for

which the adult has not yet been described. All the

specimens described have been collected from Aus-

tralian or Antarctic hosts.

The species are listed in alphabetical order under

the genus to which they were first assigned. Where a

tame change has occurred the most recent authori-

tative name is, given. Host-species names are those

used in the original descriptions.

The earliest material registered is that assembled

by T. H. Johnston who was as actively interested in

the trematodes as he was in the other helminth

groups. The group of parasitologists he established

at Adelaide University, including L. M. Angel, now

Honorary Associate of the South Australian Mus-

cum and still working in the field, provided a sig-

nificant contribution to knowledge of the Australian

trematodes, Other Australian trematologists are now

depositing type material at the Museum,

Registration numbers starting with V_ indicate

material held in the South Australian Museum, those

starting with AHC, material in the Australian Hel-

minthological Collection. Unless otherwise stated

the name Johnston refers to T. H. Johnston. The

following abbreviations were used in the text:

BANZARE = British, Australia and New Zealand

Antarctic Research Expedition, 1929-31; AHC =

* Present address: Gippsland Institute of Advanced Educa-

tion, P,O. Box 42, Churchill, Victoria 3845,

Australian Helminthological Collection; SAM =

South Ausiralian Museum; N.T. — Norther Ter-

ritory; N.S-W. = New South Wales; Old. — Queens-

Jand; S.A, = South Australia; Tas. — Tasmania:

Vic, = Victoria; W.A, = Western Australia,

CLASS TREMATODA

SUB-CLASS MONOGENEA

Anoplodiscus australis Johnston, 1930

Aust, J. Exp, Biol. Med. Sci, 7; 108-12.

Holotype V8 from the fin of Sparus australis, Sydney

Harbour, N.S.W., collector §. J, Johnston, date un-

known.

Now Anoplocotyle ausiralis (Johnston, 1930) Pa-

lombi, 1943,

Calicotyle australis Johnston, 1934.

Aust. J, Exp, Biol, Med, Sei. 12: 25-8,

Holotype V9 probably from the gills of Trygondr-

rhina fasciata, Glenelg, S.A., collector T. H. John-

ston, date unknown,

Diplasicotyle johnstoni Sandars, 1944

Trans, R, Soc. §. Aust, 68 (1): 77-81.

Types V850-1 from gills of Agonastomus forsteri,

Mandurah, Bunbury, W.A., collector probably the

author, 1943,

Note: Sandars did not specify a type Jocality for each

species, but some of the type slides have a locality

written in her handwriting.

Gonoplasius carangis Sandars, 1944

Trans. R. Soc, S. Aust. 68 (1): 75-7.

Holotype V848 from the gills of Caranx georgianus,

North Beach, Rockingham, W.A., collector probably

the author, 11/6/43,

Microcotyle agonostomi Sandars, 1945

JR. Soe, W. Aust. 29: 108-12.

Holotype V846 from the gills of Agenostemus for-

steri, Swan R., Mandurah, Bunbury, Denmark, Al-

bany, W.A., collector probably the author, 1942.

Microeotyle arripis Sandars, 1945

J. R. Soe. W, Aust, 29: 114-8,

Holotype V845 from the gills of 4rripis trutta, North

Beach, Swan River, Mandurah, Bunbury, Busselton,

Albany, Woodman’s Point, Scarborough, Whitfords

Beach, W.A., collector probahly the author,

18/6/42,

504 REC, 8. AUST. MUS. 18 (22); 503-511,

Microcotyle gerres Sandars, 1944

Trans. R. Soc. §. Aust. 68 (1): 67-8,

Holotype V843 from gills of Gerrey ovatus, Man-

durah, W.A,., collector probably the author,

14/4/43,

Microcatyle helotes Sandars, 1944

Trans, R. Soc. S, Aust. 68 (1): 72-4.

Coatypes V841 from the gills of Helotes sexlineatus,

Safety Bay, W.A., collector probably the author,

20/5/43.

Microecetyle odacis Sandars, 1945

J. R. Soc. W. Aust. 29: 123-6.

Holotype V842 from the gills of Odar semifasciatus,

Albany, W.A., collector probably the author, April

1941,

Micrecetyle parasillaginae Sandars, 1945

J. R. Sac. W. Aust. 29; 112-4.

Holotype V844 from the gills of Sillaginedes pune-

talus, Albany, W.A., collector probably the author,

14/5/42.

Microcetyle pentapodi Sandars, 1944

Trans, R, Sac, 8. Aust, 68 (1); 69-70,

Holotype V839 from the gills of Pentapodus milii,

Rockingham, W.A., collector probably the author,

26/4/43.

Microcetyle scorpis Sandars, 1944

Trans. R. Soc. S. Aust. 68 (1): 71-2.

Holotype V840 from the gills of Scorpis aequipinnis,

Safety Bay, W,A., collector probably the author,

14/1/43,

Microcotyle temnodontis Sandars, 1945

JR. Soc, W, Aust, 29; 119-23.

Cotypes V&47 from the gills of Temonodon saliator,

Bunbury, W.A., collector probably the author,

2/4/42.

SUB-CLASS DIGENEA

Apatemon (Apaiemon) vitelliresiduus Dubois &

Angel, 1972

Trans. R. Soc. Aust. 96 (4): 199-201,

Holotype V1068 from the intestine of Biziira lobata,

Purnong and Caloot, Murray R., 5S.A., collector

P. M. Mawson, 20/6/58,

Paratypes V1069, three specimens on the same slide

as the holotype.

Bancrofirema neocerotodi Angel, 1966

J, Parasitol, 52 (6): 1058-61.

Holotype V955 from the digestive tract of Neocero-

todus forsteri, Qld., collector T. L. Bancroft,

January 1911.

Paratypes. V956-8, juveniles, one sectioned,

May, 1984

Brachylecithum dacelonis Angel & Pearson, 1977

Trans, R. Soe. S. Aust. 101 (5): 122,

Holotype V89 from the liver of Dacelo novae-

euineae, Bridgewater, $.A., collector P. M. Mawson,

July 1973.

Paratypes V90, 91; AHC S15.

Brachylecithum hydromyos Angel & Pearson, 1977

Trans. R. Soc. 8S. Aust. 101 (5); 121,

Holotype V82 from the pancreatic ducts of

Hydromys chrysogaster, Lily Ck., Cairns, Old,, col-

lector C, M, Weaver, 14/2/75.

Paratypes V83, 84; AHC S14,

Brachylectthum insulare Angel & Pearson, 1977

Trans. R. Soe. 8. Aust. 101 (5): 119-21,

Holotype V76 from gall bladder and bile ducts of

Rattus fuscipes, Pearson Is., $.A., collector P. E,

Hornsby, date unknown,

Paratypes V77, 78, 79; AHC 82,

Brachylecithum latius Angel & Pearson, 1977

Trans. R, Soc, §. Aust. 101 (5): 123-4.

Holotype V102 from the gall bladder of Cracticus

tarquatus, Cowell, S.A., collector P. M. Mawson,

27/5/65,

Paratypes V103-5; AHC 824-6.

Brachylecithum pedargi Angel & Pearson, 1977

Trans. R. Soc. 8S. Aust. WA (5): 212-3,

Holotype V92 from the bile ducts of Podargus

strigoides, Moggill, Qld., collector J, C. Pearson,

6/11/62,

Paratypes V103-5; AHC $20,

Burnellus trichofurcatus Angel, 1971

Parasitology 62; 375-84.

Holotype V977 from the intestine of Tandanus tan-

danus, Murray R., S.A., collector T. H, Johnston,

L. M, Angel, dates unknown,

Paratypes V978-987, other paratype material in

AHC,

Cercariaze V930, V2772, Sporocysts Y931 as used

in the original descriptions of the juvenile stages of

the life cycle by Johnston & Angel (1940a).

Cardiacephaloides ovicerpus Dubois & Angel, 1972

Trans. R. Soe. S. Aust, 96 (4); 204.

Holotype V1070 from the intestine of an unknown

host, no collection data, from SAM.

Paratypes V1071 including sectioned material.

Catatropiy gallinule Johnston, 1928

Rec. S. Aust. Mus, 4 (1); 135-8,

Holotype V5 from the caecum of Gallinula tene-

brosa, Adelaide, S.A., collector probably T. H,

Johnston, 1926.

Cetiotrrema crassum Manter, 1970

Trans, R. Soc. 8. Aust. 94; 147-50,

HELMINTH TYPE-SPECIMENS III

Holotype V1004 from the ureter of Thunnys shynnis

muceoyil, East Bass Strait and Kangaroo Is., $.A,,

collected by CSTRO, 1939,

Ceelomotrema antichinomes Angel, 1970

Aust. J. Zool, 18: 119-24,

Holoiype V974, from the body cavity of Ante-

chinomys spenceri, Sandringham Stn. near Bedourie,

Old., collectors P. Woolley & M. Stanley, caught

18/6/68, died 22/10/68.

Paratypes V975, 976 including sectioned material

and AHC.

Cetylurus (Coetylurus) magneacetabulus Dubois &

Angel, 1972

Trans. R, Soc. §, Aust, 96 (4); 204-5.

Holotype V1072 from the lower intestine of Cygnus

atraius, Tatlem Bend, S.A., collector T. H, John-

ston, 25/10/45,

Paratypes V1072 (six on same slide as holotype),

V1073.

Cyclocoelum jaenschi Johnston & Simpson, 1940a

Trans, R, Soc, 8. Aust, 64 (2); 273-8.

Holotype V949 from the abdominal air sacs of Podi-

ceps novaelhollandiae, Tailem Bend, §.A., collectors

G. & F. Jaensch, August 1937,

Cercariaeum jaenschi Cercariae V948 as used in the

original descriptions of the juvenile stages of the

life cycle,

Note; Although a type-host is not designated in the

original description the type-slide las host data in

T_H. Johnston's handwriting.

Cylindratrema eygni Angel, 1973

Int. J. Parasitol. 3; 853-7,

Holotype V1119 from the caecum of Cygnus atratus,

Tailem Bend, S.A., collector T. H. Johnston,

20/8/51.

Diplostomum murrayense Johnston & Angel, 1941a

Trans, R, Soc, 8. Aust, 65 (1): 140-4,

Holotype V952 from the digestive tract of Childonias

leucopareia, Tailem Bend, S.A., collector G, & F.

Jaensch & L, Ellis, 7/2/40.

Paratypes V944.

Cercaria murrayensis Johnston & Cleland, 1938

Cercariae V941-2, V2768, Diplostomulum V943,

as used in the original descriptions of the juvenile

stages of the life cycle.

Diplostomuni (Diplostemum) paryulum Duhois &

Angel, 1972

Trans, R, Soc. 8, Aust. 96 (4); 206-8,

Holotype V1LO74 from the intestine of Hydroprogne

easpia, Tailem Bend, $.A., collector T. H. John-

ston, December 1939.

Paratypes V1074 (three on same slide as holotype),

VILO75,

S05

Dolichepera macalpini Nicoll, 1914

Parasitology & (4): 333-50,

From Netechis scuta(us and Denisonia superba, col-

lected in N.S.W., S.A., Vic. and Tas., now Dolicho-

peroidey macalpini (Nicoll, 1914) Johnston & Angel

(1940b).

Cereariae V924-5, metacercariae V926, sporocysts

V923, and sections of infected Ameria pyrainidata

V927 as used in the original descriptions of the

juvenile stages of the life cycle.

Derogenes johnstoni Prudhoe & Bray, 1973

BANZARE Rep, Ser. B 8 (10); 221-2,

Catypes V563 from Artedidrace shackeltoni (site in

host not known), 65°48'S, 53° 16, collector T. H.

Johnston, 24/1/30,

Note; Prudhoe & Bray (1973) did not specify type

host and type locality or designate a holotype for

any of the trematodes collected by T. H. Johnston

during BANZARE.

Echinochasmus pelecani Johnston & Simpson, 1944b

Trans R, Soc, 8. Aust, 6B (1): 113-9.

Syntypes V1059 trom the small intestine af Pele-

canus conspicillatus, Tailem Bend, S.A., collectors

G, & B. Jaensch & L, Ellis on several occasions

between 1938-44,

Cercariae V939, V2806, cysts and redia V2805,

metacercariae V940, infected Gamhusia ajfinus

V2807 as used in the original description of the

juvenile stages of the life cycle,

Note; Johnston & Simpson (1944) considered that

a more detailed description of . mordax Looss,

1899 would show that &. pelecani was a synonym

of that species. However Yamaguti (1971) still tists

E. pelecani separately with the note that i is

probably a synonym of &. mordax.

Echinaparyphium ellist Johnston & Angel, 1949

Ree. §, Aust. Mus. 9 (2): 247-54.

Type adult V902 from the duodenum of Chenepts

atraius, Tailem Bend, S.A., collectors Messrs

Jaensch & Ellis, April 1947,

Cercaria ellisi Johnston & Simpson, 1944a

Cercariae V903, 904, metacercariae V9OS, redia and

eercariae V906 as used in the original descriptions

of the juvenile stages of the life cycle.

Echineparyphium hydromyos Angel, 1967

Parasitology 57: 19-30.

Holotype V57 from the intestine of Alvdronrys

chrysegaster, Vailem Bend, S.A., collector L. M,

Angcl, December 1959,

Paratypes V58.

Cereariae fram Planorbis isingi V60, cysts from tad-

poles and Lenameria sp. V59 as used in the original

descriptions of the juvenile stages of the life cycle.

506

Echinosioma australe Johnston, 1928

Rec. S. Aust. Mus, 4 (1): 138-40,

Holotype V6 from the caecum of Gallinula tene-

brosa, Adelaide, $.A., probably collected by T. H.

Johnston, 1926.

Echinestama hancrefti Johnston, 1928

Rec. S. Aust. Mus, 4 (1) 140-2,

Holotype V7 from the intestine of Gallinula tene-

brosa, Burnett R., Eidsvold, Old,, collector T, L.

Bancroft, 29/9/19,

Folivtvema jecoris Blair, 1981

Aust. J. Zool, Suppl. Ser. 81: 31-2.

Holotype V1963 from the gallbladder and bile ducts

of Dugong dugon, Daru, Gulf of Papua, collectors

‘Dugong Project’ Wildlife Diy. Dept. Lands and

Environment, Papua New Guinea, 11/7/78.

Paratypes V1964,

Faredifix clavata Blair, 1981

Aust. J. Zool, Suppl. Ser, 81+ 41-3.

Holotype V1973 from large abscesses in the wall of

the ileum of Dugong dugen, Mornington Is,, Old,,

collected by aboriginals for food, 11/11/76.

Paratypes V1974-6 including sectioned material.

Galactosxemum angelae Pearson, 1973

Phil, Trans, R. Soc. Lond. B 266 (879); 361,

From the smal! intestine of Hydropragne caspia,

Port R., S.A., collector unknown, September 1928,

donated by L. M. Angel.

Paratypes V 1029.

Galactesomum bearupi Pearson, 1973

Phil. Trans. R. Soc, Lond, B 266 (879): 368.

From the small intestine of Aydroprogne caypia,

Townsville, Qld., collector A, J, Bearup, 20/8/68.

Paratypes V1030.

Galactosomum renincela Pearson, 1973

Phil, Trans, R. Soe. Lond. B 266 (879): 409.

From the renal ureter of Puffinus pacificus, Heron

Is,, Old, collector probably the author, 6/3/68.

Paratypes VLO31.

Galactosomum sinuilactis Pearson, 1973

Phil, Trans, R, Soc, Lond, B 266 (879): 415,

From the bursa of Phalacrocerax varius, Moreton

Bay, Qld., collectors R. Groom & R,. Greenhill,

25/7/69,

Paratypes V 1032.

Haerator caperatus Blair, 1981

Aust. J, Zool. Suppl. Ser. 81: 44-7.

Holotype V1977 from the ileum of Dugong dugon,

Mornington Is,, Qld., collected for food by abori-

ginals, 2/8/78.

Paratypes V1978-1 including sectioned material.

Haplorchis parapumilio Pearson & Ow-Yang, 1982

S-E Asian J. Trop. Med. Pub, Health 13: 37.

REC. S. AUST. MUS. 18 (22): 503-511

Muy, 1984

From Phalacrocerax pygmaeus, trom the small intes-

tine, Pulan Rambut, Jakarta Bay, Java, collector D,

Supriadi, 23/6/77.

Paratype V2752.

Haplorchis parayanissimus Pearson & Ow-Yang,

1982

S-E Asian J. Trop. Med: Pub, Health 13: 50.

From Nycticorax calendonicus, from the lower small

intestiné, Brisbane Botanic Gdns. Old.. collector

J, C. Pearson, 8/1/62.

Paratype V2752,

Harveytrema bisculatum Kruse, 1979a

J, Parasitol. 63 (6): 918-20,

Holotype V1677 from the intestine of Achoerodus

gouldi, Pearson Ts., S.A., collector P. M. Mawson,

February 1960.

Paratype V1678 (sectioned), AHC $290,

Helicametra scorpaenae Prudhoe & Bray, 1973

BANZARE Ren. Ser. B 8 (10): 213,

Cotypes V458-9, from Scorpaena truenta, 42° 40'S,

148° 27° 30”F off Maria 1, Tas,, collector T. H.

Johnston, 23/3/31.

Hemistomum infermedium §. J. Johnston, 1904

Proc, Linn, Soe, NSW, 1904; 109-110.

now 4 patemon intermedium (S, J, Johnston, 1904)

T. H, Johnston & Angel, 1951b,

Substitute type V8&70 from the duodenum of Chenv-

pis atrata, Duckmaloi R,, N,S,W,, collector T, H.

Johnston,

Cercaria lessont V872-3, 875 metacercariae V874,

876, cysts V877 as used in the original description

of the juvenile stages of the life cycle by Johnston

& Beckwith, 1947a,

Nate; The slide deposited in SAM is designated

“substitute type” in T. El. Johnston's handwriting

and is from material collected at Tailem Bend, S.A.

Attempts by Johnston & Angel, 1951b, to trace

the Original material, were unsuccessful.

Heterophyes bitequatus Pearson & Pearson, 1983

Sarawak Mus, J. 29: 171-189,

From Sternd bergii, site in host not given, Buntal,

Sarawak, collector Guan anak Sureng, 15/12/76.

Paratypes V2607.

Heterophiyes chini Pearson & Pearson, 1983

Sarawak Mus. J, 29: 171-189,

From Sterna bergti, site in host nat given, Buntal,

Sarawak, collector Guan anak Sureng, 15/12/76.

Paratypes V26(05-06,

lyoparerchis tandani Johnston, 1927

Trans. R. Soc. S. Aust. S41: 129-33,

Lectotype V3089 from the gas bladder of Tandanus

randanus, Qld., collectors ‘T, L, Bancroft & M, J.

Mackerras, date unknown,

HELMINTH TYPE-SPECIMENS IL

Syntypes V3090.

Note: This material was found in T. H. Johoston’s

slide collection. The labels indicated that it must

have been the original material, L, M. Angel desig-

nated the slides lectotype and syntypes.

Lahicola elongata Blair, 1979

Ann. Parasitol. Hum. Camp. 54 (5): 521-6,

Holotype V1679 from the upper Jip of Duong

dugen, accidentally drowned in nets used by shark-

meshing contractors and given to the dugong

research group, James Cook University, Townsville,

Old,

Paratypes V1680-4 including sectioned material.

Lankatrema minutunt Blair, 1981

Aust. J, Zool. Suppl. Ser. 81: 23-4,

Holotype V1954 from cardiac glands of the stomach

of Dugeng dugon collected for food by aboriginals,

Mornington Is., Gulf of Carpentaria, 4/8/75,

Paratypes V1946-8 including sectioned material.

Lankatrema micrecotyle Blair, 1981

Aust, J, Zool, Suppl. Ser. 81; 24-7.

Holotype V1949 from the wall of the ileum of

Dugong dugen, collected for foad by aboriginals,

Mornington Is., Guif of Carpentaria, 2/8/78.

Paratypes V1950-3 including sectioned material.

Lankalrema macrecotyle Blair, 1981

Aust, J. Zoal. Suppl. Ser. 81; 27-8,

Holotype V1954 from the wall of the ileum of

Dugong dugon, collected for food by abotriginals,

Mornington Is., Gulf of Carpentaria, 2/8/78.

Paratypes V1955-6 including sectioned material.

Lankatremoides gardneri Blair, 1981

Aust, J, Zool, Suppl, Ser. 81; 29-31,

Holotype V1957 from pancreatic ducis of Dugong

dugon, collected for food by Torres Strait Islanders,

Thursday Is., Old., 3/11/76.

Paratypes V1958-62 including sectioned material.

Lecithaster australis Prudhoe & Bray, 1973

BANZARE Rep. Ser. B 8 (10): 218.

Cotypes V503-21, site in host not given, from Noto-

thenia spp, and three other hosts, near head of

Bras Bossitre, Kerguelen and cight other localities,

callector T. H. Johnston, 20/21/29,

Syntypes V735.

Lepecreadium angelae Kruse, 1981)

Prac. Helm. Sac. Wash. 48: 195-97,

From Scerpiy geargianus, Pt Willunga, S.A., col-

lector probably ‘Tl, H, Johnston, February, 1934.

Paratypes V2753-64,

Leuehochloridium australiense Johnston & Simpson,

1940b

Trans. R, Soc. 8. Aust. 64 (1): 119-24,

Holotype V960 from the cloaca of Pomutostomues

superciliosus, Elwomple, near Tailem Bend, $.A.,

S07

collector F, Jaensch, August 1938.

Cereariae V959 as used in the original description

of the juvenile stages by Johnston & Cleland, 1938.

CLutctirema ailuroedi Angel & Pearson, 1977

Trany. R. Soc, 8. Ausi, WA (5); 127-9.

Holotype V108 from bile duct and mouth of gall

bladder of Ailuroedis crasyirostris, Mt Glorious,

Old., collector K, E, Webber, 29/10/62.

Paratypes V109-11; AHC $28,

Macropoirema pertinax Blair, Beveridge & Speare,

1979

Ann, Parasitol, 54 (6); 585-92,

Holotype V1831 (serial sections) from the cuecum

of Macropus agilis, Yabulu, Qld., collector 1, Beve-

ridge, 20/8/78.

Paratypes V1832-6 including sectioned material,

Mawsonotrema éudyptulue Angel, 1973

Int, J. Parasitol. 3: 857-9,

Holotype V1120 from ducts of liver of Eudyplula

minor, Brighton Beach, S.A., collector, from the

Instilute of Medical and Veterinary Science, S.A.,

11/7/72.

Paratypes V1121 including sectioned material; AHC

$33-5,

Mesoxtephanus neophocae Dubois & Angel, 1976

Bull. Soe, Neuchat. Sci, Nat, 99; 29-32,

Holotype V75 from intestine of Neophaca cinerea,

Gulf St Vincent, 8.A., collector G. Dubois, 29/1/75.

Paratypes, four specimens on same slide as holo-

type, AHC §31, $32,

Monostephanostamun manteri Kruse, 1979b

J. Parasitel, 65 (@): 921-3.

Holotype V1911 from intestine of Arripis gear-

giqnus, Port Willunga, S.A, (February, 1934), Gulf

St Vineent, S.A, (April, 1964), collectors T. Hi,

Johnston, and from the Adelaide Fish Market,

Paratype V1912; AHC $306.

Neodiplostomum intermedinm Pearson, 1959

Parasitology 49 (1 & 2); 111-9.

From small intestine of Rarrus assimilis, edge of ram

forest, Mt Glorious, Qld., collector probably the

author, 1956-7.

Paratypes V836-8.

Neodiplostomum (Neodiplostomum) lanceolatum

Dubois & Angel, 1972

Trans. R, Soc, §, Aust. 96 (4): 209-10,

Holotype V1076 from intestine of Ninex noevac-

seclandiae, Adelaide, S.A., collector P. M. Mawson,

April 1959,

Paratypes Y1077, four specimens on same slide “as

holotype.

Nevdiplostomunr (Trilaborchidiplosromum) <lialolt

Dubois & Angel, 1972

S08

Trans. R. Soc. §. Aust, 96 (4); 210-2,

Holotype V1078, site in host unknown, from Sar-

eophiluy Marrisii, Tas., collector B. Munday, October

1969,

Neolepidapedon antarclicus Prudhoe & Bray, 1973

BANZARE Rep. Ser. B 8 (10): 204-5.

Cotypes V406-9 [rom pyloric caeca of Corypliae-

nodes whiiwoni, 66° 218, 58° SO'R., collector T, H.

Johnston, 7/1/30,

Neolepidapedon dubius Prudhoe & Bray, 1973

BANZARE Rep. Ser. B 8 (10): 205-6,

Cotypes V410-12, V732 fram pyloric caeca of Cory-

phaenoides whitsoni, 66" 218, 58° SOE, collector

T, H. Johnston, 7/1/30.

Nevlepidapedon helicoleni Prudhoe & Bray, 1973

BANZARE Rep, Ser. B 8 (10): 208.

Cotypes V415-24, site in host unknown, from Heli-

colenis percoides and Scorpaena cruenta, 42° 40'S,

148° 27° 30°F, off Maria Is,, Tus., collector T. H.

Johnston, 23/3/31,

Neolepidapedon trematoni Prudhoe & Bray, 1973

BANZARE Rep, Ser. B 8 (101): 206,

Cotypes V413-7, site in host unknown, collected

during AAE 1911, hosts and localities Trematamus

hansonji 65° 42°8, 92° 10'E. Tremaronis scott

65° 20'S, 95° 27'E, Austrolycichthys brachycephalus

65° 6S, 95" 27'F.

Opisthotrema australe Blair, 1981

Aust, J. Zool. Suppl. Ser. BL: 9-10,

Holotype V1934 from custachian tubes and nnddle

car of Dugong dugen, collected for food by Torres

Strait Islanders, Thursday Is., Old., 3/11/76.

Paralypes V 1735-9 including sectioned niaterial,

Opecoelus scorpaenidicola Prudhoe & Bray, 1973

BANZARI; Rep, Ser, B. 8 (10): 209-10.

Cotypes V433-49, site in host unknown, from /eli-

colenus percoides, Scurpaena cruenta, and Lepide-

perca tasmanica, 42° 40'S, 148" 27' 30°F, off Maria

Is., Tas., collector T, H. Johnston, 23/3/31.

Pancreatrema meéliphagae Angel & Pearson, 1977

Trans, R. Soe. 8. Anse. LOL (5): 129-30.

Holotype V112 probably from bile duct of Meli-

phaga ernata, Blanchetown, S.A., collector P, M.

Mawson, April 1965.

Paraneocreadinm auxtraliense Kruse, 1978

J. Parasitol. 64 (3): 398-400,

Holotype V1496 from intestine of Psilocraninm

nigricans, Aldinga Reef, S.A., collector P. M. Maw-

son, 1962,

Paratypes V1497-L50L including sectioned material.

Parorchis acanthus yar. australis Angel, 1954

Trans, RB, Soe. 8. Aust. 77. 164-73.

REC. S. AUST, MUS. 18 (22): 503-511.

Muy, 19d

Holotype V988 from intestine of Larus movae-

hellandiae, Glenelg, S.A., collector T, H, Johnston,

1939,

Paratypes AFC S189,

Cercariag V991-2, metacercariae V994, redia V900,

cysts V993, and immature adult V898 as used for

the original description of the life cycle.

Pharyngosiomuides dasyuri Dubois & Angel, 1972

Trans. R. Soe, §. Aust, 96 (4); 213-4,

Holotype V1079 from small intestine of Dasyurus

Viverrinus, Leena Estate, Vas., from the collection of

J, C. Pearson, 9/11/66,

Paratypes VIORO,

Petdviger ausiralis Jahnston & Angel, [941b

Trans, R. Soe, S, Ans. 65 (2); 285-91,

Holotype V917, site probably intestine of Podieeps

poliocephalus, Vailem Bend, §.A., collectors G, & P,

Jaensch & L. Ellis, May 1941,

Cercuria giganiura V91V8-20, V2970, cysts V921

and metacereariae V922 as used for the original

description of the life cycle by Johnston & Angel.

Note: a type host was not designated in the original

description but on the slide deposited in SAM

Podiceps poliacephaluy is written as the hostin T, A,

Johuston’s own handwriting.

Plagierchis jaensehi Johnston & Angel, 1951a

Tray. R. Soc. 8. Aus, 74 (1): 49-58.

Holotype V&58 from small intestine of /ydraniyy

chrysogaster, Tailem Bend, S.A., collectors G. G.

and Bryce Jaensch between May 1938 and Deeem-

ber 1949,

Cercariage V859-60, V3794, sporocysts V&GI, cysts

V2795, V863, metacerearlae V862. Infected Lim-

nace lesson’ V2796, as used in the original deserip-

tions of the life eyele,

Postmonorchis variahilis Prudhoe & Bray, 1973

BANZARE Rep. Ser. B 8 (10): 202-3.

Cotypes V387-99, site in host unknown, from Noelo-

thenia cyanobrancha, Port Jeanne dArc, Kerguelen,

10/2/30 and Green Harbour, Kerguelen; Noto-

thenia acula, near head of Bras Bossiére, Kerguelen,

20/11/29; Notothenia rossi 49° 28'S, 70° 33'E off

entrance to Royal Sound, Kerguelen, 21/3/30, Har-

pagifer bispinus, Port Jeanne dAre, 16/2/30, col-

lector 'T. Hy Johnston.

Preboltiremea clelandi Johnston, 1934

Aust, J. Exp. Biol. Med, Sei, 12: 29-31,

Holotype V12 from hody cavity of Mustelus antare-

vicus, Encounter Bay, S.A., collector J. B. Cleland,

January, 1922.

Preéboliivema retundatuni Johnston, 1934

Aust. J. kexp. Biol. Med, Sci. 12: 28-9.

HELMINTH TYPE-SPECIMENS UI

Holotype VIO trom abdominal cavity of Trygen-

orrhina fasciata, Kangaroo Is,, S.A,, collector C. J,

Hackett, 1933.

Probolitrema simile Johnston, 1934

Aust. J. kixp. Biol. Mee. Sei. 12: 31-2,

Holotype V11 from body cavity of Mustelus antarc-

licus, Encounter Bay, $.A., collector J. B, Cleland,

January 1922,

Projolransversoirema vieert Angel, 1969

Parasitology 59: 719-24,

Holotype V970, under scales of Aldrichetia forsteri,

North Arm, Port R., S.A., collector B. Steer, July

1968,

Puratypes V971-3,

Pseudopecoelus pyriformis Prudhac & Bray, 1973

BANZARKE Rep. Ser. B 8 (10): 210-2,

Cotypes V450-3, site in host unknown, from Cal-

lanthias allporti, 42° 40'S, 148" 27° 30°R, off Maria

Is., ‘Tas,, collector 'T H, Johnston, 23/3/31.

Prosthogonimus vitellatus Nicoll, 1914

Ini. J. Parasitol. 3 p, 859—redescription by Angel,

1973,

Type host Chibia braceteata. Since the holotype,

located at the School of Public Health and ‘Tropical

Medicine Sydney, was found to be imperfect six

specimens V1122 from Gymmnerhina tihicen leuco-

nata, Tailem Bend, $.A., have been deposited in

SAM, collector T, H. Johnston, 1/6/40,

Skrjabinesonuim mawsoni Angel & Pearson, 1977

Trans, R, Soe, S. Aust, 11 (5): 124-6,

Holotype (lateral mount) V96 from liver of Mano-

rina flavigula, Port Augusta, S,A,, collector SAM,

May 1965.

Paratypes V96 (dorsal mount) on same slide as

holotype and V97; AHC S18.

Skrjabinosanum pomatastomi Angel & Pearson,

1977

Trans. R. Soc. 8S. Aust, WO1 (5); 126-7,

Holotype V99 from liver of Pomatostomus super-

ciliosus, The Bunkers, Flinders Ranges, 8.A., col-

lector SAM, 22/7/65.

Paratypes V99, six specimens on same slide as holo-

type and VIOO, VIOLI.

Stictodora diplacantha Johnston, 1942

Trans, R, Soe, S. Aust, 66 (2): 239-40,

Holotype V852, site in host not given, from Phala-

crocorax varius, Port Gawler, S.A., collector pro-

bably T, H. Johnston, date unknown,

Stenakron kerguelense Prudhoe & Bray, 1973

BANZAREL Rep. Ser. B 8 (10): 214-6,

Colypes V465-83, V733, site in host not given, from

Notothenia coriiceps Heard Is., 28/9/29; Notoe

thenia evanebrancha from various stations at Ker-

50Y

guelen between 16/11/29 and 15/2/30; Champso-

cephalus gunnari 49° 28'S, 70° 33’E off entrance

to Royal Sound, Kerguelen, 2/3/30; Zanchlorhya-

chus spinifer 54° 42’ 30°S, 158° 54° 30°, off

Lustitania Bay, Macquarie [s,, collector T. H, John-

ston, 4/12/30,

Tandanicola hancrafii T. HW. Johnston, 1927

Trans, R. See. 8. Aust. 51: 133-6.

Cotypes V3088, from the gas bladder of Tandanus

tendanis, Burnett R., Ridsvold, Old., collectors T, L.

Bancroft and M, J, Mackerras, date unknown.

CERCARIA

Cercaria avetabulopapillasa Angel & Manter, 1972

Ann, dust, Biol, Univ. Nal, Auion Mexico Al Ser.

Zool, 1; 5-9,

Cercariaec and melacercariac, V1IO07-12; redia

V1008, redia and cereariae VIOL3: infected Plu-

topsis tetrica with redia und cereariae VIO14s cysts

VIO1S,

Cercaria amerianne Johnston & Beckwith, 1947a

Rec, 8. Aust, Mus, 8 (4): 575-782.

Cercaria, V880, V¥2799; diplostomula and sporo-

cysts V&881, V882, V2880, infected Limnodynasres

sp. V2801,

Cerearia anevli Johnston & Beckwith, 1947b

Trans. R. Soe. §. Aust. 71 (2): 324-8.

Cercaria 2788, V884, sporocysts V883, infected

Ameria pyramidata V2787:

Cercaria angelae Johnston & Simpson, 19444

Trans. R. Soe. S. Aust, 68 (1): 130-2,

Cercariae V888, V2789) sporocyst V887; tetracotyle

V8S89-91. from Amerianna pyramidata and A.

fenuirosira,

Cercaria hbeckwithae Johnston & Angel, 19496

Trans, R. Soe, 8. Aust, 73 (1): 22-7.

Cercariae V2778, V895, V896; infected Planorbis

ising? V2779-80.

Cercaria clelandi Johnston & Angel, 1939

Trans, R- Soe. S. Aust. 63 (2); 200-3.

Cercaria V950, V2766; redia V951: from Planerbis

iwingi,

Cercarla gigantura var, grandior Johnston & Simp-

son, 19444

Trans. R. Soc. §. Aust. 68 (1): 128-30.

Cercuria V893, V¥2790; redia V892-3; metucerearia

V894; infected Amertanna pyramiddla V2792,

Cercaria haswelli Dolfuss. 1927

Sebrito del libro Homenaje al Dr Edwardo Caballero

y Caballero Mex, 1960; 75-85,

Taswell’s 1903 material, redescribed Angel, 1960.

Cercariae and sporocysts V1054.

S10

Cercaria jaenschi Johnston & Cleland, 1937

Trans. R. Soc, S, Aust, 61: 202-6,

Cereaniae VI057; infeeted Lenameira sp. V1055;

infected Amira pyramidaia serial sections, V 1058,

Cerearia lethargica Johnston & Muirhead, 1949

Trans. R. Soe, §. Atist, TZ (11> 106-8.

Cercariae = V868, V2783; meiaceccariae VR69;

sporocysts V867; eysts V2784: from Plotopsis tatel,

Cercaria lophosana Sohnsion & Beckwith, 1947b

Trans, R. Soc. 8. Aust 71 (2): 328-333.

Cercaria V886, V2785; sporocysts WS85: infected

Noropala hanleyi V2786,

Cercaria metadina Johnston & Angel, 1942

Trans. R. Soc. S. Aust. 66 (1): 50-9,

Cercaria V908-9, ¥2769; sporocysts V907; sporo-

cysts with diplastomula V910-3; diplostomula

V914-6: from Planorbis isingi, Ameriaana pyrami-

data and A, tenuirostra,

Cercaria natans Johnston & Muirhead, 1949

Trans. R. Soe. 8, Aust, TA (1): 103-6.

Cercariae, V865, V27815 metacereariae V866; redia

V864; infected Planorbis ising V2782.

Cercaria notopalae Johnston & Beckwith, 1945

Trans, R, Soc. S. Aust. 69 (2): 234-41,

Cercaria (fureocercaria) V879, V2775; sporocysts

V878; infected Notepala hanleyi V2776.

REC_ S. AUST. MUS, 18 (22); 503-511,

May. LO8d

Cercaria parocellata Johnston & Simpson, 1939

Trans, R. Sac. S§. Aust, 63.(1): 63-S-

Cercariae V945-6, Y2773: from Lyyinaea lessoui.

Cercaria plotiepsivy Johnstan & Simpson, 1939

Trans. R, Sec. S. Aust. 63 (1): 65-8.

Cercarise V947, V2774; from Plorepyis tatei.

Cercaria tated Johnston & Angel, 19400

Trans, R, See. §. Aust. 64 (2): 334-9.

Cercuriae W932-4, V2771; mictdeereariae V938;

sporocysts and cysts V935-7: from Ploiwpsiy tatei,

Cerearia teiradéna Johnston & Beckwith, 1945

Trans, R. Soc. S. Ausi. @9 (2); 229-33

now Cercaria tetradencidea (Johnstun & Beckwith,

1945) Johnston & Angel, 1940b,

Cercaria (furcocercaria) V928, V2777; sporocysts

V929- from Fletopsis tate.

Cercaria velesinionsis Angel, 1961

Trans. R. Soc. 8. Aust. 84+ 63-70.

Cereariae V998-99, V1003: metacereariac Y995-7,

V1002, young adults V1IO00-1: Crom Felesunio

ambiguis.

ACKNOWLEDGEMENTS

I would like to thank Mrs. P.M. Thomas (Maw-

sou), Miss L. M. Angel and Dr. S. J. Edmonds

without whose advice and assistance these lists of

helminth types could not have been prepared. This

work was funded by an Austrahan Biological

Resources Study grant.

REFERENCES

ANGEL, L, M. 1954, Purerelis acanrhis var australis,

nm. Var., with am account of the life eycle in ‘South

Australia. Trams. R. Soe. S. Anse. 172 164-74,

ANGEL, L. M. 1960. Cerearie haswelli Dollfus, 1927; a

re-examination of Haswell’s material with a discussion

of the genus Tergestia, Sohretire del libra Homenaje

al Br Eduarnda Caballera y Caballero 1960: 74-86.

ANGEL, L. M, 1961, Larval trematodes from Australian

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A REVISION OF THE GENUS APLEROTUS DALLAS (HETEROPTERA:

PENTATOMIDAE: PENTATOMINAE) WITH DESCRIPTION OF A NEW

SPECIES FROM SOUTH AUSTRALIA

BY IMTIAZ AHMAD, NASEER AHMAD KHAN AND SYED KAMALUDDIN

Summary

A new species grossi of the genus Aplerotus Dallas, 1851 is described from South Australia, with

special reference to its metathoracic scent gland ostioles and male and female genitalia. The type

species, A. maculatus Dallas, is redescribed. The relationships of the genus within the subfamily

Pentatominae are also briefly discussed.

A REVISION OF THE GENUS APLEROTUS DALLAS (HETEROPTERA; PENTATOMIDAE:

PENTATOMINAE) WITH DESCRIPTION OF A NEW SPECIES FROM SOUTH AUSTRALIA!

IMTIAZ AHMAD, NASEER AHMAD KHAN? and SYED: KAMALUDDIN

Department of Zoology-Entomology, University of Karachi, Karachi-32, Pakistan

ABSTRACT

AHMAD, |. KHAN, NL A., and KAMALUDDIN, S. 1982.

A teyision of the genus Aplerolus Dallas (Heterop-

tera: Pentatomidae:Peniatominae) with description of a

new species from South Australia, Ree. S. Aust. Mis.

18 (23): 513-518.

A new species gross’ of the genus Apleretus

Dallas, 1851 is described from South Australia, with

special reference to its metathoracic scent gland

ostioles and male and female genitalia. The type

species, A, maculatus Dallas, is redescribed, The

relationships of the genus within the subfamily

Pentatominae are also briefly discussed.

INTRODUCTION

Aplero(us Dallas is to date monotypic and appears

quite common all over eastern, western and southern

Australia. (Gross, 1976). The type species 4. snacu-

latus Dallas is known only by a short. original

description of Dallas (1851) mostly based on colour

features and a more detailed deseription of Gross

(op. cit.) including, in addition to external features,

in the male genitalia the structures of parameres,

pygophore and uninflated aedeagus. Gross (op. cit.)

noled that seven attempts, all unsuccessful, were

made to inflate ihe aedeagus. The female genitalia

of the type species are entircly unknown,

By the courtesy of Dr G. F. Gross, Principal

Curator of the South Australian Muscum, Adelaide,

Australia, the present authors were able to examine

a long series of specimens from various Jocalitics

from castern, southern and western Australia which

clearly represented a complex of two species on

the basis of consistent differences in the characters

of general size, colour patterns on the central disc of

scutelluin, (Figs. | & 11), shapes of peritremes of

the metathoracie scent gland ostioles (Figs, 2 & 12)

and structure of the male and female genitalia (Figs.

3-9 & 13-19) as noted in the present descriptions

and “comparative notes” under each species, The

holotype of A. maculatus Dallas (without abdomen)

was examined by the senior author by the courtesy

' Financially supported by an earlier PARC-USDA Research

Project No. A-17-ENT-37, FG-Pa-181 und present PR-

SEA-155, FG-Puél.

“Presently at Imperial College of Scicnee & Technology,

London,

of Drs P. Freeman, Keeper of Entomology, and

W. J. Knight, who is in charge of the Hemiptera

section, the British Museum of Natural History,

London during his visit to that museum in the year

1977-78.

The new species is named grossi in honour of

Dr G, F. Gross in recognition of his voluminous

taxonomical works on various groups of penltatomo-

morphous Heteroptera. For measurements and dia-

grams ind for dissection of male and female geni-

talia, the conventional procedures especially those of

Ahmad and Khan (1980) have pvenerally been

followed.

Genus APLEROTUS Dallas

Aplerotus Dallas, 1851, p. 256; Gross, 1976, p.

370.

Description: Coloration; Generally dark but bril-

liantly patterned; pronotum with a pale median

streak and corium hear apex with a broad trans-

verse pale stripe,

Head: Eyes only slightly stalked; paraclypei more

or less equal to clypeus; antennae four-segmented

with basal segments at least reaching or distinctly

passing beyond head apex; antenniferous tubercles

spinously produced; labium short, never passing

beyond hind coxae, with basal segment slightly pass-

ing beyond bucculae.

Thorax: Pronotum more than 2!x broader than

long, lateral margins entire; metathoracic seent vland

ostiole with peritreme clongated, blade-like, evapora-

tive area well defined.

Abdomen: Convex beneath without sulcation, stri-

dulatory vittae present forming a curved line laterally

on 2nd, 3rd and 4th abdominal segments.

Male genitalia; Pygophore broader than long,

dorsomedian surface posteriorly produced into a

rounded or blunt process, lateral lobes prominent

and broad; parameres clongate with outer margin

medially straight, anteriorly prominently inwardly

curyed and more or Jess blade-like with serrated

margins; inflated aedeagus with trilobed dorsal mem-

branous conjunctival appendage, vesica long but

shorter than distantly placed and proximally atlached

median penial plates, sclerotized conjunctival

appendages and membranous lobes present.

Sl4

Female genitalia: First goniocoxae widely separ-

ated; 91h paratergites lobe-like, passing beyond

posterior margin of Sth paratergites; triangulin, arcus

and 2nd gonocoxae quite prominent and sclerotized;

spermathecal bulb without processes, proximal and

distal flanges well developed, proximal spermathecal

duct longer than distal one,

Comparative nore: Aplerolus Dallas appears most

closely telated to Diemenia Spinola and Niarius

Stal among the Diemenfa group in having four-

segmented antennae. It resembles Diemenia and

Kalkadoona Distant in having median penial plates

and the latter in having sclerotized second paired

conjunctival appendages and other membranous and

sclerotized conjunctival appendages in the aedeagus

but can be separated from both in having the lateral

mitrgins of the head in front of the eyes without

even a reduced tooth, the pronotal lateral margins

entire, without trace of crenulations and the para-

meres unique among the entire subfamily Pentato-

ninae.

Aplerows maculatus Dallas

(Pigs. 1-10)

Aplerotus maculaius Dallas, 1851, p, 256; Gross,

1976, p. 372,

Description; Coloration: Body black, uniformly

punctate except for a faint vertical line on middle

of clypeus, anterolateral margins of pronotum on

each side and a vertical median line of pronotum; a

smal] rounded patch on each basal angle, a median

patch on mediolateral margins and a single apical

patch on scutellum; a horizontal broad streak on

FIG. 1. Apleraits muevlatas.

REC. 5. AUST, MUS. 18 (23): 513-518.

Muy, 1984

each corium and proximal portion of each connexi-

val joint pale; eyes brownish black, ocelli light

pinkish,

Head: Anteocular portion shorter than the pos-

terior portion of head including eyes, length of

antcocular portion 0,7 mm (0.7-0.8 mm); length

posterior portion of head including eyes 0.9 mm

(0.8-1.05 mm); width of head 2.3 mm (2,21-2.8

mm); interocular distance 1.2 mm (1,1-1.4 mm);

interocellar distance 0.7 mm (0.7-0.8 mm); para-

clypei with anterolateral margin nearly straight,

anteriorly narrowed; antennae with basal segment

almost equal to apex of head, 3rd segment dis-

tinctly longer than 4th, length of segments 1 0.4 mm,

2 1.8 mm (1.8-2.1 mm), 3 1.3 mm (1,.3-1.5 mm),

4 1.2 mm (1.2-1.35 mm); antennal formula

1<4<3<2; labium short, not quite reaching hind

coxac, 4th segment distinetly longer than basal

segment, Jength of segments, 1 0.6 mm (0,6-0.7

tam), 2 1.0 mm (0.9-1.0 mm), 3 0.8 mm (0,8-0,9

mn), 4 0.7 mm (0.7-0,8 mm), labial formula

<4<3<2,

Thorax and Abdamen: Pronotum slightly longer

than head; length 1.7 mm (1,6-2,0 mm); width

4.45 mm (4.2-5.2 mm), Seutellum slightly longer

than. 2x head length; length 3.3 mm (3.25-3.8 mm);

width 2.9 mm (2,7-3,3 mm); metathoracic scent

gland ostioles (Fig. 2) with apex of peritreme

curved anteriad; Jength base scutellum-apex clavuys

1,3 mm (1.2-1.4 mm); apex clavus-apex corium 1,1

min (1.0-1.1 mm); apex scutellum-apex abdomen

including membrane 3.3 mm (3,1-3,5 mm); pos-

terior margin of 7th abdominal segment in females

shallowly concave, inner lateral margin convex; total

length ¢ 9.9 mm (9.3-9.9 mm), ? 10.6 mm (10.85-

11,2 mm)

Male genitalia: Pygophore (Figs, 3 & 4) with

posteroventral margin medially notched, apex af

dorso-median process rounded, inner dorsal process

large, straight, with outer margin entire, parameres

(Fig. 5) broad, with fine serratian on middie of

inner margin and a large tooth-adjacent to it near

proximal end; theca (Figs. 6-8) with broad, knob-

like dorsolateral appendages, median lobe of dorsal

membranous conjunctival appendage short with apex

sclerotized, with lateral margin sinuate, a pair of

lobe-like dorsolateral membranous canjunctival

appendages present with apex sclerotized, frontal or

2nd pair of conjunctival appendages with apex

broad, proximolaterally broad, median penial plates

anteriorly pointed, posteriorly elongated with apex

acute.

Female genitalia: (Figs. 9 & 10), Lateral margins

of Ist gonocoxae concave; 9th paratergites with

lateral margins concave; triangulin and arcus fused

REVISION OF THE GENUS APLEROTUS

mate rste

2, metathoracic scent glind ostioles, ventral view; 3, pygophorc, dorsal view, 4, pygo-

BIGS, 2-10. Aplereris maculatus: 2,

phare, ventral view: 5, paramere, inner view; 6, uedeagus, dorsal view; 7, aedeagus, ventral yiew; 8, aedeagus;

lateral view; 9, female genitalia, ventral view; 10, spermatheca,

with posterior margin, medially indistinctly im-

pressed; 2nd yonocoxae with posterior margin

medially impressed; proctiger with posterior margin

nearly straight, posteriomedian margin fused; 8th

paratergites slightly concave; spermatheea (Fig. 10)

with pump region narrowed, tube-like, distally broad,

median selerotized duet of spermatheen) dilation

distally and medially much dilated, proximal sperm-

thecal duct slightly longer than distal spermathecal

duct.

Material examined: Holotype ? Australia; Aple-

rotus maculanis Dallas; in British Museum Natural

History. London, 9.2, 62 South Australia: Araana

Dam, Fringunda Valley; Western Australia! E.

Balladonia; on Exocarpos ciipressiformis Labil, at

light and on Exocurpes aphyllus R. Be., 19.9.1963,

193.1967, 1.12.1968, 63.1973, 17-24.11,1975;

G, PF. Gross, E. G. Matihews and N. McFarland:

South Australian Museuni, Adelaide, Australiu.

Comparative note: A. maculatus is closely related

to 4. grossr, but it can easily be separated by its

short labium not reaching the hind coxae, in males

parame¢res with a large tooth al medioinner margins

and in females, the pump region of the spermatheea

large, tube-like. equal to the length of the sperma-

thecal bulb and by other characters as noted in the

description,

Aplerotus grassi sp. nev.

(Figs. 11-20)

Description: Coloration: Body black, woifarmly

punctate except for a prominent vertical line on

middle of clypeus; anterior thin and lateral wide

portion, 4 pair of small posterior spots and a wide

yertical median band of pronotuni; a small rounded

patch on each basal angle, a broad inverted T-shaped

band on middle and a single apical patch on seutel-

lum; thin posterolateral portion of embolium;

unteriar horizontal broad band on each corium and

proximal portion of each connexival joint dull pale;

eves brownish black; oeelli light pinkish.

Head: Anteocular distance distinctly shorter than

posterior portion of head including eyes; length of

anteocular portion 0.65 mim (0.65-0,7 mm); length

posterior portion of head including eyes 0.85 mim

(0.8-1.0 mm); width of head 2.1 mm (2.05-2.3

mm); interocular distance |,15 tim (1,1-1.25 mm);

interocellar distance 0.7 mm (0.7-0.8 mm); para-

clypei with anterolateral margin prominently con-

vex, anteriorly broad, anteanae with basul segments

noticeably passing apex of head: 3rd segment about

equal to or slightly longer than 4th segment, length

of segments 1 0.55 mm (0.5-0.55 mm), 2 1,70 sun

(1.65-1,9 mm), 3 1.15 mm (1.1-1.3 mm), 4 1,15

mm (1.-1,125 mm); antennal formula <3 —

REC. 5. AUST. MUS. 18 (23); S13-318,

May, 1984

4<2; labium reaching distinctly beyond hind coxae,

4th segment equal in length to basal segment, length

ot segments | 0.6 nun, 2 0.9 mm (0,9-L.0 mm),

3.0.8 mm (0.8-0,.9 mm), 40.6 mm, labial formula

[s4<3<2,

Vhorax and Abdomen: Pronotum slightly shorter

than head, length 1.45 mm (1.35-l.7 mm); width

3.9 mm (3.7-4.5 mm); scutellum slightly shorter

than 2x head length; length 2,8 ann (2,7-3,3 mm);

width 2,4 mm (2.3-3.0 mm); metuthoracic scent

gland ostioles (Fig. 12) width apex of peritremes

narrowed and directed laterad; length base scutellum-

apex clavas 1-3 mm (1-1-1.5 mm); apex elavus-

apex corium 1.0 min (10-152 mm); apex sculellum-

upex abdomen including membrane 3.1 min (2.9-

3.2 mm); posterior margin of 7th abdominal

sternum m females deeply concave, inner margins

straight, total Jeneth 3 8.85 mm (8.4-8.85 mm);

29,85 mm (9.6-9.9 min),

Male genitalia: Pygophore (Figs, 13 & 14). with

posteraventral margin medially impressed inte a pit,

apex of dorsomedian process truncated, inner darsal

process large, curved with outer margin sjnuate;

paramercs (Pig. 15) narrowed, small dentation at

iver margin near apex; prominent theca (Tigs. 16-

18) with comparatively narrowed knob-like darso-

lateral appendages, median lobe of dorsal mem-

brinmous conjunctival appendage short with apex

sclerotized, lateral margins concuye, a pair of lobe-

like dorsolateral membranous conjunctival appen-

dage with apex sclerotized, frontal or 2nd pair of

conjunctival uppendages sclerotized with apex nar-

rowed, proximolaterally macrowed median penal

plates anteriorly rounded, posteriorly clongated with

apex blunt .and rounded,

FIG. IL. Aplerems press

REVISION OF THE GENUS APLEROTUS

<-s

FIGS. 12-20. Apleratus grassi: 12. metathoracic scent gland ostioles, ventral view; 13. pygophore. dorsal view; 14.

pygophore, ventral view; 15, paramere, inner view: 16, aedeagus, dorsal view; 17, acdeagus. ventral view; 18, acde-

agus, lateral view; 19, female genitalia, ventral view: 20, spermatheca

518

Female genitalia; (Figs. 19 & 20). Lateral margins

of Ist ygonocoxae sinuate; 9th paratergites with

lateral margins convex; triangulin and arcus fused

with posterior margin medially distinctly notched;

2nd gonocoxae with posterior margin medially con-

cave; proctiger with posterior margin deeply con-

cave; posteriomedian margin of fused 8th para-

tergites slightly convex; spermatheca (Fig, 20) with

pump region broad, barrel-shaped, shorter than

spermathecal bulb, median sclerotized duct of

spermathecal dilation much dilated near proximal

end, proximal spermathecal duct more than 4x as

long as distal sperniathecal duct.

Material examined: Holotype 3 South Australia:

Secale Bay, 16.10.1963: G, F, Gross; paratypes:

4¢ and 39 South Australia; Sceale Bay, Eritgunda

Valley; 16,10,1963, 11.11.1966, 6.3.1973, 4,12.

1974, 24,11,1975; at light; G. F. Gross, BE. G,

Matthews; all in the South Australian Museum,

Adelaide, South Australia.

Comparative nate: A. grossi is closely related to

A, maculatus Dallas, but it can readily be separated

by having the anterolateral margins of the paraclypei

strongly convex, the metathoracic scent gland

ostioles with peritremes sinuate, blade-like, apex

subacute, in males, the parameres with dentation

on the inner apical margin and in female sperma-

theca with the pump region short, barrel-shaped,

shorter than the bulb and by other characters as

noted in the description,

SYSTEMATIC POSITIONS

The members of the genus 4pleroins, by virtue

of the possession of strigose vittac, have variously

been placed in the tribes Mecideini Distant and

REC. S. AUST. MUS, 18 (23): 513-518

Way, 1984

Diemenini Kirkaldy and the Diemenia group of

Gross (1976), Gross included nine genera occur-

ring in South Australia. The three genera men-

tioned here were keyed out together as all hav-

ing four-segmented antennac. Gross (op. cit.)

described and figured the aedeagus and parameres

of representatives of Diemenia, Kalkadoona Distant

and Oncocoris Mayr, and the spermatheca of OQ,

desertus Bergroth. McDonald and Edwards (1978)

and McDonald (1978, 1979) described and figured

the male and female genitalia of all species of

Oncocoris and of one species of Kalkadoona. The

male and female genitalia of other members of the

group are virtually entirely unknown. Ahmad and

Khan (1980) described Knightiella to accommodate

Stenozygum flavifrons Distant, which also possesses

sttigose villae, Probably the strigose vittac are

shared by the members of remarkably diverse groups

in the same way as pseudoclaspers are known in

three very diverse genera of pentatomids (McDonald

1976).

On the basis of the male genitalia of the known

members of the Diemenia graup the twa species of

Aplerorns certainly share the characters of medial

penial plates, sclerotized paired second conjunctival

appendages (presently called frontal appendages)

and various membranous lobes and sclerotized

appendages of the conjunctiva, which are especially

similar to those of Kalkadoona pallida (Van Duzee),

The structure of the parameres, pygophore and other

details of the inflated aedeagus of Aplerotus appear,

however, to be entirely different and isolate it from

the rest of its group like Knightiella which also

appears to be very different [rom other Pentato-

minae (Ahmad and Khan 1980), The male geni-

talia of Niarins are uoknown, It could be that this

genus and Aplerofus would form a group.

REFERENCES

AHMAD, L, and KHAN, N, A, 1980, Two new genera

of the subfamily Pentalominae (Heleroptera:Penlulo-

midae) from the Australian Region, Ree. §. Aust. Mis,

18 (3): 83-90.

DALLAS, W. 8, 1851, List of the specimens of hemiplerous

insects in the collection of the Brilish Museum. 1

(London),

GROSS, G. F. 1976, Plant-feeding and other bugs (Hemip-

tera) of South Australia, Heteroptera, Part IL Govern-

ment Printer, Adelaide,

McDONALD, F, J. D, 1976. Revision of the genus Tri-

chopenla (Hemiptera: Pentatomidae) in’ N. America.

J. N.Y. Entomol. Sov. 84 (1): 9-22.

McDONALD, F. f. BD. 1978. Appendix. A description of

Seven new species of Oncocoris Mayr (Wemiptera:Pen-

tatommuae), wilh a key to species. Aus. J. Zool. Suppl.

Ser, No. 62: 19-53.

McDONALD, F. J. DD. 1979. Oncwceris

(HemipterasPentatomidae), J. Aust.

187-189,

McDONALD, F. J. D. and EBWARDS, P. B. 1978.

Revision of [he Genus Oneecoris Mayr (Hemiptera;

Pentatomidae), Aust, J. 4Zvel. Suppl. Ser. No. 62+

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CNUs Sp, ne

eur Soc. VB: