VOL. 128, PART |

31 MAY, 2004

Transactions of the

Royal Society of South

Australia Incorporated

INCORPORATING THE

Records of the

South Australian Museum

Contents

Berge, J. & Vader, W. Description of two new and unusual stegocephalid species

(Crustacea: Peracarida: Amphipoda: Stegocephalidae) from Heard

Island and the East ChinaSea - - - - - - = = |

Jennings, J. T., Austin, A. D. & Stevens, N. B. Species of the Wasp genus Au/acus Jurine

(Hymenoptera: Aulacidae) endemic to South Australia — - — 13

Turner, D. & Conran, J. G. The reproductive ecology of two naturalised Erica species

(Ericaceae) in the Adelaide hills: the rise and fall of two ‘would-be’

weeds? - - - - - - 23

Zbik, M. & Pring, A. The Myrtle nee meteorite: a chondrite oy from South

Australia — — - 3

Leppard, P. L., Tallis, G. M. & Pearce, C. E. M. Expected lifetime in South Australia

IGE = bys Se ee Op ae lee ee 37

Riley, I. T. & Bertozzi, T. Variation in sex ratios in four Anguina (Nematoda:

Anguinidae) species — - = - - 43

Smales, L. R. Syphacia (Syphacia) australasiensis sp. nov. (Nematoda:

Oxyuridae) from Rattus leucopus (Muridae) from Papua New

Guinea and Australia -— -— - - —- - - - = = 47

Jones, H. I. Gastric nematodes, including a new species of Abbreviata

(Nematoda: Physalopteridae) from the mangrove monitor Faranus

indicus (Reptilia: Varanidae) -— - - - - - = = = 53

Parsons, R. F. Enrichment-planting of the woody climbers Marsdenia australis

and Rhynacharrhena Linearis in north-western Victoria. - -— — 61

Spratt, D. M. & Hobbs, R. P. Breinlia (Breinlia) ventricola sp. nov. (Nematoda:

Filarioidea) from the heart of the red kangaroo, Macropus rufus, in

Western Australia - - - - - = = = = = = 67

Phillott, A. D. & Elsmore, S. A. M. Black Noddies (Anous minutus) and Wedge-tailed

Shearwaters (Puffinus pacificus) as potential hosts for fungi

invading sea turtle nests at Heron Island, Queensland - —- — 73

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

INCORPORATING THE

Records of the

South Australian Museum

VOL. 128, PART 1

EDITORIAL

This is the first issue of the Transactions of the Royal Society of South Australia to incorporate the Records of

the South Australian Museum. The Transactions have appeared uninterruptedly for 127 years, publishing

good science from and for southern Australia and the world, and the Records have a similar record of

achievement over 86 years. Why have the Society and the Museum merged these two successful journals,

each with its proud record?

The simple answer is that we believe that a single journal produced by the two institutions, which have many

other beneficial linkages, can be better scientifically, can reach a bigger audience and can offer scientists a

better platform for their research. In some fields, such as taxonomy, there has been much overlap, in others

such as anthropology, little overlap in recent years, so that there is much complementary strength in the two

journals.

What we have done so far is simply to merge the journals and to plan to send the merged Transactions out to

the subscribers to both journals. For this first year, there will be two issues. Future changes will depend

particularly on contributors and readers, the two groups who make or break any journal. If we find strong

authorial support, we shall need to consider more frequent publication, with all the benefits and costs inherent

in such a change. Or we may find that there is a need to divide the journal into parts for readers with different

interests, though for many (such as the current editor) this would be a step with the huge disadvantage of no

longer providing a journal for generalist specialists with interests in all aspects of science as {l applies to

southern Australia and related regions. The current issue, covering plant ecology, threatened species,

parasitology, taxonomy, population science, and earth science, typifies this breadth and strength within the

breadth.

The future is in your hands. Support the merged Transactions as readers and contributors, and it will go from

strength to strength.

Oliver Mayo

Editor, for the Editorial Board

TRANSACTIONS OF THE

ROYAL SOCIETY OF SOUTH AUSTRALIA INC.

CONTENTS, VOL. 128, 2004

PART 1, 31 MAY, 2004

Berge, J. & Vader, W. Description of two new and unusual stegocephalid species (Crustacea:

Peracarida: Amphipoda: Stegocephalidae) from Heard Island and the East

ChinaSea - - - - -—- = = = = =

Jennings, J. T., Austin, A. D. & Stevens, N. B. Species of the Wasp genus Aulacus Jurine

(Hymenoptera: Aulacidae) endemic to South Australia -— —- = =

Turner, D. & Conran, J.G. The reproductive ecology of two naturalised Erica species

(Ericaceae) in the Adelaide hills: the rise and fall of two ‘would-be’ weeds?

Zbik, M. & Pring, A. The Myrtle Springs meteorite: a chondrite (H4) from South Australia —

Leppard, P. I., Tallis, G. M. & Pearce, C. E. M. Expected lifetime in South Australia 1841 — 1996

Riley, I. T. & Bertozzi, T. Variation in sex ratios in four Anguina (Nematoda: Anguinidae) species

Smales, L. R. Syphacia (Syphacia) australasiensis sp. nov. (Nematoda: Oxyuridae) from

Rattus leucopus (Muridae) from Papua New Guinea and Australia - —-

Jones, H. I. Gastric nematodes, including a new species of Abbreviata (Nematoda:

Physalopteridae) from the mangrove monitor Varanus indicus (Reptilia:

eg ty a

Parsons, R. F. Enrichment-planting of the woody climbers Marsdenia australis and

Rhyncharrhena Linearis in north-western Victoria - -— - - - =

Spratt, D. M. & Hobbs, R. P. Breinlia (Breinlia) ventricola sp. nov. (Nematoda: Filarioidea) from

the heart of the red kangaroo, Macropus rufus, in Western Australia -— —

Phillott, A. D. & Elsmore, S. A. M. Black Noddies (Anous minutus) and Wedge-tailed Shearwaters

(Puffinus pacificus) as potential hosts for fungi invading sea turtle nests at

Heron Island, Queensland - - =

PART 2, 30 NOVEMBER, 2004

Appan, A., Bergfeld, J. & Beveridge, I. New species of parasitic nematodes from macropodid

marsupials in Western Australia - - - -= —~ = = = = =

Hammer, M. P. & Walker, K. F. A catalogue of South Australian freshwater fishes, including new

records, range extensions and translocations - - - - -— =

Watts, C. H.S & Humphreys, W. F. Thirteen new Dytiscidae (Coleoptera) of the genera

Boongurrus Larson 7firtudessus Watts & Humphreys and Nirripirti Watts &

Humphreys, from underground waters in Australia — - - -

Watts, C. H. S. Revision of Australian Scirtes Uliger and Ora Clark (Coleoptera: Scirtidae) —

Hodda, M. Characteristics of an Australian population of Paraxonchium orthodon (Loof

1964) Altherr & Loof 1969, with a note on evolution and biogeography of the

subfamily Paraxonchiinae (Nematoda, Dorylaimida, Aporcelaimidae) —

Beveridge, I. & Johnson, P. M. Cestode parasites of tree kangaroos (Dendrolagus spp.:

Marsupialia), with the description of two new species of Progamotaenia

(Cestoda: Anoplocephalidae) - - - - - - ~~ - =

Faucheux, M. J. Antennal sensilla of Sabatinca sterops Turner (Lepidoptera: Micropterigidae)

Barker, S. Twelve new species of Australian Buprestidae (Coleoptera) and new

synonymy - - ~- - = = = = = = = = = =

Womersley, H. B. S. Additions to the Marine Algal Flora of southern Australia - -— -— =

Smith, B. P. C. & Kokkinn, M. The use of emergence as an end-point for sediment toxicity tests

using the Australian chironomid Chironomus maddeni — -— - = =

Barton, P. S., Aberton, J. G. & Wishart, E. Spatial and temporal distribution of Culex australicus

Dobrotworsky and Drummond and Culex globocoxitus Dobrotworsky

(Diptera: Culicidae) at the Gippsland Lakes in eastern Victoria = — ~ =

Williams, C. R., Snell, A. E. & Kokkinn, M. J. Studies of temporal host-seeking patterns of Culex

Annulirostris (Diptera: Culicidae): a comparison of methods and populations

tibbs, S. E., Kemper, C. M., Byard, R. W. & Long, M. Deaths of killer whales (Orcinus orca) in

South Australia and implication of human interaction = — me

Hinchcliffe, J. & Conran, J. G. The Tennyson sand dunes: vegetation structure and conservation

status = — = 3 = Sie: ee eo a ass ee

Smith, B. B. & Walker, K. F. Reproduction of common carp in South Australia, shown by young-

of-the-year samples, gonadosomatic index and the histological staging of

ovaries - - = = = = = = = = = = = =

Brief Communications:

Souter, N. J. A comparison among three artificial substrates for aquatic macroinvertebrate

sampling - — =— RS

Matthews, E. G. New synonymy and new names in Australian Tenebrionidae (Coleoptera) = —

Insert to Transactions of the Royal Society of South Australia, Val. 128 Part 2, 30 November, 2004

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DESCRIPTION OF TWO NEW AND UNUSUAL STEGOCEPHALID

SPECIES (CRUSTACEA: PERACARIDA: AMPHIPODA:

STEGOCEPHALIDAE) FROM HEARD ISLAND AND

THE EAST CHINA SEA

By J. BERGE! & W. VADER?

Summary

Berge, J. & Vader, W. Description of two new and unusual stegocephalid species

(Crustacea: Peracarida: Amphipoda: Stegocephalidae) from Heard Island and the East

China Sea. Trans. R. Soc. S. Aust. 128(1), 1-11, 31 May, 2004.

Two new stegocephalid (Amphipoda) species are described: Mediterexis macho and

Stegocephalina wolf. One of the species, Mediterexis macho, is the very first record

of a stegocephalid from the China Sea, whereas Stegocephalina wolf was collected

North of Heard Island in the Southern Ocean.

Key Words: Amphipoda, Stegocephalidae, Mediterexis macho sp. _ nov.,

Stegocephalina wolf sp. nov.

Transactions of the Royal Society of S. Aust. (2004), 128(1), 1-11.

DESCRIPTION OF TWO NEW AND UNUSUAL STEGOCEPHALID SPECIES

(CRUSTACEA: PERACARIDA: AMPHIPODA: STEGOCEPHALIDAE)

FROM HEARD ISLAND AND THE EAST CHINA SEA.

by J. BERGE! & W. VADER?

Summary

Berce, J. & VADER, W. Description of two new and unusual stegocephalid species (Crustacea: Peracarida:

Amphipoda: Stegocephalidac) from Heard Island and the East China Sea. Trans. R. Soc. S. Aust. 128(1), 1-11,

31 May, 2004.

Two new stegocephalid (Amphipoda) species are described: Mediterexis macho and Stegocephalina wolf. One

of the species, Mediterexis macho, is the very first record of a stegocephalid from the China Sea, whereas

Stegocephalina wolf was collected North of Heard Island in the Southern Ocean.

Kry Worps: Amphipoda, Stegocephalidae, Mediterexis macho sp.nov., Stegocephalina wolf sp.nov.

Introduction

The Antarctic fauna of stegocephalid species was

reviewed by Berge ef a/. in 2000, in which a total of

19 species were recognised. Since then two

additional stegocephalid species have been recorded

from the area, excluding the new species

Stegocephalina wolf reported herein. Thus, the

Antarctic stegocephalid fauna currently consists of

22 known species belonging to I! genera. The

second new species herein described, is the first

record of a stegocephalid species from the China Sea.

A phylogenetic analysis of the family

Stegocephalidae was presented as part of a recent

revision of the group (Berge & Vader 2001). Based

upon this analysis, a new classification at the generic

level was outlined. As the classification at both

generic and subfamily levels falls outside the scope

of this paper, the two new species (Mediterexis

macho sp.nov. and Stegocephalina wolf sp.nov.)

described herein are treated within the framework of

that classification. Especially for Mediterexis macho,

the generic position may well be altered in the future

(see also below).

Material and Methods

This study is based upon material from the South

Australian Museum (SAM). All dissected

appendages were mounted in polyvinyl-lactophenol

and stained with rose-bengal. These appendages were

drawn using a Leica compound microscope equipped

with a drawing-tube, while the habitus-drawings were

‘UNIS, Department of Biology, Pb 156, 9170 Longyearbyen,

NORWAY. Email: jorgen.berge@unis.no

> Tromso Museum, Dept of Zoology, University of Tromso, 9037

Tromso, NORWAY. Email: wim@imy.uit.no

made using a Leica dissecting microscope. Mature

and immature females were distinguished from males

by the presence of oostegites. The classification of

setae and setae-groups follows that of Berge (2001).

All scales attached to the figures are 0.1 mm unless

otherwise stated.

Symbols

Al-2: Antenna 1-2; EP3: Epimeral plate 3; IP:

Inner plate; L: Labium; LBR: Labrum; LMND: Left

mandible; MX1: Maxilla 1; MX2: Maxilla 2; MXP:

Maxilliped; OP: Outer plate; P1-7: Pereopods 1-7;

PLP: palp; RMND: Right mandible; ST: Setal teeth

on the first maxilla; T: Telson; U1-3: Uropods 1-3.

Results and Taxonomy

The present study is based entirely on material

provided from the collections of the South Australian

Museum. The material comprised, in addition to the

two new species reported herein, a total of 3 species;

Glorandaniotes sandroi Berge & Vader, 2003a,

Parandania boecki Stebbing, 1888 and Tetradeion

crassum (Chilton, 1883). The three above mentioned

species were all found in Australia, whereas the two

remaining, Mediterexis macho sp.nov. and

Stegocephalinae wolf sp.nov., were collected from

the East China Sea and Heard Island, respectively.

Key to the 22 species known from the Antarctic

and sub-Antarctic regions:

1. Pereopod 6 basis expanded, posterior margin

convex (2)

- Pereopod 6 basis weakly expanded, posterior

margin straight or concave (17)

- Pereopod 6 basis not expanded, about as broad

as pereopod 5 basis (21)

to

i)

Uropod 3 rami obsolescent or absent

Stegocephalina pacis

- Uropod 3 both rami well developed (3)

3. Telson entire (4)

- Telson cleft (8)

4. Antennae subequal (5)

- Antenna 2 elongate, longer than antenna |

Parandania boecki

5. Antenna | flagellum with 5 articles Andaniexis

ollii

- Antenna | flagellum more than 10 articles (6)

6. Antennae elongate Parandania boecki (juveniles)

- Antennae not elongate (7)

7. Labrum symmetrical, both

reduced Parandania gigantea

- Labrum asymmetrical, right lobe large, left lobe

strongly reduced Parandania nonhiata

8. Rostrum weakly developed (9)

- Rostrum large and distinct Stegocephalus

rostrata

9. Coxa | anterior margin convex (10)

- Coxa | anterior margin with a deep invagination

Stegocephalus watlingi

10. Telson longer than broad, triangular and pointed

(11)

- Telson about as long as broad, not pointed (13)

11. Mouthparts not elongate and not forming a

conical bundle (12)

- Mouthparts elongate, forming a conical bundle

projecting well below coxae Stegocephalina

wolf sp.nov.

12. Epimeral plate 3 posteroventral corner acute,

with one notch Stegocephalus kergueleni

- Epimeral plate 3 posteroventral corner rounded,

crenulated Genus Pseudo (see below)

13. Antenna | flagellum article | shorter or about as

long as peduncle (14)

- Antenna | flagellum article | distinctly longer

than peduncle (15)

14. Coxae 1-3 broad and overlapping, coxa 4

posterior lobe exceeding pereon

segment 6 Stegosoladidus antarcticus

- Coxae 1-3 narrow, not overlapping, coxa 4

posterior lobe not exceeding pereon segment 6

Andaniotes linearis (immature)

15. Maxilla 2 gaping and geniculate, epimeral plate

3 posteroventral corner produced and with

teeth Genus Pseudo (see below)

- Maxilla 2 not gaping and geniculate, epimeral

plate 3 posteroventral corner weakly

produced, without teeth (16)

16. Antenna | flagellum with 4 articles Andaniotes

pseudolinearis

lobes strongly

'The validity of this genus is discussed in Berge & Vader

(submitted), To avoid any nomenclatory complications, the

classification according to Berge & Vader 2001 is utilised.

J. BERGE & W. VADER

- Antenna | flagellum with more than 10 articles

Parandania boecki

17. Uropod 3 outer ramus 2-articulate (18)

- Uropod 3. outer = ramus |-articulate

Stegosoladidus ingens

18. Telson cleft (19)

- Telson entire Andaniella integripes

19. Pereopod 6 basis posteromedially with 2-3 long

plumose setae (20)

- Pereopod 6 basis posteromedially with a row of

short robust setae Andaniotes linearis

20. Coxae 1-3 broad and overlapping, maxilliped

inner plate with 2 nodular setae

Stegosoladidus debroyeri

- Coxae 1-3 narrow, not overlapping, maxilliped

inner plate with 4 nodular setae Andaniotes

pooh

21. Telson entire (22)

- Telson cleft (23)

22. Telson short, pereopod 4 subchelate, pereopod 7

well developed Parandaniexis dewitti

- Telson long, pereopod 4 simple, pereopod 7

reduced 7etradeion crassum

23. Epistomal plate large, conspicuous

Austrophippsia unihamata

- Epistomal plate absent Schellenbergia!

vanhoeffeni

Remarks to the kev

The key is rewritten and updated from that

presented by Berge et a/. (2000) to include both the

new Stegocephalina wolf sp.nov. (see below) and the

two Pseudo Berge & Vader, 2001 species that were

described, but not given a formal scientific name in

Berge & Vader (submitted). Furthermore, all generic

names in the key are updated according to the

outlined classification of all stegocephalid taxa in the

revision of the family by Berge & Vader (2001).

Subfamily Andaniexinae Berge & Vader, 2001.

Genus Mediterexis Berge & Vader, 2001.

Mediterexis macho sp. n. (Figs 1-3)

Holotype

Male, 4 mm (SAM C6054), East China Sea (30°

06’ N, 130° 35’ E), 02.07.1966. Unique.

Etymology

The name refers to its densely setose peduncles of

the antennae, compared to all other stegocephalid

taxa.

Diagnosis

Antennae subequal, penduncles setose, accessory

flagellum longer than antenna | flagellum article 1.

Epistome laterally produced, epistomal plate present.

Mandibles with transverse smooth incisors, left

TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM

Fig. |. Mediterexis macho sp.nov. Holotype.

4 J. BERGE & W. VADER

Fig. 2. Mediterexis macho sp.nov. Holotype.

TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM 5

Fig. 3. Mediterexis macho sp.nov. Holotype.

lacinia mobilis weakly toothed. Maxilla 2 not gaping

and geniculate. Labrum shorter than broad. Pereopod

6 basis posteriorly conspicuously expanded. Uropod

outer ramus two-articulate. Telson entire.

Description

Head: Head retractable under pereonite 1. Rostrum

rudimentary.

Antennae: Accessory flagellum longer than

flagellum article 1, articulation present. Antenna 2

peduncle article 3 short, about as long as broad.

Peduncle article 4 shorter than article 5.

Epistome: Epistome laterally produced and

rectangular, with a long ridge on each side.

Epistomal plate (medial keel) produced into a small

elongate medial ridge covering the entire epistome.

Mouthparts: Mouthparts not elongate.

Mandibles with incisor transverse, smooth. Left

lacinia mobilis present, weakly toothed and distally

not produced.

Maxilla 1 palp articulation absent, distally with

long pappose setae. Outer plate distally rounded, ST

in a 6/3 arrangement with two parallel rows: First

row with ST 1-5 present, ST 6 absent and ST 7

present. Second row with ST A-C present. Inner

plate with shoulder weakly developed, setae

pappose.

Maxilla 2 outer plate not gaping and geniculate,

setae distally straight. Inner plate with setae-row A

and B appressed. Row B with first 3 setae

differentiated from the other setae. Row C present,

row D absent.

Maxilliped palp 4-articulate, article 2 not produced

distally. Articles 1-3 with long slender setae. Inner

plate distally with inner corner weakly produced, 2

nodular setae present. Outer plate with inner and

outer setae-row present, both with robust slender

setae. Distal setae-group absent.

Labrum shorter than broad, both lobes reduced.

Labium distally broad, distal finger absent.

Pereopods and coxae: Coxal plates and basis of the

pereopods smooth. Coxae 1-3 contiguous.

Pereopod | coxal plate about as deep as basis,

basis anterior margin weakly expanded. Pereopod 1

6 J. BERGE & W. VADER

propodus subovate, posterior submarginal row of

setae present.

Pereopod 2 ischium elongate, distal posterior

margin without plumose — setae. Propodus

subrectangular, posterior submarginal row of robust

setae present.

Pereopod 4 coxa locking-structure absent. Basis

with long setae on posterior margin, distally without

plumose setae. Ischium with long plumose setae on

distal posterior margin.

Pereopod 6 basis posteriorly expanded, medially

with a row of long plumose setae. Posterior margin

of basis serrate.

Pereopod 7 general appearance similar to pereopod

6. Basis medially with a row of short simple setae.

Oostegites and gills: Gills present on pereopods 2

7. Oostegites unknown.

Pleonites: Pleonites 1-3 dorsally smooth. Epimeral

plate 3 weakly produced and rounded posteriorly,

serrations absent.

Urosome: Males with urosome not enlarged.

Articulation between urosomites 2 and 3 present.

Uropod | outer ramus with short robust setae on

outer and inner margin, inner ramus with robust setae

on inner margin, Uropod 2 outer ramus outer margin

with robust setae, inner ramus inner margin with

robust setae. Uropod 3 peduncle longer than telson,

outer ramus 2-articulate. Rami without setae. Telson

entire and rounded distally, submarginal setae on

apex present.

Females

Unknown.

Distribution

Known only from the type locality,

Remarks

This species can easily be distinguished from all

other stegocephalid taxa by the combination of 1) a

laterally produced epistome, 2) the weakly

differentiated last two pairs of pereopods, 3) entire

telson and 4) the bi-articulate outer ramus of the third

uropod. The presence of a not gaping and geniculate

maxilla 2 and the smooth mandibular incisors indicate

unequivocally that the species is a true member of the

subfamily Andaniexinae Berge & Vader, 2001.

However, it is not possible to assign it unequivocally

to any of the genera of the family: its telson,

mandibles and maxilliped resemble closely those

found in the three genera Andaniexis Stebbing, 1906,

Mediterexis Berge & Vader, 2001 and Parandaniexis

Schellenberg, 1929. These three genera are also the

only genera within the subfamily that do not possess a

locking-process on the inner side anteriorly on the

fourth coxa (see Berge & Vader 2003b), but the first

maxilla and the weakly differentiated last two pairs of

pereopods have close affinities to those found in the

genus Andaniotes Stebbing, 1897. In addition, the

second maxilla, with its slightly elongate outer plate

and the arrangements of setae-rows A-D has closest

affinities to the subfamily Andaniopsinae Berge &

Vader, 2001. Thus, the assignment of this species to

Mediterexis seems, at best, a temporary decision.

However, as the phylogeny of the family and

allocation of its species to genera fall outside the

scope of this paper, this species is herein placed in

Mediterexis based on the close affinities to that genus

in the maxilliped, mandibles and epistome. This last

character (see Fig 4, Mediterexis mimonectes (Ruffo,

1975) was used in the phylogenetic analysis (Berge &

Vader 2001), but closer examination has revealed that

the epistome shows a more complex set of characters

than previously assumed (Berge, in prep.). As

currently only one single specimen of this new species

exists, its epistome was not available for SEM

examination. However, examination under the light

microscope gives reason to suspect that these two

congeners possess a very similar epistome; that of M.

mimonectes is herein illustrated.

Subfamily Stegocephalinae Dana, 1852

Genus Stegocephalina Stephensen, 1925

Stegocephalina wolf sp. n. (Figs 5-7)

Holotvpe

Male 5mm (SAM C6055), Southern Ocean just

North of Heard Island (52° 18'S, 73° 45’ E), 245 m,

col. W. Zeidler, 13.06.1990, “Aurora Australis” st.

65, Sample taken from a sponge. Unique.

Etvmology

This species is named after the collector, Dr

Wolfgang Zeidler, Adelaide.

Diagnosis

Rostrum rudimentary. Antennae — subequal,

accessory flagellum shorter than antenna | flagellum

article 1. Epistome laterally produced, epistomal

plate present. Mouthparts forming a conical bundle.

Mandibular incisors lateral, toothed. Maxilla 2

gaping and geniculate. Labrum elongate. Pereopod 6

basis posteriorly conspicuously expanded. Uropod 3

outer ramus two-articulate. Telson cleft.

Description

Head: Head retractable under pereonite 1. Rostrum

rudimentary

Antennae: Accessory flagellum shorter than

flagellum article 1, biarticulate. Antenna 2 peduncle

article 3 short, about as long as broad. Peduncle

article 4 longer than article 5.

Epistome: Epistome laterally produced, long and

triangular. Epistomal plate (medial keel) produced

TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM A

Pig. 4. Mediterexis mimonectes Ruffo, 1975. SEM picture of the epistome. A: Epistomal plate, B: Epistome laterally

produced, C: Left mandible, D: Labrum.

into a small elongate medial ridge covering the entire

epistome.

Mouthparts: Mouthparts elongate.

Mandibles with incisors lateral, toothed. Left

lacinia mobilis present, strongly toothed and distally

produced.

Maxilla | palp articulation absent, distally with

short simple setae. Outer plate distally

subrectangular, ST in a 6/3 arrangement with two

parallel rows: First row with ST 1-5 present, ST 6

absent and ST 7 present. Second row with ST A-C

present. Inner plate with shoulder well developed,

setae pappopectinate.

Maxilla 2 outer plate gaping and geniculate, setae

distally straight. Inner plate with setae-rows A and B

appressed together. Rows C and D present.

Maxilliped palp 3-articulate, dactylus absent.

Articles | and 3 long and weakly setose, article 2

short. Inner plate elongate, 2 nodular setae present.

Outer plate with inner and outer setae-row absent.

Distal setae-group present, setae short and simple.

Labrum conspicuously elongate, triangular. Left

lobe reduced.

Labium distally narrow, one distal finger present.

Pereopods and coxae: Coxal plates and basis of the

pereopods smooth.

Pereopod | coxal plate about as deep as basis,

basis anterior margin weakly expanded. Pereopod |

propodus subovate, posteriorly without submarginal

row of setae.

Pereopod 2 ischium elongate, distal posterior

margin with plumose — setae. — Propodus

subrectangular.

Pereopod 4 coxa locking-structure absent. Basis

with long setae on posterior margin, distally with

plumose setae on anterior and posterior margins.

Ischium with long plumose setae posteriorly.

Pereopod 6 basis posteriorly expanded, medially

with a row of long plumose setae. Posterior margin

of basis smooth.

8 J. BERGE & W. VADER

Fig. 5. Stegocephalina wolf sp.nov. Holotype.

TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM

Fig. 6. Stegocephalina wolf sp.nov. Holotype.

10 J. BERGE & W. VADER

Fig. 7. Stegocephalina wolf sp.nov. Holotype.

Pereopod 7 conspicuously different from pereopod serrations absent.

6. Basis posteriorly smooth, medially with a row of Urosome: Articulation between urosomites 2 and 3

short simple setae. present. Uropod 1 outer ramus with robust setae on

Oostegites and gills: Gills present on pereopods 2- outer margin, inner ramus with short robust setae on

7. Oostegites unknown. inner margin. Uropod 2 outer ramus with short

Pleonites: Pleonites 1-3 dorsally smooth. Epimeral — robust setae on outer margin. Uropod 3 peduncle not

plate 3 weakly produced and rounded posteriorly, as long as telson, outer ramus 2-articulate.

TWO NEW STEGOCEPHALID SPECIES FROM THE COLLECTIONS OF THE S.A. MUSEUM 1

Telson longer than broad, cleft and rounded

distally, submarginal setae on apex present.

Female

Unknown.

Remarks

Stegocephalina wolf is separated from all other

stegocephalid species by the combination of 1) the

absence of dactylus on the palp of the maxilliped, 2)

the fourth article of the peduncle of the second

antenna longer than the fifth, and 3) the presence of

only one distal finger on the labium.

The present species has close affinities, both

phylogenetically and morphologically, to the type

species of the genus, S. ingo/fi Stephensen, 1925.

Stephensen (1925:136) wrote that S. ingolfi is easily

recognisable e.g. by the long and narrow mouthparts;

the same is also true for S. wolf. The epistome and

mouthparts are in both species conspicuously

elongate, projecting well below the coxae. However,

the present species differs from S. ingolfi in having a

considerably shorter article one of the flagellum and

only one distal finger on the apex of the labium.

Acknowledgements

We are grateful to Dr Wolgang Zeidler at the South

Australian Museum for providing material. Also, we

are indebted to Chris Jones and Prof. Geoff Boxshall

at the Natural History Museum in London for their

help and support in preparing the SEM pictures.

The first author was funded by the Norwegian

Research Council, project number 145384/432.

References

Berar, J. (2001) Revision of Stegosoladidus Barnard &

Karaman, 1987 (Crustacea:Amphipoda: Stegocepha-

lidae); redescription of two species and description of three

new species. Journal of Natural History 35, 539-571.

, De BRoyver, C. & VADER, W. (2000) Revision

of the Antaretic and sub- Antarctic species of the family

Stegocephalidae (Crustacea: =Amphipoda) — with

description of two new species. Bulletin de l'Institut des

Sciences Naturelles de Belgique 70, 217-233.

— & VAber, W. (2001) Revision of the amphipod

te rustacea) family Stegocephalidae.

Zoological Journal of the Linnean Society 133, 531-592.

& (2003a) Stegocephalid (Crustacea:

~ Amphipoda) species from Australia and New Zealand,

with description of seven new species. Records of the

Australian Museum 55, 85-112.

= & (2003b) Metandania tordi n.sp. -

anew stegocephalid (Crustacea: — Peracarida:

Amphipoda) species from the Southern Ocean.

Proceedings of the Biological Society of Washington

116(4), 1007-1016.

& _ (submitted) Two new Antarctic

stegocephalid (Amphipoda: Stegocephalidae: Stego-

cephalinae) species, with implications for the phylogeny

and classification of the two genera Pseudo and

Schellenbergia. Deep-sea Research Part II.

CHILTON, C.

the New Zealand Crustacea. Transactions and

Proceedings of the New Zealand Institute 15, 69-86.

(1883) Further additions to our knowledge of

DANA, J. D. (1852) Conspectus crustaceorum quae in orbis

terrarum circumnavigatione, Carolo Wilkes e classe

Reipublicae Faederatae Duce, lexit et descripsit.

Jacosus D. DANA. Paris III. Proceedings of the American

Academy of Arts and Sciences 2, 201-220.

Haswett, W. A. (1879) On Australian Amphipoda.

Proceedings of the Linnean Society of New South Wales

4(3), 245-279.

Rurro, S. (1975) Nuovi Lisianassidi e Stegocefalidi del

Mediterraneo (Crustacea, Amphipoda). Bollettino del

Museo Civico di Storia Naturale di Verona 1 (1974),

439-453.

SCHELLENBERG, A. (1929) Abyssale und Pelagische

Gammariden. polio of the Museum of Comparative

Zoology LXIX (9), 193-201.

STEBBING, T. R. R. (1888) Report of the Amphipoda

collected by H.M.S. Challenger during the years 1873-

76. Report on the Scientific Results of the Voyage of

H.M.S. Challenger During the Years 1873-76 Zoology

29: 1737 pp. London: Eyre and Spottiswoode.

— (1897) Amphipoda from the Copenhagen

Museum and other sources. Transactions of the Linnean

Society of London Zoology 7, 25-45.

(1906) Amphipoda_ I.

~— Tierreich 21, 806p.

STEPHENSEN, K. (1925) Crustacea Malacostraca. VI.

(Amphipoda. II.). The Danish Ingolf-Expedition 3 (9),

101-178.

Gammaridea. Das

SPECIES OF THE WASP GENUS AULACUS JURINE

(HYMENOPTERA: AULACIDAE) ENDEMIC

TO SOUTH AUSTRALIA

By J. T. JENNINGS’, A. D. AUSTIN? & N. B. STEVENS?

Summary

Jennings, J. T., Austin, A. D. & Stevens, N. B. Species of the Wasp genus Aulacus

Jurine (Hymenoptera: Aulacidae) endemic to South Australia. Trans. R. Soc. S. Aust.

128(1), 13-21, 31 May, 2004.

This study deals with the South Australian species of Aulacus Jurine, a fauna that

comprises four geographically isolated, endemic species. Aulacus moerens Westwood

is redescribed, and three new species (A. belairensis, A. flindersbaudini and A. grossi)

are recognised. A diagnosis for the genus is presented, along with notes on taxonomic

history and species diversity for Australia.

Key Words: Evanioidea, Aulacidae, Aulacus, taxonomy, parasitic wasps.

Transactions of the Royal Society of S. Aust. (2004), 128(1), 13-21.

SPECIES OF THE WASP GENUS AULACUS JURINE

(HYMENOPTERA: AULACIDAE) ENDEMIC TO SOUTH AUSTRALIA.

by J. T. JENNINGS!, A. D, AUSTIN? & N. B. STEVENS?

Summary

JENNINGS, J. 'T., AusTIN, A. D. & Stevens, N. B. Species of the Wasp genus Au/acus Jurine (Hymenoptera:

Aulacidae) endemic to South Australia. Trans. R. Soc.

Aust. 128(1), 13-21, 31 May, 2004.

This study deals with the South Australian species of Aulacus Jurine, a fauna that comprises four

geographically isolated, endemic species. Aulacus moerens Westwood is redescribed, and three new species

(A. belairensis, A. flindersbaudini and A. grossi) are recognised. A diagnosis for the genus is presented,

along with notes on taxonomic history and species diversity for Australia.

Kry Worbs: Evanioidea, Aulacidae, 4u/acus, taxonomy, parasitic wasps.

Introduction

The Aulacidae are a family of parasitic wasps that

are endoparasitoids of wood-boring wasps

(Xiphydriidae) and beetles (Cerambycidae and

Buprestidae) (e.g. Carlson 1979; Gauld & Bolton

1988; Smith 2001). Although at various times the

Aulacidae have been treated as either a subfamily or

group of the Evaniidae s./, most recent studies regard

them as a distinct family (e.g. Naumann 1991;

Mason 1993; Gauld 1995; Jennings & Austin 2000;

Smith 2001).

Like other members of the Evanioidea, aulacids are

characterised by the high insertion of the metasoma

on the propodeum, but these wasps are also readily

distinguished by the presence of fore wing vein 2m-

cu (Gauld & Bolton 1988). Worldwide, the Aulacidae

comprise 156 valid species (Smith 2001), but this is

likely to be only one-third of the true size of the

group. Smith (2001) has provided a catalogue of the

world fauna, and included 48 species of Aulacus

Jurine, 106 species of Pristaulacus Kieffer, and two

species of Panaulix Benoit. Both Aulacus and

Pristaulacus are worldwide in their distribution, but

Panaulix is confined to sub-Saharan Africa. The

Australian fauna comprises 34 described species (18

Aulacus and 16 Pristaulacus) (Smith 2001; Jennings

2001), but most taxa are inadequately described and

date from the works of Kieffer (1911; 1912). No

taxonomic studies have been undertaken on the

Australian fauna since Crosskey (1953) described

one Aulacus and two Pristaulacus species, even

though modern collecting techniques, in particular

Malaise trapping, have added significantly to the

' Centre for Evolutionary Biology and Biodiversity and School of

Agriculture and Wine, The University of Adelaide, Glen Osmond, SA

5064, Australia. Email john.jennings@adelaide.edu.au

“Centre for Evolutionary Biology and Biodiversit

Farth and Environmental Science, The University of Adelaide, SA

5005, Australia.

y, and School of

number of specimens held in collections.

As part of a project that aims to revise the aulacid

fauna of Australia, this study deals with the unique and

isolated Adlacus fauna of South Australia. Aulacus

moerens Westwood is redescribed, and three new

species from the Fleurieu Peninsula, Kangaroo Island,

and the far north-west of South Australia are described.

At the same time, the genus is redescribed and notes

provided on taxonomic history and species diversity.

Materials and Methods

Specimens were observed under a Zeiss light

microscope or using scanning electron microscopy

(SEM). Specimens for SEM were first cleaned to

remove obvious dirt and other debris and examined

uncoated under a Phillips XL30 field emission SEM

at Ikv and a spot size of three.

Terms for general morphology follow Jennings and

Austin (1994), and for wing venation follow the

modified Comstock-Needham system, after Sharkey

(1988), but with some modifications, and using the

nomenclature of van Achterberg (1979) for cells.

Terms for surface sculpturing follow Harris (1979).

Where measurements are based on more than one

specimen, data are presented as the mean followed

by the range. The length of the ovipositor is

measured from the tip of the metasoma.

Abbreviations for institutions which are

repositories of the specimens referred to in this paper

are: Hope Entomological Collections, Oxford

(OXUM), South Australian Museum, Adelaide

(SAMA), and Waite Insect and Nematode

Collection, The University of Adelaide (WINC).

Systematics

Aulacus Jurine, 1807

Aulacus Jurine 1801: 163. [nom. nud. |

Aulacus Jurine 1807: 89. Type species: Aulacus

14 J.T. JENNINGS, A. D. AUSTIN & N. B. STEVENS

striatus Jurine 1807, by monotypy (North America

and Europe). — Blanchard 1840: 300; Schletterer

1889: 489; Kieffer 1903: 383, 453; Bradley 1908:

120; Kieffer 1912: 344, 370; Hedicke 1939: 17;

Townes 1950: 113; Townes 1951: 659; Oehlke 1983:

441: Koslov 1988: 243: Alekseev 1995: 39; Konishi

1990: 638; Smith 2001: 268.

Disphaeron Dahlbom 1837: 175. Type species:

Aulacus arcticus Dahlbom 1837, by monotypy. —

(syn. Hedicke 1939: 18).

Aulacinus Westwood 1868: 331. Erected as a sub-

genus of Aulacus. Type species: Aulacus (Aulacinus)

moerens Westwood 1868, by monotypy. — Kieffer

1903: 481; Bradley 1908: 120; Kieffer 1912: 349;

Hedicke 1939: 24. (syn. Townes 1950: 113).

Pammegischia Provancher 1882: 302. Type

species: Pammegischia burquei Provancher 1882, by

monotypy. — Kieffer 1903: 383; Bradley 1908: 120;

Kieffer 1912: 346; Townes 1938: 254; Hedicke 1939:

23. (syn. Kieffer 1902: 11).

Parafoenus Kieffer 1910: 350. No_ species

included. Type species: Parafoenus formosus Kieffer

1912, by subsequent monotypy. — Kieffer 1912: 345;

Hedicke 1939: 26. (syn. Townes 1950: 113).

Neuraulacinus Kieffer 1910: 350. Type species:

Neuraulacinus vespiformis Kieffer 1910, by

subsequent designation, see Kieffer 1912: 358.

(designated from three species included by Kieffer

1911). — Kieffer 1912: 358; Hedicke 1939: 26. (syn.

Townes 1950: 113).

Micraulacinus Kieffer 1910: 350. Type species:

Micraulacinus elegans Kieffer 1910, by subsequent

monotypy, see Kieffer 1912: 348. — Kieffer 1912:

348; Hedicke 1939: 24. (syn. Townes 1950: 113).

Disaulacinus Kieffer 1910: 350. Type species:

Disaulacinus flavimanus Kieffer 1911, by subsequent

monotypy, see Kieffer 1911: 224. — Kieffer 1912:

361; Hedicke 1939: 23. (syn. Crosskey 1953: 759).

Pycnaulacus Cushman 1929: 17. Type species:

Pycnaulacus brevicaudus Cushman 1929, by original

designation. — (syn. Townes 1950: 113).

Diagnosis based on Australian species

Head with or without occipital carina; frons with or

without a transverse carina above antennal sockets;

antenna 14-segmented in female, 13-segmented in

male; antennal insertions low on face, near lower

margin of eyes; eyes small, circular or subcircular,

remote from the mandibles; scape usually deeply

convex in lateral view, much thicker than pedicel and

flagellomeres; metapostnotum present between

propodeum and = metanotum as a_ distinct

sclerotisation; propodeum pyramidal, metasoma

inserted high on the apex; metasomal first tergite (T1)

and second tergite (T2) fused dorsally; hind coxa with

(Fig. 16) or without a groove or notch on inner ventral

surface, the apposed grooves or notches forming an

ovipositor guide; hind trochanter with a transverse

trochanteral groove; prefemur (trochantellus) present

(Figs 11-12); each tarsal claw with one basal tooth

(sometimes difficult to see); fore wings not plicate at

rest; fore wing vein 2m-cu present, vein 2r-m often

absent, largely spectral when present, vein 3r-m

present, often largely spectral (Figs 1, 3, 5); ovipositor

exserted, protruding well beyond apex of metasoma.

Comments

Aulacus was first erected as a genus by Jurine

(1801), but as a nomen nudum since no species were

included. In 1807, Jurine rectified this by describing

Aulacus striatus. Early authors did not define the

generic limits of Au/acus, so a number of genera were

erected (see above) without due consideration of

interspecific variation. This resulted in a proliferation

of genera: Disphaeron Dahlbom, Disaulacinus

Kieffer, and Micraulacinus Kieffer from Australia;

Neuraulacinus Kieffer from Australia and South

America; Parafoenus Kieffer from South America;

and Pycnaulacus Cushman from the USA. By 1912,

Kieffer had recognised some 41 species in six genera

and the subgenus Aalacinus Westwood. Of these, 17

species were Australian. Although Kieffer (1902) had

synonymised Pammegischia Provancher with

Aulacus, the mainly North American Pammegischia

were not transferred to Au/acus until much later

(Townes 1938). In his 1939 catalogue, Hedicke

synonymised Disphaeron and included a number of

additional species in the other genera. Townes (1950)

evaluated various characters such as wing venation,

and in the process, defined the generic limits of

Aulacus. This led him to synonymise Aulacinus,

Micraulacinus, Neuraulacinus, Parafoenus, and

Pycnaulacus with Aulacus. Townes erroneously

synonymised Disaulacinus with Aulacostethus

Philippi (now Pristaulacus). Disaulacinus was later

synonymised with Aulacus by Crosskey (1953).

At present, 48 species worldwide are included in

Aulacus; 18 species are endemic to Australia (Smith

2001; Jennings 2001). Most of the latter are from the

higher rainfall, forested areas of the east coast of

Australia, including Tasmania. Here we treat the

geographically isolated fauna of South Australia and

distinguish them from the known Australian taxa.

Because there are many undescribed species from

south-western and eastern Australia, a key to

Australian species would be premature.

Aulacus belairensis sp. nov.

(FIGS 1-2, 7, 11, 13, 16-17, 19)

Material Examined

Holotvpe

©, Belair N. P., I1-18.i1.1996, J.T. Jennings,

SAMA.

AULACUS OF SOUTH AUSTRALIA 15

Figs 1-6. Fig. 1. Right fore wing of Aulacus belairensis sp. nov. holotype female. The cells are: | = marginal, 2 =

submarginal, 3 = discal, 4 = subdiscal, 5 = costal, 6 = basal, 7 = subbasal, 8 = plical. Fig. 2. Right hind wing of Au/acus

belairensis sp. nov. holotype female. Fig. 3. Right fore wing of Aulacus flindersbaudini sp. nov. holotype male. Fig. 4.

Right hind wing of Aulacus flindershaudini sp. nov. holotype male. Fig. 5. Right fore wing of Aulacus grossi sp. nov.

holotype female. Fig. 6. Right hind wing of Aulacus grossi sp. nov. holotype female. Scale bars = | mm.

16 J, 1. JENNINGS, A. D. AUSTIN & N. B. STEVENS

Kips 7-12. Dorsal views of head. Fig. 7. Aidacus belairensis sp. nov, holotype female. Pig. 8.

lulacus flindershaudini sp-

nov. holotype male. Pig. 9. du/acus grossi sp. nov. holotype female. Pig. 10. du/acus moerens holotype female. Lateral

views of hind (rochanter and prefemur Pig. 11.

flindershaudini sp. nov. holotype male. Scale bars = 200 pum 7. 9; 500 pum & 10; 50 jum 11,

Paratypes

SA: 1 @, Belair National Park, 10-17.i11,1996, J.T,

Jennings, WINC; | 2, Belair National Park,

xii. 1996, J.T. Jennings, WINC.

Female

Length: 5.0 mm (4.4 — 5.5 mm), excluding

ovipositor.

Colour: Head orange, with variable amounts of

dark brown around ocelli; scape and pedicel orange

with flagellomeres dark brown; propleuron dark

tulacus belairensis sp. nov. holotype female. Pig. 12. dulacus

12

brown, body black except for variable amounts of

orange on pronotum; legs predominantly light to

dark brown; metasoma dark brown: ovipositor

sheaths black, ovipositor orange; wings fuscous,

darker apically on the marginal and submarginal

cells.

Tlead: 1.2 (1.0 — 1.3) x wider than long when

viewed dorsally; face rugose, with a few large

punctures near eye margin, pubescence short; frons

without transverse carina above toruli, rugose, with a

few punctures near eye margin, pubescence short;

IWLACUS OV SOUTH AUSTRALIA \7

Figs 13-18. Fig. 13. Mandibles and elypeus of sidacus belairensis sp, nov. holotype female. Fig. [4, Dorsal view of

mesoscutuim, scutellum and axillae of dadacus belairensis sp. nov. holotype female. Fig. 15, Dorsal view of mesoscutum.

scutellum and axillae of dudacis flindersbaudini sp. nov. holotype male. Fig, 16. Ovipositor guide on hind coxae of

Aulacuy helairensis sp. nov. holotype female. Fig. 17. Dorsal view of metasomal Tt and T2 of dalacuy hekrirensis sp.

noy, holotype female. Fig. 18. Dorsal view of metasomal TH and T2 of -liddcus grossi sp. nov. holotype female, Scale

bars = 200 pom 13,17, 18: 500 pam T4153 100 pam 16.

vertex and gena punctate-imbricate, a few radiating

striations near ocelli, with scattered short setae (Hig

7); posterior margin of head not concave in dorsal

view; malar space 0.25 x height eye; clypeus 0.37 x

as wide as high, margin sinuate with small medial

process (Fig. 13); mandibles broad, with two large

medial teeth (Fig. 13); distance from lateral ocellus

to eye margin 0.83 (0.78 — 0.90) x distance between

lateral ocelli; scape 1.6 (1.6 — 1.7) x length pedicel;

first Magellomere 1.0 (0.9 — 1.1) x as long as scape,

equal to length second flagellomere.

Mesosoma. Propleuron rugulose, pubescence long,

ventro-lateral carina weak; pronotum rugose;

mesoseutum in lateral view rounded antero-dorsally,

medial and lateral lobes rugose-punctate, with

scattered setae, admedial lines present (Fig. 14):

scutellum and axillae rugose-punctate (Fig. 14):

mesepisternum reticulate, with short pubescence,

mesepimeron broad, scrobiculate; metapleuron

rugose, with short pubescence; propodeum rugose,

Is J.T. JENNINGS, A. D. AUSTIN & N. B. STEVENS

Autacus belairensis

A, flindersbaudini

A, grossi

A. moerens

oreo

Fig. 19. Distribution of South Australian Au/acus spp.

posterior margin scrobiculate; hind coxa with

ovipositor guide ventrally on inner margin (Fig. 16);

hind trochanter and femur imbricate, pubescence

short; prefemur somewhat indistinct (Fig 11); hind

tibia imbricate, pubescence short, with scattered

emergent stout setae; hind femur 0.70 (0.68 — 0.72) x

length hind tibia; hind tibia with ventro-apical pecten

of short robust spines; hind tarsal segments 1-4 with

ventro-apical pecten of short robust spines, segment

1, 2.8 x length segment 2; segment 2, 1.4 x length

segment 3; segment 3, 1.6 x length segment 4;

segment 4, 0.6 x length segment 5; hind tarsal claw

0.5 x length segment 5; fore wing vein 2r-m and 3r-

m largely spectral (Fig. 1); hind wing with 3 hamuli

(Fig. 2).

Metasoma. Clavate, 1.67 (1.54 — 1.71) x length of

mesosoma; Tl and T2 anteriorly narrow, smooth

dorsally except for a few strigate wrinkles anteriorly

on Tl; ovipositor 5.9 (5.0 — 6.5 mm) mm.

Male

Unknown.

Remarks

This species is named after the holotype locality,

Belair National Park, South Australia (Fig. 19). All

specimens were collected by Malaise trap, and

nothing is known of its biology. This species can be

separated from other South Australian species by the

fuscous wings that are darker apically on the

marginal and submarginal cells, and the strigate

wrinkles anteriorly on metasomal Tl. Also, it is

similar in size and appearance to 4. e/egans (Kieffer)

which has been collected from several localities in

New South Wales and Victoria. The two can be

easily separated in that 4. elegans has a shiny head,

an orange metasoma, and has different sculpturing

patterns; for example, the vertex is largely smooth

with just a few scattered shallow punctures.

Aulacus flindersbaudini sp. nov.

(FIGS 3-4, 8, 12, 15, 19)

Material Examined

Holotvpe

3d, West Bay, Flinders Chase Nat. Park, Kangaroo

Is. S.A., 1.1986, A.D. Austin, SAMA. [Flagellomeres

2-11, right hind tarsal segments 4-5, and claw

missing].

Male

Length: 8.0 mm.

Colour: Body red except metasomal TI dark

brown dorsally; flagellomeres 2 and 3 black; wings

hyaline; fore wing with brown spot apically on the

marginal and submarginal cells.

Head: |.6 x wider than long when viewed dorsally;

face punctate-imbricate, pubescence short; frons

without transverse carina above toruli, areolate-

rugose, pubescence short; vertex areolate-rugose

medially, with a few radiating striations near ocelli,

with a few transverse striations posteriorly, punctate-

imbricate laterally, pubescence short (Fig. 8); gena

punctate-imbricate, with scattered short setae;

posterior margin of head slightly concave in dorsal

view; malar space 0.34 x height eye: clypeus 3.7 x as

wide as high, margin sinuate with small medial

process; mandibles broad, with two large medial

teeth; distance from lateral ocellus to eye margin

equal to distance between lateral ocelli; scape 1.8 x

length pedicel; length first flagellomere equal to

length scape, second flagellomere missing.

Mesosoma. Propleuron smooth anteriorly to

weakly rugulose posteriorly, with scattered long

setae: ventro-lateral carina weak; pronotum

rugulose; mesoscutum in lateral view rounded

antero-dorsally, medial and lateral lobes strigate with

scattered short setae, admedial lines present (Fig.

15); scutellum and axillae strigate (Fig. 15);

mesepisternum —areolate-rugose, with — short

pubescence; mesepimeron broad, scrobiculate:

metapleuron areolate-rugose, with short pubescence;

propodeum areolate-rugose, posterior margin

scrobiculate; hind coxa without groove ventrally on

inner margin, weakly rugulose to strigate dorsally,

pubescence short laterally; hind trochanter imbricate,

pubescence short; prefemur indistinct (Fig 12); hind

femur imbricate, pubescence short; hind tibia

imbricate, pubescence short, with scattered emergent

stout setae; hind femur 0.77 x length hind tibia; hind

tibia with ventro-apical pecten of short robust spines:

hind tarsal segments 1-4 with ventro-apical pecten of

AULACUS OF SOUTH AUSTRALIA 19

short robust spines, segment |, 2.0 x length segment

2; segment 2, 1.6 x length segment 3; segment 3, 2.4

x length segment 4; segment 4, 0.8 x length segment

5: hind tarsal claw 0.7 x length segment 5; fore wing

vein 2r-m incomplete, evident only by a small node

on 2r, 3r-m entirely spectral (Fig. 3); hind wing with

2 hamuli (Fig. 4).

Metasoma. Clavate, 1.5 x length of mesosoma; T1

and T2 dorsally narrow, smooth; digitus about same

length as basiparameres.

Female

Unknown.

Remarks

Aulacus flindershaudini is named to commemorate

the meeting of Matthew Flinders and Nicolas Baudin

at Encounter Bay, South Australia in 1802, This

species is known only from the holotype location

(Fig. 19), and nothing is known of its biology. This

species is readily distinguished from other South

Australian species by the presence of a brown spot

apically on the marginal and submarginal cells of the

fore wing (Fig. 3); this spot is absent in the other

species. Aulacus flindersbaudini also lacks the

eroove or ovipositor guide on the inner margin of the

hind coxae. The brown spot on the fore wing is found

in several species from south-western and eastern

Australia, such as 4. pallicaudus (Cameron), but all

of these species are largely black.

Aulacus grossi sp. Nov.

(FIGS 5-6, 9, 18-19)

Material Examined

Holotype

9°. nr. Victory Well, Everard Pk Stn, S. Aust. 2-

4.xi.1970. G. Gross, SAMA.

Female

Length: 4.5 mm, excluding ovipositor.

Colour: Head black; mesosoma, antennae, and

legs light brown; metasoma black except for basal

part of metasomal T1; wings hyaline.

Head: \.\ x wider than long when viewed dorsally:

face, frons without transverse carina above toruli:

frons and vertex rugose-punctate (Fig. 9), with

scattered short setae, a little denser on face; gena

smooth, almost glabrous; posterior margin of head

not concave in dorsal view; malar space 0.16 x

height eye; clypeus 4.0 x as wide as high, punctate,

margin sinuate with small medial process; mandibles

broad, with two large medial teeth; distance from

lateral ocellus to eye margin 0.72 x distance between

lateral ocelli; scape 1.8 x length pedicel; first

flagellomere 0.72 x as long as scape, 0.52 x as long

as second flagellomere.

Mesosoma. Propleuron weakly rugulose-punctate,

almost smooth, with a few scattered setae; ventro-

lateral carina weak; pronotum rugose-punctate;

mesoscutum in lateral view rounded antero-dorsally,

medial and lateral lobes rugose, almost strigate, with

scattered short setae, admedial lines distinct:

scutellum and axillae rugose, almost strigate;

mesepisternum rugose-punctate, with — short

pubescence; mesepimeron broad, scrobiculate;

metapleuron rugose-punctate, with short pubescence;

propodeum rugose, with several pronounced carinae,

posterior margin scrobiculate; hind coxa with groove

ventrally on inner margin, weakly strigate dorsally,

pubescence short; hind trochanter and hind femur

imbricate, pubescence short; hind tibia imbricate,

pubescence short, with scattered emergent stout

setae; hind femur 0.8 x length hind tibia; hind tibia

with ventro-apical pecten of short robust spines; hind

tarsal segments 1-4 with ventro-apical pecten of short

robust spines, segment 1, 2.9 x length segment 2;

segment 2, 1.35 x length segment 3; segment 3, 2.0 x

length segment 4; segment 4, 0.5 x length segment 5;

hind tarsal claw 0.5 x length segment 5; fore wing

veins 2r-m and 3r-m largely spectral (Fig. 5); hind

wing with 2 hamuli (Fig. 6).

Metasoma. Clavate, equal in length to mesosoma;

Tl and T2 dorsally broad, smooth (Fig. 18);

ovipositor 4.2 mm.

Male

Unknown.

Remarks

Aulacus grossi has a broad metasoma when viewed

dorsally which distinguishes it from both 4.

belairensis and A. flindersbaudini. It is also the

smallest Australian species of Aulacus known, and

can be distinguished from all other Australian

species by its colour pattern, i.e. head and metasoma

black, and mesosoma light brown. Au/acus grossi is

known from a single specimen from Everard Park

Station in the north-west of South Australia (Fig. 19),

and is named after the collector, Dr Gordon Gross,

formerly Curator of Entomology, South Australian

Museum. Nothing is known about its biology.

Aulacus moerens Westwood

(FIGS 10, 19)

Aulacus (Aulacinus) moerens Westwood 1868:

331. — Westwood 1874: 129; Schletterer 1889: 517.

Aulacinus moerens — Kieffer 1902: 12; Kieffer

1903: 481: Kieffer 1904: 6; Kieffer 1912: 349, 356;

Hedicke 1939; 25,

Aulacus maerens — Dalla Torre

(unnecessary emendation)

Aulacus moerens — Smith 2001: 273.

1902: 106]

20 J.T. JENNINGS, A. D. AUSTIN & N. B. STEVENS

Material Examined

Holotype

2, Adelaide [illeg.] 1865, OXUM. Antennae

missing, head glued to mesosoma, metasoma glued

to card, ovipositor sheaths missing.

Female

Length: 10.5 mm, excluding ovipositor.

Colour: Black, except tibiae and tarsi largely light

brown, ovipositor light brown; wings hyaline.

Head: |.18 wider than long when viewed dorsally;

face rugulose, pubescence long; frons with

pronounced transverse carina above toruli, rugulose,

with long pubescence; vertex punctulate-imbricate,

slight rugosity posteriorly near occiput, with short

scattered setae (Fig. 10); gena punctulate-imbricate,

with scattered short setae; posterior margin of head

not concave in dorsal view; malar space ().2 x height

eye; clypeus 4.0 x as wide as high, margin sinuate;

mandibles broad, with two large medial teeth:

distance from lateral ocellus to eye margin 0.9 x

distance between lateral ocelli; antennae missing.

Mesosoma. Propleuron rugulose, pubescence long,

ventro-lateral carina present; pronotum without

angular process, rugose; mesoscutum in lateral view

angular antero-dorsally, medial and lateral lobes

strigate, with scattered short setae, admedial lines

present; scutellum — strigate; axillae rugose;

mesepisternum rugose, with long pubescence;

mesepimeron broad, scrobiculate; metapleuron

rugose, with short pubescence; propodeum rugose,

with medial dorso-ventral scrobiculate groove,

posterior margin scrobiculate; hind coxa without

groove ventrally on inner margin, strigate dorsally,

pubescence long laterally; hind trochanter

punctulate-imbricate, pubescence short; hind femur

imbricate, pubescence short; hind tibia imbricate,

pubescence short, with scattered emergent stout

setae; hind femur 0.74 x length hind tibia; hind tibia

without ventro-apical pecten of short robust spines;

hind tarsal segments 1-4 with ventro-apical pecten of

short robust spines, segment |, 2.7 x length segment

2; segment 2, 1.6 x length segment 3; segment 3, 2.0

x length segment 4; segment 4, 0.4 x length segment

5; hind tarsal claw 0.6 x length segment 5; fore wing

vein 2r-m pale brown, tubular, 3r-m spectral in

middle half; hind wing with 3 hamuli.

Metasoma. Ovate, 1.25 x length of mesosoma; T1

and T2 smooth except slight rugosity medially;

ovipositor 4.0 mm.

Male

Unknown.

Remarks

Although both 4. grossi and A. moerens have an

ovate metasoma, the latter species is a much larger

and differs in a number of ways, including its largely

black colour, the lack of an ovipositor guide on the

hind coxae, and the presence of a small medial

process on the clypeus. It does not resemble any

other Australian species. This species is known only

from the holotype locality, Adelaide, South Australia

(Fig. 19), and nothing is known of its biology.

Acknowledgments

We thank the curators of the collections listed

above for the loan of specimens, and the staff of

Adelaide Microscopy, The University of Adelaide,

for access to the FESEM. This work was carried out

with the assistance of an Australian Biological

Resources Study grant and a Small ARC grant to JTJ

and ADA.

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the author” (Oxford).

THE REPRODUCTIVE ECOLOGY OF TWO NATURALISED

ERICA SPECIES (ERICACEAE) IN THE ADELAIDE HILLS:

THE RISE AND FALL OF TWO ‘WOULD-BE’ WEEDS?

By D. TURNER’? & J. G. CONRAN’?

Summary

Turner, D. & Conran, J. G. (2004). The reproductive ecology of two naturalised Erica

species (Ericaceae) in the Adelaide Hills: the rise and fall of two ‘would-be’ weeds?

Trans. R. Soc. S. Aust. 128(1), 23-31, 31 May, 2004.

The pollination biology and reproductive ecology of the newly naturalised species

Erica cinerea from Europe and E. glandulosa from South Africa (Ericaceae) are

reported from the Adelaide Hills from surveys undertaken in 1995 and 2003.

Examination of UV floral reflectance, nectar composition and floral visitors found

that E. cinerea was honeybee pollinated (although the bees also practiced nectar theft

by chewing through the corolla tube), whereas E. glandulosa was probably pollinated

by New Holland Honeyeaters (Phylidonyris novaehollandiae: Meliphagidae) which

filled the niche of the nectariniid Sunbirds which pollinate it in South Africa.

Key Words: Erica, E. cinerea, E. glandulosa, Ericaceae, weed biology, reproductive

ecology, pollination, fecundity, Australia.

Transactions of the Roval Society of S. Aust. (2004), 128(1), 23-31.

THE REPRODUCTIVE ECOLOGY OF TWO NATURALISED

ERICA SPECIES (ERICACEAE) IN THE ADELAIDE HILLS:

THE RISE AND FALL OF TWO ‘WOULD-BE’? WEEDS?

by D. TurNER!? & J. G. CONRAN!?

Summary

TURNER, D. & Conran, J. G. (2004). The reproductive ecology of two naturalised Erica species (Ericaceae) in

the Adelaide Hills: the rise and fall of two ‘would-be’ weeds? Trans. R. Soc. S. Aust. 128(1), 23-31, 31 May,

2004.

The pollination biology and reproductive ecology of the newly naturalised species Erica cinerea from Europe

and E. glandulosa from South Africa (Ericaceae) are reported from the Adelaide Hills from surveys undertaken

in 1995 and 2003. Examination of UV floral reflectance, nectar composition and floral visitors found that

EF. cinerea was honeybee pollinated (although the bees also practiced nectar theft by chewing through the corolla

tube), whereas £. glandulosa was probably pollinated by New Holland Honeyeaters (Phylidonyris

novaehollandiae: Meliphagidae) which filled the niche of the nectarintid Sunbirds which pollinate it in South

Africa. Although fecundity data in 1995 suggested that there was considerable capacity for these plants to

become weeds, the 2003 survey found that they had declined significantly, apparently as a result of competition

or shading from taller, more aggressive weeds, mainly Bracken (Preridium esculentum) and Tree Erica (E.

arborea). Nevertheless, as £. cinerea reseeds and £. glandulosa resprouts after fire, and £. cinerea can persist

in the soil seed bank for over 30 years, their populations may recover and/or expand if the area where they are

growing 1s opened up by fire.

Key Woros: Erica, FE. cinerea, E. glandulosa, Ericaceae, weed biology, reproductive ecology, pollination,

fecundity, Australia.

Introduction

Erica L. is a large genus of heaths which is centred

around South Africa (Guthrie & Bolus 1905; Baker

& Oliver 1967), with a second radiation in southern

and western Europe (Webb & Rix 1972). Much of

Southern Africa and Mediterranean Europe is

characterised by cool winters and hot dry summers

similar to South Australia and most members of the

genus grow in nutrient poor, acidic soils. The

European species, in particular, tend to be

mycorrhizal, and the majority are calcifuges

(Gimingham ef a/. 1979; Webb 1986). In many

respects the soils of the South African Cape,

Mediterranean Europe and South Australia are

similar in being nutrient poor, often acidic, and

support conspicuous sclerophyllous heath elements,

represented in South Africa and Europe by the

Ericaceae and in Australia by Epacridaceae (Specht

1979, 1981).

There are three Erica spp. listed as naturalised in

South Australia (Jessop & Toelken 1986): the

European £. arborea L. and E. lusitanica Rudolphi,

and the South African E. baccans L. Of these, the

two European species, and £. arborea in particular,

' Centre for Evolutionary Biology and Biodiversity, Environmental

Biology, School of Earth and Environmental Sciences, Darling

Building DP418, The University of Adelaide, SA 5005.

° Present address: SARDI, SA Aquatic Sciences Centre, 2 Hamra

Ave, West Beach, SA 5024.

‘Correspondence author: john.conran@adelaide.cdu.au.

are widespread weeds on nutrient-poor, acidic soils

with moderate to high rainfall (Cshures & Edwards

1998; Ojeda 1998). Erica spp. tend to coincide

growth with water availability during winter and

early spring (Gehrig 2000), flowering from late

winter into spring and summer (Jessop & Toelken

1986). However, in 1995 populations of an

additional two species (the European Bell Heather £.

cinerea L. and South African £. glandulosa Thunb.)

were found in a small area of disturbed remnant

bushland bordering Mount Lofty House on the north-

western boundary of Mount Lofty Botanic Gardens.

Erica cinerea from temperate, western Europe has

numerous ornamental cultivars grown in southern

Australia (Spencer 1997) and is a temperate, severe

cold-tolerant, generally summer-flowering species

from low, open heaths on acidic soils (Rozé 1993;

Ojeda et al. 1998; Leith et al. 1999). In contrast, E.

glandulosa is a long-lived, frost-tolerant, year-round

but mainly autumn-flowering, species from tall

fynbos on well-drained soils in the southern Cape

region of South Africa between Mosselbaai and Port

Elizabeth (Baker & Oliver 1967). E. glandulosa is

cultivated as an ornamental shrub in South Africa

and Europe (Baker & Oliver 1967; Nelson 1997),

and although not currently listed as grown in

Australian gardens (Spencer 1997), is naturalised in

Australia, but with no locality or weed status details

(AFFA 2001).

As these two species seem only to have naturalised

recently in the Adelaide Hills, the populations were

24

Spencer

Gulf

ie

ep art km / Fleurieru \ é

Mount

Lofty

Botanic

Gardens

meee

<

wee

a

*

v

on

e

¢

250m

Fig. 1. Mount Lofty Botanic Gardens site showing the

locality of the Erica cinerea and E. glandulosa colonies.

relatively small and confined to the one area.

However, as they are morphologically dissimilar and

come from different environments they may have

different establishment strategies. Accordingly, the

two aims of this study were:

To compare the reproductive biology of the two

species to assess how their strategies might relate to

any success or failure to establish.

To examine their population structure with

particular emphasis on their potential fecundity, and

to see if expansion had occurred after eight years.

Methods

The site

Mt Lofty Botanic Gardens is a 97 ha reserve

located in the Adelaide Hills 13 km SE of Adelaide

(34° 58’ S, 138° 42’ E, 670 m altitude) combining

). TURNER & J. G. CONRAN

extensive planted areas of exotics with a flora

reserve of native Eucalyptus obliqua L’Hér. forest

with sclerophyllous understorey (Haegi & Morley

1991). The area surveyed represents an

approximately 10 x 100 m area of disturbed remnant

eucalypt forest located on the NW side of the

Gardens, close to the boundary fence with the

grounds of Mount Lofty House (Fig. 1). The site

understorey is a mixture of Epacridaceae, mainly

Pink Ground Berry (Acrotriche fasciculiflora

(Regel) Benth.) and Native Heath (Epacris impressa

Labill.), other native shrubs, bracken fern (Preridium

esculentum (G. Forst.) Cockayne) and introduced

shrubby weeds including Spanish Broom (Cy/iszs

scoparia (L.) Link) and Tree Erica (Erica arborea

Lai

Floral advertisement and rewards

Advertisement for pollinators can involve several

simultaneous stimuli including visual, olfactory, and

short range tactile cues, and may be considered as

any stimulus which attracts visitors to a flower

(Facgri & van der Pijl 1979).

Visual stimulus is a combined effect of the target

colour (including combinations of colour and

pattern) and the size shape of the target. Of primary

importance is the pollinator’s visual spectrum (the

visual spectrum of insects differs from that of

humans) which affects the contrast between the

flower and the background foliage (Dafni 1992;

Dyer 1996). Flowers of the two species were

photographed using Kodak P-3200 B&W film

bracketted at f8-f22 with and without a UV

transmitting filter (Hoya U360) by a Pentax MEII

Super with macro lens and G30-rated flash. The film

was developed following — manufacturer’s

instructions for 10 minutes at 21°C, allowing

visualisation of the flowers from the perspective of

different potential pollinators.

Because floral scent may be ephemeral, flowers

were sealed in 30 mm glass vials for two hours in a

warm, well-lit location and the vials were then

opened and smelt (Bernhardt 1995). In addition,

flowers were stained in 1% neutral red for 2 hours

and rinsed in distilled water for 18 hours to detect

possible osmophore-bearing regions (Datni 1992;

Bernhardt 1995). Nectar volume was measured using

micro-sampling pipettes and electronic balances,

with ten flowers from each of the two species

sampled at various times of the day to allow for

temporal variation in nectar production. Nectar

composition of the two species was examined using

thin layer paper chromatography. Nectar from five

flowers of each species was extracted by washing

with 5 wl of deionised water, and spotted onto a

sheet of Whatman’s No. | chromatography paper.

Sugar standards were also added for glucose,

ECOLOGY OF TWO ERICA SPP. IN THE ADELAIDE HILLS 25

sucrose, fructose, maltose, and galactose and the

chromatogram run using a solution of 12:3:5 n-

butanol: acetic acid: water. After 24 hours, the paper

was removed from the solvent, dried, sprayed with

an indicator solution of 1% aniline, 1%

diphenylamine and 4% phosphoric acid in acetone

and baked at 85°C for five minutes (Neimietz &

Hawker 1988). The relative intensity of each nectar

component was coded according to the nectar

classifications of Percival (1961) and Baker & Baker

(1983).

Floral visitor behaviour was observed between 9

am and 4 pm (opening hours at Mt Lofty Gardens)

over several days in Feb.-Mar. 1995, Sept. 2002 and

Feb.-Mar. 2003 with observations mainly between 9

am and | pm, as pollinator activity was minimal in

the afternoon. Birds were monitored from about 10

m using binoculars, and insects were netted, killed in

jars with ethyl acetate fumes, examined under a

binocular microscope and then washed briefly on a

microscope slide with drops of absolute ethanol to

remove any pollen. The resulting residue was

mounted in a drop of 2.5% saturated aqueous Methyl

Green and 2% saturated aqueous Phloxine in phenol

glycerine jelly (Owezarzak 1952), and pollen

identified by light microscopy.

Population structure and fecundity

Numbers of individuals of each of the two species

were counted in order to obtain a total population

count for the infested area. Twenty individuals of

each species were then chosen at random, and the

following information recorded for each:

Number of shoots per plant

Number of inflorescences on each shoot

Number of flowers on each inflorescence

Number of ovules per ovary

Number of seeds per ovary

Results

Floral advertisement and rewards

Erica glandulosa is an erect species >] m tall with

large, orange-pink flowers in the leaf axils (Fig. 2B)

which are classed as tubular-curved (Rebelo ef al.

1985) whereas FE. cinerea is a ground hugging shrub

with inflorescences of small violet-purple flowers

(Fig. 2C) which are classified as urceolate (Rebelo ef

al. 1985).

The violet-purple flowers of E. cinerea create an

obvious contrast between the flower and_ its

background (Fig. 3A). The corollas were classed as

UV-light (pale) (Dafni 1992), with reflected shorter

wavelength light making them high-contrast to the

foliage. Reflectance was highest in buds and newly-

opened flowers, with older (possibly no-longer

receptive) flowers being less reflective and more like

the foliage.

Erica glandulosa has orange-pink flowers on a

green background making for high visual contrast,

but as the flowers reflect wavelengths outside the

normal insect visual spectrum, they are likely to be

low contrast to an insect (Richards 1986; Dyer

1996). The UV photographs showed low contrast

between the flowers and foliage, although again

newly-opened flowers were more reflective than

older ones, and the lobes of recently-opened flowers

(which visually showed a light greenish tinge),

tended to be more UV reflective.

No obvious scent or evidence of osmophores was

found in E. glandulosa suggesting that the plant’s

advertisement strategy does not involve scent.

Although no obvious odour was detected in F.

cinerea, two possible scent producing areas were

identified. The edges of the sepals stained darkly and

microscopic examination revealed specialised

tissues of uncertain function, and the toothed anther

appendages also stained heavily.

Erica glandulosa contained a mean of 0.5+0.2 ul

of nectar; whereas although minute droplets were

noticed on the annular nectary below the ovary in E.

cinerea, the volume was too minute to measure

accurately. Chromatography of FE. glandulosa nectar

revealed strong spots for glucose and fructose, with

a weak spot for sucrose making the species SFG

using the nectar composition coding system of

Percival (1961) and hexose-rich following Baker &

Baker (1983), whereas E. cinerea possessed all three

sugars, but with a slight predominance of sucrose

(SFG), thereby making it sucrose-rich.

Flowers of £. glandulosa were visited in the

mornings by New Holland Honeyeaters

(Meliphagidae: Phylidonvris —novaehollandiae

(Latham, 1790). These perched along the erect stems

approaching the flowers from below, their curved

beaks enabling them to access easily the slightly

downward-curved, tubular flowers. The only

observed visitors to E. cinerea were honey bees

(Apis mellifera L., 1758)), and inspection of captured

visiting bees found they carried only pollen from that

species. Erica cinerea flowers were also observed to

have had the corolla tubes chewed open from the

side, suggesting that bees were also engaged in

reward theft (Fig. 2D).

Population structure and fecundity

In the 1995 survey, there were 67 mature

individuals of E. glandulosa aggregated as small

groups of 2 — 5 plants and scattered across an area of

about 1000 m’, but with no evidence of juvenile or

seedling plants in the area surveyed. In addition,

there were 1,436 mature (flowering) individuals of

E. cinerea within an area of about 80 m2, as well as

numerous seedlings. Within this, there was a smaller

26 D. TURNER & J. G. CONRAN

ECOLOGY OF TWO ERICA SPP. IN THE ADELAIDE HILLS 27

27 m2? area of more open Acrotriche fasciculiflora

and) Epacris impressa heath on the eastern

(roadward) side which contained about 50 mature £.

cinerea m?, and up to about 100 seedlings m?

whereas the surrounding area contained only about

four mature individuals and 10 — 20 seedlings m=. In

contrast, by 2002, whereas there were 54 mature F.

glandulosa plants still scattered over the same area,

the numbers of EF. cinerea had fallen to only 34 adult

plants in total, all within the original 27 m? area

which had previously contained the bulk of the

population (Table 1), and there was no evidence of

seedlings.

Data from the first survey suggested that the

populations are highly fecund, with numerous seeds

per individual and high total estimated annual seed

set for both taxa (Table 1). Nevertheless, in 2003 not

only had populations of both species declined

significantly, but fecundity was also reduced, mainly

through fewer flowers and inflorescences per plant,

resulting in a dramatic decrease in estimated total

seed set for both species at the colony level.

Discussion

The area that the two species occupy is in the

north-western corner of the Botanic Gardens and

although part of a native flora reserve, is directly

below Mount Lofty House and was previously part

of those gardens. Possible scenarios for their initial

establishment are that these two Erica spp. were

planted on the bank at some time in the past and have

persisted as a remnant population, or that they spread

from the original gardens into the surrounding

bushland.

Evidence from pollination studies in a number of

ericaceous genera and Erica spp. in_ particular

suggest that they are predominantly self-sterile

(Anderson ef al. 2000; Ng & Corlett 2000) or with

limited self-compatibility (Aparicio & Garcia-

Martin 1996; Santandreu. & Lloret 1999).

Nevertheless, F. cinerea is regarded as both self-

and cross-pollinating (Knuth 1909; Bannister 1965),

with the slightly protogynous flowers effecting

pollen transfer before the buds open (Hagerup

1951). The pollination biology of the two species

studied here shows the different strategies in insect-

and bird-pollinated Ericaceae. That pollination is

successful at Mount Lofty is reflected in the high

levels of seed set, and the plants have clearly

benefited from the adaptation of local pollinators to

a new food source. This might also explain the

scarcity of nectar in the SA flowers despite reports

of copious nectar in European studies (Bannister

1965).

Erica cinerea with relatively small, pale-purple,

urceolate flowers falls into the entomophilous

syndrome common in the genus (Knuth 1909;

Rebelo ef al. 1984; 1985). Insect visitors to

European Erica spp. include honeybees,

bumblebees, flies, moths and butterflies (Knuth

1909; Aparicio & Garcia-Martin 1996; Santandreu

& Lloret 1999) and Knuth (1909) listed a wide range

of insects visiting £. cinerea, and reported corolla

chewing and nectar theft by bumblebees. He also

noted that pollination in this species was identical to

that of E. tetralix L., where honeybees chew through

the floral tubes because their c. 6 mm long

mouthparts were too short to reach the nectaries of

average-sized flowers (7 mm — the same as E.

TABLE |. Summary of fecundity parameters for the two Erica spp. naturalised at Mt Lofty.

E. cinerea

E. glandulosa

1992 2003 1992 2003

Parameter n Mean+SD Mean+SD tor x? Mean+SD Mean+SD _ t or x?

Plants per colony 1,436 34 1,428.0" 67 54 2.5 ns

Main stems per plant 20 5.0+1.7 4.741.9 0.7 ns 5.2+6.9 5.0+£7.3 0.2 ns

Inflorescences per plant 20) -18.8+8.2 10.3412.2 8.4 29.44+35.5 21.5+40.1 4.0"

Inflorescences per main stem 20 = 3.9+1.5 2.6+1.8 3.2" 6.2+1.1 4141.8 33"

Flowers per inflorescence 100 6.6+4.8 6.2+5.7 2.5" 3.2+1.0 3.041.2 1.3 ns

Ovules per ovary 10 24.041.4 22.8+2.1 2.028 ns 708.0+122.3 692.9+173.1 1.7 ns

Seeds per ovary 10 3.41.4 3.141.6 O5ns 215.0452.5 208.2+74.6 1.9 ns

Seeds per plant (estimated) 422 153 n/a 20,227 13,430 n/a

Total seeds per year (estimated) 605,992 5,202 n/a 1,355,209 725,220 n/a

‘counts compared using x?

Fig. 2. Erica cinerea and E. glandulosa habitat and habit. A. Habitat of Eucalyptus obliqua forest with sclerophyllous shrub

and bracken understorey. B. Erica glandulosa habit. C. E. cinerea habit. D. E. cinerea flower showing damage by nectar

thieves.

28 D. TURNER & J. G. CONRAN

Fig. 3. Photographs of the two Erica spp. under visible (A, C) and UV (B, D) light. A-B. Erica cinerea. C-D. E. glandulosa.

cinerea). Nectar composition was similar to other

European Erica species listed by Percival (1961) as

visited extensively by honeybees, and the sucrose-

rich nectar agrees with Baker & Baker’s (1983)

observation that bee-flowers with floral tubes > 6

mm are mainly “long-tongued” bee-flowers and

generally sucrose-rich. Most Erica spp. are

generally considered to be scentless, although there

is a number of species reported with marked honey

scents (Rebelo e/ a/. 1985). The anther appendages

in £. cinerea are thought to position the anthers for

entomophilous pollen delivery (Knuth 1909; Rebelo

et al. 1985) but the presence of apparently glandular

structures which stain strongly with neutral red

suggests that they may also serve as osmophores,

and warrants further investigation.

In contrast, £. glandulosa has orange-pink,

tubular-curved, scentless flowers with copious

nectar (Rebelo ef a/, 1984; 1985), and the hexose-

dominated nectar in E. glandulosa reflected the

results found by Barnes ef a/. (1995) for several

ornithophilous species of Erica sect. Evanthe. The

members of sect. Evanthe are thought to have co-

evolved for pollination with nectariniid Sunbirds

ECOLOGY OF TWO ERICA SPP, IN THE ADELAIDE HILLS 29

(Scott-Elliot 1890; Rebelo ef al. 1984; 1985), which

fill the same ecological niche as the honeyeaters

seen visiting it in our study, although there is

evidence also of pollination of some Evanthe spp. in

South Africa by long-proboscid Nemestrinid flies

(Manning ef al. 1999), Although, the assertion by

Baker & Baker (1983) that hexose-rich bird flowers

are associated with passerines must be treated with

some caution, given the dominance of sucrose-rich

to sucrose-dominant Evanthe spp. reported by

Barnes ef al. (1995), numerous South African

passerine-pollinated flowers do produce high

volumes of very dilute, hexose-dominated nectar

(Nicolson 2002). The orange-pink flowers showed

only limited UV-reflectance, suggesting that they

are not strongly adapted for insect pollination

(Richards 1986; Dyer 1996), although tubular red

flowers can be attractive to long-proboscid flies

(Manning ef a/. 1999). The glandular hairs which

cover all parts of this species except the corolla

possibly help to deter would-be insect nectar

thieves, similar to those ornithophilous Erica spp.

with externally viscid corollas (Scott-Elliot 1890;

Rebelo ef al. 1985). Similarly, the presence of only

anatomically rudimentary anther appendages in this

species (Palser & Murty 1967; Hermann & Palser

2000) supports the assertion by Knuth (1909) and

Rebelo e¢ a/. (1985) that they are involved in

presenting the anthers primarily for entomophilous

pollination. Nevertheless, if there is an anther

appendage - osmophore connection, the loss of

appendages in E. glandulosa would further support

pollination by passerines with a poor sense of smell.

What is most apparent from this study is that

despite initial apparent success, with high levels of

both numbers of individuals and/or annual

fecundity, neither species has managed to spread; in

fact both have declined in terms of above ground

biomass. This is particularly the case for FE. cinerea

where this previously abundant plant is now very

scarce at the site, and the remaining plants are

smaller and with lower fecundity than in 1995. The

most likely explanation for this decline in £. cinerea

is a combination of competition (Bannister 1965)

and litter accumulation (Mallik ef a/. 1984). Shading

can determine species composition in heaths and

cause removal of understorey species (Vila &

Sardans 1999), and crowding or shading by

neighbours reduced fecundity in Erica multiflora L.

(Vila & Terradas 1995, 1998). In 1995 E. cinerea

was most abundant in a small area of relatively open

epacrid-dominated heath, but by 2003 this, like the

rest of the surveyed area, had been overgrown by

Preridium and E. arborea, the latter of which

outgrows Acrotriche fasciculiflora under dry

conditions (Gehrig 2000), Erica cinerea prefers

more temperate, mesic conditions than £. arborea

(Ojeda er al. 1998) and would be disadvantaged

under the hot dry summers seen in South Australia.

Similarly, it grows mainly in low open heaths

(Bannister 1965; Rozé 1993). whereas E. arborea

generally grows better under forest cover or shade

(Ojeda e¢ al. 2000). In contrast, E. glandulosa was

apparently able to compete better for space, being of

similar size to the invading bracken and shrubs,

although crowding may have reduced fecundity, as

seed set in resprouter Erica species is correlated

with main shoot size and vigour (Riba 1998).

Establishment in £. cinerea is mainly by seed, and

flowering is improved by higher nitrogen levels

(Leith ef al. 1999), although seed set and

germination rate drops with population age (Mallik

et al. 1984). Germination occurs in the presence of

light, with 20 — 40% of the seeds germinating the

season after they are shed, nevertheless, heat shock

substantially increases germination rates (Bannister

1965), whereas litter accumulation inhibits

germination and seedling survival (Mallik e7 al.

1984). Erica cinerea grows preferentially in short

open heath, and recovers more slowly to from fire

than taller heaths (Rozé 1993), although its seeds

can survive for at least 30 — 40 years in the soil

seedbank (Mallik ef a/. 1984; Thompson & Band

1997). Erica seeds in the upper 2 — 5em soil depth

tend to survive fire better than many other

Mediterranean woody heath species (Ferrandis ef al.

1999), but although £. cinerea can produce up to a

million seeds m~, there is seedbank mortality of 20

— 40% after the first year, as well as low seedling

survival (Webb 1986).

In contrast, F. glandulosa is a resprouter (Ojeda

1998) — an unusual condition amongst Cape Erica

species — although germination in many Cape Erica

spp. is also smoke-triggered (Brown ev al. 1993),

and it is possible that the absence of seedlings of this

species, despite the large numbers of seeds

produced, may be related to the lack of a suitable

trigger, and/or the fact that many fire-dependent

heath species do not germinate well under

conditions of high litter and standing biomass (Rozé

1993: Lloret & Vila 1997). The breeding system of

this species is as yet unknown, and further studies

would be useful to determine whether it is cross-,

self-pollinated or both, as this has implications for

the ability of isolated individuals to set seed and

spread further.

In conclusion, the evidence suggests that although

the populations of these two species appear to be in

decline, despite their initial success and apparently

high potential fecundity, were a fire or other

disturbance to alter the conditions at the site, they

may well be able to recover or expand, via the

seedbank, representing a future weed problem

awaiting an environmental trigger. However, as

30 D. TURNER & J. G. CONRAN

bracken and £. arborea also recolonise quickly after

fire or other disturbances, and frequent, repeated

aerial biomass removal of the latter is needed to

reduce its aggressive regrowth capacity (Riba 1998),

it is possible that they may prevent further spread by

the two new Erica spp. although this will require

long term monitoring of the site, especially if the

area were to be burnt in the future. Nevertheless, in

either scenario, the real threat is to the native

sclerophyllous understorey vegetation, as there are

additional and apparently successful introduced

competitors for the limited resources at the site.

Acknowledgements

The former Department of Botany, now Discipline

of Environmental Biology within the School of Earth

and Environmental Sciences, The University of

Adelaide is thanked for the provision of resources for

this project, the initial phase of which was

undertaken by DT as part of a B.Sc. (Hons) degree.

The director of the Adelaide Botanic Gardens and

staff of the Mount Lofty Botanic Gardens are

thanked for permission to undertake the research in

the gardens.

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THE MYRTLE SPRINGS METEORITE: A CHONDRITE (H4)

FROM SOUTH AUSTRALIA’

By M. ZBIK* & A. PRING**

Summary

Zbik, M. & Pring, A. (2004) The Myrtle Springs meteorite: A (H4) chondrite from

South Australia. Trans. R. Soc. S. Aust. 128(1), 33-36, 31 May, 2004.

A single stone of 52.99 g was found 30 km west of the Old Myrtle Springs homestead

on the Lake Torrens Plains, (30° 27° 13.8°° S, 137° 59’ 24.6” E). The chondrules in

the meteorite are well defined and the dark matrix consists of fine olivine and

pyroxene grains. Olivine (Fa,7, , 9), orthopyroxene (Fs,57. 97W0, 8,9), clinopyroxene

(Wo,, . 29FS814 . 19), troilite and chromite are present. Radial pyroxene chondrules,

porphyritic pyroxene chondrules, granular olivine-pyroxene porphyritic and olivine-

pyroxene chondrules are abundant in the meteorite. Cryptocrystalline pyroxene

chondrules and pyroxene-olivine chondrules are rare. Based on texture and mineral

chemistry, the Myrtle Springs meteorite is classified as an H4 chondrite of shock

stage S1-2, weathering grade W4.

Key Words: Myrtle Springs, meteorite, chondrite.

Transactions of the Royal Society of S.

Aust. (2004), 128(1), 33-36

THE MYRTLE SPRINGS METEORITE:

A CHONDRITE (H4) FROM SOUTH AUSTRALIA!

by M. ZBik* & A. PRING™

Summary

ZBIk, M. & PRING, A. (2004) The Myrtle Springs meteorite: A (H4) chondrite from South Australia. 7rans. K.

Soc. S. Aust. 128(1), 33-36, 31 May, 2004.

A single stone of 52.99 g was found 30 km west of the Old Myrtle Springs homestead on the Lake Torrens

Plains, (0° 27' 13.8" S, 137° 59' 24.6" E). The chondrules in the meteorite are well defined and the dark matrix

consists of fine olivine and pyroxene grains. Olivine (Fai72 : 06), orthopyroxene (Fsis7 + 07W01s:08),

clinopyroxene (Wo i2.0Fsi4 +10), troilite and chromite are present. Radial pyroxene chondrules, porphyritic

pyroxene chondrules, ‘granular olivine-pyroxene porphyritic and olivine-pyroxene chondrules are abundant in

the meteorite. Cryptoerystalline pyroxene chondrules and pyroxenc-olivine chondrules are rare. Based on

texture and mineral chemistry, the Myrtle Springs meteorite is classified as an H4 chondrite of shock stage

S1-2, weathering grade W4.

Key Worps: Myrtle Springs, meteorite, chondrite.

Introduction

The meteorite was found by Mr Don McColl of

Glenside, South Australia on the 11th of July 2

while he was searching for tektites on Old Myrtle

Springs homestead on the Lake Torrens Plain. The

exact location as determined by GPS is 30° 27’ 13.8”

S, 137° 59' 24.6” E. The site is some 21 km due west

of the Old Myrtle Springs homestead and some 40

km WNW of Leigh Creek South township (Fig. 1).

Mr McColl undertook a detailed search of the area for

more pieces of the meteorite, but none was found.

Given that there are few geographical place names in

the area, we propose the name Myrtle Springs, the

nearest geographical name. A report of the meteorite

name and petrological details have been submitted to

the Meteorite Nomenclature Committee and approval

has been granted. Mr Don McColl surrendered the

meteorite to the Museum in accordance with the

South Australian Museum Act and was presented

with a bronze medallion to commemorate the find.

Macroscopic Description

The meteorite is a 52.99 g single stone, shaped like

a flattened rounded pebble about 4 cm in diameter

and 2 cm thick and covered with fragments of a

weathered brown crust (Fig. 2). The distinctive crust,

1 mm thick, is probably desert patina rather than a

fusion crust. The silicate minerals throughout the

interior of the stone are dark brown in colour and this

' This paper is dedicated to the late David Williams an intrepid

tektite hunter who passed away 7th February 2003.

“Tan Wark Research Institute, University of South Australia,

Mawson Lake, Adelaide, South Australia 5095.

“Department of Mineralogy, South Australian Museum, North

Terrace, Adelaide, South Australia 5000.

together with the desert patina, indicate that the

meteorite has been exposed to the weather for a

prolonged time. Metal is not present in the meteorite

and it appears to have been oxidised to goethite and

staining the matrix brown. A few unoxidized grains

of troilite however, are still present, so the meteorite

has not been completely oxidized. The extent of

oxidation of the Myrtle Springs meteorite indicates

some degree of antiquity for the fall, but it is rather

difficult to judge the terrestrial age of the fall,

possibly 20,000 to 35,000 years based on the

weathering state (Wlotzka 1993). Jull e¢ al. (1993)

found only a weak correlation between the degree of

{

coe er ees ae see L ot =

aie |

|

Kittakittaooloo |

Ke

Lf aN)

Witchelina $ coomna Bore

‘

Sone Springs sa |

Leigh aes S |

N 4 |

2

x Augusta

|

Fig. 1. Map showing the distribution of H4 chondrites in

north eastern South Australia.

34 M. ZBIK & A. PRING

Fig. 2. Upper and lower surfaces of the Myrtle Springs meteorite showing the shape of the stone and the weathered brown

desert patina crust. The scale bar is marked in centimetres.

Pig. 3. Different types of chondrules in the Myrtle Springs

meteorite. (a) Barred olivine chondrule (BO), composed

of the olivine bars and mesostasis in centre of

micrograph and granular olivine pyroxene chondrule

(GOP) to the left side (photo is 1.5 mm). (b) Elliptically

shaped cryptocrystalline pyroxene olivine chondrules

(C) display unusual wavy extinction probably due to

shock metamorphism (photo is 1.5 mm). (c) Porphyritic

pyroxene olivine (POP) chondrule with poikilitic

structure (photo is 1.5 mm).

weathering and the terrestrial age of meteorites in a

detailed study of a group of meteorites from north

western Texas, U.S.A.

Sample and Analytical Procedures

The polished thin section of the meteorite was used

for both the petrographic examination and chemical

analyses. Composition of the silicate minerals was

determined using a CAMECA SX51_ electron

microprobe, with a Moran analysis package, at

Adelaide Microscopy, The University of Adelaide.

Analyses were made using an accelerating voltage of

15 keV, a sample current of 20 nA, and beam width

of 0.1 um.

Mineralogy

In thin section the meteorite is generally brown due

to the goethite staining which permeates all of the

section. The chondrules are clearly visible but they

are extensively fractured. They are typically between

0.5 and | mm in diameter, but a few measuring more

than 2 mm in diameter were noted.

THE MYRTLE SPRINGS METEORITE: A CHONDRITE (H4) FROM SOUTH AUSTRALIA 35

The chondrules observed in the thin section are

clearly defined which is typical in low petrologic

type, but may have been additionally enhanced by

the preferential weathering and iron oxide staining of

the matrix. The chondrules and chondrule fragments

are composed predominantly of olivine and

pyroxene. Some chondrules display cryptocrystalline

structure, Using the classification system of Wasson

(1993), a number of distinct chondrule types can be

identified in the meteorite. Barred olivine chondrules

(BO). with pyroxene-plagioclase — mesostasis

between the olivine bars, are common (Fig. 3a).

Cryptocrystalline pyroxene and pyroxene-olivine

chondrules (C) are rare; those few observed vary in

size and shape and display strong wavy extinction.

These C-type chondrules are mineralogically

heterogeneous, with in some areas pyroxene and

others olivine (Fig. 3b). Radial pyroxene chondrules

(RP) are common, and display a range of crystal

sizes but nucleation appears to have occurred at a

single point on the rim of the chondrules. Abundant

are porphyritic pyroxene (PP) chondrules, granular

olivine-pyroxene chondrules (GOP) (Fig. 3a, on the

lefi-hand side), and porphyritic olivine-pyroxene

chondrules (POP). Porphyritic chondrules contain

fine and coarse grained, euhedral olivine crystals and

display a poikilitic texture (Fig. 3c). Chondrules vary

in shape, whereas the small chondrules, those below

1 mm, are generally round, the larger are often more

oval and less regular in outline. Turbid

cryptocrystalline mesostasis between olivine and

pyroxene crystals is present and has a

plagioclase\pyroxene-like composition but the

analyses were non-stoichiometric. — Distinct

plagioclase grains were not found during petro-

logical examination or microprobe analysis.

The matrix of the meteorite does not appear to

have been recrystallised and consists of fine-grained

olivine and pyroxene fragments but it is greatly

obscured by the goethite staining making tt difficult

to study. Accessory chromite and troilite grains were

detected in microprobe investigation. A number of

quartz grains, aeolian in origin, were cemented into

the secondary limonitic desert patina making crust

around meteorite.

Selected electron microprobe analyses for the

Myrtle Springs meteorite are given in Table |. The

olivine in the Myrtle Springs meteorite ts

equilibrated, with a mean fayalite content of Fajz 9.0.6.

(mean of 18 analyses). The Ca-poor pyroxene is

uniform and shows only a narrow variation in

composition with mean ferrosilite content of

Fs)5 7.9.7, and wollastonite content of Wo; s,s (mean

of 20 analyses). The Ca-rich pyroxene is somewhat

variable in composition with mean ferrosilite content

of Fsjyo-;o, and wollastonite content of Wor «i209

(mean of 15 analyses).

Classification

The Myrtle Springs meteorite has been classified

as an H4 chondrite. The olivine (Faj75.9.) and Ca-

poor pyroxene (Fsj5.7,9.7) compositions are within the

range of the H chondrites (Keil & Fredriksson 1964).

The equilibrated mineral compositions of the olivine

and Ca-poor pyroxene, the presence of about 20% of

the polysynthetically twinned Ca-rich pyroxenes, the

lack of recrystallisation of the matrix, the well-

defined chondrule boundaries and the glassy

mesostasies all indicated that the Myrtle Springs

meteorite belongs to petrological type 4

classification of Van Schmus & Wood (1967). The

wollastonite content in the low-calcium pyroxene is

within the range found in H4 chondrites (Scott e7 a/.,

1986). Van Schmus and Wood acknowledge that in

their classification scheme that petrological type 4 is

defined by relatively ambiguous characteristics as it

represents a transition between extremely variable

olivine and pyroxene compositions (type 3) and the

uniform compositions (type 5). We choose type 4

rather than type 5 because of the abundance of

clinopyroxene; type 5 chondrites have only very

minor clinopyroxene.

Most olivine and pyroxene grains display sharp

extinction and irregular fracture crystals in the

Myrtle Springs chondrite. Some pyroxene display

weak undulatory extinction. Both are indicative of

the meteorite having been unshocked or only very

weakly shocked after metamorphism. According to

the classification scheme of Stoffler ef a/. (1991), the

shock facies is estimated to be SI-2 (or i.e.)

unshocked to very weakly shocked. The oxidation of

approximately 95% of the original metal and troilite

in the meteorite indicates weathering state W4 on the

classification scheme of Wlotzka (1993).

Other H4 chondrites from

North Eastern South Australia.

There are a number of other H4 group meteorites

known from the north eastern region of South

Australia (Fig. 1). The Kittakittaooloo meteorite was

found in 1970 about I km NW _ of Lake

Kittakittaooloo (28° 2’ S, 138° 8' E) and has an

olivine composition of Fa,,, (Graham er al. 1985).

The Coonana meteorite was found in 1962. near

Coonana Bore (29° 51’ S, 140° 42’ E) and has an

Fa,y olivine composition (Graham e/ al. 1985). Both

of these meteorite localities are rather remote from

Myrtle Springs being approximately 270 km north

and 250 km ENE, respectively, of the site. The

Witchelina meteorite was found in 1920 in the area

of Witchelina homestead (30° S, 138° E), only 40 km

north of the Myrtle Springs homestead, and it is

possible that the two meteorites might be part of the

36 M. ZBIK & A. PRING

TABLE |. Representative electron microprobe analyses (wt.

%) of selected minerals in Myrtle Springs meteorite.

OXIDE Olivine Ca-poor Ca-rich

(No. Analyses) — (18) pyroxene pyroxene

(20) (15)

SiO, 39.26 56.42 49.60

TIO, 0.05 0.10 0.24

ALO, 0.04 0.41 3.41

FeO 16.13 10.38 8.83

MnO 0.48 0.52 0.47

MgO 43.41 30.58 22.86

CaO 0.04 0.94 10.77

Cr,O, 0.40 0.30 0.91

Total 99.76 99.05 97.09

same fall. Unfortunately the locality details for the

original site for the Witchelina find are sketchy and

only approximate. The Witchelina meteorite was a

3.6 kg single stone and is coated with a distinctive

fusion crust. Analysis gave an olivine composition of

Fa,,, (Graham ef al. 1985), slightly more iron-rich

than Myrtle Springs. Witchelina, however is a much

fresher stone. The metal has not been oxidized and

metal and troilite are estimated to constitute around

3% of the meteorite. This indicates a weathering

state of W1 or W2 and a terrestrial age of only a few

thousand years. The ground mass colour has a

distinctly greyish brown tone rather than reddish

brown. While it might be expected, given the

difference in size between Myrtle Springs and

Witchelina, that the former would show the effects of

weathering to a somewhat greater degree than the

larger mass, but probably not to this extent. The

difference in olivine composition, the presence of the

fusion crust, and fresh metal in Witchelina all indicate

that the two meteorites are not part of the same fall.

Acknowledgments

The authors wish to thank Mr Angus Netting and

Mr John Terlet, of Adelaide Microscopy, The

University of Adelaide, for assistance with the

electron microprobe analyses. We wish to thank the

editor and the two referees for their detailed and

helpful comments on the manuscript.

References

GRAHAM, A. L. BEVAN, A. W. R. & HUTCHINSON, R. (1985)

Catalogue of meteorites. 4th edition. British Museum

(Natural History) London. 460pp.

JuLL, A. J., DONAHUE, D. J., CIELASZYK, E. & WLOTZKA, F.

(1993) Carbon-14 terrestrial ages and weathering of 27

meteorites from the southern high plains and adjacent

area (USA) Meteoritics 28, 188-195.

Keil, K. & FREDRIKSSON, K. (1964) The iron, magnesium

and calcium distribution in coexisting olivines and rhombic

pyroxenes of chondrites. J. Geophys. Res. 69, 3487-3515.

Scott, E. R. D., TAYLor, G. J. & KEIL, K. (1986) Aceretion,

metamorphism, and brecciation of ordinary chondrites:

Evidence from petrologic studies of meteorites from

Roosevelt County, New Mexico. Proc Lunar Planet. Sci.

Conf. 17th, ELI5-E123.

STOFFLER, D., KrIL, K. & Scort, E. R. D. (1991) Shock

metamorphism of ordinary chondrites. Geochim.

Cosmochim. Acta. 55, 3845-3867.

VAN ScuMus, W. R. & Woon, J. A. (1967) A chemical-

petrologic classification for the chondritic meteorites.

Geochim. Cosmochim. Acta 31: 747-765.

Wasson, J. T. (1993) Constraints on chondrule origins.

Meteoritics 28, 14-28.

WLOTZKA, F. (1993) A weathering scale for the ordinary

chondrites. Mefeoritics 28, 460.

EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841-1996

By P. I. LEPPARD*, G. M. TALLIS & C. E. M. PEARCE

Summary

Leppard, P. I., Tallis, G. M. & Pearce, C. E. M. Expected lifetime in South Australia

1841 — 1996. Trans. R. Soc. S. Aust. 128(1), 37-42, 31 May, 2004.

For each sex, population life tables have been calculated from mortality data

associated with the age-specific population counts produced by each of the twenty-

five population censuses that have been conducted in South Australia from 1841 until

1996. Estimates of expected lifetime have been derived separately for males and

females for each census year. The computationally intensive statistical method of the

bootstrap has been used to calculate a statistical sampling error for each estimate of

expected lifetime. The results show generally increasing trends: from approximately

44 years and 48 years in 1841, to 75 years and 81 years in 1996, for males and

females respectively.

Key Words: Expected lifetime, life expectancy, bootstrap standard error, South

Australia.

Transactions of the Roval Society of S. Aust. (2004), 128(1), 37-42.

EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841 — 1996.

by P. I. Lepparp’, G. M. TALLIS & C. E. M. PEARCI

Summary

Lepparb, P.1., TALLIS, G. M. & PEARCE, C.E.M. Expected lifetime in South Australia 1841 - 1996. Trans. R. Soc.

S. Aust. 128(1), 37-42, 31 May, 2004.

For each sex, population life tables have been calculated from mortality data associated with the age-specific

population counts produced by each of the twenty-five population censuses that have been conducted in South

Australia from 1841 until 1996, Estimates of expected lifetime have been derived separately for males and

females for each census year. The computationally intensive statistical method of the bootstrap has been used to

calculate a statistical sampling error for each estimate of expected lifetime. The results show generally

increasing trends; from approximately 44 years and 48 years in 1841, to 75 years and 81 years in 1996, for males

and females respectively.

Kry Worps: Expected lifetime; life expectancy: bootstrap standard error; South Australia.

Introduction

Expected lifetime or life expectancy, meaning the

average length of life from birth until death, is often

used as a composite indicator of the health and social

conditions of a human population. Expected lifetime

is a single value that summarises the prevailing

mortality experience of a defined population of

individuals, and consequently provides a means by

which the mortality patterns of different populations

can be conveniently compared. The progress in the

development of the general well-being of a particular

population can also be assessed by a time sequence

of relevant expected lifetime values. Alternatively,

expected lifetime can be cautiously interpreted as a

prediction of future average lifetime.

Expected lifetime is conventionally calculated

separately for males and females, and is one of the

results of a (mortality or actuarial or population) life

table analysis. This type of analysis uses age-specilic

death rates calculated from age-specific population

sizes and numbers of deaths. Hence life tables are

usually calculated for census years only since

accurate estimates of population sizes are required.

Mortality data pertinent to the three consecutive

years centred on and encompassing the census year

are averaged in an attempt to smooth out random

fluctuations in the observed numbers of deaths. The

study of population mortality patterns is generally

acknowledged as beginning with Graunt (1662), and

subsequently the methodology of the life table has

evolved to become a basic tool of demographers (eg

Newell 1988) and actuaries (eg Benjamin &

Haycocks 1970). There are many standard statistical

texts covering the methodology of life table analysis

(eg Elandt-Johnson & Johnson 1980).

School of Mathematical Sciences, University of Adelaide, Adelaide,

South Australia, S005, Australia, Email: pleppard@maths.adelaide.edu.au.

Very tew official life tables and hence estimates of

expected lifetime have been produced for South

Australia. Invariably 19th century references to

Australian mortality are based on New South Wales

or Victorian data (Pell 1867, Burridge 1884). Some

official South Australian life tables have been

calculated for the period 1881-1910, but these tables

were calculated from mortality data of ten-year

periods. This methodology was discarded when the

Commonwealth Statistician (Wilson 1936) decided

to cease publication of separate life tables for each

Australian State on the basis of the “.. time (thus)

saved..”. In the 1970’s the Australian Bureau of

Statistics recommenced producing life tables

specifically for each State. Thus the complete

mortality experience of South Australia, as measured

by male and female expected lifetimes, has not been

consistently or comprehensively investigated.

Materials and Methods

The results presented in this paper have been

extracted from a thesis presented for an MSc degree

at The University of Adelaide, Leppard (2003). The

thesis can be accessed electronically at

http://thesis.library.adelaide.edu.au/public/adt-

SUA20030422.122816/.

A very brief overview of the material covered in

the thesis is given here.

The essential data requirements to calculate a life

table for a given census year are, first, the population

size at each single year of age; and second, the

number of deaths at each single year of age for that

census year, and for the calendar years immediately

preceding and following it. Population data were

obtained from official sources for each of the 25

censuses carried out in South Australia since 1841:

12. colonial censuses and 13 post-Federation

censuses. Sources for these data include the South

tad

Australian Parliamentary papers, the $A Government

Gazette, the Statistical Register of South Australia

(1850-1975), and various publications of the

Commonwealth Bureau of Census and Statistics and

its successor organisation, the Australian Bureau of

Statistics. Age-specific numbers of deaths by sex are

also available from these same sources, although the

published data for the years 1841 to 1860 are in a

statistically limiting form principally because of

various types of grouping by age of the data. This

deficiency of the official records has been overcome

by tabulating the primary data contained in death

certificates lodged with the Registry of Births,

Deaths and Marriages, which are held in the archives

of State Records of South Australia, Permission was

obtained from the Registrar for access to the

Registry’s records, and approximately 18,000 death

certificates for relevant years of the above period

were examined and the deaths they reported were

classified according to year, sex and age at death.

Indexes to the burial records of Holy Trinity church

and the West Terrace cemetery were used to obtain

data from which the age-specific number of deaths

was estimated for each sex for 1840, It is impossible

to provide within the space limitations of this paper

a comprehensive description of the characteristics of

all data sets that have been used to calculate expected

lifetime. The reader is referred to Leppard /oc cit for

a detailed description for each census year of the

process of data collection, a discussion of the

limitations of the data, estimates of expected lifetime

calculated by the life table method, and estimates of

sampling error calculated by using — the

computationally intensive statistical bootstrap

procedure (Efron & Tibshirani 1993). For each

census year a robustness analysis 1s also presented,

where an examination is made of the effect on

expected lifetime of under-reporting in the age-

specific population sizes and number of deaths. All

data files and computer programs can be publicly

accessed from the web site previously indicated.

Expected lifetime is calculated here in the

following way, conventionally for males and females

separately. Let ,p, be the number of individuals aged

between 0 and | years at a census. Let jd, be the

average number of individuals who died between the

ages of 0 and | years, determined from the number

of deaths registered in the census year and the years

immediately preceding and following the census

year. Then the quantity ,q, is calculated from ,p, and

iy as

d,

m0

170 ay ot te

Py" ay

It is sometimes recommended that, in

circumstances where extremely high infant

mortality prevails, a factor smaller than the factor

8 P. 1. LEPPARD, G. M. TALLIS & C. E. M. PEARCE

of '/2 should be used in the calculation of ,g,. This

has not been done in the calculation of expected

lifetimes shown in Table 1, and consequently some

of the values given there for the 19th century may

be slightly larger than perhaps they otherwise

would be. Newell /oc cif suggests that a factor of .3

be used instead of '/2 for “developing countries”.

Recalculation of expected lifetime for selected

census years for South Australia using this value

for the factor has produced small differences that

are less than the standard error of the expected

lifetime.

In an analogous manner, ,q, is calculated for age

group |-2 years, ,q, for age group 2-3 years, and so

on, for each yearly age group. Thus using standard

demographic notation, ,g, is a mortality rate specific

to the group aged between j years and (j+1) years.

A distribution function for lifetime values, Fy is

defined with this set of q-values as

FOX) 1-1 a )- g )O> 9a) pg)

for x = 1,2.3..., where F(0O) = 0 and here F(105) = |

by definition. Expected lifetime is calculated from

the tabular values of F’ by the trapezoidal method of

numerical integration.

It should be noted that neither / nor expected

lifetime is directly applicable to the population of

individuals in the census year from which the g-

values are determined. They are concepts that

prospectively apply to a hypothetical population of

infants born in the census year, if it is assumed that

the currently prevailing mortality rates, as measured

by the g-values, eventuate for these infants

throughout their lifetimes. There are mathematical

arguments that suggest that expected lifetime is an

underestimate of the unrealised average lifetime of

this hypothesised population of infants.

The potential effect of under-counting is examined

in the following way. Population sizes are obtained

following a census, and the numbers of deaths are

compiled from the death certificates required by law.

Both of these processes are not necessarily exact and

some individuals could be excluded. Consider jp

and jd) which are used to calculate jg», and suppose

the “correct” values of these two quantities are larger

by factors of O; and O02 respectively. Then the

“correct” value of ;qy would be

Ox ido = ido

Or ipo + $O2 ido Ov ipo+ hido

3 OD. 3

iqy( “correct”’) =

which is close to \qg) calculated by using the

supposedly under-counted values jp) and jd) if

©. ~ |; that is, jgp is an appropriate substitute for 140

(“correct”) if the extent of under-counting is

EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841 - 1996

approximately the same for both the population size

and the number of deaths. For each census year and

sex, expected lifetimes are given in Leppard /oc cif

for combinations of assumed levels of under-

counting in both population size and numbers of

deaths.

Results and Discussion

Estimates of expected lifetime (EL) for South

Australian males and females for 1841-1996 are

shown in Table |. A standard error, SE (EL), is also

given for each estimate. Table 1 also shows the total

population size and the average total number of

deaths on which each estimate is based.

The estimates of expected lifetime are also

displayed graphically in Figure |, which is annotated

to indicate

1. The first census at which the estimate of

expected lifetime for females can be statistical ly

claimed to be significantly greater than the

estimate for males.

39

over this period in the generally increasing

sequence of expected lifetime values.

. Larger time intervals between successive

estimates of expected lifetime as a consequence

of the postponement of censuses during the

Great Depression and World War II.

The estimates of expected lifetime presented in

Table | and Figure | have been derived from data

from populations that have evolved from the initial

British settlement of South Australia with

subsequent migration from Western and Southern

Europe. These estimates have been derived primarily

from routinely recorded official population and

mortality data from which Aborigines were excluded

because of a government policy that remained in

force until the second half of the 20th century.

The two series of expected lifetimes that are

presented here are the most extensive and

comprehensive available for any Australian state or

for the Commonwealth, with all estimates

consistently calculated using a currently accepted

standard methodology. The results delineate for the

2. The passing of the first South Australian Public first time the changes in mortality that have occurred

Health Act, as a response to the deterioration of — in South Australia over 150 years of European

social and public health conditions in the 1860’s settlement. The electronic provision of the basic data

and 1870’s. The effect of this deterioration on files through the previously stated web address has

population mortality is shown by the disruption — been made to assist demographic researchers.

TABLE |. Expected lifetime with standard error for South Australia 1841-1996.

Male Female

Census Population Deaths EL SE(EL) Population Deaths EL SE(EL)

1841 8195 140 44.27 2.59 6345 102 48.17 1.88

1844 9525 172 41.95 1.56 7608 128 43.49 1.60

1846 12670 236 41.57 1.55 9650 163 46.67 2.08

1851 35309 544 45.73 1.24 27730 439 47.62 1.31

1855 43716 713 47.68 1.27 41470 620 50.00 1.27

1861 64640 111 48.55 1.06 61678 951 51.88 1.20

1866 85625 1434 47.39 0.86 77975 1178 50.34 0.88

187] 95288 1438 48.69 0.67 90164 1165 52.19 0.77

1876 110410 1972 43.57 0.58 102734 1659 46.09 0.64

1881] 149530 2283 47.62 0.50 130335 1819 50.76 0.62

189] 166801 2156 52.52 0.47 153630 1787 54.98 0.52

1901 184424 2169 54.56 0.47 178182 1821 57.72 0.50

1911 207358 2269 58.81 0.41 201200 1880 62.41 0.42

192] 248267 2663 60.01 0.38 246893 2221 63.53 0.39

1933 290429 2701 65.34 0.34 289546 2386 68.04 0.35

1947 320031 3478 67.17 0.27 326042 2995 71.28 0.26

1954 403903 3958 67.72 0.25 393191 3267 73.07 0.24

1961] 490225 4412 68.62 0.22 479115 3537 74.91 0.21

1966 548530 5030 68.57 0.21 543344 4034 13,22 0.20

1971 586451 5454 68.67 0.20 587656 4408 75.58 0.20

1976 619759 5402 70.06 0.19 624595 4488 76.93 0.18

1981 635695 5473 71.49 0.19 649467 4438 78.92 0.18

1986 666159 5626 72.98 O.18 679985 4824 79.48 0.18

199] 690805 5856 74.08 0.17 709802 5156 80.30 0.17

1996 698799 5989 75.33 0.17 722673 5504 81.34 0.16

40 P. I. LEPPARD, G. M. TALLIS & C. E. M. PEARCE

pee _ _— es

Expected lifetime in South Australia 1841-1996 |

i oe |

Great Depression |

; |

bo

| A \

1 4 |

1) |

1 a ! |

7 j ae

Stattically detectable gender diferences |

'

| ' vferde

60 ‘

A

} i

| Vir ar |

s ! |

ne |

1 A 4

] a t A

/ ot 1

A t

i \ |

\

4

Public Health Act 1873

4 rs T ~~ T TO ee To

{640 {860 {880 1900 12) 10 1860 1980 2000

Year

Fig. 1. Expected lifetime in South Australia 1841-1996.

EXPECTED LIFETIME IN SOUTH AUSTRALIA 1841 - 1996

TABLE 2. Comparison of expected lifetime for males.

4|

SA Other Australia

Britain US

Sources (6) (7) (8)

1841 44.27 40.2

1855 47.68

1838-1854 39.9

1856-1861 45.58 (1)

1861 48.55

1871 48.69

1870-1881 46.47 (2)

1881 47.62

I881-1890 50.61 (3) 47.20 43.7

1891 2 ee

1891-1900 53.02 (3) 51.08 44.1

1901 54.56 48.0

1901-1910 56.76 (3) 55.20 48.5

1911 58.81

£033 65.34 63.48 58.7 62.7

1954 67.72 67.82 (4) 67.14 67.5

1976 70.06 70.27 (5) 69.56 69.9 69.9

1996 75.33 75.30 (5) 75.70 74.5 1S;

(1) Pell loc cit (NSW).

(2) Burridge loc cit (composite states, excludes SA).

(3) Commonwealth Census 1911, Vol 3. (SA).

(4) Wickens, Australian Journal of Statistics, 2, 1960. (SA).

(5) Deaths SA, Australian Bureau of Statistics.

(6) Australian Life Tables, Australian Government Actuary.

(7) English Life Tables, UK Government Actuary’s Office. (England & Wales).

(8) US National Vital Statistics Reports, Vol 51, 3. 2002.

TABLE 3. Comparison of expected lifetime for females.

SA Other Australia Britain US

Sources

1841 48.17 42.2

1855 50.00

1838-1854 41.9

1856-1861 45.58

1861 51.88

1871 52.19

1870-1881 49.64

1881 50.76

1881-1890 53.8] 50.84 47.2

189] 54.98

1891-1900 56.10 54.76 47.8

1901 $7.72 51.0

1901-1910 60.39 58.84 52.4

191] 62.41

1933. 68.04 67.14 62.9 66.3

1954 73.07 73.09 72.75 cia:

1976 76.93 77.24 76.56 76.0 TFS

79.6 79.7

1996 81.34 81.34 81.40

42 P. 1. LEPPARD, G. M. TALLIS & C. E. M. PEARCE

A selection of expected lifetimes presented here for

South Australia in Table 1 are shown in Table 2 and

Table 3 (column SA) for comparison with other

estimates and other similar populations; with

estimates for other colonies and for South Australia,

by other authors (Other sources); with Australia, by

the Australian Bureau of Statistics and by the

Australian Actuary; with Britain, by the Government

Actuary’s Office; and with the United States, by the

National Vital Statistics Office.

Generally, expected lifetimes for South Australia

have compared more than favourably with the other

estimates until the first third of the 20th century, after

which time there has been a gradual lessening of the

differences between expected lifetimes. This most

probably reflects the effect of globalisation and

standardisation in conditions of Western industrial

societies.

References

BENJAMIN, B. & H. W. HAycocks (1970). The Analysis of

Mortality and other Actuarial Statistics. Cambridge,

Cambridge University Press.

BurribGe, A. F. (1884). “On the rates of mortality in

Australia.” Journal of the Institute of Actuaries 24: 333-

358.

EFRON, B. & R. J. TIBSHIRANI (1993). An Introduction to the

Bootstrap. New York, Chapman & Hall.

ELANDT-JOHNSON, R. C. & N. L. JOHNSON (1980). Survival

Models and Data Analysis. New York, John Wiley &

Sons.

GRAUNT, J. (1662). Natural and political observations made

upon the bills of mortality (reprinted 1939). Baltimore,

Johns Hopkins Press.

Lepparn, P. 1. (2003). An analysis of population lifetime

data for South Australia. 1841- 1996. MSc (Statistics)

thesis, School of Applied Mathematics, The University

of Adelaide http://thesis.library.adelaide.edu.au/public/adt-

SUA20030422.122816/.

NEWELL, C. (1988). Methods and Models in Demography.

London, Belhaven Press.

Pett, M. B. (1867). “On the rates of mortality and

expectation of life in New South Wales as compared with

England and other countries.” Transactions of the Royal

Societv of New South Wales 1: 66-76.

WILSON, R. (1936). Official year book of the

Commonwealth of Australia, Commonwealth Bureau of

Census and Statistics, Canberra.

VARIATION IN SEX RATIOS IN FOUR ANGUINA

(NEMATODA: ANGUINIDAE) SPECIES

By I. T. RiLEy* & T. BERTOZZIf

Summary

Riley, I. T. & Bertozzi T. (200). Variation in sex ratios in four Anguina (Nematoda:

Anguinidae) species. Trans. R. Soc. S. Aust. 128(1), 43-46, 31 May, 2004.

Number and ratios of adult male and females nematodes in four Anguina species. A.

australis from Ehrharta longiflora, A. funesta from Lolium rigidum, A. microlaenae

from Microlaena_ stipoides and an _ undescribed species from Polypogon

monspeliensis, were determined and analysed. The sex ratio in A. australis and A.

funesta differed statistically from the expected 1:1, with 57% and 56% females,

respectively. The ratios for A. microlaenae and Angina sp. ex Polypogon conformed

to the expected 1:1. Galls initiated by single sex nematodes, both male and female,

were found in the four species, except that no galls containing only males were found

in A. australis. Reproduction in this group appears to be amphimitic, as single sex

galls contained no progeny and the data point to the possiblity of a mechanism that

favours female occupancy of galls.

Key Words: Anguina australis, Anguina funesta, Anguina microlaenae, seed gall

nematode, leaf gall nematode, gall initiation, sex ratios, reproduction, amphimixis.

Transactions of the Roval Society of S. Aust. (2004), 128(1), 43-46.

VARIATION IN SEX RATIOS IN FOUR ANGUINA

(NEMATODA: ANGUINIDAE) SPECIES.

by I. T. Ritey’ & T. Bertozzi'

Summary

Ritey LT. & BerrozziT. (200). Variation in sex ratios in four Anguina (Nematoda: Anguinidae) species. Trans.

R. Soc. S. Aust. 128(1), 43-46, 31 May, 2004.

Number and ratios of adult male and females nematodes in four Anguina species, A. australis trom Ehrharta

longiflora, A. funesta trom Lolium rigidum, A. microlaenae trom Microlaena stipoides and an undescribed

species from Polypogon monspeliensis, were determined and analysed. The sex ratio in A. australis and A.

yA

funesta differed statistically from the expected 1:1, with 57% and 56% females, respectively. The ratios for A,

microlaenae and Angina sp. ex Polvpogon conformed to the expected 1:1. Galls initiated by single sex

nematodes, both male and female, were found in the four species, except that no galls containing only males

were found in 4. australis, Reproduction in this group appears to be amphimitic, as single sex galls contained

no progeny and the data point to the possibility of a mechanism that favours female occupancy of galls.

Key Worps: Anguina australis, Anguina funesta, Anguina microlaenae, seed gall nematode, leat gall

nematode, gall initiation, sex ratios, reproduction, amphimixis.

Introduction

Species of the nematode genus Anguina form galls

in plants, mostly in poaceous hosts. The type species,

Anguina tritici, is considered to reproduce by

amphimixis (Triantaphyllou and Hirschmann 1966).

Galls are initiated by invasive stage juveniles

(usually J2s), which, confined to the gall, develop to

adults. So, mates can only be found amongst the

relatively small number of individuals present in the

gall. Therefore, a mechanism to increase the

likelihood of both females and males occurring in

individual galls would be advantageous. The

mechanism would need to operate before or during

gall initiation, unless sex in Anguina spp. can be

environmentally determined.

In two species, Anguina australis and Anguina

microlaenae, most galls contain only small numbers

of adults, with some galls containing only females

(Riley ef al, 2001; de Silva and Riley 2002). It was

suggested that this may indicate a mechanism

whereby only genetically female juveniles could

initiate galls or that sex was environmentally

determined (de Silva and Riley 2002). In both cases,

no progeny were found in the single sex galls, so

there was no evidence of parthenogenetic

reproduction.

As sex ratios and adult populations in galls for only

two leaf-gall forming species of the genus have been

examined in detail, similar data were collected for

two seed-gall forming species, Anguina funesta and

“Plant and Pest Science, School of Agriculture and Wine, The

University of Adelaide, PMB 1, Glen Osmond, South Australia

5064, Australia, Email: ian.riley(@adelaide.edu.au,

‘Evolutionary Biology Unit, South Australian Museum, North

Terrace, Adelaide South Australia 5000, Australia.

Anguina sp. ex Polypogon monspeliensis and further

data for A. microlaenae. These data were examined

for further evidence of mechanisms that determine

sex ratios and first generation populations in

Anguina galls.

Materials and Methods

Anguina australis. Data on sex of A. australis

adults in galls in Ehrharta longiflora came from a

published study (Riley ef a/. 2001)

Anguina funesta. Lolium rigidum plants were

grown (about 100 plants per pot) in a peat/sand mix

(UC mix; Baker 1957) in 280 mm pots outdoors at

Urrbrae, South Australia. Seed and nematode galls

(about 100 per pot) were applied early in July 2001

and inflorescences harvested for collection of galls in

late October 2001. Ten inflorescences were dissected

and 203 galls collected. Gall positions (spikelet and

floret) were recorded. Each gall was opened in water

under a dissecting microscope to release and count

adult female and male nematodes. The presence of

eggs and juveniles was noted.

Anguina microlaenae. A further eighty galls from

each of two populations of Microlaena_ stipoides

from Toowoomba, Queensland were assessed as for

the A. fiesta galls from L. rigidiwm and combined

with data previously obtained from these populations

(de Silva and Riley 2002).

Anguina sp. ex Polypogon. Anguina sp. galls from

Polypogon monspeliensis were obtained from plants

collected in 1992 and 1993 from two sites in south-

eastern South Australia. Galls in these plants had

formed in primary shoot meristems, pedicel

meristems and ovaries and in this report are termed

apical, pedicel and seed galls respectively. A total of

44 I. T. RILEY & T. BERTOZZI

TABLE |. Gall types, sex and ratio of adults for populations of four Anguina species.

A. australis* A, funesta A. microlaenae** Anguina sp. ex Polypogon

Gall type Leaf Seed Leaf Apical Seed Pedicel Total

Number examined 194 203 199 95 122 14 232

Females

Range 1-5 0-13 0-6 0-7 0-8 1-3 0-8

eantSE 1.68+0.07 2.66 + 0.14 1.65 + 0.08 1.744014 1.3940.08 1.64+£0.23 1.55 +0.07

Median | 2 | I | |

Males

Range 0-3 0-9 0-8 0-8 0-11 0-3 0-11

ean+SE 1.96+0.04 = 1.98 +£0.10 1.75 + 0.10 1.904018 1.374010 1.214019 1.58 + 0.09

Median | 2 | | | | ]

Adults

Range 1-7 1-19 0-12 1-15 1-19 2-5 1-19

eantSE 2.88+0.08 4.6440.22 3.40+0.15 3.63 £0.29 2.754017 2.864033 3.12+0.15

Median 3 4 3 2 2 2 2

Proportion female

Range 0.33 -1 0-1 0-1 0-1 0-1 0.4—1 0-1

ean+ SE 0.573 + 0.009 0.556 + 0.011 0.504 40.014 0.497+ 0.016 0.504 + 0.011 0.567 + 0.039 0.505+ 0.009

Median 0.50 0.57 0.50 0.50 0.50 0.50 0.5

*Data from Riley ef al. (2001)

**Data from de Silva and Riley (2002) combined with further data collected on the same population

12] A. australis A. funesta

°0C)

6 1% 5% 25% 50%

3 oe

Gon |

42| A. microlaenae

Males per gall

Females per gall

Fig. |. Proportion of galls formed by four Anguina spp.

with various combinations adult nematodes (dotted line

represents 1:1, area of the circle represents the proportion

of the population at each combination and scale indicated

by circles labelled 1, 5, 25 and 50%.)

232 galls were assessed consisting of 95 apical, 122

seed and 14 pedicel galls. Galls were dissected and

adults counted, as above.

GENSTAT 5. (Lawes Agricultural — Trust,

Rothamsted Experimental Station) was used to run

statistical analyses.

Results

Table | gives details of the adults in the galls of

each of the four nematode species. Infestation of the

Lolium rigidum was heavy, nearly all florets in the

spikelets examined containing galls. The contents of

the galls from the two populations of A. microlaenae

did not differ statistically and were combined.

Differences between the three classes of gall

produced by Anguina sp. in P. monspeliensis could

not be reliably tested because of inequality of

variance and no variance-stabilizing transformation

was effective, so they are presented individually and

as a total. Two galls with exceptionally large

numbers of adults, one gall formed by 4.

microlaenae with | female and 38 males and one

apical gall formed by Anguina sp. in P.

monspeliensis with 24 females and 31 males, were

excluded from analysis as aberrant.

The sex ratio for each species was tested for the

whole sample by chi-square test (Table 2). The

numbers of females per male in galls formed by both

A. australis and A. funesta were greater than the

expected ratio of 1:1. This was consistent with the

mean proportion of females found in the galls of both

species (Table 1) being more than half. The ratio for

A. microlaenae and Anguina sp. ex Polypogon did

not differ statistically from the expected 1:1. Despite

the divergence of the sex ratio from 1:1 in two

species, the only species with a median proportion of

females that differed from this ratio was A. funesta

with a median of 0.57 (i.e. 1.33:1),

The proportion of galls with each combination of

VARIATION IN SEX RATIOS IN FOUR ANGUINA (NEMATODA: ANGUINIDAE) SPECIES 45

TABLE 2. Chi-squared test of conformity of Anguina sex ratios to an expected 1:1.

Anguina sp. Observed ratio

Chi-squared Probability

(female/male) (df=1)

A. australis 1.41 15.91 < 0.001

A. funesta 1.36 22.01 <0.001

A, microlaenae 6.95 0.53 0.47

A. sp. ex Polypogon 0.98 0.07 0.79

adults for the four species is shown in Figure 1. The

bias in female occupancy of galls can be seen for A.

australis and A. funesta with 40% and 51% of galls

having more females than males, respectively. By

comparison, only 25% and 19% of A. microlaenae

and Anguina sp. ex Polypogon galls had more

females than males.

Galls occupied by single adult males and females

were found in the four nematode species except

that no galls containing only males were found for

A. australis. The greatest proportion of single sex

galls was found for A. microlaenae (10%). No eggs

or juveniles were found in any single sex gall.

Analysis of gall position (spikelet and floret) for

A. finesta in relation to numbers of adults and

progeny and sex ratio revealed only one significant

relationship (data not shown). A_ significant

regression (P<0.001) was found between sex ratio

and spikelet (the latter being the independent

variable). The lower the position of the spikelet in

the inflorescence the greater proportion of females

in the galls. However, only about 6% of the

variation in sex ratio was attributable to spikelet

position. No effect of floret position within the

spikelet was found.

Discussion

Reproduction in the species examined appears to

be exclusively amphimitic, as no progeny were

found in galls containing single adults. The sex

ratio of 1:1 found in 4. microlaenae and Anguina

sp. ex Polypogon is consistent with amphimitic

reproduction. However, the overall sex ratio of

about 1.4 females/male for A. australis and A.

funesta points to the existence of mechanism that

controls the sex of adults in galls.

The data arising from the earlier, more limited,

examination of A. microlaenae galls and the 4.

australis galls led to postulation (de Silva and

Riley 2002), based on single adults in galls being

only female, that galls might only be initiated by

genetically female juveniles or that environmental

sex determination was involved. The additional

"Chit, W. (1971) “Biology of Anguina mobilis n. sp. on capeweed

(Cryptostemma calendulaceum)” PhD thesis, The University of

Adelaide (Unpubl.).

data presented here revealed that galls formed by

single males occur in three of the species,

including A. microlaenae and provide no support

for either proposition.

Therefore, the significantly higher number of

females in A. australis and A. funesta points to a

possible mechanism involving relative rather than

absolute differences in behaviour between the

sexes. Although both sexes can initiate galls,

female occupancy is quantitatively more likely.

Given that host invasion and gall initiation rates

were high in the L. rigidum examined (mean of

4.6 adults per gall compared to 2.2 to 2.6 in field

populations, McKay er al. 1981), it is possible

that such a mechanism operates to maximise

potential reproduction of the species when there

is more competition for gall sites. This cannot be

tested from the data obtained, as they were not

derived from a controlled experiment. However,

the tendency for more females in the lower galls

in L. rigidum might be consistent with this

proposal, if there was more competition for sites

at the time these galls were formed. Chit! also

found that galls formed by Mesoanguina mobilis

in Arctotheca calendula had 2 to 7 first generation

adults, most commonly with 3 females and 2

males, thus supporting a mechanism to favour

females occurring more broadly in anguinid

genera.

A consistent pattern in the data is the low

frequency of galls containing only single adults

for all species and a strong bias towards galls

containing two adults (one female and one male),

in three species. It is possible that galls initiated

by single nematodes largely fail to mature and or

that gall initiation is favoured by the combined

efforts of multiple juveniles. Juveniles must

secrete plant growth regulators to induce galls, so

it is possible that single juveniles might have

difficulty achieving a critical concentration to

effect the necessary change in the plant. The

approximate balance of male and females in most

galls also points to mechanism other than chance.

The occurrence of galls containing multiple and

variable numbers of adults in all species indicates

that this process is not tightly regulated and must

be to some degree a function of number of

nematodes that invade the plant relative to the

46 I. T. RILEY & T. BERTOZZI

number of potential gall sites. Examining gall

occupancy under a range of inoculation levels could

provide more evidence of the underlying pattern.

Acknowledgments

Tony Debicki helped with the culture 4. funesta in L.

rigidum. Drs K. Owen and P. Williamson provided

infested M. stipoides from Toowoomba. Ms M. Lorimer

provided statistical advice. I. Riley’s position is in part

funded by the Grains Research and Development

Corporation. During the period the P monspeliensis

galls were collected, T. Bertozzi was supported by the

Australian Meat and Livestock Corporation.

References

Baker, K. F. (1957). The U.C. system for producing

healthy container-grown plants through the use of clean

soil, clean stock and sanitation. California Agricultural

Experiment Station Manual 23 (University of California,

Berkeley, USA.).

bE SILVA, P. & Rivey, I. T. (2002). Aspects of the survival

and reproduction of Anguina microlaenae (Nematoda:

Anguinidae). Trans. R. Soc. S. Aust. 126: 45-49,

McKay, A. C., Fisuer, J. M. & Dubé A. J. (1981).

Ecological field studies on Anguina fiesta, the vector in

annual ryegrass toxicity. Australian Journal of

Agricultural Research 32, 917-926.

Riey, 1. T., Scumirz, A. & DE SILVA, P. (2001). Anguina

australis, a vector for Rathayibacter toxicus in Ehrharta

longiflora. Australasian Plant Pathology 30, 361-364.

TRIANTAPHYLLOU, A. C. & HIRSCHMANN, H. (1966).

Gametogenesis and reproduction in the wheat nematode,

Anguina tritici. Nematologica 12, 437-442.

SYPHACIA (SYPHACIA) AUSTRALASIENSIS SP. NOV.

(NEMATODA: OXYURIDAE) FROM RATTUS LEUCOPUS

(MURIDAE) FROM PAPUA NEW GUINEA AND AUSTRALIA

By L. R. SMALES*

Summary

Smales, L. R. (200) Syphacia (Syphacia) australasiensis sp. nov. (Nematoda:

Oxyuridae) from Rattus leucopus (Muridae) from Papua New Guinea and Australia.

Trans. R. Soc. S. Aust. 128(1), 47-51, 31 May, 2004.

Syphacia (Syphacia) australasiensis sp. nov. is described from the caecum of Rattus

leucopus (Gray) (Rodentia: Muridae) from Papua New Guinea and Queensland,

Australia. The new species is distinguished from congeners with an oval cephalic

plateau by the lack of lateral alae, a longitudinal ridge along the egg and a

combination of measurements including spicule length, tail length, distance to

anterior mamelon and size of eggs. The origins of the genus and the relationships of

the species in the Australian region are discussed.

Key Words: Nematoda, Muridae, Rattus, Syphacia, new species.

Transactions of the Roval Society of S. Aust. (2004), 128(1), 47-51.

SYPHACIA (SYPHACIA) AUSTRALASIENSTIS SP. NOV. (NEMATODA: OXYURIDAE)

FROM RATTUS LEUCOPUS (MURIDAE) FROM PAPUA NEW GUINEA

AND AUSTRALIA

by L. R. SMALES*

Summary

SMALES, L.R. (200) Svphacia (Svphacia) australasiensis sp. noy. (Nematoda: Oxyuridae) from Rattus leucopus

(Muridae) from Papua New Guinea and Australia. Trans. R. Soc. S. Aust. 128(1), 47-51, 31 May, 2004.

Syphacia (Syphacia) australasiensis sp. nov. is described from the caecum of Rattus leucopus (Gray)

(Rodentia: Muridae) from Papua New Guinea and Queensland, Australia. The new species is distinguished from

congeners with an oval cephalic plateau by the lack of lateral alae, a longitudinal ridge along the egg and a

combination of measurements including spicule length, tail length, distance to anterior mamelon and size of

eggs. The origins of the genus and the relationships of the species in the Australian region are discussed.

Key Worbs: Nematoda, Muridae, Rattus, Syphacia, new species.

Introduction

The pin worm genus Syphacia (Nematoda:

Oxyuridae) is a cosmopolitan genus occurring in the

rodent families Cricetidae and Muridae (Hugot

1988). All Australian rodents are contained within

the family Muridae, either a few relatively recent

arrivals, representative of the genus Ratrus

(Murinae), or, the majority of — species

(Hydromyinae), that arrived in Australia from

Indonesia via New Guinea no more than 5-8 million

years ago (Strahan 1995). The currently known pin

worm species associated with the murids from

Australia and the island of New Guinea are

Syphacia muris Yamaguti, 1935, a cosmopolitan

species; Syphacia darwini Quentin & Hugot, 1988

known only from Australian representatives of the

hydromyine genus Melomys and Syphacia

longaecauda Smales, 2001 known only from New

Guinean representatives of the genus Melomys.

Rattus leucopus (Gray) the Cape York rat is one of

only six species of murids that occur in both

Australia and Papua New Guinea (Flannery 1995;

Strahan 1995), Examination of material collected

from both locations revealed a new species of

Syphacia that is described in this paper.

Materials and Methods

Nematodes, previously dissected from R. leucopus

and stored in 70% ethanol in the South Australian

Museum (SAM) or the CSIRO Wildlife Collection,

Canberra (CSIRO), were cleared in lactophenol for

examination. Specimens identified as Syphacia sp.

“School of Biological and Environmental Sciences, Central

Queensland University, Rockhampton Qld 4702, Australia.

Email: Lwarner@cqu.edu.au

were measured using an ocular micrometer and

drawn with the aid of a drawing tube attached to an

Olympus BH microscope. Measurements for 10

individuals of each sex are presented in zm as the

range followed by the mean in parentheses.

Syphacia (Syphacia) australasiensis

(FIGS 1-11)

Holotype

3 from caecum of Rattus leucopus (Gray) Brown

River, Papua New Guinea, 25.vi.1968 coll. W,

Ewers, SAM AHC 32142.

Allotvpe

2 same data SAM AHC 32143.

Paratypes

Same data SAM AHC 5141.

Other material examined

From caecum of Rattus leucopus, E. Mcilwraith

Ra., Cape York Peninsula, North Queensland,

Australia 10.viii.1990, coll. P. Catling and P.

Haycock, CSIRO N3325, 5 9°, 3 d6; Brown

River, Papua New Guinea, 25.vi.1968, coll. W.

Ewers, AM AHC 5201, 5152, 5149, 95 92,16.

Site in host

Caecum and colon.

Description

Small nematodes, typical oxyurid shape, with

transverse cuticular striations. Cephalic inflation

distinct. Cephalic plateau oval, elongated laterally;

distance between amphids 30, amphids situated

between cephalic papillae; cephalic papillae on

lateral projections; six labial papillae; mouth

48 L. R. SMALES

Figs 1-11 Syphacia (Syphacia) australasiensis sp. nov. 1. Male, anterior end, dorso-ventral view. 2. Male, lateral view.

3. Female, lateral view. 4. Anterior end, optical section, dorso-ventral view. 5. En face view. 6. Anterior end, optical

section, lateral view. 7. Spicule and gubernaculum, lateral view. 8. Vagina, lateral view. 9. Male, posterior end, ventra

view. 10. Excretory pore, lateral view. 11. Egg. Scale bars 1, 8,9, 10, 11 25m; 2, 502m; 3, 100pm; 4, 5, 6, 7, 10m.

NEW NEMATODE FROM AUSTRALIA AND NEW GUINEA 49

opening simple, three small distinct pseudolabia.

Oesophagus with isthmus, terminating in spherical

bulb. Excretory pore posterior to oesophageal bulb.

Nerve ring surrounding anterior oesophagus.

Deirids not seen. Alae absent.

Male

Total length 690 — 1250 (810), maximum width

80 — 102 (95). Oesophagus 142 — 208 (192) long,

oesophageal bulb 46 — 55 (51) in diameter. Nerve

ring 75 — 120 (94), excretory pore 228 — 462 (259)

from anterior end. Three mamelons with prominent

annulations, spines not observed; first mamelon 297

583 from anterior end, second mamelon close

behind first, third mamelon close behind second:

mamelons all 35 — 40 long. Tail 142 — 181 (155)

long. Spicule needle shaped 61 — 68 (65) long;

gubernaculum 29 — 36 (31) long, ventral barb not

observed. Two pairs preanal, one pair large post anal

caudal papillae.

Female

Total length 1900 — 2900 (2400), maximum width

160 — 268 (170). Oesophagus 187 — 355 (298) long,

oesophageal bulb 73 — 82.5 (78) in diameter. Nerve

ring 107 — 135 (122), excretory pore 315 — 570

(460). Vulva with prominent knob, 436 — 804 (655)

from anterior end. Tail 335 — 436 (385) long. Eggs

with longitudinal ridge, 82.5 — 95 (87.5) by 29.5 —

36.3 (33).

Etymology

The species is named to reflect that it is found in

both Papua New Guinea and Queensland.

Discussion

The nematode species described above belongs to

the genus Syphacia Seurat, 1916 because it

possesses the suite of characters proposed by Hugot

(1988, see also figs 23, 24 and 39) to distinguish the

genus. It belongs within the subgenus Syphacia

Seurat, 1916 because it has neither the rectangular

cephalic plateau and well developed triangular

shaped lateral alae of the subgenus Crictoxyuris

Hugot, 1988 nor the short conical tail characteristic

of the subgenus Seuratoxvuris Hugot, 1988. Rather,

it has an oval cephalic plateau, lacks cervical alae

and well developed deirids.

Syvphacia (Syphacia) australasiensis n. sp. in en

fuce view most clearly resembles a cluster of species

from murid hosts (Rattus species) from the

Australasian biogeographic region, namely S. muris

Yamaguti, 1935, 8. darwini Hugot & Quentin, 1985,

S. fongaecauda Smales, 2001 and S. sulawesiensis

Hasegawa & Tarore, 1996. It differs from each of

these species in the form of the vulva, having a

longitudinal ridge in the egg and males lacking

lateral alae. It further differs from S. muris in having

an oval rather than square cephalic plateau; from

S. longaecauda in having a smaller cephalic plateau,

30u.m between the amphids rather than 45 — 50pm;

from S. darwini in having three, not two, mamelons

and from S. su/awesiensis in having the females

lacking lateral alae. Syphacia australasiensis differs

from S. /ophuromys Quentin, 1966 and S. megaloon

Quentin, 1966, also lacking lateral alae in the

morphology of the egg and vulva and in en face

aspect. S. lophuromys and S. megaloon have a

cephalic plateau elongated laterally with papillae

and pseudolabia characterised as Group VIII by

Quentin (1971). S. /ophuromys and S. megaloon are

found only in African murids. Further, S.

australasiensis differs from each of the above

species in one or more measurements of

oesophagus, tail, spicule gubernaculum or eggs

(Table 1).

The oxyurids, subfamily Syphaciinae are

generally considered to have coevolutionary

relationships with their hosts (Hugot, 1988). The

similarities between S. australasiensis, S. darwini,

S. longaecauda and S. sulawesiensis are therefore

not surprising. Each has an oval cephalic plateau

with distinct pseudolabia, “lips”, and mouth

surrounded by 6 labial papillae; each occurs in an

endemic murid host, Raflus species or Melomys

species; each has evolved within the Australasian

biogeographic region, Sulawesi being to the east of

Wallace’s line, the boundary between the Asian and

Australasian faunal regions (Raven, 1935).

The relationship between these four species and S.

muris also occurring in Rafts species hosts, but

cosmopolitan in distribution, is not clear. Svyphacia

muris is found in R. ratttus (Linnaeus) and R,

norvegicus (Berkenhout) (see Smales, 1997) both of

which are recent arrivals in Australia. Given that

Hugot & Quentin (1985) found that S. mauris from

endemic Rattus spp. in Australia, corresponded

morphologically and morphometrically to the

description from cosmopolitan Rattus spp., the

suggestion of Hasegawa & Tarore (1996) that

infections in Australian endemic murids have been

acquired from the recent arrivals seems sound.

The endemic Svphacia species may therefore have

been derived from syphaciine populations

introduced into the region as their rodent hosts

invaded the Island of New Guinea and then

Australia (Flannery, 1995). Subsequently speciation

took place in both regions. Syphacia longaecauda

has been found only in Me/omys spp. endemic to

New Guinea while S. darwini is known only from

Melomys cervinipes hosts endemic to Australia. This

suggests that there has been no migration of

Melomys between the two islands. Very few data are

L. R. SMALES

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NEW NEMATODE FROM AUSTRALIA AND NEW GUINEA 51

available from rodent hosts from New Guinea

localities however, so there may be populations of S.

darwini present there yet to be discovered. Rattus

leucopus occurs in both regions and in both cases

harbours S. australasiensis. This suggests that S.

australasiensis migrated with R. leucopus into

Australia. More data from murids across the region

is needed before relationships can be derived and

firm conclusions drawn.

Acknowledgments

Sincere thanks to Jan Forrest from the South

Australian Museum and Dr. David Spratt from

CSIRO Sustainable Ecosystems for lending me the

material.

References

FLANNERY, T. (1995) Mammals of New Guinea (Reed

Books, Chatswood).

HAsecawa, H. & TARORE, D. (1996) Syphacia (Svphacia)

sulawesiensis n. sp. and S. (S.) muris (Yamaguti, 1935)

(Nematoda: Oxyuridae) collected from Rattus xanthurus

(Graf, 1867) (Rodentia: Muridae) in Northern Sulawesi,

Indonesia. Trop. Zool. 9, 165-173.

Hucot, J.-P. (1988) Les nématodes Syphaciine parasites de

rongeurs et de lagomorphes. Mem. Mus. Nat. Hist. Nat.

Sér A 141, 1-148. ‘

& QUENTIN, J.-C. (1985) Etude morphologique

de six espécies nouvelles ou peu connnues appartenant

au genre Syphacia (Oxyuridae: Nematoda), parasites de

Rongeurs Cricétidés et Muridés. Bull, Mus. Nat. Fist.

Nat. Ser: 47, 383-400.

QUENTIN, J.-C. (1966) Oxyures de Muridae africans.

Ann.Parasit. 41, 443-452.

_ (1971) Morphologie comparée des structures

céphaliques et génitales des oxyures du genre Syphacia.

Ann. Parasit. Hum. Comp. 46, 15-60.

RAVEN, H. C. (1935). Wallace’s Line and the distribution of Indo-

Australian mammals. Bull, Am. Mus. Nat. Hist. 68, Article 4.

SMALES, L. R. (1997) A review of the helminth parasites of

Australian rodents. Aust. J. Zool. 45, 505-521.

— (2001) Svphacia longaecauda nn. sp.

(Nematoda: Oxyuridae) Syphacinea from Me/omys spp.

(Muridae: Hydrominae) from Papua New Guinea and

Irian Jaya, Indonesia. Parasite 8, 39-43.

STRAHAN, R. (1995) The Mammals of Australia (Reed

Books, Chatswood).

GASTRIC NEMATODES, INCLUDING A NEW SPECIES OF

ABBREVIATA (NEMATODA: PHYSALOPTERIDAE) FROM

THE MANGROVE MONITOR VARANUS INDICUS

(REPTILIA: VARANIDAE)

By H. I. JONES

Summary

Jones, H. I. (000) Gastric nematodes, including a new species of Abbreviata

(Nematoda: Physalopteridae) from the mangrove monitor Varanus indicus (Reptilia:

Varanidae). Trans. R. Soc. S. Aust. 128(1), 53-59, 31 May, 2004.

Three species of nematode, Tanqua tiara (von Listow, 1879), Heliconema longissima

(Ortlepp, 1922), and Abbreviata melanesiensis sp. nov., were recovered from the

gastrointestinal tracts of 124 Varanus indicus (Daudin, 1802). Tanqua tiara occurred

in two-thirds of the lizards, at a mean prevalence of 7.2 worms per infected host. It is

suggested that the low intensity and geographically variable prevalence of this

nematode may relate to the distribution of the invertebrate intermediate host in the

discrete and discontinuous insular habit of the lizard. Abbreviata melanesiensis sp.

nov. was recovered from two lizards, and is distinguished from other species in this

genus principally by the enlarged tip of the right copulatory spicule, the thick sheath

which envelops the retracted left spicule, the relatively conspicuous phasmids in

females, and the elongated, thin-walled eggs. A single male Heliconema longissima

was recorded.

Key Words: Varanus indicus, Tanqua tiara, Abbreviata melanesiensis, Heliconema

longissima, Solomon Islands, Papua New Guinea, Indonesia, Australia.

Transactions of the Royal Society of S. Aust. (2004), 128(1), 53-59.

GASTRIC NEMATODES, INCLUDING A NEW SPECIES OF ABBREVIATA

(NEMATODA: PHYSALOPTERIDAE) FROM THE MANGROVE MONITOR

VARANUS INDICUS (REPTILIA: VARANIDAE)

by H. IL. JONES

Summary

Jonrs, H. 1. (000) Gastric nematodes, including a new species of Abbreviata (Nematoda: Physalopteridac) from

the mangrove monitor Varanus indicus (Reptilia: Varanidae). Trans. R. Soc. S. Aust. 128(1), 53-59, 31 May, 2004.

Three species of nematode, Zanqua tiara (von Linstow, 1879), Heliconema longissima (Ortlepp, 1922), and

Abbreviata melanesiensis sp.noy., were recovered from the gastrointestinal tracts of 124 Varanus indicus

(Daudin, 1802). Tangua tiara occurred in two-thirds of the lizards, at a mean prevalence of 7.2 worms per

infected host. It is suggested that the low intensity and geographically variable prevalence of this nematode may

relate to the distribution of the invertebrate intermediate host in the discrete and discontinuous insular habitat of

the lizard. Abbreviata melanesiensis sp. nov. was recovered from two lizards, and is distinguished from other

species in this genus principally by the enlarged tip of the right copulatory spicule, the thick sheath which

envelops the retracted left spicule, the relatively conspicuous phasmids in females, and the elongated, thin-

walled eggs. A single male //eliconema longissima was recorded.

Key Worps: Varanus indicus, Tanqua tiara, Abbreviata melanesiensis, Heliconema longissima, Solomon

Islands, Papua New Guinea, Indonesia, Australia.

|

Introduction

The mangrove monitor, Varanus indicus (Daudin,

1802), has a wide and discontinuous distribution

from Sulawesi in eastern Indonesia, east through

Papua New Guinea and northern Australia to the

Solomon, Caroline and Marshall Islands (De Lisle

1996). The taxonomy of the monitor lizards in the

Varanus indicus complex has recently been revised

(Boehme ef a/. 1994, Philipp ef al. 1999) to include

several closely-related species. Varanus indicus

sensu lato is found close to water, on beaches or

riverbanks, and in swampy and mangrove areas

(Cogger 1992). It is largely a terrestrial feeder, and

its diet principally comprises crabs (especially

grapsoid), smaller reptiles, mammals and birds, and

occasionally turtles’ eggs (McCoy 1980). It is a

diurnal lizard, and when alarmed will take to water

(Cogger 1992), although in the Solomon Islands it

will invariably climb a tree (McCoy 1980).

Confusion in the nomenclature of Varanus lizards

has resulted in a number of nematodes being

attributed erroneously to Varanus indicus trom India

(Ortlepp 1922, Mirza 1934, Sharief 1957, Deshmukh

1969, Ali & Ilyas 1969). Since these records are far

to the west of the known geographical range of

V. indicus, and as all state or imply that the lizards

were locally obtained and were not exotic specimens

held in captivity, it is probable that these nematode

Microbiology, School of Biomedical and Chemical Science,

University of Western Australia, Nedlands, Western Australia 6009,

Phone: (08)9346-1270 or (08)9386-6 140.

Email: hjones@cyllene.uwa.edu.au.

records pertain to V. bengalensis (Daudin, 1802), or

possibly V. flavescens (Hardwick and Gray, 1827) or

V. griseus koniecznyi Mertens, 1954 (De Lisle 1996).

Little is known of the internal parasites of Varanus

indicus. The nematode Kalicephalus megacephalus

Schad, 1962 was described from this host from

Florida Island in the Solomon Islands by Schad

(1962). Tangua tiara (Von Linstow, 1879) is

primarily a nematode parasite of lizards in the genus

Varanus, and has been recorded from a number of

Varanus spp. from Africa to southeast Asia and

northern Australia, in aquatic or coastal habitats, but

has not hitherto been reported from /. indicus (see

Gibbons and Keymer 1991). The present study was

therefore undertaken to ascertain the nematode fauna

of Varanus indicus sensu lato, as part of a study of

the gastrointestinal nematodes of varanid lizards.

Materials and Methods

The results of dissections of 124 Varanus indicus

are presented. Thirty seven Vi) indicus held in the

collection of the Australian Museum, Sydney, and

one from the collection of CSIRO Sustainable

Ecosystems, Canberra, were examined. Dennis King

forwarded nematodes recovered from stomach

contents of 20 i. indicus collected from islands and

small archipelagos of Bandaneira, Aru, Kai Besar,

Yamdena and Selaru in the Moluccas in south and

eastern Indonesia during Western Australian

Museum/Museum Zoologicum Bogoriense

expeditions in 1992/1993. In addition, nematodes

from stomachs of sixty-six V. indicus collected at

Maningrida in the Northern Territory by Tim Schultz

54 H. 1. JONES

during 2001 were removed by Alain de Chambrier

and identified by lan Beveridge and the author. The

snout-vent length (SVL) of 58 hosts examined varied

from 135 and 550 mm (mean = 335mm).

Measurements were not available from lizards from

Maningrida. The viscera were removed, and

stomachs (and the intestines from the Australian

Museum specimens) examined for helminths. Food

residues were noted. All nematodes collected were

cleaned, and stored in 70% alcohol with 10%

glycerine. They were subsequently cleared in

chlorolactophenol for examination. All specimens

have been returned to the Australian Museum,

Western Australian Museum, CSIRO and_ the

Australian Helminth Collection respectively.

Results

Helminths recovered. Three species of nematode

were recovered. The predominant species was

Tanqua tiara, which was present in 75/124 lizards

(60.5%, Fig. 1); specimens lodged as Australian

Museum W 28660-W 28674; CSIRO Canberra

N5294: Western Australian Museum V 4374 and V

4375, and in the Australian Helminth Collection.

Prevalence was highest in the lizards from

Maningrida (84.8%), and lowest from the Moluccas

and Solomon Islands (32.8%). Total mean intensity of

T. tiara was 7.2 worms/host. Four Vo indicus,

collected from Adelaide River and Maningrida in the

Northern Territory, contained >20 7: tiara (22 — 123);

in the remaining 71 infected hosts 7. tiara numbers

ranged from | — 16 (mean: 4.2). In four hosts some

nematodes were attached with their heads buried in

the stomach mucosa. A single male //eliconema

longissima (Ortlepp, 1922) was recovered from a

lizard from Maningrida in the Northern Territory. A

new species of Abbreviata was recovered from two

hosts collected from New Britain and from Woodlark

Island, Papua New Guinea.

There was no relation between the size of the host

lizard, type of food residues in the stomach, and the

presence or intensity of infection with 7) tiara. Both

prevalence and intensity of this worm from lizards

collected from the Moluccan islands in south-eastern

Indonesia were lower (2 of 20 infected with one and

two worms, respectively) than those collected from

Papua New Guinea or the Solomon Islands, with

highest prevalence and intensity in lizards from

northern Australia. Neither of the two lizards with

the new species of Abbreviata was concurrently

infected with 7. tiara. One of the two lizards infected

with this species of Abbreviata was one of the largest

dissected (500 mm SVL).

Food residues. In the 37 V. indicus from Papua New

Guinea and the Solomon Islands, crustacea, in three

I I

130 E 140

NORTHERN

TERRITORY

SOLOMON

ISLANDS &

Fig. 1. Map showing distribution of Zangua tiara in Varanus indicus. Open circle, lizard without infection; black circle,

lizard with infection. M: Maningrida, where 56 of 66 lizards were infected. (No location data available for two lizards).

PARASITIC NEMATODES IN V4ARANUS INDICUS

cases identifiable as crabs. formed the most

commonly recovered food source, and occurred in 16

lizards. In addition, a snake (Candoia sp.; Boidae), an

agamid lizard skull, an unidentified lizard, a frog,

reptile eggs, cicadas, a grasshopper, and a beetle were

each recovered from a single host stomach, and

vertebrae from an unidentified animal, and molluscs

from two hosts each. Stomach contents from the 20

lizards from the Indonesian islands included 10 with

insect remains, one with a gecko, one with a bird, one

with reptile eggs, one with a crab, and four with plant

residues. Twenty one of these fifty seven stomachs

had no food residues; no fish remains were identified.

Taxonomy

Order Spirurida

Superfamily Physalopteroidea

Family Physalopteridae

Genus Abbreviata Travassos, 1920.

Abbreviata melanesiensis sp. Nov.

Holotype

Male, Australian Museum W 28675, from stomach

of Varanus indicus, AM R129614, Amelei village,

West New Britain, Papua New Guinea (06° 06’ S,

150° 37’ E), collected 27.x11. 1988.

Allotvpe

Female, AM W 28676, same data as for holotype.

Paratypes

16 males, 9 females, 4 larvae/immatures, AM W

28677, same data as for Holotype. Additional non-

type specimens: three males, four females and two

immatures, AM W 28678, from /. indicus R124815,

collected Guasopa, Woodlark Island, Milne Bay

Province, Papua New Guinea (09° 15’ S, 152°, 56’

E), collected 09.viil.1987.

Diagnosis

With characteristics of the genus, viz. two large

lateral pseudolabia, with externolateral tooth present

on each pseudolabium, a bifid internolateral tooth

and two double pairs of submedian teeth; male with

wide ornamented caudal alae united anteriorly on

ventral surface of body, supported by four pairs of

pedunculate papillae, markedly unequal spicules,

females with vulva in anterior portion of body.

Mouth with dorsal and ventral corner denticles.

Spicules well sclerotised, left four times length of

right; left spicule in thick walled sheath, terminating

in a very fine point, right spicule curved ventrally,

with pointed, weakly-sclerotised enlargement at tip.

Female tail slightly attenuated, vulva flush with body

wall without extensions; eggs thin-shelled, elongate,

length approximately twice width.

Wn

al

Description (Fig. 2)

Small to medium worms, males not greatly smaller

than females, tapering at both ends, fine transverse

cuticular striations. Mouth surrounded by two

pseudolabia, each bearing large externolateral apical

tooth and small bifid internolateral tooth. Bifid

submedial tooth on dorsal and ventral border of each

pseudolabium. Four to 5 small, regular denticles at

dorsal and ventral median surface of each

pseudolabium. Two sessile papillae and amphid on

external surface of each pseudolabium. Cervical

collar present. Nerve ring surrounds muscular

oesophagus near its posterior end. Glandular

oesophagus of uniform width, wider than muscular

portion. Cervical papillae and excretory pore on

external surface posterior to origin of glandular

oesophagus.

Male

Caudal alae meet anteriorly, extend just beyond

tip of tail posteriorly. Alae supported by four pairs

of pedunculate pericloacal papillae and three pairs

of very short pedunculate or sessile caudal papillae,

of which the central pair is about one-third the

distance between the first and second pair. Papillae

on ventral surface sessile, 3 immediately anterior to

the cloaca, and 2 pairs immediately posterior to the

cloaca. Caudal tubercles arranged in rows,

extending from anterior border of alae, converging

towards cloaca, and running parallel to one another

lateral to cloaca on ventral surface of tail and

adjoining alae; diminishing in size posteriorly,

terminating as small scattered tubercles at level of

posteriormost caudal papillae. Spicules dissimilar,

unequal. Left spicule approximately 4 times length

of right, uniformly sclerotised, terminating in very

fine tapering point, enclosed in conspicuous thick

sheath. Right spicule thicker than left, heavily

sclerotised, curved ventrally, and with a weakly-

sclerotised pointed enlargement, slightly concave

on one side, at the tip.

Female

Tail short, terminating in a slight attenuation;

phasmids relatively conspicuous at two-thirds

distance along tail. Vulva a transverse slit flush with

body wall, without extensions or altered adjacent

cuticle, posterior to commencement of intestine,

about one-quarter of distance from anterior end of

worm. Two ovaries, uterus with four branches, one

of which often extends a variable distance anteriorly

beside the posterior portion of glandular oesophagus.

Eggs with smooth thin shells, elongate, embryos not

visible, length almost twice width.

Measurements (mm; range with mean in brackets)

Males (N:6): length 18 — 24 (21.7); maximum width

0.36 — 0.54 (0.45); muscular oesophagus length 0.26

56 H. I. JONES

Fig. 2. Abbreviata melanesiensis sp. noy. A, anterior end, male paratype, lateral view; B, anterior end, en face, female

paratype; C, male tail, slightly-oblique ventral view; D and E, distal tip of right spicule, male paratype; F, tail, female

paratype, lateral view; G, eggs in oviduct and vulva, female paratype, lateral view; H, egg. Scale bars, E and H: 100um,

all other figures, 200um.

PARASITIC NEMATODES IN VARANUS INDICUS 37

— 0.36 (0.33); muscular oesophagus width 0.10

0.14 (0.12); glandular oesophagus length 2.8 — 3.3

(3.2); glandular oesophagus width 0.24 — 0.28 (0.26);

nerve ring* 0.26 — 0.38 (0.32); cervical papillae*

0.36 — 0.68 (0.53); excretory pore* 0.56 — 0.72

(0.61); tail 0.96 — 1.32 (1.16); distance between

caudal papillae | and 2, 0.056 — 0.16 (0.11); distance

between caudal papillae 2 and 3, 0.14 — 0.19 (0.16);

left spicule 1.320 — 1.680 (1.470); right spicule 0.280

— 0.360 (0.320). Females (N: 6) length 26 — 33

(29.3); maximum width 0.62 — 0.74 (0.70); muscular

oesophagus length 0.36 — 0.44 (0.41); muscular

oesophagus width 0.14 — 0.18 (0.15): glandular

oesophagus length 3.36 — 4.40 (3.93); glandular

oesophagus width 0.26 — 0.41 (0.32); nerve ring*

0.34 — 0.38 (0.36); cervical papillae* 0.48 — 0.60

(0.54); excretory pore* 0.62 — 0.86 (0.70); tail 0.34 —

0.44 (0.38); vulvat 1.9 — 3.7 (2.78); 19.4% — 27.8%

(23.3%) of distance from anterior end (* distance

from anterior end; + distance posterior to oesophago-

intestinal junction).

Discussion

Taxonomy

Abbreviata melanesiensis sp. nov. is distinguished

from other species of Abbreviata occurring in

reptiles from the Australo-Papuan region by a suite

of characters. The size, general form, characteristics

of the anterior end and the male copulatory bursa are

shared with a number of other species. However, the

weakly-sclerotised enlargement at the tip of the

right spicule 1s unique; a similar feature occurs in

A, hastaspicula Jones, 1979, in which species

however it is well sclerotised (Jones 1979).

Furthermore, in that species the left spicule is

considerably shorter (0.620 — 0.700 mm) and the

right spicule longer (0.590 — 0.670 mm: Jones 1979)

than in A. melanesiensis sp. nov., and the

sclerotisation of the left spicule often appears

discrete and discontinuous; in addition, in A.

hastaspicula the female bears a tubular extension

from the vulva. Eggs of A. me/anesiensis sp. nov. are

elongate and thin walled; most species of Abhreviater

in this region possess eggs with thicker and denser

shells (Jones 1983b, 1988) except A. hastaspicula

and 4. perenticola Jones, 1985, whose thin shelled

eggs are subspherical. Other characteristics which

differentiate this species include less disparity in

length between males and females, the thick sheath

that envelops the retracted left spicule, and the

readily visible phasmids on tails of females. In other

species of Ahbbreviata described from Australia and

Papua New Guinea, except A. /evicauda Jones, 1983,

from V. tristis (Schlegel, 1839) the penultimate

caudal papillae are closer to the anterior papillae

(Jones 1986) than in A. melanesiensis sp. noy. The

enlargement at the tip of the right spicule

distinguishes 4. melanesiensis sp. noy. from six

other species of Abbreviata recorded from Papua

New Guinea, viz. 4. oligopapillata (Kreis, 1940)

(see Jones 1979), A. multipapillata (Kreis, 1940),

A. natricis (Kreis, 1940), A. heterocephala

(= Kreisiella) (Kreis, 1940), A. confusa Johnston &

Mawson, 1942, (see Jones 1983a) and A. katilensis

(Jones, 1979). A. borneensis Schad, 1959 from V.

rudicollis Gray, 1843 in Sarawak also has the

posterior portion of the tail free from tubercles, as in

A. oligopapillata (Schad 1959). In A. melanesiensis

sp. nov. the delicate enlargement at the tip of the

right spicule is not visible unless this is extended

through the cloaca, and if this spicule is retracted,

dissection is necessary to ascertain its character; all

specimens in the type host had the right spicules

retracted, but in all four males in the second host the

right spicules were extended and this character was

clearly visible. The small denticles at the dorsal and

ventral mouth margins are often not visible if

obscured by the cervical collar.

Biology

The absence of a relationship between host size

and numbers of 7. tiara contrasts with the findings of

Shine e¢ al. (1998), who found that 7) tiara were

present at a higher intensity in juvenile Mi sa/vator,

They also found a significant difference in infection

with this worm between two sites in Sumatra, though

the intensity of worms was similar to findings from

V. indicus in the present study (mean 6.1 worms per

host). The arthropod intermediate host for 7. tiara is

not known, but as Varanus specimens infected with

this worm inhabit swampy, mangrove or riverine

habitats, a crustacean intermediate host is possible.

Although V. indicus feed on a wide range of prey, the

fact that crustacean remains were found in 16 of 37

from Papua New Guinea and the Solomon Islands,

and in only 2 of 20 from the Moluccas, (food data not

available from lizards from Maningrida) illustrate

local or regional differences in prey availability. The

distribution of the worm in the present study may be

related to the discontinuous nature of insular

habitats. In the present study highest intensity of 7.

tiara, 22 123 worms, occurred in indicus

examined from northern) mainland Australia

(Adelaide River and Maningrida), which may

provide both more opportunities for the spread of

worms and intermediate hosts than from small

isolated habitats. Furthermore, 7: fiara have been

reported from V. panoptes Storr, 1980, Vo gouldii

(Gray, 1838) and Vo mertensi Glauert, 1951, from

aquatic habitats in northern Australia at mean

intensities of 18.9 (max. 120 worms), 20.5 (max. 50

worms) and 7.2 (max. 30 worms) respectively; an

absence of correlation with Abbreviata spp. infection

58 H. L. JONES

was also noted in that study (Jones 1988). The

similar species 7) ophidis Johnston & Mawson,

1948, described from the aquatic file snake

Acrochordus sp. was present in all eight Acrochordus

sp. examined from north and northwest coastal

Queensland, at numbers ranging from seven to 179

per host (mean, 51); (Jones 1978). As K. indicus,

Acrochordus arafurae McDowell, 1979 and

A. granulatus (Schneider, 1799) are sympatric over

much of their range, it is possible that these two

species of Zanqua are also sympatric.

The single male #7. /ongissima reported in the

present study was probably an accidental infection,

taken with infected prey. The type host and type

locality of Heliconema longissima (Ortlepp, 1922)

are given as ‘snakes, Australia’, although all other

specimens have been recorded from anguilliform

fish, and Ogden (1969), considered that the type host

identification is probably in error. However, H.

longissima occurred in 3/5 specimens of the aquatic

colubrid snake Fordonia leucobalia (1, 23 and 41

nematodes per host; Jones 1978).

The scarcity of Abbreviata melanesiensis sp. nov.

in Ki indicus contrasts with findings of prevalence

and intensity of Abbreviata spp. in other species of

Varanus and other large terrestrial reptiles in

northern and arid Australia, in several species of

which infection with Abbreviata spp. occurs at high

prevalence and intensity (Jones 1983b, 1988).

Epidemiological evidence suggests that termites

might have a role to play in the life-cycles of species

of Abbreviata from arid regions (Jones 1995), Since

only two K. indicus were infected with Abbreviata

sp., no conclusions can be drawn from the absence of

concurrent infection with 7) tiara, Since Varanus

indicus feeds on a wide variety of invertebrate (and,

in the larger specimens, vertebrate prey; McCoy

1980) the nematodes present may relate to

differences in diet; the predominance of aquatic prey

in these lizards suggests that the intermediate hosts

of T. tiara may be aquatic invertebrates. The fact that

one of the two hosts infected with A. melanesiensis

was particularly large may be significant. The larger

of the two infected lizards contained a Candoia sp.

snake prey item, and though it is possible that this

Abbreviata infection was spurious, this is unlikely as

several of the nematodes were already attached to

and apparently feeding on the external surface of the

snake.

Conclusion

This study demonstrates that despite the wide

range of prey items consumed by indicus, this

lizard supports gastric nematodes in low numbers,

and at a moderate prevalence, with only three species

recorded. No intestinal nematodes were recovered. It

is possible that the large range of prey types may

inhibit the development of parasite cycles,

particularly in parasite species with narrow

intermediate-host specificity. The low intensity of 7.

fiara and geographically uneven distribution may

relate to the insular and discontinuous nature of the

region from which FV. indicus was examined; in this

regard the higher intensities of this nematode

recorded from other species of Varanus in northern

mainland Australia may be significant. Further

conclusions cannot be drawn without knowledge of

the arthropod intermediate hosts required by this

nematode to complete its life-cycle. Studies on the

gastrointestinal parasites of other larger reptiles in

this region may reveal a wider range of hosts for

A. melanesiensis.

Acknowledgements

| thank Allen Greer for allowing me access to the

lizards under his care in the Australian Museum, and

for providing facilities, Ross Sadlier for practical

assistance, David Spratt for allowing me to examine

a lizard in the collection of CSIRO Sustainable

Ecosystems, Canberra, and Brad Maryan for

assistance at the Western Australian Museum. | am

grateful to Alain de Chambrier for collecting

nematodes from lizards collected at Maningrida; to

lan Beveridge for identifying and forwarding these

specimens, and to the late Dennis King for collecting

nematodes during the course of his own studies, and

to Mrs. Ruth King for making his papers available

for my perusal.

References

Al, S. M. & Ityas, R. (1969) Neoxysomatium

longicaudatum n, sp. from Varanus indicus in

Marathwada, India. Marathwada Univ. J. Science 8, 73-

75.

BOEHME, W., HorN, H. G. & ZIEGLER, T. (1994) On the

taxonomy of the Pacific monitor lizards (Varanus indicus

complex): resurrection of Varanus doreanus (A. B.

Meyer, 1874) and description of a new species.

Salamandra 30, 119-142.

CoacGcpr, H. G. (1992) Reptiles and Amphibians of

Australia. (Sth Edition, Reed, Chatswood, NSW 2067; I-

775).

Dr Liste, H. F. (1996) The Natural History of Monitor

Lizards. Krieger Publishing Company, Malabar, Florida,

1-201.

DesHMukH, P. G, (1969) A new species of the genus

Herpetostrongvlus (Nematoda) from Varanus indicus.

Vest. Cesk. Spol. Zool. 33, 211-213.

PARASITIC NEMATODES IN VARANUS INDICUS 59

Gippons, L. M. & Krymer, I. F. (1991) Redescription of

Tanqua_ tiara (Nematoda, Gnathostomatidae), and

associated lesions in the stomach of the Nile monitor

lizard (Varanus niloticus). Zool. Scr. 20, 7-14.

Jones, H. I. (1978) Gastrointestinal nematodes from aquatic

Australian snakes. Mem. Queens! Mus.18, 243-254.

(1979) Nematodes from Papua New Guinean

snakes. Mem. Queensl. Mus.19, 393-397.

(1983a) A collection of nematodes from snakes

from Papua New Guinea. Syst. Parasitol. 5, 131-134.

— (1983b) Abbreviata (Nematoda: Physalop-

teroidea) in lizards of the Varanus gouldii Complex

(Varanidae) in Western Australia. Aust. J. Zool. 31, 285-

298.

: (1986) Differences in caudal morphology in

male Abbreviata levicauda (Nematoda: Physalop-

teridae)in two sympatric species of Varanus (Reptilia:

Varanidae). J. Parasitol. 72,185-186.

Varanus (Reptilia) from tropical northern Australia, with

particular reference to the genus Abbreviata

(Physalopteridae). Aust. J. Zool. 36, 691-708.

(1995) Gastric nematode communities in

lizards from the Great Victoria Desert, and an hypothesis

for their evolution. Aust. J. Zool. 43, 141-164.

(1988) Nematodes from nine species of

MeCoy, M. (1980) Reptiles of the Solomon Islands. Wau

Ecology Institute. Handbook 7. Wau Ecology Institute,

Wau, Papua New Guinea.

Mirza, M. B. (1934) Sciurus palmarum als ein interessante

Wirt von Physaloptera sp. Z. f. ParasitKde 6, 638-641.

OGpEN, C. G. (1969) A revision of the genus Heliconema

Travassos, 1919, Physalopteridae (Nematoda). J. Nat.

Hist. 3, 423-431.

OrtLepp, R. J. (1922) The nematode genus Phvsaloptera

Rud. Proc. Zool. Soc. Lond. 999-1107.

Puiuipe, K. M., BorumMe, W. & ZEIGLER, T. (1999) The

identity of Varanus indicus; redefinition of a sibling

species coexisting at the type locality (Sauria, Varanidae,

Varanus indicus group). Spixiana 22, 273-287.

ScHAbD, G. A. (1962) Studies on the genus Kalicephalus

(Nematoda: Diaphanocephalidae). Il. A taxonomic

revision of the genus Kalicephalus Molin 1861. Canad.

J. Zool. 40, 1035-1065.

SHARIFF, A. (1957) On a new species of trichostrongylid

nematode from Hyderabad, India. Ann. Mag. Nat. Hist.

ser.12, 10, 705-709.

SHINE, R., AMBARIYANTO, HARLOW, S. & MUNPUNI (1998)

Ecological traits of commercially harvested water

monitors, Varanus salvator, in northern Sumatra.

Wildlife Res. 25, 437-447.

ENRICHMENT-PLANTING OF THE WOODY CLIMBERS

MARSDENIA AUSTRALIS AND RHYNCHARRHENA LINEARIS

IN NORTH-WESTERN VICTORIA

By R. F. PARSONS*

Summary

Parsons, R. F. (2000). Enrichment-planting of the woody climbers Marsdenia australis

and Rhyncharrhena linearis in north-western Victoria. Trans. R. Soc. S. Aust. 128(1),

61-66, 31 May, 2004.

Marsdenia australis and Rhyncharrhena linearis are woody climbers (family

Asclepiadaceae) found in semi-arid to arid areas of all mainland states of Australia.

Where studied, they are declining drastically, probably due to grazing. In Victoria,

their conservation status is vulnerable. In 1991, in Hattah-Kulkyne National Park, a

site was found carrying single plants of each species. At this site, 384 seedlings and

1,741 seeds of Marsdenia and 240 seedlings of Rhyncharrhena were planted

underneath trees of Eucalyptus socialis in 1991-1992, half of the seedlings in

mammal-proof cages. After high seedling mortality due to mammal grazing and

especially to drought in the first summer, only six of the planted seedlings, all of

Marsdenia, survived by the end of 1992. All of these, and a further 15 Marsdenia

plants derived from seed sowing, have survived until present. Most of them have

climbed up nets and ropes provided onto the stems of three Eucalyptus host plants and

at least two of them have produced fruits. Marsdenia is likely to be an obligate

outbreeder, in which case the genotypes introduced to the site will be vital in allowing

future seed production there. Implications of the work for conservation and

management are stated.

Key Words: Drought, enrichment planting, kangaroo grazing, mallee vegetation,

Marsdenia australis, Rhyncharrhena linearis, seedling mortality, woody climbers.

Transactions of the Roval Society of S. Aust. (2004), 128(1), 61-66.

ENRICHMENT-PLANTING OF THE WOODY CLIMBERS

MARSDENIA AUSTRALIS AND RHYNCHARRHENA LINEARIS

IN NORTH-WESTERN VICTORIA

by R. F. PARSONS"

Summary

PARSONS, R. F. (2000). Enrichment-planting of the woody climbers Marsdenia australis and Rhyncharrhena

linearis in north-western Victoria. Trans. R. Soc. S. Aust. 128(1 ), 61-66, 31 May, 2004.

Marsdenia australis and Rhyncharrhena linearis are woody climbers (family Asclepiadaceae) found in semi-

arid to arid areas of all mainland states of Australia. Where studied, they are declining drastically, probably due

to grazing. In Victoria, their conservation status is vulnerable. In 1991, in Hattah-Kulkyne National Park, a site

was found carrying single plants of each species. At this site, 384 seedlings and 1,741 seeds of Marsdenia and

240 seedlings of Rhvacharrhena were planted underneath trees of Eucalyptus socialis in 1991-1992, half of the

seedlings in mammal-proof cages. Afier high seedling mortality due to mammal grazing and especially to

drought in the first summer, only six of the planted seedlings, all of Marsdenia, survived by the end of 1992. All

of these, and a further 15 Marsdenia plants derived from seed sowing, have survived until present. Most of them

have climbed up nets and ropes provided onto the stems of three Eucalyptus host plants and at least two of them

have produced fruits. Marsdenia is likely to be an obligate outbreeder, in which case the genotypes introduced

to the site will be vital in allowing future seed production there. Implications of the work for conservation and

management are stated.

Kry Worps: Drought, enrichment-planting, kangaroo grazing, mallee vegetation, Marsdenia australis,

Khyncharrhena linearis, seedling mortality, woody climbers.

Introduction

Two species of lianes (woody climbers) in the

family Asclepiadaceae, Marsdenia australis and

Rhyncharrhena linearis, occur in semi-arid and arid

areas of all mainland states of Australia. In Victoria,

both species are declining drastically, probably due

to grazing. While small sucker shoots can be found

when grazing pressure is low, no seedlings at all can

be found, even in the absence of vertebrate grazing

(Nichols, Browne & Parsons, 1991). The primary

aim of the work was to investigate seed and seedling

survival in the field to assist the conservation

management of the species, both of which have a

conservation status currently rated as vulnerable

Victoria-wide (Victoria: Department of Natural

Resources & Environment, unpublished). However,

we hoped that any plants surviving from the

experiments would be left to augment naturally-

occurring populations. The work began as a B.Sc.

(Honours) project from February to September 1991

(Smith, 1991).

Methods

Hattah-Kulkyne National Park was chosen for the

work, being an extensive biological reserve carrying

both species. At present, only 12 naturally-occurring

Marsdenia plants are known there (Fiona Murdoch,

" Botany Department, La Trobe University, Bundoora, VIC 3086,

Australia.

pers. comm., 20 December 2002). The work was

carried out near one of these, which has a

Rhyncharrhena plant nearby.

The study site in Hattah-Kulkyne National Park is

within the Mournpall Block, 0.6 km along Jasmine

Track from its N end, then 0.2 km WSW of the Track

(Fig. 1). The fenced Rhyncharrhena plot no. 9E is

about 50 m away.

The area is Eucalyptus socialis tall shrubland

(‘mallee scrub’) on sandy loam topsoils. After

various episodes of heavy grazing by stock and

rabbits since 1847, stock grazing was stopped in

1974 (Parks Victoria records). Subsequent rabbit

control meant that rabbit numbers were ‘very low’

during the first year of the project (D. Major personal

communication). There was also grazing by a

population of western grey kangaroos, estimated at

800 in the 5,700 ha Mournpall Block during 1991

(D. Major personal communication).

All seed used was collected between 1988 and

1991 from the Red Cliffs area (50 km from the

plots), Marsdenia from sites | and 3 and

Rhyncharrhena from site | of Nichols, Browne &

Parsons (1991), Germinability of seed lots was

established in a growth cabinet using a 10 hour

photoperiod at 25/15° C after plume removal.

Seedlings to transplant into the field were produced

by planting germinated seed into sandy loam,

vermiculite and perlite (2:1:1) in pots in the same

growth cabinet. At four weeks of age they were

placed outside to harden and then planted at the field

site.

62 R. F. PARSONS

Hattah ,

Camping

Ground

HATTAH - KULKYNE

NATIONAL

x

PARK

+ 34°50'S

142° 20E NS

> eS .

: vv,

Stockyard Track

HIGHWAY

Fig. |. Location of study area in north-western Victoria. Study site shown as cross west of Jasmine Track.

For both species, as fruiting seems only to occur in

plants climbing in shrubs or trees (Nichols, Browne

& Parsons, 1991), all seedlings were planted beneath

low branches of mature Eucalyptus socialis trees.

Four trees growing in an area of 50 m radius were

used (Fig. 2) with four sites selected beneath each

tree crown. For Marsdenia seedlings, at each of four

planting dates from March to July 1991, one site

under each tree was chosen at random and planted,

providing four replicates. Each site consisted of a

caged and an uncaged plot (for seedlings) side-by-

side. Cages were 40 x 40 x 30 cm. Thus for

Marsdenia the design was 4 trees x 4 planting times

x 2 treatments (caged/uncaged) x 12 seedlings = a

total of 384 seedlings planted. The Rhyncharrhena

plantings were similar, but only 240 seedlings were

available.

Before planting, litter and understorey plants were

removed and plots made level by rake. Seedlings

were planted 10 cm apart, the species intermingled.

The wire mesh cages were flanged at the base to

deter rabbit entry via burrowing.

The plots were watered to try to minimize seedling

deaths from water deficiency. At each planting date,

enough water to saturate the surface 5 cm of soil was

added, both to the plots to be planted and to all plots

planted previously. Water poured onto the sites was

stopped from running off by infiltrometer rings put

in place until infiltration was complete.

At all planting and inspection dates, all herbs other

than Marsdenia and Rhyncharrhena were identified,

counted and removed from the plots by manual

uprooting. The plots were monitored in September

1991, four times in 1992-4 and finally in May 2002

(Table 2).

Eleven Marsdenia fruits containing a total of 1741

filled seeds were left over from the 1991 experiment.

We removed their plumes and sowed them under a

shallow soil covering in three of the cages on 2

March 1992 (rather than discard them). At the same

time, surviving seedlings were marked with stakes to

distinguish them from plants of seed origin. On 5

May 1994, nets or ropes made of natural fibres were

positioned to provide all surviving plants with a

pathway so that they could twine upwards into

Eucalyptus crowns. This necessitated cutting the

tops out of the cages.

Results

Survival of planted seedlings inside cages, after a

winter of mild temperatures (coldest single night of

1° C) and before the severe water deficit of summer,

was high (84 — 85%) for both species (Table 1).

Judging from plant symptoms and climatic data, by

far the main cause of mortality in the cages up until

September 1991 was soil water deficit.

Analysis of bite marks suggested that the much

lower survival outside cages (49 — 52%) was due to

either rabbit or kangaroo grazing.

ENRICHMENT-PLANTING OF MARSDENIA 63

TABLE 1. Number of survivors on 3 September 1991 of

seedlings planted from March to July 1991,

Number Number Percentage

planted surviving survival

Marsdenia

Caged 192 164 85

Uncaged 192 100 52

Rhyncharrhena

Caged 96 81 84

Uncaged 144 71 49

TABLE 2. Height, leaf number and survival at five dates of

384 seedlings of Marsdenia australis planted from March

to July, 1991. na = not available.

Date Sep Mar Dee Dec May May

91 92 92 93 94 02

Age (yr) <l | 1.5 2.5 3 U1

(approx.)

Number of 264 9 6 6 6 6

survivors

Mean height na 2 5 19 na 163

(cm)

Mean leaf na 6 8 21 na na

number

Pee cole 2, 75m S of

x plot corner

K Tree 1, 15m from

edge of plot

Marsdenia plot

a

_—

i 10m

ae Tree 3, 53m E of

plot corner

Fig. 2. Location of the three trees where Marsdenia has

been established near Jasmine Track.

TABLE 3. Height, leaf number and survival at four dates of

plants which germinated from 1741 seeds of Marsdenia

sown on 2 March 1992. na = not available.

Date Dec Dec May May

92 93 94 02

Age 9mo lyr9mo 2yr3mo_ 10 yr3 mo

Number of 22 67 77 15

plants alive

Maximum 3 7 na 270

height (cm)

Maximum 6 14 na na

leaf number

The alien annual weed Brassica tournefortii

(Mediterranean Turnip) was always by far the most

common species appearing in the plots as seedlings

(cover from 45 to 80%) followed in order by the

natives Sclerochlamys brachyptera (Short-wing

Saltbush) and Zygophyllum spp. (Twinleaf) and the

alien Salvia verbenaca (Wild Sage).

After water deficits in summer, by March 1992 all

152 Rhyncharrhena seedlings and all except nine of

the 264 Marsdenia seedlings had died (Table 2),

presumably mostly from drought and to a lesser

extent from grazing (all but one of the nine were

inside cages). Number of survivors fell to six caged

plants by December 1992 but all six were still alive

more than 11 years later (Table 2).

The seeds sown in March 1992 germinated at

various times between then and May 1994, when 77

plants were seen. Fifteen of these were still present

ten years later (Table 3). One tree crown had no

surviving Marsdenia seedlings and is not shown in

Fig. 2.

In the absence of other support, it is common for

Marsdenia stems to twine around themselves to form

tightly plaited ropes which can be up to 6 m long and

35 mm in diameter (see Fig. 7 in Nichols ef a/.,

1991). This occurred beneath all three host trees

which had surviving Marsdenias under them in the

present study, each plait involving from two to seven

individual plant stems (Table 4). In fact all plants

which reached from the soil surface to a host tree

were members of such plaits or systems of stems.

Marsdenia is an apical stem twiner, a behaviour

said to be effective in climbing tree trunks of up to

100 mm diameter at breast height but seldom bigger

ones (Hegarty and Clifford, 1984). In the present

study, by May 2002, only one Marsdenia stem had

completely encircled the stem of a host tree: a stem

of 50 mm diameter belonging to host tree 3, where

the second system of Marsdenia stems is also firmly

attached (at a host stem fork; Table 4). However, the

only other two systems of stems to have reached a

stem of their host tree (at host trees | and 2; Figures

3 and 4; Table 4) were less securely attached, with a

64 R. F. PARSONS

Fig. 3. Cage with top cut out at host tree 1. Marsdenia has Fig. 4. Cage with top cut out at host tree 2. Marsdenia has

climbed the net provided to reach stem of host climbed the rope provided to reach stem of host. Scale as

(Marsdenia not clearly visible). Scale graduated in 10 cm in Fig. 3. 14 May 2002.

intervals. 14 May 2002.

TABLE 4. Height, stem diameter and notes on attachment and reproduction of Marsdenia plants on 14 May 2002, including

plants derived from planting both seeds and seedlings.

Location No. of | Maximum Maximum stem — Attachment Reproduction

plants height (m) — diameter (mm) to host

a) Original (non-planted) | 6 8 Fully attached; 28 suckers <20 cm

plant in fenced plot climbing into high up to 8 m

crown. from parent plant.

b) Host tree 1 4 1.2 2 Three plants twined Two suckers climbing

together up net and in dead annual

attached to bark of plants in cage

big host stem.

c) Host tree 2 2 1.8 3 The 2 plants twined None

together up rope;

supported by upper

knot in rope. Two

shoot tips inside

bark of host.

d) Host tree 3

(i) from seedlings 3 1.6 4 The 3 plants twined One young,

together up net. Now attached fruit

firmly attached to

stem fork of host.

(ii) from seeds 12 2.7 4 Seven plants twined Four attached,

together up net. mature fruits.

Firmly twined around Three suckers

host stem.

ENRICHMENT-PLANTING OF MARSDENIA 65

few shoot tips held between attached dead bark and

the main stem of the host (Table 4). Because of this,

and because the host stems may be too large for easy

Marsdenia encirclement (tree | stem diameter 230

mm; tree 2, 170mm), the Marsdenia stem systems

have since been tied to these host trees (Fiona

Murdoch, pers. comm., 5 December 2002).

While stem diameters of long-established

Marsdenia plants reach 8 to 10 mm (Nicholls e¢ a/.,

1991; Table 4), the I1l-year-old plants have only

reached a maximum of 4 mm (Table 4). Some young

plants have already produced shoot suckers from the

roots as 1s normal in adult plants of Marsdenia

(Table 4). In the absence of support, such shoots

remain small and fail to produce fruit (Nichols e7 al.,

1991). Two plants carried fruit in May 2002 (Table

4) although it 1s not known if they represent the first

fruit crop.

Discussion

In an area which carried a single naturally-

occurring Marsdenia plant, the present work has

established an additional 21 individuals on three host

trees. The conditions provided included complete

protection from vertebrate grazing for three years

and supplementary watering and removal of

competing seedlings in autumn-winter of year one.

The huge mortality late in year one was not

surprising given that, in this climate, summer

drought is the main cause of death for perennial

seedlings in their first year (see e.g. Parsons, 1994).

For efficiency, future enrichment-plantings in this

climate should allow for summer watering in case of

drought, despite the possible favouring of unsuitable

genotypes.

Hobbs and Atkins (1991) also recorded the

presence of dense, non-native annuals including

Brassica tournefortii.. Future workers in- mallee

vegetation should note that such annuals are capable

of severely inhibiting regeneration by native

perennials (Hobbs and Atkins 1991). In the present

study, it is not clear to what extent clearing the plots

influenced density of alien annuals.

In Marsdenia as in other asclepiad vines, root

suckering allows clones to establish and it is possible

that such clones may be capable of persisting more

or less indefinitely (e.g. see Penalosa, 1984).

However, it 1s common for species in family

Asclepiadaceae to be obligate outbreeders. In such

species, clone formation can cause extensive self-

pollination and hence cause fruit set to be low or

absent (see references in Nichols ef a/., 1991). In the

present case, the genotypes we introduced from

outside the area may be vital in allowing outbreeding

and hence seed production.

The greater success of Marsdenia planting than

Rhyncharrhena may be because the tuberous root of

Marsdenia gives it greater drought avoidance than

the latter (see Nichols e7 a/., 1991).

Work is now needed to check the conservation

status of Marsdenia and Rhynacharrhena in other

areas of inland Australia. Although Marsdenia was

recorded from as many as 19 out of 1,143 sites in the

Stony Deserts of South Australia (Brandle 1998), we

need to know to what extent grazing and other

factors are allowing current recruitment both here

and elsewhere. In Victoria, the populations of both

species recorded at Pink Lakes State Park and three

other reserves by Nichols ef a/. (1991) should be

relocated and monitored. In particular, it should be

noted that the present study and Nichols e¢ al. (1991)

only saw fruiting of both Marsdenia and

Rhyncharrhena when they were climbing in shrubs

or trees. The only Rhyncharrhena plants seen at Pink

Lakes were 22 suckers in an area lacking any

potential host plants. Because installing tree guards

300 mm high produced increased Rhyncharrhena

shoot growth without fruit production (Nicholls e¢

al., 1991), host plants should now be established

there. Other work on vine enrichment-planting, on

the depleted vine Pararistolochia praevenosa, also

shows that a supporting host plant is a requirement

for fruiting (Sands e/ a/. 1997; Sands, pers. comm.).

As for many other native plant species, the

palatability of Marsdenia and Rhyncharrhena needs

to be considered in managing mammal numbers at

Hattah-Kulkyne National Park and other areas.

Acknowledgements

I thank J.H. Browne for providing seed, P. and

F. Murdoch, T. Phillips and M. Bartley for help and

Parks Victoria for financial support. The work was

carried out under permit numbers 901/701, 912/076

and 10001922.

66 R. F. PARSONS

References

BRANDLE, R. (1998) “A Biological Survey of the Stony

Deserts of South Australia 1994-1997” (Department for

Environment, Adelaide).

Hecarty, E. E. and Ciirrorp, H. T. (1984) Climbing

angiosperms in the Australian flora. pp. 105-120 Jn

Werren, G. & Kershaw, A.P. (Eds) “The Rainforest

Legacy. Australian National Rainforests Study. Vol. 2”

(Monash University, Melbourne).

Hosss, R. J. & ATKINS, L. (1991) Interactions between

annuals and woody perennials in a Western Australian

nature reserve. J. Veg. Sci. 2, 643-054.

NICHOLS, K. M., BRowne, J. H. and Parsons, R. F. (1991)

Ecology of two asclepiad lianes in semi-arid Victoria.

Proc. R. Soc. Vict. 103, 93-112.

PARSONS, R. F. (1994) Eucalyptus scrubs and shrublands.

pp. 291-319 Jn Groves, R.H. (Ed) 9 “Australian

Vegetation” 2nd ed. (Cambridge University Press,

Cambridge).

PENALOSA, J. (1984) Basal branching and vegetative spread

in two tropical rainforest lianes. Biotropica 16, 1-9.

Sanps, D. P. A., Scott, S. E. & Morratt, R. (1997) The

hreatened Richmond birdwing butterfly (Ornithoptera

richmondia|Gray]): a community conservation project.

Mem. Mus. Vict. 56, 449-453.

Smitu, B. C. (1991) Ecological studies of seasonal and

biotic effects on seedling recruitment of two asclepiad

lianes. B.Sc. (Honours) thesis, Botany Department, La

Trobe University.

BREINLIA (BREINLIA) VENTRICOLA SP. NOV., A NEMATODE

PARASITE FROM THE HEART OF THE RED KANGAROO,

MACROPUS RUFUS, IN WESTERN AUSTRALIA

By D. M. SPRATT* & R. P. HOBBST

Summary

Spratt, D. M. & Hobbs, R. P. (2004) Breinlia (Breinlia) ventricola sp. nov.

(Nematoda: Filarioidea) from the heart of the red kangaroo, Macropus rufus, in

Western Australia. Trans. R. Soc. S. Aust. 128(1), 67-71, 31 May, 2004.

Breinlia (Breinlia) ventricola sp. nov., is described from the right ventricles and

pulmonary arteries of the red kangaroo, Macropus rufus (Desmarest, 1822), from the

Pilbara region of Western Australia. It is the largest and most robust species of

Breinlia (Breinlia) known to date and is distinguished from all other species by the

close similarity in size between males and females, the presence of filamentous

membrane surrounding the distal tip of the left spicule and the presence of a pair of

caudal glands secreting a material which adheres to the caudal end of some fixed

specimens, masking external cuticular features. The new species is most similar to B.

(B.) trichosuri, B. (B.) boltoni and B. (B.) mundayi but distinguished from all 3 by its

much greater size and the absence of internolateral cephalic papillae.

Key Words: Nematoda, marsupials, kangaroo, Breinlia, new species, Macropus.

Transactions of the Royal Society of S. Aust. (2003), 128(1), 67-71.

BREINLIA (BREINLIA) VENTRICOLA SP. NOV., A NEMATODE PARASITE

FROM THE HEART OF THE RED KANGAROO, MACROPUS RUFUS,

IN WESTERN AUSTRALIA.

by D.M. SPRATT’ & R. P. HOBBS*

Summary

Sprarr, D. M. & Hopps, R. P. (2004) Breinlia (Breinlia) ventricola sp. noy. (Nematoda:Filarioidea) from the

heart of the red kangaroo, Macropus rufus, in Western Australia. Trans. R. Soc. S. Aust. 128(1), 67-71, 31 May,

2004.

Breinlia (Breinlia) ventricola sp. nov. is described from the right ventricles and pulmonary arteries of the red

kangaroo, Macropus rufus (Desmarest, 1822), from the Pilbara region of Western Australia. It is the largest and

most robust species of Breinlia (Breinlia) known to date and is distinguished from all other species by the close

similarity in size between males and females, the presence ofa filamentous membrane surrounding the distal tip

of the left spicule and the presence of a pair of caudal glands secreting a material which adheres to the caudal

end of some fixed specimens, masking external cuticular features. The new species is most similar to B. (B.)

trichosuri, B. (B.) boltoni and B. (B.) mundayi but distinguished from all 3 by its much greater size and the

absence of internolateral cephalic papillae.

Kry Worbs: Nematoda, marsupials, kangaroo, Breinlia, new species, Macropus.

Introduction

The genus Breinlia Yorke and Maplestone, 1926,

subgenus Brein/ia represents a group of parasitic

filarioid nematodes occurring in the body cavities,

subcutaneously and in the vascular system of murid

and sciurid rodents and primates in Asia (Chabaud

and Bain, 1976; Bain ef a/., 1983), in dasyurid,

peramelid, petaurid, pseudocheirid, phalangerid,

phascolarctid, potoroid and macropodid marsupials

in Australia and Papua New Guinea (Spratt and

Varughese, 1975; Spratt ef a/. 1991), and in native

murid rodents in northernwestern Western Australia

(Beveridge and Spratt, 1996). Microfilariae occur in

the peripheral circulation and in the four life cycles

that have been investigated (B. booliati Singh and

Ho, 1973, B. manningi Bain, Petit, Ratanaworabhan,

Yenbutra and Chabaud, 1981,and B. sergenti Mathis

and Léger, 1909 in Southeast Asia and B. macropi

Spratt and Varughese, 1975 in Australia) mosquitoes

act as intermediate hosts (Ho ef a/., 1973; Bain ef al,

1981: Ramachandran and Dunn, 1968; Zaman and

Chellappah, 1968; Beveridge and Spratt, 1996).

There are many species of Breinlia but only one. B.

woerlei (Spratt and Varughese, 1975) Bain, Baker

and Chabaud, 1983 occurs in the heart. However, in

the filarioid genus, Dirofilaria Railliet and Henry,

1911 a number of species occur in the heart, the best

known example in Australia being the dog

“CSIRO Sustainable Ecosystems, GPO Box 284, Canberra ACT

26001.

‘School of Veterinary and Biomedical Sciences, Murdoch

University, Perth, WA 6150.

heartworm, D. immitis (Leidy, 1856). An inspector in

a wild meat export abattoir in Perth encountered

large filarioid nematodes in the right ventricles and

pulmonary arteries (Fig. 1) of red kangaroos,

Macropus rufus, from the Pilbara region of Western

Australia, which represents a new species of

Breinlia(Breinlia) described below.

Materials and Methods

Nematodes were fixed at room temperature in 70%

ethanol with 5% glycerine and cleared in lactophenol

for examination. £7” face preparations of the cephalic

extremity were prepared by the method of Anderson

(1958). The same glycerine jelly hanging-drop

\

Fig. |. Breinlia (Breinlia) ventricola sp. nov. in right

ventricle of red kangaroo, Macropus rufus, scale bar

10 mm.

D. M. SPRATT & R. P. HOBBS

NEW NEMATODE FROM RED KANGAROO 69

technique was used to study the posterior ends of

some males and females.

Measurements of 9 adult males and 9 adult females

are presented in millimetres as the mean followed by

the range in parentheses, those of 10 microfilariae

from the vagina wterina of female worms are in

micrometres. Drawings were made with the aid of a

drawing tube. Type-specimens are deposited in the

South Australian Museum (SAM) Australian

Helminth Collection (AHC), Adelaide and the

Queensland Museum (QM), Brisbane. Additional

material is in the Western Australia Museum

(WAM), in CSIRO Sustainable Ecosystems and in

the School of Veterinary and Biomedical Sciences,

Murdoch University

Breinlia (Breinlia) ventricola sp. nov.

(FIGS 2-14)

Material examined:

Holotvpe

d from right ventricle of Macropus rufus

(Desmarest, 1822), Mindaroo Station, 50 km

southwest of Onslow (21° 41’ S, 115° 08’ E)

Western Australia, coll. F. Stephens, 9 May, 2002,

SAM AHC [IDW]32174.

Allotvpe

2 same data, SAM AHC [IDW]32175.

Paratypes

43.4, from right ventricle and pulmonary arteries

of M. rufus, Warrawagina Station (20° 51’ S, 120°

41' E) Western Australia, coll. F. Stephens, 4 June,

2002, SAM AHC 32176, 422 same data AHC

32177, 435 same data, QM G222333-222336,

42 9 same data, QM G222337-222340.

Other material examined

Fragments of specimens from type localities in the

Parasite Collection of CSIRO Sustainable

Ecosystems, Canberra, W/L HC# N5259; 12 from

M. rufus near Mt. Augustus (24° 19’ S, 116° 50’ E)

in WAM V4382, 22 2 same data in Department of

Veterinary Biology and Biomedical Sciences,

Murdoch University, Perth, X02/07.

Prevalence

Observed in 19 of 550 (3.5%) red kangaroos

examined.

Site in host

Right ventricle and pulmonary arteries.

Etymology

The suffix -icola means “inhabiting” and the

specific epithet reflects the location of the nematodes

in the right ventricle of the host.

Description

Long, robust nematodes with attenuated anterior

and helically coiled posterior extremities. Oral

opening small, round, not bounded by prominent

refractile membrane. Cephalic extremity large, oval.

Four pairs of submedian papillae arranged in outer

circle of 4 large and inner circle of 4 slightly smaller

papillae. Rectangular cuticular peribuccal field

present, appearing as slight elevation of cuticle

joining bases of papillae of inner circle. Distance

between papillae on lateral surface greater than on

dorsal and ventral surfaces. Second cuticular

peribuccal field present, rectangular in shape, formed

by slight elevation of cuticle joining bases of papillae

of outer circle. Distance between papillae on lateral

surfaces greater than on dorsal and ventral surfaces.

Internolateral papillae absent. Amphids lateral, large

but not raised, opening at level of outer circle of

papillae; amphidial pouches large. Buccal capsule

minute, narrow, with delicate wall. Small, refractile

pre-pharyngeal ring present. Oesophagus divided

into short anterior muscular and long posterior

glandular regions. Excretory pore not observed.

Cuticle with conspicuous transverse striations.

Refractile cuticular bosses observed on ventral

surface of males only. Lateral cords and hypodermal

nuclei inconspicuous. Lateral alae absent. Deirids

not observed.

Male

Total length 195 (180 — 220); maximum width 0.84

(0.76 — 0.89) in mid-body region; nerve ring 0.40

(0.37 — 0.42) from anterior extremity; muscular

oesophagus 0.79 (0.71 — 0.90) long, glandular

Pigs 2-14. Breinlia (Breinlia) ventricola sp. nov. 2. Anterior end female, dorsal view. 3. Cephalic extremity female, apical

view. 4. Female tail, lateral view. 5.Cuticular rugosities on ventral surface of male in posterior one-third of body. 6.

Female vulva and vagina, lateral view. 7. Microfilaria. 8. Tail of microfilaria showing terminal nuclei. 9. Diagrammatic

representation of seven patterns of distribution of cloacal papillae in males. 10. Male tail, lateral view. 11. Distal tip of

left spicule. 12. Caudal end of male showing pair of lateroventral lappets and medial openings of ducts of caudal glands,

apical view. 13. Caudal end of male, latero-ventral view. 14. Caudal end of male with caudal glands and secretion on

cuticle, ventral view. Scale bars = 0.05 mm 4, 6, 7, 11, 12, 13; 0.1 mm 1, 2, 9, 10; 0.2 mm 3, 5.

70 D. M. SPRATT & R, P, HOBBS

oesophagus 2.1 (1.9 — 2.4) long. Spicules unequal,

dissimilar, heavily sclerotized, left spicule 0.59 (0.56

0.61) long, calamus 0.26 (0.25 — 0,28), lamina 0.16

(0.15 — 0,17), filament 0.17 (0.16 — 0.19) terminating

in fine sclerotized rod surrounded by filamentous

membrane; right spicule 0.33 (0.32 — 0.34) long,

with calciform, spatulate distal extremity.

Gubernaculum present, 0.06 (0.06 — 0.07) long, U-

shaped in ventral view with deep lateral walls

appearing to wrap around lateral edges of right

spicule. Tail helically coiled, 1.2 long, terminating in

two large lateroventral lappets located close to one

another. Two large caudal glands present, opening

adjacent and median to lappets. Some males with

material secreted from glands adhering to caudal

cuticle and masking view of external features of

caudal end. Cloacal papillae highly variable in

number, ranging from 6 16 with 8 the most

frequently occurring number, and in pre- and post-

cloacal positions, the majority clustered around

cloacal aperture; aperture slightly elevated. Some

males with one or two subterminal medio-ventral

papillae, some with a subterminal medio-dorsal

papilla. Cuticle with longitudinally elongate,

refractile, cuticular bosses on ventral surface,

commencing posterior to oesophago-intestinal

junction and terminating approximately 3 mm from

cloacal aperture.

Female

Total length 233 (205 — 260); maximum width 1.60

(1.30 — 2.20) in mid-body region; nerve ring 0.40

(0.34 — 0.45) from anterior extremity; muscular

oesophagus 0.75 (0.63 — 0.87) long, glandular

oesophagus 1.50 (0.91 — 2.40) long. Vulva well

posterior to oesophago-intestinal junction, 7.20 (5.20

— 9,50) from anterior extremity; vagina 2.40 (1.90

2.70) long, thick-walled, muscular. Tail helically

coiled, tapering characteristically on the dorsal

surface and terminating in two large lateroventral

lappets located close to one another. Two large

caudal glands present, opening adjacent and median

to lateroventral lappets. Some females with material

secreted from glands adhering to caudal cuticle and

masking view of external features of caudal end.

Phasmids not observed. Cuticular bosses absent.

Microfilariae from vagina uterina

Body length 265 (263 — 267); maximum width 6 in

mid-body region; nerve ring 51 (49 — 53) and

excretory vesicle 79 (78 — 80) from anterior extremity;

anal vesicle 208 (205 — 210). Body tapering behind

vesicle; nuclear column terminating in single row of 3

small, elongate nuclei; tail filamentous. Distance from

last nucleus to tail tip 26 (25 — 28). Cuticle with

conspicuous transverse striations. Microfilaria

unsheathed, presumed to occur in blood of host.

Discussion

Two subgenera of Breinlia were defined by

Anderson and Bain (1976), B. (Breinlia) Yorke and

Maplestone, 1926 (not Chabaud and Bain, 1976 as

listed by Anderson and Bain, 1976) recognised by

spicules unequal in length, the presence of a divided

oesophagus and the presence of a gubernaculum, and

B. (Johnstonema) (Yeh, 1957) recognised by

spicules equal or subequal in length, the presence of

an oesophagus not clearly divided in females, the

presence of external labial papillae arranged in the

form of a dorsoventrally expanded rectangle, a

buceal cavity with thin walls and the absence of a

gubernaculum. However, the morphology of the

spicules with the left spicule clearly divided into

calamus, lamina and filament and the right with a

calciform, spatulate distal extremity is more typical

of the subgenus Brein/ia than of Johnstonema and on

this basis particularly, we have placed this species

therein.

Breinlia (B.) ventricola is distinguished from all

other known species of B. (Breinlia) (n = 19) and B.

(Johnstonema) (n= 4) by its great length and breadth

(males twice as long and broad and females one

eighth as long and nearly three times as broad as

largest species known previously), the close

similarity in size between males and females, the

presence of a filamentous membrane surrounding the

distal tip of the left spicule and the presence of a pair

of caudal glands in male and female worms secreting

a material which adheres to the caudal end of some

fixed specimens masking external cuticular features.

In addition, the spicules in species of B.

(Johnstonema) are stout, subequal and the left

spicule is not divided into calamus, lamina and

filament. The new species most closely resembles

Breinlia (B.) trichosuri (Breinl, 1913) from. the

peritoneal cavity of the common brushtail possum,

Trichosurus vulpecula (Kerr, 1792) in Queensland,

B. (B.) mundayi Spratt and Varughese, 1975 from the

pericardium of the red-necked wallaby, Macropus

rufogriseus (Desmarest, 1817), in Tasmania and B.

(B.) holtoni Spratt and Varughese, 1975) from the

peritoneal cavity of the agile wallaby, /. agilis

(Gould, 1842), from northern Australia. Breinlia (B.)

ventricola is distinguished from all three by its much

greater size and the absence of a pair of internolateral

cephalic papillae. Previously, B. (B.) macropi trom

the quokka, Setonix brachyurus (Quoy and Gaimard,

1830), was the only filarioid nematode known from

marsupials in Western Australia.

The two subgenera of Breinlia occurring in

Australia are most common in macropodoid

marsupials with 14 species previously described

from the pleural and peritoneal cavities, pericardial

sac, right ventricle and subcutis (Spratt and

NEW NEMATODE FROM RED KANGAROO 71

Varughese, 1975) in addition to a number of

undescribed species (Spratt ef a/. 1991). Additional

species of the subgenus occur in dasyurid,

peramelid, petaurid, pseudocheirid, phalangerid and

phascolarctid marsupials. The subgenera have been

interpreted as a group of australian parasites, having

migrated relatively recently from Asia in murid

rodents and subsequently radiating widely in

marsupials (Bain ef al, 1983). Species of the

subgenus Breinlia are known to occur in six species

of “old endemic” murid rodents but only in the

Kimberley region of Western Australia, all

morphologically indistinguishable from species

described from pseudocheirid and macropodid

marsupials. No species of the subgenus

Johnstonema are known from murid rodents in

Australia. In Australian marsupial hosts, species of

both subgenera appear relatively host specific, but

those occurring in macropodoids, especially in the

family Macropodidae, exhibit the least host

specificity. Consequently, one might expect to

encounter B. (B.) ventricola in other species,

particularly of the larger kangaroos and wallaroos.

However, it is surprising that such a large nematode

has not been encountered previously, particularly

given the large export trade in kangaroo meat. This

suggests that the species may not only be specific to

the red kangaroo but also may be limited to the

geographic region of northwestern Western

Australia.

It is expected that many aspects of pathology seen

in canine dirofilariasis would occur in red kangaroos

infected with B. ventricola, because Dirofilaria

immitis is similar in size and is found in the same

sites. Dirofilariasis is characterised by pulmonary

hypertension caused primarily by an intense tissue

reaction (Knight, 1977). Heavy infections would

also be expected to result in impaired heart valve

function and physical blockage (Soulsby, 1982).

Exercise intolerance is a feature of dirofilariasis and

would be likely to result in greater susceptibility to

predation in red kangaroos infected with B.

ventricola,

Acknowledgements

Thanks are due to Fran Stephens for her efforts in

collecting and forwarding the material for study. lan

Beveridge and Hugh Jones offered valuable

comments on an earlier draft of the manuscript.

References

ANDERSON, R. C. (1958) Méthode pour Vexamen des

nématodes en vue apical. Ann. Parasitol. hum. comp. 33,

171-172.

— & BAIN, O. (1976) Keys to the genera of the

Order Spirurida. Part 3. Diplotriaenoidea, Aproctoidea

and Filarioidea. No. 3. pp. 59-116 /n: Anderson R.C.,

Chabaud A.G. & Willmott S. (Eds.) “CIH Keys to the

Nematode Parasites of Vertebrates” (Commonwealth

Agricultural Bureaux, Farnham Royal, Bucks, England).

BAIN, O., BAKER, M. & CHABAUD, A. G. (1982 issued 1983)

Nouvelles données sur la lignée Dipetalonema

(Filarioidea: Nematoda). Ann. Parasitol. hum. comp. *7,

593-620.

, Perit, G., RAYANAWORABHAN, N., YENBUTRA,

S. & CuaBpaup, A. G. (1981) Une nouvelle filaire

d@ecureuil en Thailande, Breinlia (B.) manningi n. sp. , ct

son développement chez Aedes. Ann. Parasitol. hum.

comp. 56, 193-201.

BEVERIDGE, L. & Sprarr, D. M. (1996) The helminth fauna

of Australasian marsupials: origins and evolutionary

biology. Adv. Parasitol. 37, 135-254.

CuaABAUb, A. G. & Bain, O. (1976) La lignée

Dipetalonema. Nouvel essai de classification. Ani.

Parasitol. hum. comp. 51, 305-397.

Ho, B. C., Sincu, M. & Lim B. L. (1973) Observations on

the development of a new filaria (Breinlia booliati Singh

and Ho, 1973) of a rat Rattus sabanus in the mosquito

Aedes togoi. J. Helminthol. 47, 135-140.

KNiGHtT, D. H. (1977). Heartworm heart disease. Adv. ver.

Sci. comp. Med. 21, 107-149.

RAMACHANDRAN, C. P. & Dunn, F. L. (1968) The

development of Breinlia sergenti (Dipetalonematidae) in

Aedes mosquitoes. Ann. trop. Med. Parasitol. 62, 441-

449.

Soutspy, E. J. L. 1982. “Helminths, Arthropods and

Protozoa of Domesticated Animals” 7th ed. (Baillicre

Tindall, London),

Spratt, D. M. & VARUGHESE, G. (1975) A taxonomic

revision of filarioid nematodes from Australasian

marsupials. Aust. J. Zool. Suppl. Ser. 35, 1-99.

—_, Brveripcr, 1. & WaALrer, E. L. (1991) A

catalogue of Australasian monotremes and marsupials

and their recorded helminth parasites. Rec. S. Aust. Mus.,

Monogr. Ser. No. 1, 1-105.

ZAMAN, V. & CHELLAPPAH W. T. (1968) Studies on vector

susceptibility and larval morphology of a filaria of the

slow loris, Nycticebus coucang. Ann. trop. Med.

Parasitol. 62, 450-454.

BLACK NODDIES (ANOUS MINUTUS) AND WEDGE-TAILED

SHEARWATERS (PUFFINUS PACIFICUS) AS POTENTIAL

HOSTS FOR FUNGI INVADING SEA TURTLE NESTS AT

HERON ISLAND, QUEENSLAND

By A. D. PHILLOTT*? & S. A. M. ELSMORE*

Summary

Phillott, A. D. (200) Black Noddies (Anous minutus) and Wedge-tailed Shearwaters

(Puffinus pacificus) as Potential Hosts for Fungi Invading Sea Turtle Nests at Heron

Island, Queensland. Trans. R. Soc. S. Aust. 128(1), 73-76, 31 May, 2004.

Black noddies (Anous minutus) and wedge-tailed shearwaters (Puffinis pacificus) on

Heron Island (Capricorn Group, Queensland) were surveyed to determine if they

harboured fungi that had been implicated in sea turtle egg mortality in situ. Isolated

from the cloacal exterior of these seabirds were Penicillium citrinum, Fusarium

semitectum, F. trichothecioides, Aspergillus unguis, Aspergillus candidus,

Acremonium sp., Alternaria sp. and Drechslera sp. These isolates are common soil or

plant fungi and their most likely origin is therefore the nest or burrow material. None

of these species have been isolated from sea turtle eggs that failed to hatch.

Key Words: Seabird, fungi, sea turtles.

Transactions of the Royal Society of S. Aust. (2004), 128(1), 73-76.

BLACK NODDIES (ANOUS MINUTUS) AND WEDGE-TAILED SHEARWATERS

(PUFFINUS PACIFICUS) AS POTENTIAL HOSTS FOR FUNGI INVADING SEA

TURTLE NESTS AT HERON ISLAND, QUEENSLAND.

by A. D. PHiILLoTT’ & S. A. M. ELSMORE®

Summary

Putott, A. D. (200) Black Noddies (4nous minutus) and Wedge-tailed Shearwaters (Puffinus pacificus) as

Potential Hosts for Fungi Invading Sea Turtle Nests at Heron sland, Queensland. 7rans. R. Soc. S, Aust. 128(1),

73-76, 31 May, 2004.

Black noddies (Anous minutus) and wedge-tailed shearwaters (Puffinus pacificus) on Heron Island (Capricorn

Group, Queensland) were surveyed to determine if they harboured fungi that had been implicated in sea turtle

egg mortality in situ. Isolated from the cloacal exterior of these seabirds were Penicillium citrinum, Fusarium

semitectum, FE) trichothecioide

Aspergillus unguis, Aspergillus candidus, Acremonium sp., Alternaria sp. and

Drechslera sp. These isolates are common soil or plant fungi and their most likely origin is therefore the nest or

burrow material. None of these species have been isolated from sea turtle eggs that failed to hatch.

Kry Worbs: Seabirds, fungi, sea turtles.

Introduction

Heron Island (23° 26’ S, 151° 55’ E) is a small

coral cay of the Capricorn Group in the southern

Great Barrier Reef, lying approximately 80km_ off

the Queensland coast, north-east of Gladstone. The

island is dominated by a central forest of Pisonia

grandis and common stands of Ficus opposita and

Pandanus sp. Exposed fringing vegetation consists

of Argusia argentea, Casuarina equisetifolia, Cordia

subcordata and Pandanus sp. Fosberg and Thorne

(1961), Gillham (1961), Cribb (1969, 1976),

Chaloupka and Domm (1986) and Rogers (1989)

present descriptive lists of vascular plants from the

island,

The Capricorn Group of islands have four of the

major colonies of black noddies, Anous minutus, in

the Great Barrier Reef and the largest breeding

population of wedge-tailed shearwaters, Pu/finus

pacificus, in the world (Hill e¢ al. 1995). Colonies of

both species at Heron Island have been subject to

intense surveillance.

Black noddies reside on the island year round.

Breeding occurs between October and March with a

peak in November/December. A single egg is laid

(Kikkawa 1970) in rough nests constructed from

excreta and shed P. grandis leaves (Barnes and Hill

1989: Hill and Rosier 1989). The majority of nesting

occurs in the P. grandis forest, with correlations

between bird density and vegetation species and

physiognomy (Shipway 1969; Dale ef al. 1984;

Hulsman e¢ al. 1984; Barnes and Hill 1989; Ogden

1979, 1993).

"School of Biological and Enyironmental Sciences, Central

Queensland University, Rockhampton QLD 4702.

' Corresponding author. Email: a.phillott(@iequ.edu.au.

Numbers of black noddies on Heron Island have

been extensively documented (Campbell and White

1910; MacGillivray 1928; Cooper 1948: Shipway

1969; Kikkawa 1970; Bingham 1977; Ogden 1979;

Hulsman 1983, 1984; Barnes and Hill 1989; Ogden

1993; Hill ef al. 1997). From 53 nests in 1910

(Kikkawa 1970), the population increased to 63,000

£7,000 pairs in 1992 (Ogden 1993), Hence there has

been an almost geometrical increase in the last 75

years (Barnes and Hill 1989) of approximately 7%

per annum (Ogden 1993). The rate of increase

predicted 116,615 breeding pairs by “the turn of the

century” (Ogden 1993), though high adult mortality

in 1997 due to lack of prey (P. O’Neill, pers.comm.)

resulted in reduced numbers over the past years

(Phillott, pers.ob.). Population growth has been

accompanied by an expansion of the nesting area

into areas inhabited by humans and the coastal

woodlands since and an increase in the number of

nests per tree (Barnes and Hill 1989).

The migratory wedge-tailed shearwaters arrive at

Heron Island in October (Campbell and White 1910;

Cooper 1948: Moulton 1961; Gross ef al. 1963;

Kikkawa 1970; Bingham 1977; Ogden 1979),

Nesting commences between mid-November and

mid-December (Dyer and Carter 1997) and peaks in

late-December (Gross ef al. 1963). A single egg

(Dyer and Hill 1990) is incubated in a burrow (Hill

and Rosier 1989; Dyer and Hill 1990) or in protected

hollows on the ground among tree roots (Dyer and

Hill 1990). Adults and fledglings depart the island by

May-June (Miles 1964; Kikkawa 1970).

Nesting by wedge-tailed shearwaters on Heron

Island has been thoroughly described (MacGillivray

1928; Shipway 1969; Kikkawa 1970; Ogden 1979;

Hulsman 1983, 1984; Hill and Barnes 1989; Ogden

1994; Carter ef al. 1996: Hill et al. 1996). Though

74 A. D. PHILLOTT & S.A. M. ELSMORE

TABLE 1. Occurrence of Fungi in Seabird Cloacal Swabs.

y

Occurrence of Fungi (%

Fungi A. minutus (n=20) _ P. pacificus (n=20)

P. citrinum 95 90,

F. semitectum 10 10

F. trichothecioides 0 5

A. unguis 5 0

A. candidus 5 0

Acremonium sp. 0 5

Alternaria sp. 0 5

Drechslera sp. 0 5

the literature appears to report an increasing

population from 8,300 burrows in 1965 (Shipway

1969) to 13,381+1,556 in 1996 (Hill ef al. 1996),

Dyer ef al. (1995) corrected for different sampling

methodology and deduced a mean of approximately

15,000 active burrows over the past 35 years.

Within the southern Great Barrier Reef, green sea

turtles, Chelonia mydas, nest almost exclusively on

coral cays such as Heron Island, while the Capricorn

Group is the major nesting area for loggerhead sea

turtles, Caretia caretfa, in the South Pacific Ocean

(Limpus ef al. 1983). The fungi Fusarium

oxysporum, Fusarium solani and Pseudallescheria

hoydii have been isolated from the exterior of green

and loggerhead sea turtle eggs at Heron Island that

failed to hatch (Phillott et a/, 2001; Phillott e¢ al.

2004). Fungi within turtle nests have been implicated

with variation in egg mortality between coral cays

(Heron Island and the adjacent Wreck Island, 23° 20’

S, 151° 57’ E), and the mainland (Limpus ef al.

1983; Phillott 2002).

F. oxysporum and F) solani have been detected in

wild and domestic bird nests (Mazen ef al. 1994),

and F) oxysporum has been isolated from feather

samples (Hubalek ef a/. 1995). They are known also

as cellulolytic fungi (Mazen e/ al, 1994). Therefore,

it is possible the substantial seabird colonies of

Heron [sland may function as potential reservoirs of

fungi invading sea turtle nests.

Materials and Methods

The cloacal exteriors of twenty each black noddies

and wedge-tailed shearwaters were individually

swabbed to compare mycobiota with that isolated

from failed sea turtle eggs. Adult noddies were

approached while on the nest, briefly removed,

swabbed and replaced. Animals with eggs or young

chicks were not sampled. Shearwaters were captured

by hand during the nightly courtship or pre-dawn

congregations prior to leaving the island for feeding.

Swabbing occurred at the point of capture and

animals were released immediately.

Swabs (MW170 TRANSTUBE", Amies Clear

Transport Media) were refrigerated at 3 — 5°C prior

to incubation. Fungi were incubated on specific

media and identified according to Booth (1971),

Ellis (1971), Pitt (1979), Carmichael ef al. (1980)

and Klich and Pitt (1988).

Results

All 20 of the P. pacificus and 19 of the 20 A.

minutus swabs were positive for cloacal mycobiota.

Isolates were Penicillium citrinum, Fusarium

semitectum, F) trichothecioides, Aspergillus unguis,

Aspergillus candidus, Acremonium sp., Alternaria sp.

and Drechslera sp. (see Table 1). Most swabs

produced monocultures of P citrinum (80%

A. minutus; 75% P. pacificus) or F) semitectum (5% P.

pacificus), however mixed cultures of P. citrinum +

F. semitectum (10% A. minutus; 10% P. pacificus),

P. citrinum + EF. trichothecioides (5% P. pacificus),

P. citrinum + Drechslera sp. (5% P. pacificus),

2 citrinum + Aspergillus candidus + Aspergillus

unguis (5% A. minutus) and Alternaria sp. +

Acremonium sp. (5% P. pacificus) were also obtained.

~S

Discussion

All of the genera isolated from seabirds are

commonly associated with soil or agricultural crops

(Booth 1971; Ellis 1971; Pitt 1979; Carmichael e7 al.

1980: Klich and Pitt 1988) and some (Acremonium,

Alternaria, Aspergillus, Fusarium, Penicillium) are

known to be cellulolytic fungi from birds nests

(Mazen ef al. 1994). None are true coprophiles (see

Webster 1970) or keratinophiles (Pugh and Evans

1970; Rees 1977), although F! semitectim has been

isolated from the keratinaceous scutes of a tortoise

carapace (Rose ef a/. 2001). Therefore, the most

likely source of the isolates is from the nesting

material or the burrow substrate. None are known

invaders of sea turtle eggs (see Phillott ef a/. 2001;

Phillott ef al. 2004).

While the seabirds of Heron Island do not appear

to be harbouring fungi known to invade sea turtle

eggs, it is possible they may play an accessory role

in maintaining resident soil microbiota. Staunton

Smith and Johnson (1995) calculated a guano

deposition of approximately 107t from A. minutus

and approximately 22t from P. pacificus on Heron

Island in 1992. Total annual deposition of guano is

expected to contain 9.4t of nitrogen and 1.9t

phosphorus. While much of the nitrogen is leached

by the high rainfall (Heatwole ef a/. 1981),

phosphorus is expected to be incorporated into the

soil and is important in maintaining island vegetation

(Allaway and Ashford 1984). Increased soil nutrients

may also assist in maintaining a high microbial load.

The relationship between phosphorus levels and soil

SEABIRDS AS FUNGAL HOSTS 7

mycobiota of Queensland turtle rookeries is under

further investigation.

Acknowledgments

This work was conducted under the Queensland

Parks and Wildlife Service Scientific Purposes

Wi

Permit No. C6/000114/99/SAB and Central

Queensland University Ethical Clearance Certificate

No. 99/11-82. K. Coufal assisted with seabird

capture and handling. Dr. K. M. Harrower assisted

with identification of the Drechslera, Acremonium

and Alternaria sp. Dr. C. J. Parmenter reviewed a

copy of the manuscript.

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VOL. 128, PART 2

30 NOVEMBER, 2004

Transactions of the

Royal Society of South

Australia Incorporated

INCORPORATING THE

wT D)

JReCor alt

Contents

Appan, A., Bergfeld, J. & Beveridge, I New species of parasitic nematodes from

macropodid marsupials in Western Australia - - - - — 77

Hammer, M. P. & Walker, K. F. A catalogue of South Australian freshwater fishes,

including new records, range extensions and translocations — — 85

Watts, C. H. S & Humphreys, W. F. Thirteen new Dytiscidae (Coleoptera) of the

genera Boongurrus Larson Tjirtudessus Watts & Humphreys and

Nirripirti Watts é& Humphreys, from underground waters in

Agile) Se ee 99

Watts, C. H. S. Revision of Australian Scirtes Illiger and Ora Clark z eebieopiees:

Scirtidae) - - - - =) = = = = = She 131

Hodda, M. Characteristics of an Australian population of Paraxonchium

orthodon (Loof 1964) Altherr & Loof 1969, with a note on

evolution and biogeography of the subfamily Paraxonchiinae

(Nematoda, Dorylaimida, Aporcelaimidae) - - - -— - 169

Beveridge, I. & Johnson, P. M. Cestode parasites of tree kangaroos (Dendrolagus spp.:

Marsupialia), with the description of two new species of

Progamotaenia (Cestoda: Anoplocephalidae) - - - - —- 175

Faucheux, M. J. Antennal sensilla of Sabatinca sterops Turner (Lepidoptera:

Micropterigidae) -— - - - = = =~ ~ = = = 187

Barker, S. Twelve new species of Australian Buprestidae (Coleoptera) and

new synonymy —- -—- -— - = = =~ =~ = = = = 195

Womersley, H. B. S. Additions to the Marine Algal Flora of southern Australia - - 205

Smith, B. P. C. & Kokkinn, M. The use of emergence as an end-point for sediment

toxicity tests using the Australian chironomid Chironomus maddeni pale)

Barton, P. S., Aberton, J. G. & Wishart, E. Spatial and temporal distribution of Culex

australicus Dobrotworsky and Drummond and Culex globocoxitus

Dobrotworsky (Diptera: Culicidae) at the aa Lakes in

eastern Victoria - - ee 2)

Williams, C. R., Snell, A. E. & Kokkinn, M. J. Studies of temporal host-seeking

patterns of Culex Annulirestris (Diptera: Culicidae): a comparison

of methods and populations - - - = —~ -~ = = Bas

Gibbs, S. E., Kemper, C. M., Byard, R. W. & Long, M. Deaths of killer whales

(Orcinus orca) in South Australia and implication of human

jimieretetney ae ee 231

Hinchcliffe, J. & Conran, J. G. The Tennyson sand dunes: vegetation structure and

conservation status - - - — = 239

Smith, B. B. & Walker, K. F. Reproduction of common carp in South Australia, shown

by young-of-the-year samples, gonadosomatic index and the

histological staging of ovaries = - - - - = = - 249

Brief Communications:

Souter, N. J. A comparison among three artificial substrates for aquatic

macroinvertebrate sampling - - - - -~ = = = 259

Matthews, E. G. New synonymy and new names in Australian Tenebrionidae

(Coleone) = = = = a Se 261

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INCORPORATING THE

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VOL. 128, PART 2

NEW SPECIES OF PARASITIC NEMATODES FROM

MACROPODID MARSUPIALS IN WESTERN AUSTRALIA

By A. APPAN*, J. BERGFELD* & I. BEVERIDGE*

Summary

Appan, A., Bergfeld, J. & Beveridge, I. (2004) New species of parasitic nematodes

from macropodid marsupials in Western Australia. Trans. R. Soc. S. Aust. 128(2), 77-

84, 30 November, 2004.

Two new species of parasitic nematodes are described from macropodid marsupials

from Western Australia. Rugopharynx setonicis sp. nov. from the stomach of the

quokka, Setonix brachyurus, belongs to the Rugopharynx australis complex, but

differs from related species in the anterior position of the nerve ring, a shorter spicule

length and the anterior position of the deirids. A second new species, from the

oesophagus of Macropus irma, is described. Cyclostrongylus irma sp. nov. is

distinguished from congeners by a buccal capsule which is wider near its anterior

extremity, longer spicules and a short oesophagus. The descriptions of the two new

species provide additional evidence in support of current hypotheses for the mode of

evolution of the respective genera.

Key Words: Nematoda, marsupials, Macropus, Setonix, new species, Rugopharynx,

Cyclostrongylus.

Transactions of the Royal Society of S. Aust. (2004), 128(2), 77-84.

NEW SPECIES OF PARASITIC NEMATODES FROM MACROPODID

MARSUPIALS IN WESTERN AUSTRALIA

by A. AppAN", J. BERGFELD® & I. BEVERIDGE®

Summary

ApPpaN, A., BERGFELD, J. & BeveribGe, I. (2004) New species of parasitic nematodes from macropodid

marsupials in Western Australia. Trans. R. Soc. S. Aust. 128(2), 77-84, 30 November, 2004.

Two new species of parasitic nematodes are described from macropodid marsupials from Western Australia.

Rugopharynx setonicis sp. nov. from the stomach of the quokka, Setonix brachyurus, belongs to the

Rugopharynx australis complex, but differs from related species in the anterior position of the nerve ring, a

shorter spicule length and the anterior position of the deirids. A second new species, from the oesophagus of

Macropus irma, is described. Cyclostrongylus irma sp. nov. is distinguished from congeners by a buccal capsule

which is wider near its anterior extremity, longer spicules and a short oesophagus. The descriptions of the two

new species provide additional evidence in support of current hypotheses for the mode of evolution of the

respective genera.

Kry Worbs : Nematoda, marsupials, Macropus, Setonix, new species, Rugopharynx, Cyclostrongylus.

Introduction

Kangaroos and wallabies harbour an extremely

diverse fauna of helminth parasites (Spratt ef al.

1991), dominated numerically by species of the

strongyloid subfamily Cloacininae Stossich, 1889

which occur principally in the stomachs and to a

lesser extent the oesophagi of these hosts (Beveridge

& Spratt 1996; Beveridge & Chilton 2001). In spite

of the fact that some 36 genera and 256 species have

been described thus far within the subfamily

(Beveridge & Chilton 2001), substantial numbers of

species remain undescribed (Spratt et a/. 1991;

Beveridge & Chilton 2001). Apart from their

numerical importance and possible role in disease

(Beveridge & Presidente 1978), the nematodes are

also of interest due to the fact that some genera

inhabit unusual sites within the gastrointestinal tract

such as the oesophagus (e.g. Cyclostrongylus

Johnston & Mawson, 1939) and in some genera,

large numbers of cryptic species are present (e.g.

Rugopharynx Moénnig, 1927), suggesting recent and

rapid evolutionary expansions within the parasite

genus (Beveridge & Chilton 1999, 2001). In

addition, recent phylogenetic studies (Beveridge &

Chilton 2001) indicate that colonization has been the

major mode of diversification rather than

cospeciation. Thus the nematode parasites of

macropodids are of particular biological as well as of

taxonomic interest.

The nematode parasites of eastern Australian

macropodid species have been studied to a much

greater degree than those of western Australia (see

summary in Beveridge & Chilton 2001), such that

“Department of Veterinary Sciences, University of Melbourne,

Parkville Vic, 3052

much descriptive work remains to be carried out in

hosts from the latter region, In this paper, two new

species of nematodes are described from Western

Australian marsupials, a new species from the

Rugopharynx australis (Ménnig, 1926) species

complex, which has apparently undergone a recent

and dramatic evolutionary expansion in macropodids

(Beveridge & Chilton 2001) and a new species of the

genus Cyclostrongylus, a genus which is highly

unusual in occurring coiled around papillae in the

oesophagus of its host.

Materials and Methods

Nematodes were collected from macropodids

obtained as road-kills. Host animals, which had been

frozen following collection, were thawed and

stomach or oesophageal content containing

nematodes was fixed in 10% formalin for

transportation to the laboratory. In the laboratory,

nematodes were extracted from fixed stomach

content, washed in water to remove formalin and

stored in 70% ethanol.

All specimens were examined after clearing in

lactophenol. Drawings were made with the aid of a

drawing tube attached to an Olympus BH2

microscope. Apical views of the anterior end and

body sections are oriented with the dorsal aspect

uppermost; extended bursae are oriented with the

ventral lobes uppermost. Measurements were made

with an ocular micrometer and are presented in mm

as the range followed by the mean in parentheses.

Morphological terminology follows Beveridge &

Chilton (1999) for the description of the new species

of Rugopharynx and Beveridge (1982) for the

description of the new species of Cyclostrongylus.

Types of new species have been deposited in the

78 A. APPAN, J. BERGFELD & I.BEVERIDGE

Figs 1-10. Rugopharynx setonicis sp. nov. from the stomach of Setonix brachyurus. 1. Anterior end, lateral view. 2. Buccal

capsule, lateral view. 3. Transverse optical section through buccal capsule. 4. Genital cone dorsal view. 5. Apical view of

cephalic extremity. 6. Spicule tip, lateral view. 7. Gubernaculum, ventral view. 8. Terminal female genitalia, lateral view.

9. Bursa, apical view. 10. Female tail, lateral view. Scale bars = 0.1 mm, I, 8, 10; 0.01 mm, 2-7, 9. Legend : A, amphid;

BC, buccal capsule; C, cementum; CC, cephalic collar; D, deirid; E, excretory pore; EB, extrabuccal support; LC, labial

collar; N, nerve ring; O, ovejector; S, submedian papilla; V, vagina.

NEW NEMATODES FROM MARSUPIALS 79

Sn

oe cease

———

i

Figs 11-14. Scanning electron micrographs of Rugopharynx setonicis sp. nov. from the stomach of Setonix brachyurus. 11.

Apical view of anterior end showing collars and stoma. 12. Submedian cephalic papilla armed with seta with bifurcate

extremity. 13. Bursa with everted spicule tips. 14. Bosses on internal surface of lateral lobe of bursa. Scale bars = 0.1 mm,

11-13, 0.01 mm, 14.

collections of the South Australian Museum,

Adelaide (SAM), the Natural History Museum,

London (BMNH) and the United States National

Parasite Collection (USNPC).

Several specimens of each species were examined

by scanning electron microscopy (SEM). Specimens

were dehydrated in an ethanol series, transferred to

hexamethyldisilasane, allowed to dry, then sputter-

coated with gold and examined with a Phillips 505

SEM using an accelerating voltage of 5-1OKV.

Results

Rugopharynx setonicis sp. nov.

(FIGS 1-14)

Types

Holotype d from stomach of Setonix brachyurus

(Quoy & Gaimard, 1830), Wellington Dam, Western

Australia (33° 24’ S 116° 00’ E), coll. R. Brazelle,

SAM 32182. Allotype 2, same data, SAM 32183.

Paratypes: 5d, 59, same data, SAM 32184; Id,

12, BMNH 2003.2.7.1-2: 146, 12, USNPC 93586.

Description

Chabertiidae (Popova, 1952); Cloacininae Stossich,

1899. Small worms; body covered with numerous fine

transverse annulations. Prominent cephalic collar

present (Fig. 2). Collar with paired lateral amphids on

prominent elevations, and 4 conical, sub-median

papillae; each papilla bearing single, medially-

directed seta; setae only visible in apical preparations

of head (Fig. 5); scanning electron micrographs

(Fig. 12) show seta arising from bulbous expansion on

medial aspect of papilla; seta bifid distally with one

branch much shorter than other. Prominent,

continuous labial collar internal to cephalic collar

(Figs. 5, 11); collar indented medial to amphids and

sub-median papillae. Stoma circular in apical views of

head; labial collar continuous internally with lining of

buccal capsule. Prominent extra-buccal supports

present surrounding anterior extremity of buccal

80 A. APPAN, J. BERGFELD & I.BEVERIDGE

capsule (Fig. 2). Buccal capsule short, thick-walled,

variable in shape, longer than wide, thicker at anterior

extremity with slight constriction at one sixth of

length. Buccal capsule with numerous transverse

striations; striations sinuous, faint, branch and

anastomose frequently; near anterior extremity,

striations faint and irregular; in optical transverse

section, striae radially arranged (Fig. 3). Oesophageal

corpus elongate, sub-cylindrical, slightly wider

towards posterior extremity (Fig. 1); isthmus short,

leading to elongate, clavate bulb. Deirids in anterior

oesophageal region. Nerve ring encircles oesophageal

corpus at '/; length, anterior to isthmus. Excretory

pore between nerve ring and isthmus (Fig. 1).

Male

Measurements of 10 specimens, types. Total length

5.30 — 6.48 (6.15); maximum width 0.28 — 0.35

(0.30); buccal capsule 0.030 — 0.040 (0.037) long by

0.015 (0.015) wide; oesophagus 0.70 — 0.85 (0.79)

long; nerve ring to anterior end 0.34 — 0.40 (0.38);

excretory pore to anterior end 0.40 — 0.50 (0.48);

deirids to anterior end 0.09 — 0.13 (0.11). Bursa short;

dorsal lobe with slight indentation in margin; dorsal

lobe equal in length to lateral lobes; lateral lobes with

crenulate margins with radially directed striae close to

margin, Internal surfaces of ventral and lateral lobes

with numerous refractile bosses; bosses overlie

ventral and lateral rays, extend in marginal band along

lateral lobes between postero-lateral and externo-

dorsal rays (Fig. 9). Ventro-ventral and latero-ventral

rays slender, apposed, reaching margin of bursa.

Medio-lateral and postero-lateral rays stout, reaching

margin of bursa; externo-lateral ray shorter, divergent,

originates close to base of lateral trunk, slightly

reflexed near distal extremity, terminating in elevation

of cuticle close to margin of bursa. Externo-dorsal ray

originates close to lateral trunk, stout, straight, almost

reaches margin of bursa. Dorsal ray broad at origin,

divides at mid-length; internal branchlets arcuate,

elongate, directed posteriorly, reach margin of bursa;

external branchlets arise at main bifurcation, slightly

shorter than internals, recurrent, terminate in

elevations of cuticle on internal surface of bursa.

Genital cone prominent; anterior lip large, conical;

posterior lip with pair of bilobed appendages and ring

of smaller projections around base (Fig. 4). Spicules

1.00 — 1.34 (1.20) long, alate; proximal tips knobbed,

shafts cylindrical, distal tips blunt with slight terminal

enlargement; alae broad, diminish in width towards

tip, lose striations (Fig. 6). Gubernaculum poorly

sclerotised, visible in lateral views but rarely visible in

dorso-ventral views (Fig. 7); cordate and paired lateral

thickenings of spicule sheaths present.

Female

Measurements of 10 specimens, types. Total length

6.75 — 7.90 (7.36); maximum width 0.28 — 0.36

(0.32); buccal capsule 0.03 — 0.05 (0.04) long by

0.015 (0.015) wide; oesophagus 0.83 — 0.96 (0.89)

long; nerve ring to anterior end 0.37 — 0.42 (0.39);

excretory pore to anterior end 0.43 — 0.57 (0.48);

deirids to anterior end 0.09 — 0.11 (0.10); tail

elongate, tapering, 0.35 — 0.50 (0.44) long; vulva

0.50 — 0.62 (0.58) from posterior end, frequently

surrounded by mass of cementum (Fig. 10); vagina

straight, short, 0.34 — 0.55 (0.43) long; ovejector

J-shaped; vestibule longitudinally disposed,

differentiation between sphincters and infundibula

indistinct (Fig. 8); egg ellipsoidal 0.11 — 0.14 (0.12)

long by 0.05 — 0.06 (0.055) wide.

Cyclostrongylus irma sp. nov.

(FIGS 15-28)

Synonyms

Cyclostrongylus wallabiae Johnston & Mawson,

1939 of Spratt et al, 1991, p. 51.

Types

Holotype 3d from oesophagus of Macropus irma

(Jourdan, 1837), Collie, Western Australia (33° 22'S

116° 09’ BE), 11.xii.2001, coll. R. Brazelle, SAM

32185. Allotype 2, same data, SAM 32186.

Paratypes: 10d, 202, same data, SAM 32187; 16,

12, BMNH 2003.2.7.3-4; 1d, 19, USNPC 93587.

Other material examined: from oesophagus of M.

irma : Western Australia: 146, 142, Perth, Feb.

1981, coll. P. Christensen (SAM 8324); 23d, 22°,

Jandakot, April, 1982, coll. L. Jue Sue (SAM 11545).

Description

Chabertiidae (Popova, 1952); Cloacininae

Stossich, 1899, Small nematodes up to 12mm long.

Mid-region of bodies coiled around oesophageal

papillae of host; males usually with a single coil

(Fig. 25), female nematodes with 2 or more coils

(Fig. 26). Body covered with numerous transverse

annulations. Single triangular ala present on the

ventral surface (Figs. 20, 27), extending along mid-

body region. Cephalic collar distinct, with 2 slightly

domed amphids and 4 domed submedian papillae.

No labial crown or collar. Mouth opening and buccal

capsule circular in apical view (Fig. 17) and optical

cross-section respectively (Fig. 18). Buccal capsule

short, thick-walled, transversely striated; all of

buccal capsule thicker anteriorly; transverse

striations regular, extend along entire buccal capsule

(Fig. 16). Extra-buccal supports prominent.

Prominent, refractile ring present between buccal

capsule and oesophagus. Oesphagus short, corpus

widening slightly before constriction at isthmus;

ovoid oesophageal bulb with 3 scelerotised plates

(Fig. 15). Excretory pore at level of deirids, latter at

NEW NEMATODES FROM MARSUPIALS 81

Figs 15-24. Cyclostrongylus irma_ sp. nov. from the oesophagus of Macropus irma 15. Anterior end, lateral view. 16.

Cephalic end, lateral view. 17. Cephalic end, apical view. 18. Transverse optical section through buccal capsule. 19.

Distal tip of spicule, lateral view. 20. Transverse section through mid-region of body showing ventral ala. 21. Bursa of

male, apical view. 22. Genital cone, dorsal view. 23. Gubernaculum and thickenings of spicule sheath, ventral view. 24.

Posterior end of female, lateral view. Scale bars = 0.1 mm, 15, 21, 23, 24, 0.01 mm, 16-20, 22. Legend : A, amphid; AL,

ala; BC, buccal capsule; C, cementum; CE, central cordate thickening of spicule sheath; CC, cephalic collar; D, deirid;

E, excretory pore; EB, extrabuccal support; G, gubernaculum; L, lateral thickening of spicule sheath; N, nerve ring; O,

ovejector; R, post-buccal sclerotised ring; S, submedian papilla; V, vagina.

82 A. APPAN, J. BERGFELD & I.BEVERIDGE

level of mid-oesophageal bulb. Nerve ring encircles

oesophageal isthmus.

Male

Measurements of 10 specimens, types. Total length

8.78 — 9.84 (9.60); maximum width 0.22 — 0.28

(0.24); buccal capsule 0.035 — 0.045 (0.042) long by

0.030 (0.030) wide; oesophagus 0.65 — 0.75 (0.71)

long; nerve ring 0.49 — 0.56 (0.52) from anterior end;

excretory pore 0.57 — 0.69 (0.62) from anterior end:

deirid 0.59 — 0.72 (0.66) from anterior end. Bursa

short, lobes of equal size (Fig. 21), Separation of

lobes indistinct; no striae or bosses on internal

surface of bursa. Ventro-ventral and ventro-lateral

rays slender, apposed, reaching margin of bursa.

Lateral rays stout; medio-lateral and postero-lateral

rays apposed, reaching margin of bursa; externo-

lateral ray divergent, joins lateral trunk near origin,

does not reach margin of bursa. Externo-dorsal ray

originates close to lateral trunk, not reaching margin

of bursa, curves slightly near tip. Dorsal ray

bifurcates '/4-length, each major branch with small

laterally-directed branch arising more than half-way

along its length; 2 major branches of the dorsal ray

reaching margin of bursa. Spicules alate, 1.25 — 1.48

(1.38) long; proximal extremity knobbed; shaft

cylindrical; distal tip blunt, slightly recurved (Fig.

19); ala diminishes in width towards tip, loses

transverse striations. Gubernaculum approximately

quadrangular in dorsoventral view (Fig. 23), 0.10 —

0.20 (0.14) long. Genital cone prominent; anterior lip

large, conical; posterior lip with 2 lobe-like

appendages (Fig. 22). Central cordate and lateral

paired thickenings of spicule sheaths present (Fig.

23).

Female

Measurements of 10 specimens, types. Total length

9.55 — 11.58 (10.81); maximum width 0.30 — 0.34

(0.32); buccal capsule 0.040 — 0.050 (0.045) long by

0.030 — 0.040 (0.034) wide; oesophagus 0.73 — 0.84

(0.79) long; nerve ring 0.52 — 0.57 (0.55) from

anterior end; excretory pore 0.61 — 0.70 (0.65) from

anterior end; deirid 0.63 — 0.71 (0.68) from anterior

end. Tail short, conical, 0.14 — 0.18 (0.16) long;

vulva immediately anterior to anus, 0.23 — 0.36

(0.29) from posterior end, frequently surrounded by

mass of cementum (Fig. 24); vagina straight, 0.08 —

0.22 (0.14) long; ovejector longitudinally disposed.

Egg thin shelled, ovoid, 0.06 — 0.09 (0.08) long by

0.04 — 0.05 (0.05) wide.

Discussion

The specimens described from the stomach of

Setonix brachyurus are allocated to the genus

Rugopharynx, due to the presence of two pairs of

branches to the dorsal ray, a cylindrical buccal

capsule and the externo-dorsal ray arising with the

lateral rays, all characters of the subfamily

Cloacininae. The prominent tranverse striations of its

buccal capsule, short oesophageal corpus and lack of

petaloid or lobed labial crown elements and alae

place it in the genus Rugopharynx. Within the genus,

the sub-cylindrical buccal capsule, lacking

prominent subdivisions, places it within the

Rugopharynx australis complex (Beveridge 1982;

Beveridge & Chilton 1999),

Within this complex, the specimens cannot be

attributed to any of the known species (Beveridge &

Chilton 1999). The nerve ring encircles the anterior

part of the oesophageal corpus rather than the

isthmus, hence differentiating them from all species

except R. petrogale Beveridge & Chilton, 1999, R,

rosemariae Beveridge & Presidente, 1978 and R. rho

Beveridge & Chilton, 1999. The specimens

described here lack the small sclerotised bosses

lining the oesophagus which are characteristic of R.

rosemariae and the length of the spicules, 1.00 —

1.34 (1.20) mm, is significantly shorter than those of

R. rosemariae (3.85 — 4.35 mm). R. rho has a non-

lobed buccal capsule but its spicules (1.65 — 1.92

mm) are significantly longer. R. petrogale has three,

very weakly-developed lobes to the wall of the

buccal capsule, compared with the current specimens

which have the buccal capsule wall slightly

thickened at its anterior extremity. In addition, the

dorsal ray in R. petrogale is longer than the lateral

rays while in the specimens described here, the rays

are of similar length. The proximity of the deirids to

the anterior end of the nematode also differ

significantly from specimens of R. petrogale. For

these reasons, the material described is considered to

represent a new species, and is named R. sefonicis sp.

nov. as the quokka, Sefonix brachyurus, is its only

known host.

Beveridge & Chilton (2001) examined the host

range of members of the R. australis complex and

concluded that the distribution of nematode species

bore no relationship to the phylogeny of the hosts.

They suggested that members of this species

complex may have evolved recently, due to the

relatively minor morphological differences between

their constituent members, and that colonization of

hosts had occurred rather than co-speciation. The

addition of a new member of the complex from

Setonix is consistent with this hypothesis given the

relatively basal (though uncertain) position of

Setonix in the phylogeny of the Macropodidae

(Flannery 1989).

Two genera of cloacinid nematode are known to

occur coiled around the oesophageal papillae of

wallabies, namely Cyclostrongylus Johnston &

Mawson, 1929 and Spirostrongylus Yorke &

NEW NEMATODES FROM MARSUPIALS 83

Figs 25-28. Cyclostrongylus irma

sp. nov. from the oesophagus of Macropus irma. 25. Male nematode. 26. Female

nematode. 27. Internal (ventral ) surface of body coil showing central ala (a). 28. Mouth opening, amphids and sub-

median papillae. Scale bars = 1.0 mm, 25, 26, 0.1 mm, 27, 0.01 mm, 28.

Maplestone, 1926 (see Beveridge 1982).

Spirostrongylus possesses a lobed labial collar, while

in Cyclostrongylus, a labial collar is totally lacking

(Beveridge 1982). The specimens described here

therefore belong to Cyc/lostrongylus and have

previously been allocated to C. wallabiae (Johnston

& Mawson, 1939) (see Spratt et a/. 1991), a species

which occurs commonly in the swamp wallaby,

Wallabia bicolor (Desmarest, 1804).

The genus Cyclostrongylus currently comprises six

species (Beveridge 1982). Morphologically, the

species of Cyclostrongylus occurring in the

oesophagus M. irma is most similar to C. wallabiae

in possessing a buccal capsule which is wider in

diameter towards the anterior end, rather than being

perfectly cylindrical and in having regular transverse

striations extending to the anterior extremity of the

buccal capsule. It was presumably for this reason

that specimens of Cyclostrongylus from M. irma

have hitherto been identified as C. wallabiae (see

Spratt ef al. 1991). However, the specimens

described from M. irma differ in the length of

spicules in the males (0.94 — 1.10 mm in C.

wallabiae; 1.25 — 1.48 mm in specimens from /.

irma). In addition, the bursa of C. wallabiae is

greatly elongated dorsoventrally (Beveridge 1982,

Fig. 324), while in the specimens from M. irma, the

bursa is wider than long (Fig. 21). As a consequence

of the difference in shape of the bursa, the stem of

the dorsal ray is relatively much longer in C.

wallabiae (see Beveridge 1982, Fig. 324) than in

material from M. irma. The length of the oesophagus

in specimens from ™. irma is also shorter than those

of C. wallabiae (0.65 — 0.75 mm in males and 0.73 —

84 A. APPAN, J. BERGFELD & I.BEVERIDGE

0.84 mm. in females from M. irma, mean lengths 9.6

and 10.8 mm respectively; 0.85 — 0.97 mm in males

and 0.87 — 1.02 mm in females of C. wallabiae,

mean lengths 10.8 and 11.6 mm respectively).

The data presented here indicate that the specimens

in M. irma are very similar but morphologically

distinct from C. wallabiae and are therefore

considered to represent a new species, herein named

C. irma sp. nov.

The presence of the ventral ala in C. irma may

assist the nematode in attaching to the papillae on the

surface of the oesophagus of its host as has been

suggested for other species (Beveridge 1982).

Beveridge & Chilton (2001) examined the

relationships between species of the genus

Cyclostrongylus and the phylogeny of their hosts.

Major problems encountered in their analysis were

lack of resolution both in host and parasite

phylogenies. However, comparisons of the

phylogenies of hosts and parasites were generally

consistent with an hypothesis of co-speciation in part

because all the host species are macropodids with an

oesophagus lined with prominent papillae (Obendorf

1984). Macropus dorsalis (Gray, 1837) parasitised

by C. leptos (Mawson, 1965), M. eugenii

(Desmarest, 1817) parasitised by C. kartana

(Mawson, 1955), M. parma Waterhouse, 1845

parasitised by C. parma (Johnston and Mawson,

1939), M. parryi Bennett, 1835 parasitised by C.

elegans Beveridge, 1982 and M. rufogriseus

(Desmarest, 1817) parasitised by C. alatus

Beveridge, 1982 and C. perplexus Beveridge, 1982

belong to a single subgenus, Notamacropus , while

C. wallabiae occurs in the related genus Wallabia.

Two potential examples of colonization rather than

host speciation were noted (Beveridge & Chilton

2001), one of which was the purported occurrence of

C. wallabiae in both W. bicolor and M. irma. The

description of the new species in M. irma thus

potentially excludes one of these examples of

colonisation and further supports the hypothesis of

co-speciation between hosts and parasites in the

genus.

Beveridge & Chilton (2001) argued that the

cloacinine nematodes of macropodid marsupials

may represent excellent models for the study of the

way in which mechanisms such as co-speciation and

colonization interplay in the evolution of parasitic

nematodes, with their study suggesting that

colonization is the more significant mechanism.

However, critical for such studies is a sound

taxonomic basis. The description of the two

nematodes presented in this paper provides support

on the one hand for the colonization hypothesis

advanced for species of the R. australis complex and

on the other hand for the co-speciation hypothesis

advanced in the case of the genus Cyclostrongylus

(see Beveridge & Chilton 2001).

Acknowledgements

We wish to express our sincere thanks Rob Brazelle

for making specimens available and to Joan Clark for

assistance with scanning electron microscopy.

References

BeveripGe, I. (1982) A taxonomic revision of the

Pharyngostrongylinea Popova (Nematoda; Strongy-

loidea) from macropod marsupials. Aust. J. Zool. Suppl.

Ser. No. 83, 1-150.

& CuiLtton, N. B. (1999) Revision of the

Rugopharynx australis (Moennig, 1926) complex

(Nematoda: Strongyloidea) from = macropodid

marsupials. /nvert. Taxon. 13, 805-843.

& (2001) Co-evolutionary

relationships between the nematode subfamily

Cloacininae and its macropodid marsupial hosts. Jn. J.

Parasitol. 31, 976-996.

& PRESIDENTE, P. J. A. (1978) Rugopharynx

rosemariae new species (Nematoda: Pharyngo-

strongylidae) from grey kangaroos (Macropus giganteus

and M. fuliginosus) with life cycle stages and associated

pathology. /bid. 8, 379-387.

& Sprart, D. M. (1996) The helminth fauna of

Australasian marsupials: origins and evolutionary

biology. Adv. Parasitol. 37, 135-254.

FLANNERY, T. F. (1989) Phylogeny of the Macropodoidea: a

study in covergence. /n G. Grigg, P. Jarman, I. Hume

(Eds) “Kangaroos, Wallabies and Rat-kangaroos”

(Surrey Beatty & Sons : Chipping Norton) pp. 1-46.

Openporr, D. L. (1984) The macropodid oesophagus I.

Gross anatomical, light microscopic, scanning and

transmission electron microscopic observation of its

mucosa. Aust. J. Zool. 32, 415-435.

Spratt, D. M., BeveripGe, I., & WALTER, E. L. (1991) A

catalogue of Australasian marsupials and monotremes

and their recorded helminth parasites. Rec S$. Aust. Mus.

Monogr. Ser. 1, 1-105.

A CATALOGUE OF SOUTH AUSTRALIAN FRESHWATER

FISHES, INCLUDING NEW RECORDS, RANGE EXTENSIONS

AND TRANSLOCATIONS

By M. P. HAMMER*} & K. F. WALKER*

Summary

Hammer, M. P. & Walker, K. F. (2004) A catalogue of South Australian freshwater

fishes, including new records, range extensions and translocations. Trans. R. Soc. S.

Aust. 128(2), 85-97, 30 November, 2004.

Published data, recent surveys and studies of museum specimens are combined to

provide a list of 84 fishes for South Australia in five drainage divisions. The list

includes 58 native species (44 restricted to freshwater) and 26 alien species. Seven

endemics are recognised, namely Chlamydogobius eremius (Zeitz), Chlamydogobius

gloveri Larson, Craterocephalus dalhousiensis Ivanstoff & Glover, Craterocephalus

eyresii (Steindachner), Craterocephalus gloveri Crowley & Ivanstoff, Mogurnda

thermophila Allen & Jenkins and Neosilurus gloveri Allen & Feinberg.

Key Words: Freshwater fishes, conservation, management, taxonomy.

Transactions of the Royal Society of S. Aust. (2004), 128(2), 85-97.

A CATALOGUE OF SOUTH AUSTRALIAN FRESHWATER FISHES, INCLUDING

NEW RECORDS, RANGE EXTENSIONS AND TRANSLOCATIONS

by M. P. HAMMER" & K. F. WALKER*

Summary

HAMMER, M. P. & WALKER, K. F. (2004) A catalogue of South Australian freshwater fishes, including new

records, range extensions and translocations. Zrans. R. Soc. S. Aust. 128(2), 85-97, 30 November, 2004.

Published data, recent surveys and studies of museum specimens are combined to provide a list of 84 fishes

for South Australia in five drainage divisions. The list includes 58 native species (44 restricted to freshwater)

and 26 alien species. Seven endemics are recognised, namely Chlamydogobius eremius (Zeitz),

Chlamydogobius gloveri Larson, Craterocephalus dalhousiensis Ivanstoff & Glover, Craterocephalus eyresii

(Steindachner), Craterocephalus gloveri Crowley & Ivanstoff, Mogurnda thermophila Allen & Jenkins and

Neosilurus gloveri Allen & Feinberg. New records are reported for Craterocephalus stercusmuscarum ?stercus-

muscarum (Giinther), Galaxias truttaceus Valenciennes and Neochanna cleaveri (Scott), and a terapontid of

uncertain status also is noted. Range extensions are reported for Nannoperca obscura (Klunzinger), Nannoperca

australis Giinther and an undescribed species of Hypseleotris, and the presence of Galaxias olidus Giinther and

Galaxias brevipinnis Giinther in particular regions is confirmed. Possible extirpations are reported for Ambassis

agassizii Steindachner, Gadopsis marmoratus Richardson, Galaxias rostratus Klunzinger, Maccullochella

macquariensis (Cuvier), Macquaria australasica Cuvier, Mogurnda adspersa (Castelnau), Neochanna cleaveri

and Prototroctes maraena Giinther. There is need for further evaluations of fish distributions, better systematic

frameworks, clarifications of conservation status, reviews of the introduction and impacts of alien species and

development of protective measures for fish species and communities and their ecosystems.

Key Worpbs: Freshwater fishes, conservation, management, taxonomy

Introduction

Despite a generally dry landscape, South Australia

harbours a diverse array of aquatic habitats including

artesian mound springs, swamps, lakes, episodic

streams and the River Murray and associated

wetlands. These habitats, and the effects of

biogeographical isolation (e.g. Unmack 2001),

sustain a corresponding diversity of freshwater biota.

The term “fresh water” here includes inland saline

waters (>3000 mg L"!), as these are common in the

state (e.g. Williams 1967; EPA 1998; Hammer

2002a).

Freshwater fishes in South Australia display a

variety of physical forms and life histories. The

dwarf galaxias Galaxiella pusilla is remarkable for

its ability to survive dry periods in seasonal swamps,

where it takes refuge in swamp-crayfish burrows

(Geocharax: Beck 1985). Large species like the

Murray-Darling golden perch Macquaria ambigua

ambigua may cover long distances (for example, a

tagged fish is known to have travelled 2300 km

along the Murray and Darling rivers: Reynolds

* Cooperative Research Centre for Freshwater Ecology, School of Earth

and Environmental Sciences DP312, The University of Adelaide,

Adelaide, SA 5005. Email: michael.hammer@adelaide.edu.au

' Evolutionary Biology Unit, South Australian Museum.

' HAMMER, M. (2001) Molecular systematics and conservation

biology of the southern pygmy perch Nannoperca australis

(Giinther, 1861) (Teleostei: Percichthyidae) in south-eastern

Australia. Unpub. BSc(Hons) Thesis, Department of

Environmental Biology, The University of Adelaide.

1983), whereas small species like the southern

pygmy perch Nannoperca australis are much less

vagile (Hammer'). Other species need to move

between fresh water and marine habitats, although

even diadromous species like the galaxiids Galaxias

maculatus and G. brevipinnis may occur in

‘landlocked’ populations (Pierce ef al. 1985;

Hammer 2002a; SKM 2002). In addition, there are

euryhaline species like the small-mouthed hardyhead

Atherinosoma microstoma, found in fresh or salt

water (Molsher ef a/. 1994; Hammer 2002a).

This catalogue lists 84 species in the freshwater

fish fauna of South Australia. It updates earlier work

(Waite 1923; Scott et al. 1974; Sim 2000), corrects

and amends records of species and _ their

distributions, and is designed to assist in research

and planning for management and conservation.

Methods

Drainage divisions

Five of the 13 principal drainage divisions in

Australia (AWRC 1976) occur wholly or partly in

South Australia, and provide a biogeographic

framework (Fig. 1):

* South East Coast (SEC), including the Millicent

Coast and Glenelg River (part) river basins,

* Murray Darling (MD), part of the Lower Murray

River Basin,

¢ South Australian Gulf (SAG), the only division

contained wholly within the state (the shared

86 M. PD. HAMMER & K. F. WALKER

oe ees |< || om etres

0 LOO, 200 400

~

AUSTRALIAN DRAINAGE DIVISIONS

North-east Coast

South-east Coast

Tasmanian

Murray-Darling

South Australian Gulf

South-west Coast

Indian Ocean

Timor Sea

Gulf of Carpentaria

Lake Eyre

Bulloo-Bancannia

Western Plateau

Distant Islands (not shown)

Fig. 1. Drainage divisions in Australia and South Australia (AWRC 1976).

FRESHWATER FISH SPECIES RICHNESS IN SOUTH AUSTRALIA 87

boundary with MD is west of the Murray Mouth,

but SAG includes the coastal streams of Fleurieu

Peninsula),

* Lake Eyre (LE), draining toward lakes Eyre and

Frome, and

* Western Plateau (WP), containing sparse coastal

lakes and some ephemeral waters.

Records of species were obtained from the

literature and examination of specimens at the South

Australian Museum, Adelaide (SAMA), including

material from recent collections by the senior author,

Information on rare or doubtful species was

scrutinised with special care.

Nomenclature

The systematic framework and nomenclature

employed here follow Eschmeyer (1998) and

subsequent updates (see Californian Academy of

Sciences on-line “Catalogue of Fishes”, March

2003"), except that the lamprey families Geotriidae

and Mordaciidae replace Petromyzontidae (Strahan

1980), subspecific status is recognised for M. a.

ambigua (after Musyl & Keenan 1992) and six

informal taxa and a species complex are recognised.

The informal taxa include dwarf flathead gudgeon

Philypnodon sp. (Larson & Hoese 1996), Lake Eyre

golden perch Macquaria sp. (Musyl & Keenan 1992)

and western chanda perch, an undescribed species

referred to in earlier literature as “Ambassis muelleri

Klunzinger” (syn. A. agassizii), but lacking a formal

name since “4. mueller” was invalidated by Allen et

al, (2002). The carp gudgeon genus Hypseleotris

awaits a formal review but, following Allen er al.

(2002), this catalogue recognizes Midgley’s carp

gudgeon H. “sp. 1” sensu Hoese et al. (1980) and

Murray-Darling carp gudgeon H. “sp, 3” sensu

Unmack (2000). In addition, a species complex of

hybrids and possible semi-clonal hybridogenic forms

are recognised (Bertozzi et a/. 2000), including

Lake’s carp gudgeon 7. “sp. 2” sensu Hoese ef al.

(1980). Following Allen and Jenkins (1999), prior

records of northern purple-spotted gudgeon

Mogurnda mogurnda (Richardson) in South

Australia should be referred to Dalhousie purple-

spotted gudgeon M. thermophila or Flinders Ranges

purple-spotted gudgeon M. clivicola (these were

described from within the range of M. mogurnda).

Criteria for inclusion

A “freshwater” species here includes obligate

freshwater and diadromous species and _ select

euryhaline taxa known to complete their lifecycle in

fresh water. “Alien” species include exotic species

(not native to Australia) and native Australian

* http://www.calacademy.org/research/ichthyology/catalog/fishcatmain.asp

species translocated outside their natural range.

Alien fishes in natural waterways are regarded as

established species if their populations are self-

sustaining or if they are continually stocked, and as

introduced species if records are few and isolated or

confined to artificial waterbodies (and potentially

could become established). The latter include

interstate translocations within drainage divisions.

Results

Native fish richness

A total of 58 native freshwater fish species in 15

families is recorded for South Australia (Table 1). All

are shared with other states, except for seven

endemics in isolated areas of LE. Mogurnda

clivicola may be another endemic, as only small

populations of uncertain affinity occur outside the

state (Allen & Jenkins 1999; Wager & Unmack

2000).

Forty-four native species are confined to fresh

water. One of these, Australian smelt Retropinna

semoni, may occasionally occur in the Coorong

(Eckert & Robinson 1990), but is not strictly

diadromous. Four euryhaline taxa meet the

aforementioned criteria of “freshwater” species,

namely A. microstoma, flathead gudgeon

Philypnodon grandiceps, western bluespot goby

Pseudogobius olorum and lagoon — goby

Tasmanogobius lasti (e.g. Wedderburn & Hammer

2003). Thirteen of the 44 obligate freshwater species

occur in more than one division, and none is

common to all. Most obligate freshwater species

occur in LE (24) and MD (24, plus 11 diadromous

and euryhaline taxa). Diadromous and euryhaline

species generally occur in more than one division.

Remarkably, three diadromous species are recorded

for WP, although data there are sparse (Table 1).

New records for South Australia

Fly-specked hardyhead

Craterocephalus stercusmuscarum

?stercusmuscarum (Giinther)

This taxon was identified in samples collected

from the northern Flinders Ranges in 1994-95

(SAMA F7331, F9002, F9078). It is distinguished

from the Lake Eyre hardyhead Craterocephalus

eyresii (Steindachner), which occurs in the same

region but not the same habitats, by fewer transverse

scale rows (7-8 cf. 11-14 in C. eyresii) and dark

lateral banding (Ivanstoff et a/. 1987; Crowley &

Ivanstoff 1990a). Subspecific identification is

tentative owing to taxonomic problems and_ the

isolated nature of the population (the nearest known

conspecifics are from Aramac Springs in the remote

M. P.- HAMMER & K. F. WALKER

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89

HNESS IN SOUTH AUSTRALIA

FRESHWATER FISH SPECIES RIC

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UOISIAIG, oueu UOUWOZ UOXeL Aqrure J

90 M. PD. HAMMER & K. F. WALKER

upper reaches of Cooper Creek, Queensland). A

molecular revision of Craterocephalus in progress

indicates that sub-species within the C.

stercusmuscarum species complex remain confused

(P. Unmack, Arizona State University, pers. comm.)

and further morphological and molecular analyses

are required.

Spotted galaxias

Galaxias truttaceus Valenciennes

This species was first reported in 1999 from karstic

springs in coastal south-eastern South Australia (e.g.

Ewens Ponds: Hammer ef a/. 2000; SAMA F9217,

F10111) representing a minor westward range

extension into South Australia. Another single

specimen from the same area occurred among

specimens of G. maculatus collected in 1979

(SAMA F10109). Note that a prior report of G.

truttaceus from SAG (Scott ef al. 1974) was based

on misidentified specimens (SAMA F3094, F3188).

Tasmanian mudfish

Neochanna cleaveri (Scott)

This species is known in South Australia only from

a single specimen collected from Bool Lagoon in

1974, and previously registered as G. maculatus

(SAMA F4919). Recent surveys have failed to locate

others (Hammer 2002a). The new record is

noteworthy as the species is cryptic, with an ability

to survive extended dry periods by burrowing into

mud or hiding under rocks and wood, and otherwise

is native to Tasmania and Victoria (Fulton 1986;

Koehn & Raadik 1991).

A possible new terapontid

A form of grunter (Terapontidae) resembling a

deep-bodied Welch’s grunter Bidvanus welchi or a

hybrid B. welchi x Barcoo grunter Scortum barcoo is

known from Coongie Lakes (J. Puckridge,

University of Adelaide, pers. comm. 2001). This

form is listed as the ‘Cooper grunter’ by Sim (2000).

It was also reported near Goyder Lagoon on the

lower Warburton River in 2002 (Costelloe ef al.

2003).

Range extensions

Surveys in the Mount Lofty Ranges (Hammer')

have provided three new drainage division records,

namely a genetically distinct sub-population of

Nannoperca australis from the Inman River

Catchment (SAG), Hypseleotris sp. 3 from the same

location, and Yarra pygmy perch Nannoperca

obscura from Lake Alexandrina (MD). The review

uncovered other, previously misidentified specimens

of N. obscura in the museum collection dating from

1915 (SAMA F572), suggesting the species is native.

The presence of mountain galaxias Galaxias olidus

(a species complex presently under systematic

review: Raadik 2001) recently was confirmed from

the South Australian section of SEC (Mosquito

Creek: Hammer 2002a). Despite its inclusion in a

south east regional list by Glover (1983), no

specimens of the species were previously known. In

addition, Glover mistakenly referred to the Mosquito

Creek population as G. maculatus. The presence of

G. brevipinnis in MD is also confirmed (SAMA

F153: Angas River, 1914; previously registered as G.

maculatus), a record predating the Snowy Mountains

Hydroelectric Scheme which appears to be the

source of G. brevipinnis in the upper Murray

catchment (Waters et a/. 2002).

A report of R. semoni from SAG (SKM 2002) is

suspect because voucher specimens are not available

and no other records exist for the division (e.g.

McDowall 1979; Unmack 2001). Other SAG reports

of bony herring Nematalosa erebi and spangled

grunter Leiopotherapon unicolor in the Lake Torrens

catchment, and western carp gudgeon Hypseleotris

klunzingeri as native to the Broughton River (Pierce

et al. 2001) are also discounted in the absence of

voucher specimens or other data, There is an

uncertain report of fish resembling C. eyresii in the

remote, isolated Durkin Swamp (WP), following

exceptional rainfall (Ehmann & Tynan 1997).

Finke goby Chlamydogobius japalpa Larson,

Finke hardyhead Craterocephalus centralis Crowley

& Ivanstoff and Finke purple-spotted gudgeon

Mogurnda larapintae (Zeitz) potentially could

colonise the ephemeral, lower reaches of the Finke

River in South Australia, following floods from the

headwaters in the Northern Territory, but they have

not been formally recorded.

Alien species

There are records of 26 alien species in South

Australia (Tables 2-3), although two may prove to be

natives (Philypnodon sp. from the Onkaparinga

River (SAG) (SAMA F10087, April 2002), and

Murray rainbowfish Melanotaenia fluviatilis from

SEC (SAMA F2409, dated 1903)). Most alien

species records are for SAG (20 species, including

13 established alien species). There are high numbers

also for MD and SEC, but few in the remote LE and

WP (Table 2).

Fourteen alien species are established in South

Australia. These include seven exotic taxa and seven

translocated native taxa. Another 12 alien species

have been introduced, but are not established or

present only in artificial waterways (Tables 2-3).

These include barramundi Lates calcarifer in the

River Torrens and Australian bass Macquaria

91

S RICHNESS IN SOUTH AUSTRALIA

“TE

FRESHWATER FISH SPE!

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M. P- HAMMER & K. F. WALKER

SAMA F6199 (1986); Reid & Puckridge (1990)

SAMA F7331, F9002, F9078 (1994/95)

Glover (1985); Hammer pers. obs. 2002

Glover (1980); Pierce ef al. 2001

SAMA F5496 (1981), F7405 (1982)

SAMA 2615 (1947), F4789 (1984)

SAMA F1388 (1929)

Wager & Unmack (2000)

SAMA F10056 (1947)

Pierce (1990)

Glover (1979)

Source

Leigh Creek retention dam. Poisoning attempted, but still present in 1999 Pierce ef al. (2001)

(Hammer pers. obs.)

Cooper Creek near Innamincka. Population small, may not be viable.

Stocked into Clayton Bore.

Introduced to Moro George, Flinders Ranges. Now probably absent.

WESTERN PROVINCE DRAINAGE DIVISION

Several regional records (e.g. spring at L. Hamilton; L. Newland).

Streaky Bay (not strictly freshwater habitat but included),

Specimens from the North Flinders Ranges previously identified as

most westerly record.

C. eyresii. Taxonomic status requires further investigation.

First record Neales R. (1984). Common there in 2002.

Coongie Lakes/ Cooper Creek.

Dams, reservoirs at Woomera (with Gambusia holbrooki).

Davenport Creek near Ceduna and Laura Bay.

Spring at L. Hamilton (extant?).

1

2

3

4

4

4

2

2

3

4

Record Details

Type

LAKE EYRE DRAINAGE DIVISION

Macquaria ambigua ambigua and 4

Bidyanus bidyanus

Craterocephalus stercusmuscarum

Perca fluviatilis

?stercusmuscarum

Maccullochella peelii peelii

Atherinosoma microstoma

Pseudaphritis urvillii

Gambusia holbrooki

Pseudogobius olorum

Carassius auratus

Amniataba percoides

Carassius auratus

Cyprinus carpio

Species

novemaculeata, sleepy cod Oxyeleotris lineolata and

Atlantic salmon Sa/mo salar in the River Murray.

Gambusia Gambusia holbrooki and goldfish

Carassius auratus were recorded in all drainage

divisions.

Four large native MD species (silver perch

Bidvanus bidyanus, Murray cod Maccullochella

peelii peelii, freshwater catfish Tandanus tandanus,

M. a. ambigua) are spawned in commercial

hatcheries in other states and are commonly

introduced to South Australia (Tables 2-3), including

undocumented stockings in farm dams in MD and

SAG.

Translocations in drainage divisions within South

Australia are not considered in detail here, but have

reportedly included transportation of M. clivicola in

the Flinders Ranges region and fish from Cooper

Creek to a retention dam at Leigh Creek (see Pierce

et al. 2001).

Extirpations and species decline

Museum records are not necessarily a_ true

indication of range and abundance, but indications

from all sources combined are that there have been

significant declines in the range of several species.

Records for some species may represent occasional

stray individuals on the fringe of their geographic

range, but these could not be distinguished from

established species due to a paucity of detailed

historic surveys and/or temporal replication.

There is historical evidence (Table 3) that Murray

galaxias Galaxias —_rostratus, trout — cod

Maccullochella macquariensis and Macquarie perch

Macquaria australasica formerly occurred in MD in

South Australia. Ambassis agassizii was last

recorded from the Marne River mouth (MD) in 1983

(Lloyd & Walker 1986), and state-wide extirpation

appears confirmed for the southern purple-spotted

gudgeon Mogurnda adspersa (last record in MD

1973: SAMA F3727; no sightings in SAG for >50

years). The river blackfish Gadopsis marmoratus

can be considered extirpated from SAG (it may

persist on Kangaroo Island, but the record is

dubious: Table 3) and has undergone significant

range contraction in MD (Sim ef al. 2000),

exacerbated since 1997 by the loss to irrigation

diversions of more than half of the spring-fed

habitats in the Marne River, one of few remaining

refuges (Hammer 2002b). Similarly, range

contraction and on-going local extirpations have

been recorded for N. australis (Hammer'). Estuary

perch Macquaria colonorum was once more

widespread in the lower Murray prior to the

construction of barrages near to the Murray Mouth

(Sim ef al. 2000). For SEC, N. cleaveri and the

Australian grayling Prototroctes maraena have not

been reported since 1974 and 1982 respectively and

FRESHWATER FISH SPECIES RICHNESS IN SOUTH AUSTRALIA 95

other SEC species including G. pusilla have likely

suffered large range reductions coinciding with

massive loss of wetland habitat (Hammer 2002a).

Other species are confined to small areas, including

five endemic species in Dalhousie Springs (LE)

(Wager & Unmack 2000), M. clivicola (recorded only

from Balcanoona Creek in the Flinders Ranges (LE):

e.g. SAMA ~ F3042), Murray — hardyhead

Craterocephalus fluviatilis (very few sites in the

lower Murray (MD): Lloyd & Walker 1986;

Wedderburn & Hammer 2003); N. obscura (three

habitat fragments in SEC and a small section of MD:

Hammer 2002a; Wedderburn & Hammer 2003) and

the variegated pygmy perch Nannoperca variegata (a

4-kim? spring-fed area in SEC: Hammer ef a/. 2000).

Discussion

This catalogue is a contribution toward an

inventory of state and regional biodiversity. Well-

maintained historic collections and voucher

specimens are critical to record information, validate

doubtful records and sustain progress in taxonomy,

ecology and conservation. Ideally, this information

should be updated frequently, as work progresses.

Although surface waters in South Australia are

limited (NLWRA 2001), the state harbours about one

fifth of the continental freshwater fish fauna. As the

state borders intersect, rather than enclose, some

drainage divisions, and as most divisions allow

access to the sea, the number of endemic species is

comparatively low. Some ‘new’ records here arise

from minor re-alignments of physiographic

boundaries between drainage divisions or states (e.g.

South Australian Gulf Drainage Division: N.

australis; South East Coast Drainage Division: G.

truttaceus), but others represent significant range

extensions (e.g. Murray Darling Drainage Division:

N. obscura; South East Coast Drainage Division: N.

cleaveri; Lake Eyre Drainage Division: C. s,

?stercusmuscarum).

Biodiversity assessments and monitoring should

favour obligate freshwater fishes isolated within

particular drainage divisions or regions, because they

are most likely to have diverged (cf. Crowley &

Ivanstoff 1990a,b; Musyl & Keenan 1992; Larson

1995; Allen & Jenkins 1996; Allen & Feinberg 1998;

Hammer'). These studies may gain impetus from

assessments of ecosystem ‘health’, as fishes are

* LLoyp, L. N. (1987) Ecology and distribution of the small native fish

of the lower River Murray, South Australia, and their interactions with

the exotic mosquitofish, Gambusia affinis holbrooki. Unpub. MSc

Thesis, Department of Zoology, The University of Adelaide.

potential indicators (e.g. Harris 1995). Clarifications

are needed in regard to the taxonomy of undescribed

taxa, species complexes and the biogeographic status

of some species, especially where there are few

historical data. Fine-scale molecular markers may

help to distinguish natural and translocated

populations (e.g. Waters ef al. 2002).

Alien freshwater fishes are ubiquitous in South

Australia. They are most apparent in areas directly

affected by human industry, particularly in the

Murray Darling and South Australian Gulf drainage

divisions. All such species are potential vectors for

pathogens and parasites (e.g. Langdon & Humphrey

1987). Predators like brown trout Salmo trutta,

rainbow trout Oncorhynchus mykiss and European

perch Perca fluviatilis are implicated in the decline

of small native fishes (e.g. Crowl ef al. 1992;

Morgan ef al. 2002). Gambusia holbrooki is an

aggressive, highly fecund competitor that

undoubtedly has affected native species (e.g.

Lloyd’). The feeding behaviour and high abundance

of common carp Cyprinus carpio have contributed to

destruction of wetlands associated with the River

Murray (e.g. Sim ef al. 2000), and thereby affected

native fishes. There is also some risk of genetic

contamination of native stocks by translocated native

species (Arthington 1991).

The preservation of native biota is a management

priority in South Australia (e.g. Kahrimanis ef al.

2001; EPA 2003), and avenues for the introduction of

non-native fishes such as the government-sanctioned

releases of salmonids, sales of fingerling angling

species to the public, “conservation” stocking,

releases of unwanted aquarium fishes and inter-basin

transfers from the River Murray all need review

within broadly-based programs of flow and habitat

protection, particularly where small isolated

populations of native fish occur,

Acknowledgements

We gratefully acknowledge access to the

collections of the South Australian Museum,

financial support to MPH from the Nature

Foundation (SA) and technical help from S.

Wedderburn. T. Raadik, Arthur Rylah Institute,

Melbourne, kindly identified several galaxiids and

assisted with information on G. rostratus and T.

Trnski confirmed the identity of hardyheads at the

Australian Museum, Sydney. Our thanks also to

members of Native Fish Australia (SA) for field

assistance and to A. George for advice on GIS data.

Comments on a draft manuscript by two referees

were greatly appreciated.

96 M. P. HAMMER & K. F. WALKER

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(2001) Biogeography of Australian freshwater

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WAGER, R. & UNmack, P. J. (2000) “Fishes of the Lake

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Walrte, E. R. (1923) “The Fishes of South Australia”

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THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE

GENERA BOONGURRUS LARSON, TJIRTUDESSUS WATTS &

HUMPHREYS AND NIRRIPIRTI WATTS AND HUMPHREYS,

FROM UNDERGROUND WATERS IN AUSTRALIA

By C. H. 8S. WATTS* & W. F. HUMPHREYST

Summary

Watts, C. H. S. & Humphreys, W. F. 2004. Thirteen new Dytiscidae (Coleoptera) of

the genera Boongurrus Larson Tjirtudessus Watts & Humphreys and Nirripirti Watts

& Humphreys, from underground waters in Australia. Trans. R. Soc. S. Aust. 128(2),

99-129, 30 November, 2004.

Thirteen new species of stygobitic Dytiscidae from inland Western Australia are

described: Tjirtudessus hillviewensis sp. nov., T. microocula sp. nov., T. occidentalis

sp. nov., T. padburyensis sp. nov., T. wogarthaensis sp. nov., Nirripirti arachnoides

sp. nov. N. bulbus sp. nov., N. byroensis sp. nov., N. copidotibiae sp. nov., N.

dingbatensis sp. nov. N. eurypleuron sp. nov. N. innouendyensis sp. nov., and N.

verrucosus sp. nov. This brings the total of stygobitic Dytiscidae decribed from

Australia to 55, derived from 33 discrete groundwater calcretes in 8 palaeodrainages.

Key Words: Coleoptera, Dytiscidae, Stygobitic, Descriptions, New Species, Water

chemistry.

Transactions of the Royal Society of S. Aust. (2004), 128(2), 99-129.

THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS

LARSON, TJIRTUDESSUS WATTS & HUMPHREYS AND NIRRIPIRTI WATTS AND

HUMPHREYS, FROM UNDERGROUND WATERS IN AUSTRALIA.

by C. H. S. WatTs* & W. F. HUMPHREYS*

Summary

Warts, C. H. S. & Humpnreys, W. F. 2004. Thirteen new Dytiscidae (Coleoptera) of the genera Boongurrus

Larson Tjirtudessus Watts & Humphreys and Nirripirti Watts & Humphreys, from underground waters in

Australia. Trans. R. Soc. S. Aust. 128(2), 99-129, 30 November, 2004.

Thirteen new species of stygobitic Dytiscidae from inland Western Australia are described: Tjirtudessus

hillviewensis sp. nov., T: microocula sp. nov., T. occidentalis sp. nov., T. padburyensis sp. nov., T. wogarthaensis

sp. nov., Nirripirti arachnoides sp. nov., N. bulbus sp. nov., N. byroensis sp. nov., N. copidotibiae sp. nov., N.

dingbatensis sp. nov. N. eurypleuron sp. nov., N. innouendyensis sp. nov., and N. verrucosus sp. nov. This brings

the total of stygobitic Dytiscidae described from Australia to 55, derived from 33 discrete groundwater calcretes

in 8 palacodrainages. One species, 7. microocula sp. nov., has partial eyes and wings that are only slightly

reduced. Boongurrus occidentalis sp. nov. was collected from both surface and underground water and is little

modified for an underground existence. Geographically the new species extend the range of stygobitic

Dytiscidae in Australia to the Murchison and Moore drainage systems both of which drain to the Indian Ocean.

Chemical and physical characters of the groundwater in some of the calcretes in which the new species were

found are given.

Key Worbs: Coleoptera. Dytiscidae. Stygobitic. Descriptions. New species. Water chemistry.

Introduction

This is the sixth paper in our series describing the

stygobitic Dytiscidae of Australia (Watts and

Humphreys 1999, 2000, 2001, 2003; Balke e¢ al.

2004). In it we describe the new species found during

fieldwork in Western Australia in winter 2002. Four

new species in the Bidessine genus Tjirtudessus

Watts & Humphreys, and 8 in the Hydroporine genus

Nirripirti Watts and Humphreys, are described from

the westerly-draining Murchison and Moore

paeleodrainage systems. A new species of the genus

Boongurrus Larson is described from material

collected mainly from interstial sand/gravels in

seasonal creeks in the Pilbara but also from three

bore holes accessing two different underground

calcretes. This new Boongurrus species is fully

sighted and winged and shows little apparent

physical adaptation to a hyporean existence yet was

found together with a true stygobitic fauna in deep

calcrete.

For the first time in Australia two stygobitic

species (only one described here due to lack of a

male specimen in one species) were discovered with

a true eye remnant, as distinct from the usual small

sclerite or short suture line, and wings that, although

rather small, still retain veins and folded tips

“C. H. S. Warts, South Australian Museum, North Terrace,

Adelaide, South Australia 5000.

* W. F. Humpnreys, Western Australian Museum, Francis Street,

Perth, Western Australia 6000.

suggesting that they are at an earlier stage of

adaptation to underground life than the other species

so far discovered.

Geographically the new finds extend the known

distribution of stygal Dytiscidae some 300

kilometres to the west. We also recollected sites in

the Northern Territory that yielded stygal Dytiscidae

in 2001 without discovering any additional species.

In addition, areas of groundwater calcretes near The

Granites, Tennant Creek and in the Amadeus basin in

the Northern Territory were sampled extensively

without finding any Dytiscidae and only a sparse

stygobitic fauna (Syncarids, Copepods, Amphipods)

at the occasional site. Although too early to be sure

the results suggest that the Australian stygobitic

dytiscid fauna is restricted to inland Western

Australian and the Ngalia basin in central Australia.

The prime aim of this series of papers is to

formally describe the dytiscid fauna and to provide a

preliminary indication of the ground water

characteristics in which they are found. Companion

papers by and with co- workers are starting to

address questions of phylogeny and evolution (Balke

et al. 2003, Cooper et al. 2002, Leys et al. 2003) and

the taxonomic composition of the rich fauna

associated with the beetles (Taiti and Humphreys

2001, Karanovic and Marmonier 2002, Karanovic

2004).

Materials and Methods

The collection methods and measurements of

100 C.H.S. WATTS & W. F. HUMPHREYS

physico-chemical parameters in the water largely

follow those used previously (Watts and Humphreys

2000) except that the use of a Quanta-G (Hydrolab

Corporation, Austin, Texas) water quality monitoring

system attached to a 50 m cable permitted the

measurement of various physico-chemical water

quality parameters (temperature, specific

conductance (or TDS), pH, dissolved oxygen (%

saturation or mg L-'), oxidation-reduction potential

(redox), and depth, the latter facilitating the

determination of any vertical stratification present in

the water column in some boreholes. The instrument

was calibrated against the standards recommended

for the instrument.

Abbreviations used:

BES Prefix for field numbers, WAM

Biospeleology.

SAMA South Australian Museum, Adelaide.

WAM __ Western Australian Museum, Perth.

MB Groundwater monitoring bore.

Systematics

Key to Australian species of stygobitic

Dytiscidae

1 — Scutellum well developed; length 4.5 mm

Copelatus abditus Balke et al.

— Scutellum absent; length 1.0 to 4.9 mm..2

2(1) — Paramere one-segmented; metatibia

approximately the same — width

throughout; without pronotal plicae

(Hydroporint) .......cseecesesseeeseeteeteeeenees 34

— Paramere two-segmented; metatibia

narrow at base then strongly expanding

towards apex; usually with pronotal

plicae (Bidessini) «0.0... 3

Ba(Z)v "With: YES. ei, esceectenareeeeeprzenperisisneaseonsd +

— Without eyes, may have a_ small

chitinized plate or suture line where eyes

Hortinally Ave ccs seass cet svececeegeeeeter sessed 5

4 (3) — Eyes of normal size; with elytral plicac ..

pubs egies Boongurrus occidentalis sp. nov.

— Eyes approximately one-fifth normal

size; without elytral plicae........... ee

Hants Tjirtudessus microocula sp. nov.

5 (4) — Body length approximately 1.0 mm; legs

stout, without swimming-hairs on fore

and midlegs ...........05 Kintingka kurutjutu

Watts and Humphreys

— Body length > 1.2 mm; legs normal, all

with swimming-haifS ...........:00eceeeeeee 6

6 (5) — Mesofemur with spines on hind edge

approximately the same strength as those

on mesotrochanter; length > 3.0 mm...29

— Mesofemur with spines on hind edge

much more robust than those on

7(6) —

11(9) —

nny —

13 (8) —

14 (13) —

15 (14) —

mesotrochanter; length 1.4 — 3.6 mm....7

Normal ventrites 1 and 2 without suture

between them (ie. number of visible

abdominal segments reduced to four)

(Fig. 82); length 3.2 — 3.6 mm............0+

Sebwatboptanaces Tjirtudessus sweetwatersensis

Watts and Humphreys

Ventrites 1 and 2 with suture between

them, at least in inner portion (Figs 83-

86); length 1.3 — 3.2 MM... eee 8

Pronotal plicae strong, well marked,

excavated On INSIdC.........cceeeeeeeeeeees 9

Pronotal plicae weak, difficult to trace,

may be absent, not excavated on inside .

sia doneppfthepaen emencdey-=ocapenqer ania tore eaStoaT Me eter ke 13

Mesosternum with posterior portion

triangular in midline (Figs 77, 79)....... 10

Mesosternum with posterior portion

rounded in midline (Fig. 78)............+ 11

Prosternal process rounded at tip (Fig.

75); tip of metatrochanter pointed; lobe

on apical segment of paramere short.......

wineaap Phas Tjirtudessus morgani (Watts and

Humphreys)

Tip of prosternal process pointed (Fig.

76); apex of metatrochanter rounded;

lobe on apical portion of paramere long..

angered Tjirtudessus bialveus Watts and

Humphreys

Head broad, deflexed, metatrochanter

round; setae on mesofemur long.............-

.Tjirtudessus silus Watts and Humphreys

With none of above characters ............ 12

Combined length of first two segments of

metatarsus > rest; eye remnant present as

small oval or triangular structure;

paramere with long apical lobe...............

sia vie kag Mls saa ea bgeated he Bey Tjirtudessus pulpa

(Watts and Humphreys)

Combined length of first two segments of

metatarsus approximately equal to rest;

eye remnant reduced to single short

suture; paramere with small apical lobe .

Ho baie Settee Tjirtudessus cunyuensis

Watts and Humphreys

Elytron with row of large punctures

adjacent to SULUTC 1.0... eee ete eteteees 26

Elytron without sutural punctures, other

than a few weak ones near base .......... 14

Eye remnant present as a small oval or

triangular Structure .........c cece 23

Eye remnant reduced to single short

SUTLE ater teense aed dal Mt beeberedenedeeneeste i)

Mesofemur with 5 to 7 spines on hind

edge in basal half... cece 16

Mesofemur with 2 to 4 spines on hind

edge in basal half ...........cceeeeeeteeeees 19

THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON

16 (15) —

17 (16) —

fea ee

19 (15) —

20 (19) —

21 (20)—

22 (21)—

23 (14)—

Protibia thick (Fig. 73); protarsus

moderately expanded, mesotarsus less so;

mesotibia slightly angular......Bidessodes

gutteridgei Watts and Humphreys

Protibia thin (Fig. 74); protarsus and

mesotarsus approximately the same size;

mesotibia not angular... eee 17

Length 2.5 — 2.7 mm; suture between

ventrites | and 2 complete (Fig. 83)........

susan hs Tjirtudessus padburvensis sp. nov.

Length 1.6 — 2.0 mm; suture between

ventrites | and 2 obliterated laterally

(BISS:84-86). 2. cscarteevterieelaaeticcceieicte teed 18

Paramere with lobe as wide as rest of

apical segment, flat on top, expanded

slightly at tip .....7jirtudessus masonensis

Watts and Humphreys

Paramere with lobe shorter than rest of

apical segment, rounded on top, tip

pointed.......... Tjirtudessus yuinmeryensis

Watts and Humphreys

Mesofemur with four spines near base;

antenna with segments | and 3 of similar

length, segment 11 approximately 1.5x

length of segment 10; length 2.1 — 2.4

160100 Eee ore eo Tjirtudessus cueensis

(Watts and Humphreys)

Mesofemur with two to three strong

spines on hind edge near base; antenna

with segment 2 large, oval; segment 3

much smaller and thinner, segment 11

approaching 2x length of segment 10;

lengthrl3. 2 PSS es h.c.rPeves. ese sche 20

Mesofemur with two strong spines on

hind edge near base; paramere with apical

segment with two finger-like projections

ein Tjirtudessus pinnaclesensis

(Watts and Humphreys)

Mesofemur with three strong spines on

hind edge near base; paramere with apical

segment with one finger-like projection..

SRAM URS fA sae Se, icc Rc 21

Mesofemur with three spines grouped

together near base... Tjirtudessus

Jridaywellensis (Watts and Humphreys)

Mesofemur with two spines near base and

One More distant... eects 22

Pro and mesotibia club-shaped; antenna

with middle segments enlarged a little on

INSIdE eee Tjirtudessus hinkleri

(Watts and Humphreys)

Pro and mesotibia elongate/triangular in

shape; middle segments of antenna

virtually symmetrical............ Tjirtudessus

karalundiensis Watts and Humphreys

Pronotum not constricted at base:

prosternal process reaching or almost

24 (23) —

Das

Douay.

27 (26)—

I06)——

29 (6) —

30 (29) —

31 (30) —

reaching mesosternum; 1.4 mm long.......

dee calls Peas aes Tjirtudessus wilunaensis

Watts and Humphreys

Pronotum moderately constricted at base;

prosternal process not reaching

mesosternum; 2.3 — 3.2 mm long........ 24

Mesofemur with 6 spines close to base on

hind edge.......... Tjirtudessus bigbellensis

(Watts and Humphreys)

Mesofemur with 3 to 6 spines spread out

along basal half of hind edge............... 29

Suture line between ventrites | and 2 well

marked (Fig. 83); aedeagus with medial

lobe parallel sided, apex not upturned.....

bee eweecasMenetearctee 5 Tjirtudessus challaensis

(Watts and Humphreys)

Suture lines between ventrites | and 2

weak (Fig. 84), usually obsolete in lateral

half; aedeagus with medial lobe distinctly

narrower in middle, apex upturned

pies te teiat Ie 8 Tjirtudessus jundeeensis

Watts and Humphreys

Distinct oval eye remnant present ....... 27

Eye remnant reduced to single short

SUC OM la aad meat ca. ce teeh tet oa Ateas ei retnal 28

Metatrochanter with tip slightly pointed

(Fig. 11); sutural lines between ventrites

1 and 2 complete, distinct ....7jirtudessus

hillviewensis sp. nov.

Metatrochanter with tip rounded; suture

between ventrites 1 and 2 obliterated

laterally......... Tjirtudessus windarraensis

(Watts and Humphreys)

Mesofemur with 3 spines on hind edge

(Fig. 28); suture between ventrites | and

2 complete; metatrochanter elongate (Fig.

29)...Tjirtudessus wogarthaensis sp. nov.

Mesofemur with 5 to 6 spines on hind

edge; suture between ventrites 1 and 2

obliterated laterally; metatrochanter

rounded ............ Tjirtudessus lapostaae

(Watts and Humphreys)

Mesofemur with spines arranged in two

comb-like rows along hind edge from

base to apex; mesotibia thin, curved .......

De ee ee ere Bidessodes limestoneensis

Watts and Humphreys

Mesofemur with spines on hind edge

spaced out, not dense and comb-like;

mesotibia straight

Pro and mesotarsus with basal segment

much more expanded than other

SCOMOMS techs. coer wetescecisy teh Mesenetton 3]

Pro and mesotarsus with basal segment

only moderately expanded compared

with other segments

Antenna with segments 8 to 11 noticeably

102

32 (30) —

33 (32) —

34 (2) —

35 (34) —

36 (35) —

37 (36) —

38 (37) —

39 (34) —

C.H.S. WATTS & W. F. HUMPHREYS

thinner than others, segment 3 longer

than segment 2...Tjirtudessus magnificus

Watts and Humphreys

Antenna with segments 8 to 10 not

noticeably thinner than others, segment 3

same length as segment 2.....7jirtudessus

macrotarsus. Watts and Humphreys

Pronotum a little narrower than elytra;

length 3.5 — 5.0MmM.........cccseseereeeeeeees 33

Pronotum wider than elytra; length 3.2 —

SST ssseecss seston Tjirtudessus eberhardi

Watts and Humphreys

Metatrochanter rounded at tip; aedeagus

with central lobe straight, tip pointed; eye

remnant small ..7jirtudessus raesideensis

Watts and Humphreys

Metatrochanter pointed at tip; aedeagus

with central lobe twisted, tip knobbed;

without eye remnant...7jirtudessus hahni

Watts and Humphreys

From the Northern Territory ... ‘

From Western Australia ...........0c 39

Head short, very broad, strongly

deflexed; pronotum strongly narrowed at

base; prosternal process anvil-shaped......

sdeaggiteartnendsteecesy Nirripirti macrocephalus

Watts and Humphreys

Head variably shaped, not deflexed, base

of pronotum variably shaped; prosternal

process “normally” shaped ...........++ 36

Protarsus with segment 3 not bilobed;

pronotum not constricted at base; antenna

thin, segments | and 2 subequal..............

FRR onseseysaned usncondecty Nirripirti pentameres

Watts and Humphreys

Protarsus with segment 3 bilobed;

pronotum weakly to moderately

constricted at base; antenna thick,

segment 2 much broader than segment |

fs To yng os tag dhagd Tas T EA FEE RaSaalca nosh nad nese agRRE? 37

Length 1.8 mm; body well-chitinized .....

suse bates civeee ty sep yes Nirripirti napperbyensis

Watts and Humphreys

Length 1.2 — 1.6 mm; body weakly

CHILMIZEM «...ervsrerapsateaaderradacnssdcdernedicebiand 38

Length 1.2 mm; body only slightly

constricted at junction of pronotum and

SL yIPA NY, Aseetyyenovehaest Nirripirti wedgeensis

Watts and Humphreys

Length 1.5 mm; body quite strongly

constricted at junction of pronotum and

elytra wee Nirripirti newhavenensis

Watts and Humphreys

Pronotum (and head) about half width of

elytra (Fig. 36) .....Nirripirti arachnoides

sp. Nov.

Pronotum > three quarters width of elytra

40 (39) —

41 (40) —

42 (41) —

43 (42) —

44 (43) —

45 (40) —

46 (45) —

47 (46) —

48 (47) —

Elytron with visible ventral portion

extensive except close to apex (eg. Fig.

BS) its Aap eee egptterca ye lpanaed 41

Elytron with visible ventral portion

narrow except in basal quarter (eg. Fig.

BG) resscypedennteeditccatecaetussaenees

Length 3.6 — 3.8 mm Nirripirti

stegastos Watts and Humphreys

Length 1.5 — 2.5 MM....... eee 42

Antenna with segments 6 to 8 greatly

expanded, much broader than segments 9

and 10 (Fig. 42)

hesberseeyitgnge theguszte Nirripirti bulbus sp. nov.

Antenna with segments 6 to 10 of

approximately equal SiZe ..........:ee 43

Meso and metatibia elongate triangular;

body strongly boat-shaped, pronotum

much narrower in front «2.0... 44

Meso and metatibia cylindrical (Figs 61,

62); front and rear of pronotum same

width (Fig. 63) .....eccceeseseeeeees Nirripirti

eurypleuron sp. nov.

Length 2.1 to 2.3 mm, metatrochanter

with tip sharply pointed ............ Nirripirti

skaphites Watts and Humphreys sp. nov.

Length 1.5 to 1.9 mm; metatrochanter

with tip rounded ....Nirripirti killaraensis

Watts and Humphreys

Antenna with segment 2 larger and more

oval than segment 1; 1.2 — 2.1 mm long.

deseontuasnyatbernadnebeesiedabeoddeveanganuresieettesdey etd 52

Antenna with segment 2 more or less the

same shape as segment | or smaller; 2.5 —

BO mtn OME ete seessreadss de speasdonaterends 46

Mesofemur with row of about 20 closely

placed small spines along hind edge (Fig.

AB): seaeerdeosstete Nirripirti byroensis sp. nov.

Mesofemur with 10 or fewer weak to

very strong spines along hind edge .....47

Metasternal plate parallel sided (Fig. 80);

mesofemur with 8 to 10 spines, closely

placed, very strong; metatrochanter long

and thin about 4 x as long as wide..........

Aiyssgigshdugeciccbes sbeupaeny rs Nirripirti fortisspina

Watts and Humphreys

Metasternal plate narrowing towards rear

(eg. Fig. 81); mesofemur with 4 to 8

spines, weak to moderately strong;

metatrochanter moderately elongate 2 to

2.5 x as long as wide

Metatarsus with segment | as long as

others combined, with confluent group of

5 strong spines in middle on outside (Fig.

Si) ecsaetdene Nirripirti copidotibiae sp. nov.

Metatarsus with segment | much shorter

than others combined, without confluent

THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 103

group of spines on outside... 49

49 (48) — Metasternal plate without wings (Fig. 81)

eptqbiid tua peels Midgenad Nirripirti plutonicensis

Watts and Humphreys

— Metasternal wings obvious but short (eg.

Eee OW) atc itscy eau dia dullest eit aad maida ng 50

50 (49) — Metafemur with 2 to 4 small lumps on top

edge (Fig. 71)........ Nirripirti verrucosus

sp. Nov.

— Metafemur with smooth top edge........ 5]

51 (50) — Mesofemur with moderately strong

spines; metacoxal plate nearly reaching

MESOCOXAC. 0. eee eeeee Nirripirti hinzeae

Watts and Humphreys

— Mesofemur with thin spines; metacoxal

plate at least the width of metafemur from

MESOCOXAE.... eee Nirripirti darlotensis

Watts and Humphreys

52 (45) — Elytron with shoulder flared outwards

CPig. 86). «2. ciealaetereed Nirripirti hamoni

Watts and Humphreys

— Elytron with shoulder not flared (eg. Fig.

OO) stasis wechecemntant lis upivalptensidinaaaeds 53

53 (52) — Metatrochanter produced into long strong

Point. (Figs 68) eveseeseeeeeeder verses Nirripirti

innouendyensis sp. nov.

— Metatrochanter rounded, at most bluntly

POSH seers ig eh IEA io ia cas Shae 54

54 (53) — Eye remnant absent; metatrochanter

large, squat; hind leg stout; length 1.2

TMM, 0... eeeeeeeeeeeeeeees Nirripirti milgunensis

Watts and Humphreys

— Eye remnant represented by a short suture

at side of head; metatrochanter elongate;

hind leg elongate; length 1.8 - 2.0 mm....

a cMoshters concrete ners? Nirripirti melroseensis

Watts and Humphreys

Descriptions

The following species descriptions are grouped in

alphabetical order under genus which are placed in

the order Boongurrus, Tjirtudessus, Nirripirti.

Boongurrus Larson (Hydroporinae, Bidessini)

Boongurrus occidentalis sp. nov.

FIGS 13-18

Holotype

m: ‘10 K NW Eerala Stn CHS Watts 23/5/01’,

WAM 34222. Card mounted.

Paratypes

25; 8, as for holotype, SAMA; 12, ‘Wittenoom

Gorge Town Pool CHS & GA Watts 26/5/01’, 10

SAMA, 2, Lars Hendrich collection (Berlin); 1, ‘BES

9277 Killara Station north, unused water bore, 26°

03’ 55” S 118° 41’ 58” E, 6/6/2002 W. F.

Humphreys and R Leys’, SAMA; 1, ‘BES 9318

Moorarie Stn nr calcrete quarry site 419, 25° 52’ 26S

117° 27’ O9E, 8/6/02, W. F. Humphreys & R. Leys’,

SAMA; 2, ditto except ‘BES 9320’, | WAM 34177,

| SAMA; 1, ‘BES 9246 Wagga Wagga Stn, mineral

exploration bore 28° 26' 36” S 116° 38’ 9" E,

4/6/02, W. F. Humphreys & R. Leys’, SAMA.

Description (number examined, 24)

Habitus. Length 1.9 — 2.3 mm, relatively flat,

weakly constricted at junction of pronotum/elytra;

elongate oval; uniformly light testaceous, elytra

uniformly darker; hindwing not reduced; eyes of

normal size.

Head. Narrower than elytra; smooth, shiny, weak

reticulation towards rear, punctures small, sparse;

without cervical stria, subparallel in posterior half,

widest just behind eye, antenna stout, segments! and

2 cylindrical, segment 3 as long as segment 2,

narrower, narrowing slightly towards base, segments

4 to 8 becoming progressively slightly broader,

segments 9 and 10 a little narrower than segment 8,

segment 11 about twice length of segment 10, each

segment, except segment 1, with some very small

setae on inside apically. Maxillary palpus elongate,

segment 4 as long as segments | to 3 combined.

Pronotum. A little narrower than elytra;

anteriolateral angles projecting strongly forward;

base weakly constricted, posteriolateral angles

square, surface slightly rugose, with moderately

dense, moderately sized punctures and a row of

stronger punctures along front margin; basal plicae

strong, excavated somewhat on inside, converging

slightly towards front, reaching to about two-thirds

way along pronotum.

Elytra. Not fused, with weak inner ridges near

apex (ligula), lacking in some; elongate, nearly

parallel-sided in middle; rugose, quite densely

covered with moderate sized punctures; plicae well

impressed, straight, about as long as pronotal plicae.

Epipleuron well differentiated from rest of elytra

particularly anteriorly, lacking basal carina,

relatively broad in anterior quarter then

progressively narrowing to near apex.

Ventral surface. Prosternal process strongly

narrowed between coxae, reaching mesothorax,

apical half elongate triangular, weakly arched in

lateral view with highest point (viewed ventrally)

between coxae. Mesocoxae separated. Metathorax

strongly triangular in front in midline; wings short;

broadly rounded in midline behind. Metacoxal plates

large, quite strongly punctate, moderately rugose,

metacoxal lines distinct, moderately widely spaced,

reaching to metasternum, weakly diverging; closely

adpressed to ventrite 1. Ventrites | and 2 fused,

sutural lines distinct, ventrites 3 to 5 mobile, rugose,

104 C. H. S. WATTS & W. F. HUMPHREYS

well covered with moderate sized seta-bearing

punctures.

Legs. Protibia triangular, outer edge bow-shaped,

widest towards apex where it is about four times its

basal width; protarsus weakly expanded, segment |

as broad as long, segment 2 as wide as segment | and

about half its length, segment 3 as long as segment 1|

and a little narrower, deeply bifid, segment 4 very

small, hidden within lobes of segment 3, segment 5

narrow, cylindrical, about 1.5x length of segment 3,

segments | to 3 with dense covering of adhesive

setae; claws short and simple. Mesotrochanter

elongate-oval with a few relatively long, thin setae

on inner edge; mesofemur with 4 to 5 moderately

strong setae in basal half, only slightly stronger than

those on other parts of the femur (Fig. 16),

mesotarsus slightly less expanded than protarsus.

Metatrochanter tip rounded (Fig. 17); metafemur

relatively stout, lacking spines; metatibia narrow,

moderately curved, widening towards apex;

metatarsi relatively stout, segment | longest,

segment 5 longer than segment 4, segments | and 2

in combination about as long as others; claws weak.

Male

Little external differentiation between the sexes.

Median lobe of aedeagus broad in middle narrowing

to sharp point; paramere broad, apical segment with

relatively stout apical lobe well separated from rest

of segment. Figs 13-14.

Remarks

This new species of Boongurrus closely resembles

B. rivulus (Larson) from the Atherton tableland

region of north Queensland differing externally only

in the larger eyes, slightly less rugose body, slightly

weaker tarsi and thinner antennae. The main

differentiating characters are the size of the eyes

which in the new species are of normal epigean size

in contrast to those of B. rivulus which are about

three quarters normal size and in the male genitalia.

The aedeagus of B. occidentalis is broader medially

and tapers to a longer and sharper point. The apical

segment of the paramere is longer than in B. rivulus

and the apical lobe smaller but much more distinct

(Larson 1994),

Figs 1-6. Tjirtudessus hillviewensis: |, lateral view of central lobe of aedeagus; 2, ditto dorsal view; 3, paramere; 4,

mesotrochanter and mesofemur; 5 metatrochanter and metafemur; 6, dorsal view. Scale bar represents 1mm (habitus

only).

THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 105

Most of the specimens have been collected at the

edge of pools in sandy riverbeds and interstitially to

at least two meters from the water’s edge in an

upstream direction. It has also been recorded as flying

to light (in Australian Insect Collection CSIRO

Canberra). A little surprisingly a few specimens were

collected during this year’s fieldwork from boreholes

in two separate calcretes together with a true

stygobitic fauna. The Killara North site was an open

unused water bore ca 3 m to water and a water depth

of ca 9 m in a calcrete area and the associated

stygofauna included Bathynellacea, crangonytoid and

Ceinidae Amphipoda, Harpacticoida, Cyclopoida.

The Moorarie site was a sealed monitoring bore in

calcrete within 30 m of a calcrete quarry that

penetrated the groundwater; the borehole contained a

stygobitic fauna comprising Bathynellacea and

Amphipoda. DNA studies (Remko Leys pers. com.)

found no differences between B. occidentalis

specimens collected from epigean and_ stygal

habitats.

Etymology

Latin. ‘Occidental’ — western. A reference to its

distribution.

Tjirtudessus Watts & Humphreys

(Hydroporinae, Bidessini)

Tjirtudessus hillviewensis sp. nov.

FIGS 1-6

Holotype

m. BES 9399, Hillview Station, bore at Camel

Well, 26° 58’ 20S 117° 27’ 09" E, 13/6/02, W. F.

Humphreys & R. Leys’, WAM 34178. Slide

mounted.

Paratype

I (partial), as for holotype except ‘BES 9398’,

SAMA.

Description (number examined, | + | partial)

Habitus. Length 2.35 mm; relatively flat, weakly

constricted at junction of pronotum/elytra; elongate

oval; uniformly light testaceous; hindwing reduced,

broad, about half length of elytron, tip slightly

folded; weakly sclerotized.

Head. A little narrower than elytra; smooth,

reticulation weak, punctures sparse, very small;

subparallel in posterior half, widest just behind eye

remnant, eye remnant reduced to small triangular

area. Antenna moderately stout, segments | and 2

broad, oval, segment 3 as long as segment 2,

narrower, narrowing towards base, segments 4 to 10

subequal, segment 11 about twice length of segment

10, each segment, except segment 1, with some very

small setae on inside apically. Maxillary palpus,

elongate, segment 4 as long as segments | to 3

combined.

Pronotum. About same width as elytra;

anteriolateral angles projecting strongly forward;

base weakly constricted, posteriolateral angles

obtuse, overlying elytra somewhat; smooth, with

sparse, very weak punctures and a row of stronger

punctures along front margin; basal plicae

moderately impressed, converging slightly towards

front, reaching to about half way along pronotum;

numerous long setae laterally in anterior half.

Elytra. Not fused, tightly closed, lacking inner

ridges; elongate, almost parallel sided, smooth, very

weakly reticulate, sparsely covered with small

punctures, a few widely spaced larger punctures

close to inner edge; row of long setae near lateral

edge, a few additional larger punctures with long

setae, more frequent towards sides and apex.

Epipleuron moderately differentiated from rest of

elytra, moderately broad in anterior quarter,

progressively thinner till near apex.

Ventral surface. Prosternal process strongly

narrowed between coxae, not reaching mesothorax,

apical half broad, sides slowly converging to

rounded apex, strongly arched in lateral view with

highest point (viewed ventrally) between coxae.

Mesocoxae in contact at midline. Metathorax

broadly triangular in front in midline; wings stout,

narrow; narrowly rounded in midline behind.

Metacoxal plates large, weakly reticulate, metacoxal

lines indistinct, moderately widely spaced, reaching

to about halfway to metasternum, quite strongly

diverging; a few small setae-bearing punctures

towards midline; closely adpressed to ventrite 1.

Ventrites 1 and 2 fused, sutural lines distinct,

ventrites 3 to 5 mobile, sparsely covered with small

seta-bearing punctures, ventrites 3 and 4 with a long

central seta or bunch of long setae.

Legs. Protibia bow-shaped, relatively narrow,

widest towards apex where it is about four times its

basal width; protarsus weakly expanded, segment |

as broad as long, segment 2 as wide as segment | and

about halfits length, segment 3 as long as segment 1,

narrower, deeply bifid, segment 4 very small, hidden

within lobes of segment 3, segment 5 narrow,

cylindrical, about 1.5x length of segment 3,

segments | to 3 with very dense covering of adhesive

setae; claws short and simple. Mesotrochanter

elongate/oval with a few thin setae on inner edge;

mesofemur with 4 evenly spaced spines in basal half

(Fig. 4) mesotarsus a little more elongate than

protarsus. Metatrochanter tip angular (Fig. 5);

metafemur relatively stout, lacking spines; metatibia

narrow, moderately curved, widening towards apex;

metatarsus elongate, segment | longest, segment 5 a

little longer than segment 4, segments | and 2 in

combination about as long as others; claws weak.

106 C.H. S. WATTS & W. F. HUMPHREYS

Figs 7-12. Tjirtudessus microocula: 7, lateral view of central lobe of aedeagus; 8, ditto dorsal view; 9, paramere; 10,

mesotrochanter and mesofemur; 11, metatrochanter and metafemur; 12, dorsal view. Scale bar represents 1mm (habitus

only).

Male

Female not known. Median lobe of aedeagus with

unusually long thin apical portion, slightly

crenulated on top; paramere broad, apical segment

with long, narrow, apical lobe separated from rest of

segment except at its apex which overlaps rest of

segment. Figs 1-2.

Etymology

Named after the pastoral station on which it was

found.

Remarks

Most closely resembles 7) windarraensis in size,

presence of an eye remnant, row of large punctures

on elytron adjacent to the suture and pointed median

lobe of the aedeagus, but differs in the shorter apical

segment of the paramere and larger apical lobe, as

well as having a more oval eye remnant, and a

complete suture between the first and second

ventrites which is obliterated laterally in 7:

windarraensis.

Tjirtudessus microocula sp. nov.

FIGS 7-12

Holotype

m. ‘BES 9223, Bunnawarra Station, bore nr.

shearing shed, 28° 36’ 35” S 11° 34’ 25” E, 3/6/02,

W. F. Humphreys & R. Leys’, WAM 34179. Slide

mounted.

Paratypes

37: 19 + 5 partial specimens, as for holotype, 8 + 5

partial specimens WAM 34180 - 34192, 11 SAMA;

4, as for holotype except ‘BES 9224, well near

shearers quarters, 28° 36’ 36” S 116° 34’ 18” EB’,

WAM 34193 - 34196; 4, as for holotype except ‘BES

9225’, SAMA.

Description (number examined, 38)

Habitus. Length 2.2 — 2.3 mm; relatively flat,

weakly constricted at junction of pronotum/elytra;

elongate oval; uniformly light testaceous; eyes

reduced to about a fifth normal size; hindwing

reduced, about length of elytron, tip folded.

THIRTEEN NEW DYTISCIDAE (COLEOPTERA) OF THE GENERA BOONGURRUS LARSON 107

17

Figs 13-18. Boongurrus occidentalis: 13, lateral view of central lobe of aedeagus; 14, ditto dorsal view; 15, paramere; 16,

mesotrochanter and mesofemur; 17 metatrochanter and metafemur; 18, dorsal view. Scale bar represents Imm (habitus

only).

Head. Narrower than elytra; smooth, reticulation

weak, punctures sparse, small; subparallel in

posterior half, widest just behind eye; eye reduced to

about a fifth normal size, lacking individual facets,

darkly pigmented. Antenna relatively thin, segments

1 and 2 cylindrical, segment 3 as long as segment 2,

narrower, narrowing towards base, segments 4 to 10

subequal, segment 11 elongate, a little less than twice

length of segment 10, each segment, except segment

1, with some very small setae on inside apically.

Maxillary palpus, elongate, segment 4 as long as

segments | to 3 combined.

Pronotum. A little narrower than elytra;

anteriolateral angles projecting strongly forward;

base moderately constricted, posteriolateral angles

bluntly pointed, smooth, with sparse, very weak

punctures and a few stronger punctures along front

margin; basal plicae weak, strongly slanting inwards,

reaching to about half way along pronotum; with row

of long setae laterally in anterior half.

Elytra. Not fused but tightly closed, lacking inner