VOL. 113,

PARTS 1 & 2

31 MAY, 1989

Contents.

Transactions of the

Royal Society of South

Australia

Incorporated

Lee, D. C. & Birchby, C. M. Decoribatula gen. nov. from Singapore, with notes on allied

Reticuloppia (Acarida: Cryptostigmata: Oribatulidae) from Tropical

Australia - - - - - - - - - - -

Hirst, D. A new genus of huntsman spider (Heteropodidae: Araneae) from south eastern

Australia - - - - - - - - - - -

Tyler, M. J. A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of

Queensland, with a redefinition of the ilial characteristics of the

genus - - - - - - - - - - - -

Boulton, A. J. Over-summering refuges of aquatic macroinvertebrates in two intermittent

streams in central Victoria - - - - - - - -

Southcott, R. V. The larva and nymph instars of Odontacarus (Leogonius) adelaideae

(Womersley) (Acarina: Trombiculidae: Leeuwenhoekiinae) - - -

Tiver, F., Sparrow, A. D. & Lange, R. T. The composition and distribution of the vegetation

of north-west Eyre Peninsula - - - - - - - -

Bourman, R. P. & Lindsay, J. M. Timing, extent and character of Late Cainzoic faulting

on the eastern margin of the Mt Lofty Ranges, South Australia - -

Ainslie, R. C, Johnston, D. A. & Offer, E. W. Intertidal communities of northern Spencer

Gulf, South Australia - - - - - - - - -

Koste, W. & Shiel, R. J. Rotifera from Australian inland waters. III. Euchlanidae,

Mytilinidae and Trichotriidae (Rotifera: Monogononta) - - -

Brief Communications:

Davies, M. Rediscovery of Uperoleia orientalis (Parker) (Anura: Myobatrachinae) - -

Nobbs, J. The occurrence of plant parasitic nematodes in the arid region of South

Australia - - - - - - - - - - -

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

VOL. 113, PART 1

2 zz

—-

Se 4

——

S im

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TRANSACTIONS OF THE

ROYAL SOCIETY OF SOUTH AUSTRALIA INC.

CONTENTS, VOL. 113, 1989

PARTS | & 2, 31 May

Lee, D. C. & Birehby, C. M. Decoribatula gen. noy. from Singapore, with notes on allied

Reticuloppia (Acarida: Cryptostigmata: Oribatulidae) from Tropical

Australia - - + > = 6 = + > . "

Hirst, D. A new genus of huntsman spider (Heteropodidae: Araneae) from south eastern

Australia - - - - - - - ~ - - -

Tyler, M. J. A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of

Queensland, with a redefinition of the ilial characteristics of the

genus - - - - - - - - : - -

Boulton, A. J. Over-summering refuges of aquatic macroinvertebrates in two intermittent

streams in central Victoria - ~ - - - - - -

Southcott, R. ¥. The larva and nymph instars of Odontacarus (Leogonius) adelaideae

(Womersley) (Acarina: Trombiculidae: Leeuwenhoekiinae) - *

Tiver, F., Sparrow, A. D. & Lange, R. T. The composition and distribution of the vegetation

of north-west Eyre Peninsula - - - - - - -

Bourman, R. P. & Lindsay, J. M. Timing, extent and character of Late Cainzoic faulting

on the eastern margin of the Mt Lofty Ranges, South Australia -

Ainslie, R. C., Johnston, D. A. & Offler, E. W. Intertidal communities of northern Spencer

Gulf, South Australia 2 3 = e ; 3 e 2

Koste, W. & Shiel, R. J. Rotifera from Australian inJand waters, III, Euchlanidae,

Mytilinidae and Trichotriidae (Rotifera: Monogononta) - -

Brief Communications:

Davies, M. Rediscovery of Upero/eia orientalis (Parker) (Anura: Myobatrachinae) +

Nobbs, J. The occurrence of plant parasitic nematodes in the arid region of South

Australia - - - - - - - - - - -

115

117

PARTS 3 & 4, 30 November

Koste, W. & Shiel, R. J. Rotifera from Australian inland waters IV. Colurellidae roar:

Monogononta) - - - - - - -

Short, A. D., Buckley, R. C., & Fotheringham, D, G, Preliminary investigations of beach

ridge progradation on Eyre Peninsula and Kangaroo Island - -

Pledge, N.S. The occurrence of a diprotodontid marsupial referable to Euowenia grata

De Vis in western New South Wales — - - - - -

Austin, A. D, & Allen, G, R. Parasitoids of Uraba lugens Walker (Lepidoptera:Noctuidae)

in South Australia, with description of two new species of

Braconidae - - - - - - - - - -

Barker, S. Contributions to the taxonomy of Australian Buprestidae (Coleoptera): New

species of Astraeus and Stigmodera (Castiarina) and a key to Astraeus

(s.s) - - - - - - - - - - >

Jenkins, J. F. & Hasenohr, P. Trilobites and their trails in a Black Shale: Early Cambrian

of The Fleurieu Peninsula, South Australia - - - - -

Gowlett-Holmes, K. L. & Holmes, N. J. CG. Rediscovery of Primovula (P) heleneae Cate

and description of a new species of Crenavolva from South Australia

(Mollusca: Gastropoda: Ovulidae) - - - - - -

Davies, M. Developmental biology of the Australopapuan hylid frog Litoria eucnemis

(Anura: Hylidae) - - - - - - - - -

Brief Communication:

Beardsell, G. R. Hybridisation of Litoria chloris and L. xanthomera. (Anura:

Hylidae) ; - = e = 3 = = = =

Insert to Transactions of the Reval Society af Sauth Avstralia, Voi. 113, parts 2 .& 4, 30 November, 1989

145

163

169

185

195

215

221]

DECORIBATULA GEN. NOV. FROM SINGAPORE, WITH NOTES ON

ALLIED RETZCULOPPIA (ACARIDA: CRYPTOSTIGMATA:

ORIBATULIDAE) FROM TROPICAL AUSTRALIA.

BY DAVID C. LEE & CAROLYN M. BIRCHBY*

Summary

Decoribatula pustulata gen. nov., sp. nov. on an orchid from Singapore, intercepted at Adelaide

Airport, is described. It is similar to Reticuloppia reticulata Balogh & Mahunka, 1966 from

rainforest litter in Queensland, the description of which is extended. Both mites are unusual in

having a deficient chaetotaxy on femora I and 11.

KEY WORDS: Acarida, Oribatulidae, Decoribatula pustulata, new genus, new species,

Reticuloppia reticulata Balogh & Mahunka, Singapore, Queensland, leg chaetotaxy.

DECORIBATULA GEN. NOV. FROM SINGAPORE, WITH NOTES ON ALLIED

RETICULOPPIA (ACARIDA: CRYPTOSTIGMATA: ORIBATULIDAE) FROM

TROPICAL AUSTRALIA.

by DAVID C. LEE & CAROLYN M. BircHay’

Summary

Lee, D, C, & BircuBy,C.M, (1989) Deeoribunela gen- noy. from Singapore, with notes on allied Reticaloppia

(Acarida: Cryptostigmata: Oribaiulidae) from tropical Australia, Trans. R. Soc. S. Aust, 13 (2), 1-5,

31 May, 1989.

Decaribatila pustulata gen. nov., 5p. nov. on an orchid from Singapore, intercepted at Adelaide Airport,

is described. It ix similar to Reticuloppia reticulata Balogh & Mahunka, 1966 from rainforest fitter in

Queensland, the description of which {8 extended, Both mites are unusual in haying a deficient chaclotaxy

on femora { and II.

Key Worns Acarida, Ovibatulidae, Decaribaiula pustufate, new genus, new species,

Reticuloppia reticulata Balogh & Mahunka, Singapore, Queensland, leg chactotaxy,

Introduction

The mites considered here have been studied

because of their relevance to an ongoing study of

sarcoptiform mites of South Australian soils,

sampled from nine florally diverse sites. An

introduction to relevant work on the advanced

oribate mites (Planofissurae), with comments on

the Oripodoidea, which include the Oribatulidae,

is provided by Lee (1987), These mites and a group

of seven species of Oribatula-like mites from the

South Australian study aré atypical within the

Oripodaidea in that they havea deficient chaetotaxy

on femora I and []. Oripodoid leg chaetotaxy will

be discussed more fully when the South Australian

species are described (Lee & Birchby in preparation).

The chaetotaxy for femur I and II on oripodoid

mites is 0 anterior; 2 dorsal/2 ventral, I posterior

setae (0,2/2,1), In the deficient chaetolaxy one or

two setae are missing, resulting in one of three

chsctotaxies as follows: (0,2/2,0; 0,2/1,0; 0,2/1,1. The

Oribatula-like 5, Aust, mites differ superficially in

having only short or medium length hysteronotal

setae and they were collected from only the arid,

semiarid or mallee sites; whilst the two species

considered here have long hysteronotal setae and

are from moist, tropical sites.

The notation and methods of measurements

follow Lee (1981) with modifications made by Lee

{1987}. Measurements are in micrometers (jum). The

type of the new specics is deposited in the South

Australian Museum, Adelaide (SAMA) whilst rhe

type of Reticufoppia reticulata is deposited in to. the

Hungarian National Museum, Budapest,

* South Australian Museum, North ‘Terrace, Adelaide.

S. Aust. S000

etlewloppia Balogh & Mahunka

Reticuloppio Balogh & Mahunka, 1966; p. 564

Type-species. Reticuloppia reticulata Balogh &

Mahunka, 1966.

Diagnosis; Oribatulinae. Hysteronoium with 14

pairs (SJ, 62, 3S) of long setae. Lamella absent.

Hysteronotal lenticulus present near dorsosejugal

furrow. Integument of hysteronotum with extensive

reticulate sculpturing and cerotegument forming a

thick, columnar refractile exudate. Femora I and

L) with deficient chactotaxy, lacking posterior setae:

I — 0,2/2,0; 1] — 0,271.0. Legs slim and long, with

leg IV longest (Jeg If] ts lost). Pretarsal claws short

(céntral claw {1 less than 0,33 x length of tarsus 11)

and lateral claws much slimmer than central. cliew

(depth less than 0.5 x depth of central claw I),

Remarks: Reticuloppia is based on a single female,

grouped in the Oribatulinae, and distinguished by

the hysteronotum having a large number of Jong

setae and teticulate sculpturing around a lenticulus

(Balogh & Balogh 1984). In addition, the

proteronotum lacks ridges, having only a weak

sublamejla, and the leg chactotaxy is deficient in

a way that is unique amongst esiablished oripodoid

species.

Reticuloppia reticulata Balogh & Mahunka

FIG. t

Reticulappia reviculata Balogh & Matunka, 1966; p. 564,

Female: \digsomal length, 475 (470 in original

description). Leg lengths (femur-tarsus); I — 270,

M -— 254, 1f —?, (V—311. Tibial maximum heights:

1 = 25,5, 11 — 20.5, Hi — 7, LY — 18. Yellow brown

colour. Thick (Maxitnum depth equal to distance

between setae 2I-/2), whitish, columnar refractile

DC, LEE & C.M. BIRCHBY

Figs 1-2: Posterior aspects of femur-pretarsus of right legs I and 11, all setae on femora illustrated, p = posterior,

= ventral. t, Retieuloppia reticulata Balogh & Muhunka. 2, Decoribatula pustulata sp. nov.

cerotepument on proteronotum anterior to /2,

thickest lateral to seta zl; also covering

hysteronotum except around lenticulus and near

pleural margins, with (hick patches posterolateral

to seta Sl, lateral to $5 and between setae J6—J6.

Proteronotum with faint cosrate sublamella.

Sensory seta (32) with long stalk (length = x 3

caput), Margin of bothridium raised into turret-like

structure, height subequal to length of <2 caput.

Seta s2 short (0.5 x or less distance between setae

j2-zi1), fine, smooth, without cilia. Hysteronotum

with fine reticulate sculpturing under columnar

exudate. Five multiporose foramina, foramen F°3

divided in F3a and F3b, Hysteronotal seta Sl

confirmed similar to and nearly as long as 21.

Podosternal chactotaxy: 34, 177, 3777, 37 with all

third rank setae as long as /7/3 (illustrated in the

wrong position by Balogh & Mahunka 1966, fig.

15) and 73 on pedotectum I, 7//3 posterior to

pedotectum II (as for Decoribatula pustulosa sp,

nov., Fig.4), /¥3 on discidium, Seta Sal shorter than

Sa2 and Sa3. Opisthosternum with foveolate

sculpturing, Four subcylindrigal eggs, exochorion

. smooth, 80 x 230 (mean). Legs long (mcan femur-

tarsus: 59% of soma) and slim (mean maximum

tibial height: 27% of mean length).

Material examined: Holotype female, rainforest

litter, Mt Spec (18°S7’S, 146°1 VE), Townsville, Qld,

DECORIBATULA, ORIBATLILID MITES a

19,17,1965, J. Balogh. Some appendages lost, no

femur-tarsus Hl on either side,

Remarks: The further description here of

Reticuloppia reticulata adds to or corrects the

original description, with a few important

conlirmations. Reference to the relative dimensions

of the legs is distorted by legs 11] being lost (usually

# short, slits leg amongsr the oripadoids).

Decoributula gen, nov.

Tipe-species: Decoribatula pusrulata sp. nov.

Diggnosixs: Oribatulinae. Hysteronotum with 14

pairs (5J, 6Z, 3S) of mainly long setae (51 is short).

Lamella with only costate antenor part present, not

reaching bothridium to seta 72, Smooth, paler mid-

dotsal anterior area on hysteronotum, bot not

clearly delineated as lenticulus, Integument of

hysteronotum with extensive foveolate sculpturing,

cerolegument inconspicuous, Femora | and U1 with

deficient chactotaxy, but with posterior setac: | -

0,2/1,1; U—0,2/1,1, Legs of medium girth and long,

with Icg TV longest. Pretarsal claws long (central

claw {1 more than 0.33 « length of tarsus El) and

tateral claws only slightly slimmer than central claw

(depth mare (han 0.5 x depth of central claw 11),

Remarks: Decoribatula is based on # single male,

It is similar to Reticu/oppia in having # delicient

femoral chaetotaxy, long hysreronotal setae, a

divided anterior hysteronotal foramen (#3u, 43h)

and a turret-hke bothridium to 22. It particularly

differs, in having a lamella, a different type of

ceTotegument, a different type of setal loss on

femora | and I] and unusually large pretarsal claws.

On the basis of this it is considered to be a sister-

group to Reviculoppia bur different enough to be

regarded a6 a Separate genus.

Decortbatula pustulata sp. nov.

FIGS 2-4

Male; Idiosomal length, 527, Leg lengths (femur-

tarsus): 1-298, 11-280, LIL—285, [V —323. Tibial

Maximum heights: | — 346, 1 — 31, IE — 23, IY -

23, Red brown colour, Incomspicucus (depth less

than diameter of hysteronoral setal bases}

cerplegument, coucentrated in round of uval

pusiwes on hysteranotum,

Proteronotum with partial costare lamelta,

superficially inconspicuous bul backed by relractile

infernal apodeme, not reaching back to turret-like

bothridium (0 seta s2, Two multiporose

foramina (#1, F2a) recognisable dorsally, Central

setae (1 J2, 21) with 3 or 4 files of cilia, /2 longest.

Sensory seta (22) with caput and exposed stalk

subequal in length, Seta s2 short (0,6 % or less

distance j2-21), fine, smooth, without cilia.

Hysteronorum with smooth central triangulate

area {rom anterior margin to level with seta J3, but

no clearly delineated pale lenticulus, Tive

multiporose foramina (F3a, £30, F4, FS, F6) each

backed by a disc-Nhaped chamber i) integument.

Hysteronotal setae smooth, S| short, subequal tu

s2, whilst J5 and §5 very long with whip-like end.

Podosternum with reticulate sculpturing.

Subpodal ridge in three parts, pedotectum II and

discidium large, extending laterally beyond level ol

pedotectum |. Lateral setae (especially 2273) longer

than central setae. Opisthosternum with three setuc

(JZe4, /Z25, Se) represented only by bases on both

sides and assumed broken off. Scia Sal Junyer than

Sa? and Sa3. Shield dark, centrally horizontal and

punctate with curving upwards, smooth marginal

strip.

Legs long (mean femur-tarsus: 56% of somah,

with leg f second longest, and of mediuni-girth

(mean maximum tibial height: 35% of mean

length), Ventral flange on trochantera (11 and LV.

Femora with ventral incrustation.

Material examined: Holotype male (SAMA

N1988474), on orchid from Singapore, intercepted

at Adelaide International Airport, 13.iv. 1987, Grex

Raker (S, Aust, Department of Agriculture),

Remarks: The male of D. pustutata is described,

although oripodoid species are usually based on

females, because it is unlikely that more material

will be collected and [his species is importan! to the

classification af S. Aust, mites being studied (Lee

& Birchby in preparation). There is a lack of sexual

dimorphism amongst oripodoid character states

that are considered important in distinguishing

sptoes. The female is likely to be bigger and have

the thickening around the genilal orifice separated

from the ventrosejugal apodeme, Its relationships

are considered under the remarks on Decoribatnda.

Acknowledgments

We thank the Australian Biological Resources

Study for a grant to DCL. funding the salary of

C.M.B., Ms Kathy Bowshall foi the notation and

presentation of the figures and Mrs Debbie Lowery

for typing the manuscript.

4 DC. LEE & C.M. BIRCHBY

100um

Fig. 3: Decoribatula pustulaia sp. nov., notum of soma.

DECORIBATULA, ORIBATULID MITES 5

Fig. 4: Decoribatula pustulata sp. nov, sternum of soma.

100um

References

BALOGH, J. & BALOGH, P. (1984) Review of the

Oribatuloidea Thor, 1929 (Acari: Oribatei), Acta zool.

hung. 30, 257-313.

& MAHUNKA, S. (1966) New Oribatids (Acari)

from Australian Soils. Folia ent. hung. 33, 553-568.

Lee, D. C. (1981) Sarcoptiformes (Acari) of South

Australian soils. I. Notation. 2. Bifemorata and

Ptyctima (Cryptostigmata). Rec. 5S. Aust, Mus. 18,

199-222.

_—__. (1987) Introductory study of advanced oribate

mites (Acarida: Cryptostigmata: Planofissurae) and a

redescription of the only valid species of

Constrictobates (Oripodoidea). Ibid. 21, 35-42.

A NEW GENUS OF HUNTSMAN SPIDER (HETEROPODIDAE:ARANEAE)

FROM SOUTH EASTERN AUSTRALIA

BY DAVID HIRST*

Summary

A new heteropodine genus, Keilira, gen. nov. and two new species, K. sparsornaculata sp. nov. and

K. sokoli sp. nov. are described from South Australia and Victoria respectively. Keilira differs from

other Australian genera in labium shape and the relatively short, thick embolus of the male. The

arrangement of cheliceral teeth is similar to that of Heteropoda Latreille, 1804, but the general

appearance resembles Australian species attributed to Olios Walckenaer, 1837, here considered as

belonging in Neosparassus Hogg, 1903.

A NEW GENUS OF LIIUNTSMAN SPIDER (HETEROPODIDAE:ARANEAE)

FROM SOUTH EASTERN AUSTRALIA

by Davin Hirst*

Summary

Hrest, D, (1989) 4 new genus of huntsman spider (Hereropedidae Araneae) trom south eastern Australia.

Trans. R, Soo S Aust, (13, 7-23, FL May, 1989.

A new heteropodine genus, Aeiliru, gen. nov. and twa new species, KL sparsarmmacnidta sp. nov. ad

K, sokuli sp. nov. are duseribed from South Australia and Victoria respectively. Nef/ing differs from other

Australian genera in labium shape and the relatively short, thick embolus of the male. The arrangement

of cheliceral teerh is sumilar ro that of Heferopodg Latrenle, 1804, bit the general appearance resembles

Australian species attributed to Olios Walvkenaer, 1837, here considered as belonging in Neosparassus Hage,

1903,

KEY WorRDS, Arancae, Heteropodidae, Keslird gen. nov,, new taxa.

Introduction

This paper forms the first part of a laxonomic

study on the Australian Heteropodidae A new

heteropodine genus is defined for undescribed

species previously mis-identified as Neosparassus

(sublamily Deleninae Hogg, 1903), which they

resemble in general colouring and shape,

Neosparassus will be discussed fully later (Hirst in

prep.), but it muy be mentioned here that despite

its revalidation by Jarvi (1914) all relevant species

hive been referred to Olfos in subsequent literature.

Subfamilies of the Heteropodidae are poorly

defined. Some useful characters of the

Heteropodinae Keyserling, 1880 are referred to here

in associating the new genus with that subfamily.

Similarly, characters of the Deleninac are mentioned

in comparing Neasparassus with Olios. Allhough

limits of that subfamily are uncertain, it is retained

here as separate from the Busparassinae (Jarvi 1912)

on the basis of genitalic $uructure.

Materlals and Methods

All measurements are in millimetres, made with

an eyepiece praricule. Eve diameters, interspaces and

MOQ nieasurements are expressed as relative to the

diameter of an AME. Drawings were done with the

aid of a camera lucida on a Wild microscope. All

abbreviations are standard for rhe Araneae. Bilateral

variability of leg spine counts are indicated in

parenthesis, Types are deposited in the South

Australian Museum, Adelaide (SAMA), and the

Australian Museum, Sydney (AM).

Keilira pen- nov.

Diagnosis: Labium twice as broad as long. Male

embolus short, thick relatively strateht with a curved

* South Australian Museum, North Terrace, Adelaide,

S.Aust, 5000,

tip. Female with small fossa and hood-like anterior

marein.

Definition: Carapace raised, flattish or convex in

profile, highest in the fovea region, sloping wently

to clypeus, Foveal groove weak or indistinct. Eyes

subequal. Anterior eye row slightly recurved.

Pusterior eye row straight to slightly recurved.

Clypeus half width of an AME or slightly less.

Cheliceral fang groove with three teeth on

promargin, three or four leeth on retromargin and

one of more median rows of minute, pointed teeth.

Labium ai least. twice as broad as long, Sternum

barely longer than broad or equal, slightly convex.

Legs 2143. Anterior legs with four ventral pairs of

spines on tibia, without an extreme distal pair.

Reduced prolateral {cg spination, patellae withour

prolateral spines. Female palp tarsal claw with four

small teeth, Female genitalia with paired uncoiled

spermathecae. Small fossa with weakly sclerotized

hood-like anterior margin. Male embolus short,

thick, relatively straight with a curved (ip. Palpal

ubia with a small prodorsal distal extension and a

triangular spur rising from a retrolateral apophysis,

Type species: Keilira sparsomaculata sp. nov.

Camments

Although resembling Weosperassis in body shape

and coloration, the presence of a distal prodorsal

extension an the tibia of the male palp.and three

teeth on the promargin of the chelicerae followed

by one of more rows of median tecth, places the

genus close lo Meteropoda and Pundercetes L,

Koch, 1875, the Australian representatives of the

Heteropodinac. Neosparassus and other genera af

the Deleminae lack median teeth and a well defined

extension on the male palp tbia.

The retromarginal tecth of Aeslira differ trom

those of Hereropoda and Panderceres in that both

the latter have four long teeth which are evenly sized

and close together with a small space between the

two middle teeth. In Keilira greater spacing occurs

A NEW GENUS OF HUNTSMAN SPIDER 9

between the middle and distal teeth (ig. 8), This

placement remains unchanged when a fourth tooth

is present, this being minute and found al the

proximal end of the row (Fig, 12).

The teeth of the female palp tarsal claws of

Fleferupoda and Puridercetes are elongated and

curved, as Jong of Jonger than the curved claw tip,

and comb-like. This condition is approached in at

least one other Australian heteropodid, Oltus

Aernitis Hoge, 1914, the correct generic status of

which is unknown 4s no mature specimens have

been scen. The clongated teeth on |he palpal claw

are nut found in Xe/ire, but as all other Austratian

genera narmally possess at least five teeth, it is

unusual in having four.

Keilira and Pandercetes have four ventral spine

pairs on anterior ibiae but lack a distal pair, While

four ventral spine pairs are found in Hererapoda,

one pair is shorter and positioned distally near the

articulation with the metatarsi, This character was

used by Kaston (1980) to distinguish Heferoyada

from Olies, the latter having two pairs and lacking

an extreme distal pair. Neosparassus and most other

Deleninac genera have three pairs with rhe distal

pair near the articulation, The exceptions are some

species of Pediana Simon, 1880 and juvenile

specimens of at least Isapeda L. Koch, 1875 and

Neasparassus (juveniles of other genera have not

been examined). There, the distal spine 1s

represented as 2 bristle barely distinguishable from

these adjacent on the tibia margin,

Keilira is similar to Heferopeda in carapace

profile und in having leg 1 shorter than leg LU

(Panderceres has a conesve carapace profile and leg

T longer than lee tI), but differs in eye position and

relative size of the anterian median eyes: The male

bulb, in the upexpanded slate, resembles thar of

Panderceres and, to a lesser extent, Heternpoda,

Keilira lacks the long tapering embolus typical of

other Australian heteropadid genera, having a

relatively straight, short, thick embolus which tapers

at the tip to a blunt point, For much of its length

the embolus runs adjacent to the bulb In

Pandercetes and Heteranada the embolus is well

clear of the bulb for its entire length.

It can be concluded that Keilira is referable to

the Heteropodinge, yet is distiner from Hereropode

and Pandercetes, However it is Known that

Heteropodinge genera of uncertain status occur in

Queensland with live or six ventral spine pairs on

anterior tibia. These are not congeneric with Xeilire

(unpubl, dala) but appear to be closely related and

may prove to be w sister group.

Keifira contains the smallest known Australian

heteropodids. Its representation by a handful of

specimens from the south-eastern parts of both

South Australia and Victoria extends the known

range of the Australian Heteropodinae, Little is

known of their biology but they appear to have

similar habits to Neosparassus (Main 1976), Several

specimens ot Keilira sparsomaculata were collected

whilst foraging at night on low flowering shrubs

(Leptaspermum junipermum Sm.), which were

attracting abundant jnsects, a habitat three species

of Neasparassus have also been found sharing. One

juvenile specimen of Keilire was foutid in litter at

the base of shrubs during the day,

Etymology: The generic name Ketilra is derbved

from Keilira Station, Jt km N.E, of Kingston, SE,

South Australia.

Keilira sparsomacufata sp, Wo,

FIGS 1-10

Holalype; SAMA NIYSTS, o. 13 km N, OF

Keilira Station, 36° 37°S_ 140° 11° E, §. Aust, Feb.

1974, D. Hirst,

Allowpe SAMA NIQR76, 9, same data as

holotype,

Paratypes: SAMA NI9877, 9, same dala as

helotype: SAMA NI9S7B, 9, Cortina Station,

Coorong, 369 20'S, 1399 46° FE. S. Aust. 201.1968,

AW. Forbes.

Other miaterial examined: SAMA NI9872,

Juvenile, Cortina Station, same data as above;

SAMA N1L9873, Juvenile, Kendall Station, Coorong,

S. Aust., Mar. 1968, A.W. Forbes; SAMA N19874,

Juvenile, AWinga Scrub, S, Aust., 16, ix. 1987, D.

Hirst,

Description of holotype (Figs 1-4) Carapace length

2.90, width 2.85, Abdomen length 3.40, width 2,20.

Colour in alcohol: Yellow-brown carapace with

pair of median black, shart linear markings with

adjacent spots, anterior of fovea. Caput with darker

suffusion in ocular region, eyes with black rims.

Chelicerae reddish-yellow, Sternum yellowish, Legs;

anterior pairs yellow-crange, posterior pairs yellow,

palps yellow. Abdomen creamish-whire with small

scattered spots of black pigment, fewer verctrally

Carapace: Convex, highest between !! coxae

sloping gradually tw. clypeus, steeply to posterior.

Fovea indistin¢t, marked by darker pigment. Setae

sparse, short (ca 0.2), moderately long (ca 0,4)

bristles around ocular region, few on caput between

eyes and fovea and along lower margin of clypeus.

Eyes: In two rows, from above anterior row

slightly recurved, posterior row straight, AME

Rigs. 1-4, eilira sparsomeculata zen. ct 3p. nov. Holotype (hale) | bady alld (ght pal, dorsal; 2-4 Lelt palp. 2

ventral; 3 prolateral; 4 retrolateral. Scale line; = O.Smm,

a) D, HIRST

Figs, 5-10. Keilira sparsomaculata gen. et sp. nov. Allotype female, 58: 5 body, dorsal; 6 right leg [, prolateral; 7

tarsal claw; 8 chelicerae, maxillae and labium. Paratype female (SAMA N19878), 9-10: 9 epigyne; LO vulva, ventral,

Scale lines: Figs 5 -— 6, 8 = 0.5mm; figs 7, 9-10 = 0.1mm,

A NEW GENUS OF HUNTSMAN SPIDER i}

diameter 0.20, AME - ALE: PME ; PLE = 1: 1:

0.9 = 0.0. Interspacess AME - AME 0.8 ; AME =

ALE 0.5 ; PME - PME 1.55; PME — PLE 1.55 ;

AME — PME |.1; ALE- PLE 1. MOQ: anterior

width ; posterior width ; length = 2,75 53.4: 2.63,

Chelicerae; Length 1,20, width 1.69, geniculate,

almost glabrous, setae long (ca 0.6), shorter on

retromargins. Three rows of tecth on fang groove;

three teeth on promargin, middle toorh largest, three

teeth on retromargin, middle and distal teeth about

vqual, median row of about cleven minute pointed

teeth, placed posterior ty middle tooth of promargin

row. Labium: Length 0.30, widrh 1.70, emarginate

posterio-laterally, six shart setae in recurved row

across. anterior half with oné near anterior margin

midway: towards each lateral edge. Longer setae on

anterior margin, few setue on posterior half.

Maxillac: Lengih 0.75, width 0.63, slightly

Vonverging, setae long. Sternum: Lenzth 1.25, width

1,22, slightly convex, setae long.

Legs: (Table 1), Scopula sparse. Splnes: Leg f, fe

d2 p2 r3 (2 on right fe), ti rl v8, mt v4; Leg El, fe

G2 p3 rl, ui v8, me v4; Lee IO, fe d2 p3, ti v4, mt

p2 v4; Leg IV, fe d2 pl, ii pl v3. mr p2 v4; Palp,

fe d2 pl all distal, cl p2. Claws: Teeth small with

about 6 on praclaw, 4—5 on reiroclaw.

Abdomen: Setac of shore to moderale length,

Spinnercts: ALS basal segment 0.30, apical segment

0.08, PLS 0,20, 0,08.

Palps: (Fies 2-4) Tibia retrolaterally with

apophysis carrying a short triangular blunt-tipped

spur. Embolus thick, short, straight for the most

part along prolateral side of eymbium, then curved

inwards with a short blunt (ip.

Description of allatype (Figs 3-10): As in holotype

but for the following: Carapace length 3.51, width

3.50, Abdomen length 5.40, width 3.80,

Colour in alcohol: Abdomen creamish with larger

black spots,

Eyes: AME diameter 0.24. AME: ALE: PME:

PLE = |: 1.04: 0,92: ().92 Interspaces: AME —

AME 0,67; AME — ALE 0,5; PME — PME 1.3;

PME - PLE 1.5; AME - PME 1; ALE - PLE 0,92.

MOO: Anterior widti: posterior wideh: length =

28: 3,2 :.2,6,

Chelicerae: Length 1.60, width 2.10, Lubium:

Length 0.36, width 0.80. Maxillae: Leneth 0.94,

width 0.71. Steroum: Length 1,80, width 1.80,

Legs: (Table L). Spines: Lee |, fe (3 on right), ri

r2; Leg IL, fe 2 on right); Leg TV, fe p2, mt pd el,

Spinneretss ALS basal segment 0.36, apical

segment 0.08, PLS 0.30, 0.10.

Genitalia: Epigyvnum a low mound, highest

posteriorly, small fossa with weakly sclerotiyed

hood-like anterior margin (Fig. 9). Vulva paired,

copwatory openings inside hood, spermatheca

leading to a curved receptaculum shorily joined to

a smaller receptaculum (Fig. 10),

Etymology; The specific name (efers to the widely

scattered spots an. the abdomen,

Keilira sokali sp. nov.

FIGS I-13

Holotypes AM KSI9141, o, Holey Plains State

Park, 38° 13'S, 46° 51'E, Vict,, 29. ill. 1980, A.

Sokol.

Description of Holotype: Similar ta &.

spursoruculata except as follows: Carapace length

3,44, width 3.22. Abdomen length 3.90, width 2,20,

Colour in alcohol: With blackish markings

laterally on carapace, Abdomen with numerous

spots.

Carapace: Raised byt flattish above, highest in

foveul region, fovea distinct.

Eyes; Both rows slightly recurved, AME diameter

0.20. AME: ALE: PME: PLE = 1-0.9; 0.8 -

0,9, Interspaces: AME —- AME 1; AME-ALE G9

> PME - PME 1.7; PME- PLE I.h! AME-PME

0.9; ALE — PLE 8. MOQ: Anterior width -

posterior width ; length = 3.0 <> 3.4: 2,3.

Chellcerae: Length 1.44, width 1,72, Relramargin

with four teeth, irregular median rows ol & — 13

minute teeth arranged in triangular area with apex

pointing distally, Labium: Length 0.36, width (.76.

Short setae on posterior half, Sternom: Length 1,76,

width 1.70.

lees; (Table 2). Spines: Leg ] fe dl p2 3, ri 2

v8, tnt pl r2 v4; Leg IL, fe dl} p2 12, el vB, mi

p2 r2 v4; Leg IL, fe d2 p2, ti p2 r2 v4, mt p2 12

v4; Leg IV, fe d) pl ti v4, nit pl 13 ¢2-0n right) v4;

Palp, fe d2 pl, all disral, ti dl p2 rh,

Palp: Tegulum positioned more basally in

eymbium than in K. sparsomaculata and enlarged

Tetrolaterally at proximal end, Embolus extending

further io prolateral side, distal half straighter.

Tibial spur longer and more upright on larger

apophysis.

Enyniotogy: The species is named after the collector,

A, Sokol.

Remarks.

Known only from the holotype, K. sekali is

separated from &, sparsomaculaia by the flatter

carapace, four retrolateral teeth on the fang margin,

several irregular rows of median teeth and abdomen

with more numerous spots. Its habits are not

known.

Acknowledgments

The complenon of this paper was supported by

an Australian Biological Resources Sjudy grant,

Special thanks go to Dr D, €, Lec for his helpful

comments on manusctipt preparduon, Alsu io Dr

M. Gray and C, Horseman (AM) for the loan of

material and assistance.

. D. HIRST

Figs 11-15. Keilira sokoli gen. et sp. nov. Holotype maie: 1L Abdomen, dorsal; 12 cheliceral tecth, left retrolateral;

13-15 left palp; 13 ventral; 14 prolateral; 15 retrolateral. Scale lines: Figs 11, 13 = 15 = 0.5mm; Fig. 12 = 0.1mm,

A NEW GENUS OF HUNTSMAN SPIDER 13

TaBLe |, Leg measurements of Keilira sparsomaculata gen. et sp. nov. values for holotype male with allotype female

in parentheses.

Leg Femur Patella Tibia Metatarsus Tarsus Total

I 2.85 (3.32) 1.55 (1.84) 2.58 (2.82) 2.50 (2.92) 0.90 (1,12) 10.38 (12.02)

II 3.21 (3.68) 1,69 (2.00) 2.92 (3.16) 2.92 (3.16) 0.98 (1.24) 11.72 (13.24)

Ill 2.32 (2.56) 1,12 (1.38) 1.70 (1.96) 1.69 (1.81) 0.80 (0.95) 7.63 ( 8.86)

IV 2,45 (2.84) 1.15 (1.36) 2.00 (2.24) 2.1 (2.40) 0.88 (1.00) 8.63 ( 9.84)

Palp 1.00 (1.00) 0,52. (0.74) 0,49 (0,80) - - 1.50 (1.52) 3.51 ( 4.06)

TABLE 2. Leg measurements of Keilira sokoli gen. et sp. nov. of holotype male.

Leg Femur Patella Tibia Metatarsus Tarsus Total

{ 3.48 1.83 3,30 3.20 1,04 12.85

II 4,20 1.95 3.84 3.58 1.14 14.71

Hl 2.88 1.38 2.18 2.04 0.84 9.32

IV 3.34 1.30 2.52 2.64 0.92 10,72

Palp 1.10 0.66 0.67 - 1.50 3.93

References

Hoae, H. R, (1903) On the Australasian spiders of the Kocn, L. & KEYSERLING, E. (1871-1883) Die Arachniden

subfamily Sparassinae. Proc. Zool. Soc. Lond. 1902 (2), Australiens. Vols 1 & 2. (Bauer & Raspe, Nurnberg).

414-466. .

r X LATREILLE, P. A. (1804) Tableau methodique des Insects.

Fadl bee Pe od CO oe NO Mslands. Proc. Nouv. Dict. Hist. Nat. 24, 129-200.

Jarvi, T. H. (1912-1914) Das Vaginalsystem der Main, B. Y. (1976) “Spiders”. (Collins, Sydney).

Sparassiden. Ann. Acad. Sci. Fenn. (A) 4, 1-235. at . J

Kaston, B. J. (1980) “How to know the spiders”. 3rd Edin Pe ee. ) beh phe aa re Sperone

(W. C. Brown & Co., Dubuque, lowa). : 4 is ? heel °

KEYSERLING, E. (1880) Die Spinnen aus Amerika. WALCKENAER, C, A, (1837) “Histoire Naturelle des

Laterigradae, Nurnberg, 1880, 1, 1-283. Insects”, Apteres. I. (Paris),

A NEW SPECIES OF LECHRZODUS (ANURA: LEPTODACTYLIDAE)

FROM THE TERTIARY OF QUEENSLAND, WITH A REDEFINITION OF

THE ILIAL CHARACTERISTICS OF THE GENUS

BY MICHAEL J. TYLER

Summary

A large collection of frog ilia from the Tertiary site of Riversleigh Station in northwestern

Queensland, includes 190 specimens of Lechriodus intergerivus sp. noy. The new species is

described, compared with the four extant congeners, and shown to be the smallest member of the

genus. A revised definition of the ilium of Lechriodus is provided, and the significance of the find

as a link between the geographically isolated extant species is discussed. The rainforest habitat

preferences of all extant species suggest a similar environment at Riversleigh Station in the Late

Oligocene to Mid-Miocene.

KEY WORDS: Lechriodus, ilia, Leptodactylidae, Tertiary, Queensland.

A NEW SPECIES OF LECHRIODUS (ANURA; LEPTODACTYLIDAE) FROM THE

TERTIARY OF QUEENSLAND, WITH A REDEFINITION OF THE ILIAL

CHARACTERISTICS OF THE GENUS

by MICHAEL J. TYLER*

Summary

Tyrer, M, J. (1989) A new species of Lechriodus (Anura: Leptodactylidae) from the Tertiary of Queensland,

with a redefinition of the ilial characteristics of the genus. Truris, R. Soc, 8, Aust, 113, (5-21, 3) May, 1989.

A large collection of frog ilia from the Tertiary site of Riversleigh Station in northwestern Queensland,

includes 190 specimens of Lechriodus intergerivus sp. nov, The new species is described, compared with

the four extant congeners, and shown to be the smallest member of (he genus, A revised definition of the

ili of Lechriodus is provided, and the significance of the find asa link between the geographically ixolated

extant species is discussed. The rainforest habitat preferences of all extant species suggest.a similar environment

at Riversleigh Station in the Late Olizacene to Mid-Miocene:

Kry Worps: Lechriodus, ilia, Leptodactylidae, [ertiary, Queensland,

Introduction

Lechriodus Boulenger comprises a group of

vrownd-dwelling leptodactylid [rogs confined to the

Australian Geographical Region. In addition to the

morphological aticibures thal support its

recognition, it is umigue in the nature of its

distribution: of the five leptodactylid genera that

have been reported to occur both m Australia and

New Guinea, itis the only one which is represented

by more species in New Guinea than in Australia

(Zweifel 1972),

McDonald & Miller (1982) clarified the

geographic distribution of Léechriodus in Australia,

demonstrated that a previous record from north

(Queensland was in error, and confirmed that there

is a major disjunction between the Australian and

New Guinea populations. Thus, whereas other

shared genera occupy the Cape York Peninsula of

north Queensland (Tyler 1972), Lechriodus does

not, and in Australia it is confined to the eastern

seaboard along the Great Dividing Range between

Qurimbah, N.SW, and Brisbane, Qld, All that can

be interpreted about guch allopatry is that at some

time in the past Lechriodus must have occurred In

the intermediate area.

Here | report a new species of Lechriodus from

the Tertiary, The marerial comes from Riversleigh

Station in northwest Queensland, which js

intermediate between the distribution of the extant

species, The finding is significant for several

reasons: it constitutes the link in the distribution

of the extant species; it is the first record of fossil

Lechriodus, and it ts the first Occasion on which

an Australian frog fossil has been reported from a

site that js beyond the modern geographic range of

its genus,

* Pepartinent of Zoology, Universiry of Adelaide, G.P.O.

Flax 498, Adelaide, S. Aust. SOQ,

Material and Methods

The material is deposited in museums and other

collections abbreviated in the text as follows:

Australian Museum, Sydney — AM; Department of

Zoology, University of Adetaide —- AU?Z;

Queensland Museum, Brisbane =~ QM; South

Australian Museum, Adelaide - SAM, and

American Museum of Natural History, New York

— AMNH. Letters following the abbreviations are

departmental identifications.

Comparative studies were based on the

osteological collections of thé Department of

Zoology, University of Adelaide, supplemented by

skeletal material of New Guinea taxa borrowed

from the Amerivan Museum of Natural History.

Osteological nomenclature follows Tyler (1974).

The following measurements were taken with dial

callipers: ilial length - measured from the superior

extremity of the dorsal acetabular expansion to the

distal end of the shaft; dorsal acelabular expansion

to ventral acetabulat expansion (DAE-VAB)

measured as the distance between their extremities,

and acetabular fossa diameter measured at the

proximal, external edge of (he peripheral acetabular

rim.

Fig, 1, Onemation of lium to horizontal plane durine

preparation of descriptions.

All measurements and deseriptive features are

derived from the lateral surface, Expressions of

relative extent of features are obtained with the

\lium positioned in such a way that the proxwnal

edge of the bone is maintained at 45° to the

horizontal (Fig, lL). Concepts of relative size of

16 M. J. TYLER

component features to one another are perceived

with regard to their proportional size in other

anuran taxa.

Scanning electron micrographs were taken on an

ETEC Autoscan SEM at 20 Kv.

Systematics

Family: Leptodactylidae Werner, 1896.

Sub-family: Limnodynastinae Lynch, 1971

Genus: Lechriodus Boulenger, 1882

The definition of the generic characteristics of

the ilium by Tyler (1976) was based solely on L.

melanopyga (Doria). Now that representatives of

all extant species and the fossil species have become

available, the definition requires modification and

expansion:

Ilial shaft slightly to moderately curved, bearing

large, fanlike dorsal crest extending along at least

three-quarters its length. Maximum depth of dorsal

crest approximately one-third from proximal end.

Crest concave to varying extents proximally,

flattened distally.

Acetabular fossa diameter varies with size of

individual from which it is derived: proportionally

larger in larger species. Acetabular fossa bordered

by rim in large species; rim confined to inferior half

in smaller species. Location of dorsal margin of

acetabulum ranges from base of ilial shaft to

midway up shaft.

Pre-acetabular zone narrow, meeting inferior

border of ilial shaft in gentle curvature, commonly

forming quadrant. Ventral acetabular expansion

moderately developed, commonly truncate

inferiorly.

Dorsal prominence small, extending superiorly

above superior margin of ilium, or not. Dorsal

prominence moderately to well developed, oval and

horizontal or vertical.

Dorsal acetabular expansion acutely angled, its

superior margin on a level with or above the

maximum extent of the dorsal crest.

Lechriodus intergerivus sp. nov.

FIG. 2

Holotype: QM F16614. An almost entire left ilium

collected at C.S. Site, Riversleigh Station,

Queensland.

Description of holotype: lial shaft slightly curved

and bears enlarged, fanlike dorsal crest whose

maximum depth is at position approximately one-

third from proximal end of shaft. Proximal one-

half of crest concave on lateral surface, progressively

becoming less pronounced distally. Distal half of

crest flat.

Acetabular fossa small, deep and with prominent

rim bounding inferior half. Dorsal margin of

acetabular fossa situated slightly superior to inferior

margin of ilial shaft.

Fig. 2. S.E.M. of Lechriodus intergerivus (SAM P29771). A left ilium, but image reversed to aid comparison with

Fig. 3.

ANEW SPECIES OF LECHRIODUS (ANURA: LEPTODACTYLIBAR) "

Pre-acetabular zone evenly rounded with narrow

separation from acetabular fossa, Ventral acetabular

expansion poorly developed and incomplete

inferiorly,

Dorsal acetabular expansion narrow bul well

developed superiorly, extending to position

approximately on a level With maximum extent of

superior margin of (lia) crest

Dorsal prominence poorly developed and scarcely

deteclable: in profile visible as slight superior

evlension upon dorsi! marein of illum, Dorsal

protuberance ovoid, lateral, well developed and

projecting: laterally, Anterior margin of dorsal

prominence extends to position slightly anterior to

antenor margin of acetabuluny,

Length of ilium 13.6 mm; DAB-VAE 3.7 mm:

acetabular fossa maximum diameter 1.4 mm,

Paraivpes: Henk’s Hollow Site: SAM P29742: Two

‘Trees Sites AM F76951-32, AMNH 25351-53: Last

Minute Site: SAM P29764-65; Gag Site AM

F76957, AMNH 25355-58, SAM P27968; Upper

Site; AM F76953-44, 76958, AMNH 25354, OM

F14640-50_ 17036-39, SAM P29734, 29743~44,

29757-62; CS, Site : AM F76955-56, T6959.

AMNH 25359-60, QM FL6615-18, 16674,

1703|-32, SAM P29746-50; Wayne's Wok Site; AM

F 76960, OM Fl6636-39, 1703435, 17040-47, SAM

P29756, 29766-67; Outasite Sites SAM P27929,

2Y751-55; R,S.O. Site: QM F16619-35, 17033, SAM

P29735-41, 29745, 29771 (subject of SE,M.),

Variation: The complete ilia range in length (rom

5.3 mm to 14.4 mm, A paratype is shown in Fig.

2. Nereis minimal difference in overall shape, and

variation ts largely eenlined to the posttion of the

dorsal prominence in relation to (he acetabulum.

Thus some specimens agree with the holotype in

having the anterior (nargin of the prominence on

a level With the anterior margin of the acetabulum,

i Others Lhe anterior half of the prominence pay

project beyond rhe acetabulum.

The extent and degree of the curvature of che

dorsal crest varies in terms of its elevation, proximal

limit and medial protrusion,

Referred specimens: Small portions of an additional

79 specimens gre sufficiently complete co permit

\denttlication, but $0 fragmentary that they do nor

eontribute 10 an understanding of the nature op

variability of (he species. For those reasons they are

considered here “referred specimens” rather than

Paratypes. «All of the referred specimens are lodged

in the Queensland Museum; Henk’s Hollaw Site:

Fl6652; Two Trees Sits Fl4688-89; Las) Minute

Sites FL6673, 16675, 1647879, 16685; Gag Site;

FL665|, F16676-77, 16680-83, LAGBA-R7, 1670-01;

Upper Sites F16658-89, 16665-72, 170S0—53> 0,8,

Sites F16660-64, 16684, 16690, 17048: R.S.O, Site:

FL6653-57, 16691-99, 17049; Wayne's Wok Sire:

Fl6702-14, 17054~683.

Coniparison with other species: ia of all extant

species have been examined; L. flescheri (Boulenger)

(AUZ 8 uncat,): L. melunopyen (Doria) (AMNE

81223, AUZ 2 uneals; L. agenaposiv Zweitel

(AMNH 74646), £, placveeps Parker (AMNH

74178). Representatives are shown in Fig, 3.

The comparisons assume that {he sample size of

& intergerivus is sufficielit ' reflect with reasonahle

confidence the size of tha attvined by the species,

With that assumption il is relevant te nore that the

absolute sizes of the iia examined of each of the

adults of New Guinea species is substantially lurger

than any of the £, inrereerivus,

The relationship between jlial length aod snour

to vent length of extant species is almast Ifnear,

Thus given ilial length alone it is possible to

extrapolate snout co vent length, The lonvest iliues

of L. intergerivus is 14.4 mm, The corresponding

snout 16 vent length derived from the elosely lineal

relationship ol congeners is approximately 37.5 wim

(Fig. 4). This, the apparent maximum is fess Urea

two-thirds of the size of the smallest adult £.

aganoposis and L. platyceps, and is slightly below

the range of L, meloriopigaand L. fletchert (Table

1). Clearly £. intergerivus is the smallest member

af (he genus.

As indicated by the genene definition, (he itinm

an this genus is conservative and inferspeeitic

variation is not pronouliced. For (hat reason barn

reluctant to ascribe differences in ilial vharucters

greater significanve than Ley inay merit. However,

given the data on size, the integrity of £. ilergerivus

is nol in question.

Sraligraphy and lithology: The assumed

statigraphic sequence of the sites at Riversleigh is

that followed in the list of paratypes, OF the sipes

named to dale those containing L, interger(yus form

an almost unintercupted sequence, Ole of the fossil

bearing sediments is described by Fland (1985) as

fine-grained arenaveous freshwater limestones, and

possibly a facies within the Carl Creck Limestone.

The ages of the sediments containing £. dnterperivus

are currently understood tu he between Late

Oligocene and Middle Miovene (M. Areler pers.

conim,),

Elymefogy: Latin for “placed between”, $6 alluch ny

to the geographic position of the fossil population

belween those of the extant species,

Phylogeny: Zweitel’s (1972) Proposed phyligenctic

relationships is reproduced here ag Fig, 5. Giventhe

ecographic lovation af L, /ntergerivus {Fig 6), the

age of the Riversleigh Station deposits, aid the fact

ihat the lower estimarlon of age is CONLeM POTN eoUs

wih the emergence and subsequent uplift of ret

M. J. TYLER

Fig. 3. Ilia of extant species with or without ischium and pubis: A. Lechriodus fletcheri (AUZ uncat.); B. L. melanopyga

(AMNH 81223); c. L. platyceps (AMNH 74178); D. L. aganoposis (AMNH 74646),

A NEW SPECIES OF LECHRIODUS (ANURA: LEPTODACTYLIDAE) 19

tal)

7 /

a e

= 40

2S ae

3 Y = 4.48 + 2.98x

5 \ r 0.96

c {

30+ |

{

fe) am | T 1 a —- —

5 10 15 20 25 30

tum Length (mm)

Fie. 4, Length of ilia of Lechriadus species plotted against

snout to. vent length, Assumed snout tu vent length of

fossi! species indicated by broken lines, t-value for slope

10.891, p < .00L. For x = 4, y = 37.51 (95%

confidenve limits = 33,75-44,25). Square = L.

platyceps, circles = L. melanopyga; triangles =L.

Fletcher,

FLETCHER! MELANOPYGA ®LATYGEPS AGANOPOSIS

fig. 5. Phylogenctic relationships of extant species of

Lechriodus as postulated by Zweifel (1972),

of New Guinea, L. intergerivus is a candidate as

the ancestral stock from which the extant species

evolyed_

With the existence of L. intergerivus, Zweifel’s.

phenetic scheme would require two invasions into

New Guinea: one deriving L. melanopyga from an

intermediate ancestor that also gave rise to L.

fletcheri, and the other leading to L. platyeeps and

L. aganoposis. A slightly simpler scenario would

derive L, fletcheri directly from L. intergerivus, with

asecond stock giving rise to the three New Guinea

species, Further systematic studies are required to

test the validity of that option,

Discussion

The fossil frog fauna of Australia includes 22

species and nine genera from 10 sites of Quaternary

ot Tertiary age (Tyler 1989). Riversleigh Station

appears likely to be the richest fossil frog site yet

discovered, in terms of the number and diversity

of species. This trend is: common to other vertebrate

classes, and Archer, Hand & Godthelp (1988) state

thal more than 200 new species have been recovered

there. The frog fauna will increase the total.

Amongst the frog material from Riversleigh

Station now available, L. intergerivus is the

predominant species. OF 379 ilia now known from

the site 190 (50%) represent that species.

Extant Lechriodus aré predominantly inhabitants

of temperate and sub-tropical rainforest and, in the

absence of any data to the contrary, it can be

inferred that the habitat of L. intergerivus would

have been raintorest. A second point relevant to

interpreting the mid-Miocene environment is the

fact that a high proportion of the ilia of other frog

species found in Riversleigh Station deposits are

from small creatures. Because frogs lose water

readily through the skin, the body mass/surface

area ratio is such that small frogs are particularly

vulnerable to dehydration. In consequence they are

predominant only in areas which are moist

throughout the year. In communities of frogs in

Australia high frequency of small frogs occurs only

in areas of high and seasonally reliable rainfall, such

as the extreme southwest of Western Australia and

the northern periphery of Arnhem Land in the

Northern Territory (Tyler 1989),

Acknowledgments

This investigation was made possible as a result

of a Suggestion from, and the subsequent

encouragement-of, Dr Michael Archer. I am further

indebted to him for provision of laboratory space,

and yarious help from his colleagues Mr Henk

Godthelp and Dr Suzanne Hand at the University

of New South Wales.

The research program was funded by the

Australian Research Grants Scheme. Veronica Ward

M. J. TYLER

@} RIVERSLEIGH

L, melanopyga

L. aganoposis

L. platyceps

L. fletcher!

>meOe®

Fig. 6. Geographic distribution of Lechriodus.

A NEW SPECIES OF LECHRIODUS (ANURA: LEPTODACTYLIDAE) 21

TABLE I, Size af Lechriadus species in millimetres. Snout to vent length data mostly obtained from Zweifel (1972).

Snout to vent length of donor L. aganoposis unknown, and maximum length of £, intergerivus extrapolated from

Fig. 4.

species snout-vent Jength

ao GQ

aganaposis 64-73 66-77

Jletcheri 42-48 45-54

intergerivus max. 41

melanopyga 38-47 46-60

platyceps 64-78 72-95

largest jlium examined

size of ilium DAEF- acer.

donor length VAE fossa

diam

= 24,2 8.1 3.5

AZ 16.3 42 1.9

237.5 14.4 40 1.7

§2 20.7 6.0 2,6

67 24.0 74.8 38

played a vital role in that she undertook the

cataloguing and initial sorting of material, and

prepared Figures. ]-2 and 4-6.

Dr R. G, Zweifel and Dr C. W. Myers (American

Museum of Natural History) lent ila of extant

species and provided data about them, and Mr J.

I, Menzies (National Museum & Ari Gallery,

Konedobu) provided distribution records from

Papua New Guinea. Mr P. Kempster prepared

Figure 3.

I am also indebted to Dr Keith Walker and Dr

Margaret Davies for valuable discussions, and to

the University of Adelaide for the provision of

research facilities.

The materials upon which this study was based

were obtained through the support of the following

funding bodies and organisations to M. Archer, S.

Hand and H, Godthelp; Australian Research Grants

Committee; Department of Arts, Sport, the

Environment, Tourism and Territories; National

Estate Program Grant Scheme; Wang Computers

Pty Ltd; ICI Australia Pty Id; Australian

Geographic Society Inc.; Mount Isa Mines Pty Ltd;

the Queensland Museum; the Australian. Museum:

the Royal Zoological Society of NSW; the Linnean

Society of NSW; Ansett/Wridgways Pty Ltd;

Mount Isa Shire Council; the Riversleigh Society

and the Friends of Riversleigh,

References

Arcuer, M., Hanp, 8: & GoorHeip, H. (1988) A new

Order of Tertiary zalamibdodont marsupials, Science

239, 1528-1531.

Hann, S. J. (1985) New Miocene megadermatids

(Chiroptera: Megadermatidae) from Australia with

comments on megadermatid phylogenetics. Aust.

Mamm. 8, 43-54.

McDona.p, K. R, & MILLER, J. D, (1982) On the status

of Lechriedus fletcheri (Boulenger) (Anura:

Leptodactylidae) in northeast Queensland. Trans. R.

Soc, S, Aust. 106, 220.

Tyrer, M. J. (1972) An analysis of the lower vertebrate

faunal relationships of Australia and New Guinea. In

D. Walker (Ed,) “Bridge and barrier: the natural and

culiural history of Torres Strait", Dept of Biogeography

& Geomorphology, Research Schoo! of Pacific Studies,

Australian National University, Canberra.

(1976) Comparative osteology of the pelvic girdle

of Australian frogs and description of a new ‘fossil

genus. Trans. R, Soc. S. Aust. 100, 3-14.

(1989) “Australian frogs’, (Viking O'Neil,

Melbourne),

ZWEIFEL, R, G. (1972) A review of the frog genus

Lechriodus (Leptodactylidac) of New Guinea and

Australia. Am, Mus, Navit, (2507), 1-41.

OVER-SUMMERING REFUGES OF AQUATIC MACROINVERTEBMTES

IN TWO INTERMITTENT STREAMS IN CENTRAL VICTORIA

BY A. J. BOULTON*

Summary

Eight potential refuges for macroinvertebrates were sampled in two intermittent streams in central

Victoria, Australia, during summer 1982-83 and summer 1983-84. Ninety-one aquatic taxa, mostly

insects, were recorded. Receding pools harboured nearly three-quarters of these taxa; comparatively

few were collected from the hyporheos or the water in crayfish burrows. Almost half the taxa were

from refuges that did not hold free water. Macroinvertebrates persisted as desiccation-tolerant eggs

(mayflies), larvae (chironomids and some beetles) or adults (beetles).

There was remarkable similarity between the broad taxonomic representation in these refuges and

those described for intermittent streams in Ontario, Canada.

KEY WORDS: Intermittent streams, over-summering refuges, aquatic macroinvertebrates, Victoria,

Australia, pholeteros, hyporheos.

OVER-SUMMERING REFUGES OF AQUATIC MACROINVERTEDRATES IN TWO

INTERMITTENT STREAMS IN CENTRAL VICTORIA

by A. J. BouLToN*

Summary

Bouton, A. J, (1989) Over-summering refuges of aquatic macroinvertebrates in two intermitient streams

in Vigloria, Trans. R, Soc. §. Aust, 113, 23-34, 31 May, 1989.

Eight potential refuges for macroinvertebralws were sampled in two intermillent streams in central Victoria,

Austraha, during summer 1982-83 and summer 1983-84, Ninery-one aquatic taxa, mosily insects, were

recorded, Receding pools harboured nearly Uiree-quarters of these taxa; coniparatively few were collected

from the hyporheos or the water in crayfish burrows. Almost half the taxa were from refuges thal did not

hold tree water. Macroinvertebrates persisted as desigeation-tolerant eggs (mayflics}, larvae (chironomids

and some beetles) or adults (beetles).

There was remarkable similarity between the broad janonaniic representation.in these refuges and those

described lor intermittent streams ip Ontario, Canada.

KEY WORDS. Intermittent streams, over-summering tefuges, aquatic macromvertebrates,

Victoria, Australia, pholeteros, hypotheos:

Introduction

Ephemeral (episodic) and intermittent rivers and

streams drain over half of the Australian maintand

(W,D, Williants 1983) but despite their ubiquily ard

scientific interest, these systems have attracted little

limnological attention (Boulton & Suter 1986;

Boulton & Lake 1988), The situation is little better

elsewhere (Williams £987),

In intermittent streams, loss of water during the

dry season is probably the most influential

environmental parameter affecting the aquatic biota

and has led to a wide range of physiological and

behavioural adaptations (reviewed by Williams

1987). Bebavioural avoidance appears to play a

major part in the survival of many stream

invertebrates during drought. Williams & Hynes

(1977) recognized eight distinct types of refuges that

were used by the fauna of a tempvrary stream in

Ontario during summer atid suggested that

members of certain major taxonomic groups tended

to over-suminer as similar stages in their life cyele.

For example, Ephemeroptera and most

Chironomidae over-summered us exgs whereas

Gastropoda, some Odonata, Hemiptera. and

Coleoptera survived the dry period as adults

(Williams & Hynes 1977),

In temperate Australian intermittent streams,

tecolonization pathways and potential over-

summiering tefuges have neyer been investigated and

lidle is known about the physiological or

behavioural adaptations exhibited by the aquatic

* Centre for Stream Ecology, Department of Botany and

Zoology, Monash University, Claytan, Vic. 3168.

Present Address; Departinent of Zoology, University

of Adelaide, G.PO. Box 498, Adelaide, S. Aust. 5004

biota (Boulton & Lake 1988), This study was aimed

ar elucidating over-summering strategies of aquatic

macroinvertebrates in two Intermitient streams in

Victoria, 1 also was interested to see how closely

the strategies used by biota in the intermittent

Victorian streams matched those described by

Williams & Hynes (1977) based on work done on

intermittent streams in the northern hemisphere

Materials und Methods

Study Arey

Two study sites were Jocated on the upper reaches

of the Werribee River and two mare on its main

tributary, the Lerdetderg River (Fig. 1). Both rivers

arise on the southern edge of the Great Dividing

Range approximately 100 km north-west of

Melbourne andl flow south-east belore joining near

Bacchus Marsh, north-west of Melbourne. Details

of flow regime atid catchment vegetation are given

in Boulton & Smith (1985); other physicochemical

data are presented in Boulton & Suter (1986),

‘The two rivers differ in permanency: the Werribee

River ceases flow almost annually whereas the

Lerderderg River flows throughour sunimer for one

year in three. On average, the Werribee River does

not flow for nine weeks while the Lerderderg ceases

flow for ax. The Werribee River did not Mow at

all at one site (Spargo Creek, 8C) during the 1982

drought and only flowed for five mooths (late June

‘to Jate Novetnber) at the site downstream (Werribee

Picnic Spot, WPS), The study pool at WPS dried

up completely during the cnsuing summer, The

following year, flow commenced in late June and

continued for seven and a half months at both sites.

Flow started in late May 1982 at both sites

(Fireplace Ford, FF and Wheeler Road, WR) on the

24 A.J, BOULTON

Lerderderg River and ceased in early December, At

FF, the study. pool dried to a moist patch of leaves

and water did nol appear in the depression until

mid-March 1983. At WR, the study poal was dry

for six weeks before it also refilled in mid-March;

flow resumed at both sites in early May, continuing

over the following summer.

Physicochemical Monitoring of Potential Refiiges

On 20.1.1983, diel ranges of ternperature in and

around a receding pool at WPS were measured

using 2 9-channel Miniature Intermittent Temper-

alure Recorder (Grant Instruments, Cambridge,

England). Thermistors were placed in shallow (5

cm) and deep (45 cm) water, below leat’ litter,

beneath a large flat rock, in (he water of a crayfish

(Engaeus sp.) burrow and in exposed grass in direct

sunlight (regarded as. “air emperature” cf. normal

meteorological practise). Recording commenced at

5,00 a.m, and ceased at midnight.

At other times, spol walter temperatures (mercury:

thermometer), dissolved oxygen (oxygen probe,

Model 514A. Y.S.1., Yellow Springs, Ohio),

To ’

Deyiestara

“ Blakeville

’

Fig, | Map of the study urea showing the locations of the four study sites: Werribee Picnic Spot (WPS) = |

conductivity (conductiviry meter, Radiometer,

Denmark). and pH (Metrohm pH meter, Model

CH9100, England) were measured when potential

refuges containing free water were sampled.

Conductivity data were converted to values at (8°C

(Ky3) (Bayly & Williams 1973) whereas dissolved

oxygen was expressed as percentage saturation using

the conversion table in Bayly & Williams (1973) and

an appropriate correction factor for altitude,

Biological Sampling af Potential Refuges

A variety of collecting techniques was necessary

to sample the diverse range of potential refuges:

(a) An F.B.A. pond-net (300 zm mesh) was used

to sample fauna in the receding pools. I vigorously

shuffled along the bottom of the pool, sweeping

the net from side to side across the disturbed path

for 30 seconds for each sample. The size of the pool

limited the number of samples that could be

collected; while I was keen to ascertain the relative

abundance of the fauna, | did not want to deplete

the remnant populations, A nearby permanent lake

(Shaws Lake, Fig. 1) was sampled similarly,

To Trentham

DIVIDING

37°27 5,

Spargo Creck (SC) = 2, Fireplace Ford (FF) = 3, Wheeler Road (WR) = 4.

OVER-SUMMERING REFUGES IN INTERMITTENT STREAMS 25

(b) Animals residing in the water in crayfish

(Engaeus sp.) burrows (“pholeteros” sensu Lake

1977) were collected by carefully excavating the

burrow opening and lowering a flexible plastic tube

(6 mm internal diameter) into the burrow water

which was sucked oul into a plastic bag, In. the

taboratory, this waler was sieved (50 pm) with

frequent washing to remove fine silt,

inthe dry stream bed and sweeping a small hand-

net (50 zm mesh) through the seepage. In some

cases, it Was Necessary tO use a plastic lube to collect

the water, Quantitative sampling of this habitat was

not attempted.

(d) Other potential refuges thar were qualitatively

sampled while the streams were dry included the

humid microhabitats beneath rocks, stumps and

mats of leaf litter and dried filamentous algae, and

among the roots of riparian plants (c.g, Cerex spp.,

Leptospermum flanigerum) and stranded

Myriophyllum propinqguum (water milfoil) and

Trelochin procera (water ribbon), Rotting wood

was broken open and examined, and strips of bark

were peeled from cxposed water-logged snags.

Pieces.of decomposed wood were brought back to

the laboratory for microscopic inspection.

(e) Plastic bags were filled with surface (upper

10 cm) substrata and organic matter collected from

the dry beds of riffles and pools. The water vontent

of a subsample of the substratum was determined

by subtracting the oven-dry weight (constant weight

achieved after 48 h at 100°C) from the initial weight

and expressing the value as a percentage. It was

was tecorded. This process was repeated until no

further species were seen. Subsequently, samples

were taker) al wregular mlervals over the next

fortnight, always returning the specimens to che

tank. Some aquatia were marmraincd for several

months to rear hatchlings through to adults to assist

identification,

Were identificd as far as practicable (sce

Acknowledgments), Abundance was expressed

qualitatively ias “present” {1-2 individuals),

“common” (3-10) or “abundant™ (> 10); given the

variety of collecting methods and the uneven

sampling effort, more precise quantification was

inappropriate.

Results

Physicochemical conditions in potential refuges

Means atid ranges of spot water temperature, pH,

dissolved oxygen and conductivity in two refuges

that held free water when they were sampled are

lisied in Table J, Not surprisingly, the ranges of

these variables were grcater in {he teceding pools

than in the burrow water of crayfish (Table 1),

Hyparheic water was too disturbed during sampling

to obtain reliable physicochemical data and dala

trom Shaws Lake are too few to be useful,

Continuous records of water temperature in a

teceding poal al WPS illustrated the diel Nuctuation

of temperatures in various refuges (Fiz, 2), Air

temperature in direct suntight near the pool ranged

from 3.5°C at dawn to 35°C early in the aflerncon

(Fig. 2). The day was fine and clear with a light

south-easterly breeze starting at 2.30 p.m. Sunrise

TARE }. Means and ranges of waler lemperalure, pil, dissolved mxyzen and condueriviry in two. porentis!

oversuntmering refuges, based upon n spot measurements

Refuge Water

Temperature

PC)

Receding x 3

pants range 7-25

n 25

Cray fist x 43

burnow range 7-12

waler 0 8

pH Dissolved Canduchivity

Oxygen (Ky

(% saturation) (Sem)

64 30.4 154

47-72 6-78 290

23 25 25

5.8 63 RR

5,2-6.2 45-79 66-J)0)

8 8 8

assuined that all weight lost during incubation was

die to the evaporation of water.

The test of the sample was emptied into an

aquarium immediately upon return to Lhe

laboratory and flooded with dechlorinated tap

water, Within 30 minutes of immersion of the

substrata, a hand-net (SOum mesh) was swept

vigorously through the tank and the live contents

examined using an Olympus stereomicroscope,

and the rank. abundance of all invertebrates present

was at 6.2) a,m, but the pool was shaded by

surrounding forest until about 9.00 a.m. Sunset was

at 8.41 p.m.

Water temperature [i (he shallows (5 cm) lagged

closely behind air Lemperature (Fig. 2a) while in

deeper water (45 cm) the daily range was far less

(Fig. 2b). Water in a crayfish burrow exhibited a

diel range of less. than 3°C (Fig, 2b). The insulative

capacity of several centimetres of eucalypt litter and

dried filamentous algal mat approximated that of

26 AJ. ARULTON

(a)

Temperature (4G)

ears ~~ Below leat ttter

eee ee ee

0600 7200 1800

(b)

t 4

te an bans

[Deepwater 1-7 Burrow water

410

Below rock

4. a ae _ oe 4

2400 0800 1200 1380 2400

Kig, 2 Variation in water temperature of several potential oyer-summering refuges in and around the study pool

at WPS on 20,i.1983. See text tor details.

a flar rock 1§ cm thick (cf. Figs 2a and b), During

the day, conductivity remained at 270 »S/cm and

pFI ranged from 5.5 to 6.1 (n = 6 determinations,

mean pH = 5,6).

Aquatic macroinvertebrates collected from potential

refuges

A total of 91 taxa was collected from the cight

over-summering refuges sampled al four sites on

the Werribee and Lerderderg Rivers (Table 2).

Nearly three-quarters of these taxa over-summered

in receding pools along the river beds. These pools

also harboured tadpoles of Litoria ewingi (Dumeril

& Bibron) (Hylidae) and Neobarrachus sudelli

(Lamb) (Leptodactylidae); Jarge numbers of the

latter perished when the pool at WPS dried up over

the summer 1982-83. Several specimens of Galaxiux

olidus Gunther (Pisces: Galaxiidac) were collected

from the same pool.

Few taxa were recorded from the hyporheos or

crayfish burrow water (lable 2); individuals in Lhese

habitats were usually tiny. Decomposed wood

yielded smail oligochaetes, tipulids and boring

chironomids (Orthocladiinae) while amphipods

(Austrochiltonia australis) were found in curled up

strips of Eucalyprus spp. bark near the margins of

dried out pools. These habitats were not sampled

intensively and it is likely that other taxa (e.g,

helminthid beetles) may also use rotting, wood as

a refuge over summer,

Almost half of the taxa recorded were collected

from refuges that did not hold free water when

sampled (Table 2). Some of these (e.g. Nousia spp.

and ?Dinoloperla (hwaitest) apparently hatched

from desiccation-resistant eggs whereas others

survived as larvae (chironomids, psephenids,

helminthids and case-building caddisflies) or ss

adults (dytiscids and helminthids) in moist

microhabitats below rocks or mats of leaf litter and

dried filamentous algae. Large numbers of

Mmicrocrustacea (ostracods, harpacticoid and

cyclopoid copepods) were recovered from dry

substrata flooded in the laboratory. The water

content of these samples was usually less than 10%

by weight (range = 0,19 -—74,00%, mean = 7.71%,

S.E. = 0.93%, m = 53) and did not differ

significantly between samples collected from the

beds of the pool and riffle habitats (Mann-Whitney

U-test, p > 0.05).

Comparisons among the refuges should be made

cautiously because the collecting techniques and the

numbers of samples taken from each habitat

differed substantially. For this reason, J did not

atlempt calculations of coefficients of similarity of

faunal composition among refuges,

Discussion

The classification of over-summering refuges

used by aquatic macroinvertebrates in temporary

streams in Ontario (Fig. 3 in Williams & Hynes

1977) may be applied usefully to the data from the

Werribee and Lerderderg Rivers. At a broad

taxonomic level, the taunal elements over-

summering in the various refuges are almost

identival and du so al similar stages of their hfe

cycle. This may seem unexpected given the

differences in latitude, source of stream flow

(snowmelt in Ontario, rainfall in this study) and the

difference in the degree of species overlap with

nearby permanent streams (Boulton & Suter 1986;

OVER-SUMMERING REFUGES IN INTERMITTENT STREAMS 27

TABLE 2. Aquatic taxa recorded from potential refuges sampled over the 1982-1983 summer when the streams had

ceased flow. Dead specimens were omitted. Abundance categories are qualitative (+ = present (1-)2 individuals),

C = common (3-10 individuals), A = abundant (more than 10 individuals)) because of the different sampling frequency

(nj and methodology (see text). * These individuals did not grow to identifiable size in the laboratory. ** (NMV

sp.n) refers to the number of the specimen held in the voucher collection of the Museums of Victoria.

Refuge Below Crayfish Receding Shaws Wood Dry substrata Hyporheos

dried (Engaeus sp.) pools Lake and flooded in

litter burrows bark laboratory

riffle pool

(n) (7) (12) (23) (2) (10) (47) (14) (5)

TURBELLARIA

Neorhabdocoela

?Mesostoma sp. + c Cc +

Tricladida

Cura pinguis (Weiss) Cc Cc

NEMATODA

Nematoda spp. + Cc A Cc A A Cc

GASTROPODA

Hydrobiidae

Angrobia sp. A Cc +

Glacidorbis hedleyi \redale -

Ancylidae

Ferrissia spp. Cc +

Planorbidae

Physastra gibbosa (Gould) A A

BIVALVIA

Sphaeriidae

Sphaerium tasmanicum Cc

(Tenison-Woods)

OLIGOCHAETA

Oligochaeta spp. Cc Cc A A

CRUSTACEA

Janiridae

Heterias sp. c +

Ceinidae

Austrochiltonia australis A A Cc Cc

(Sayce)

Atyidae

Paratya australiensis Kemp Cc

Parastacidae

Cherax destructor Clark +

Engaeus sp. C Cc Cc

HYDRACARINA

Limnesiidae

Limnesia spp. + Cc

Hygrobatidae

Australiobates spp.

Corticacarus spp.

Mesostigmata

Aquatic sp. ss

EPHEMEROPTERA

Leptophlebiidae

Nousia spp.

Atalophlebia sp.

ODONATA

Lestidae

Austrolestes Vio (Selys) 4

Corduliidae

Hemicordulia 21au Setys +

28 A.J. BOULTON

Refuge Below Crayfish Receding Shaws Wood Dry substrata Hyporheos

dried (Engaeus sp.) pools Lake and flooded in

litter burrows bark laboratory

riffle pool

(n) (7) (12) (23) (2) (0) (47) ~~ (14) (5)

PLECOPTERA

Austroperlidae

Acruroperila utra (Samal) +

Notonemouridae

Austrocerca tasmanica (Tillyard) A

Gripopterygidae

?Dinotoperla thwaitesi *Kimmins

c C +

HEMIPTERA

Veliidae

Micrevelia dubia Hale

M. distincta Matipatil

Notonectidac

Anisops deanei Brooks

A. Thackeri Brooks

Corixidae

Micronecta annae illiesi Cc

Wroblewski

M. a. tasmanica Wroblewski +

O+

COLEOPTERA

Dytiscidae

Antiporus blakei (Clark)

A, femoralis (Boheman)

Chostonectes johnsoni (Clark)

Chostonectes spp. larvae

Copelatus australiae Clark

Hyderodes schuckardi Hope

Lancetes lanceolatus (Clark)

Liodessus shuckhardi (Clark)

Necterosoma penicillatum (Clark)

Necterosoma sp. larvae

Platynectes decempunctatus c

(Fabricius)

Rhantus suturalis (Macleay)

Sternopriscus mundanus Watts

Gyrinidae

Macrogyrus sp. +

Hydraenidae

Hydraena luridipennis Macleay +

Hy, Vtricantha Zwick +

Helodidae

Helodidae sp. larvac '

Psephenidae

Sclerocyphon striatus

Lea larvae +

Helminthidae

Austrofimnius hebrus Hinton +

A, maro Hinton '

A, “mermo” larvae (NMV sp,

Ho)**

Austrofimnius sp. larvae

Simsonia tasmanica (Blackburn)

larvae +

DIPTERA

Tipulidae

Limonia sp. +

Ormosia sp. + + +

Psychodidae

Psychoda sp. (NMV sp. 3)** p+ ©€

Q+ en

C a“

++ D+ 4+

A> FO

AM at

OVER-SUMMERING REFUGES IN INTERMITTENT STREAMS 29

Refuge Below Crayfish Receding Shaws Wood Dry substrata Hyporheos

dricd (Engaeus sp.) pools Lake and _ flooded in

litter burrows bark laboratory

riffle pool

(n) (7) (12) (23) (2) (10) (47) (14) {5)

Culicidae

Aedes spp. A Cc

Anopheles annulipes Walker Cc

Culex fatigans Weidmann Cc

C. annulirostris Skuse +

C. australicus Skuse +

Chironomidae

Ablabesmyia sp. 1 (NMV sp.

7E)** Cc +

Ablabesmyia sp. 2 (NMV sp.

66E)** -

Paramerina spp, (nr NMY sp.

32E)**

Chironomus nr februarius

(NMV sp, 136E)** Cc +

Etnfeldia sp. +

nr Dicrotendipes sp, (NMV sp.

34E)** Cc

Riethia sp. (NMV sp, 5E)** + +

Stenachironomus sp. (NMY sp.

3E)** +

Calopsectra sp. (NMV sp. 22E)** Cc 4

Stempellina nr bausei

or Monodiamesa sp. Cc +

Orthocladius-Cricotopus complex

(includes NMV spp. 12E and

160E).

Heterotrissocladius sp. F Cc c

Tiny chironomids* ‘ Cc A + + +

Ceratopogonidae

Bezzia sp. +

Nilobezzia sp. +

Stratiomyidae

Stratiomyidae spp. + c + Cc Cc

Empididae

Empididae spp. + Cc 4

Dolichopodidae

Dolichopodidae sp, 4

Muscidae

Muscidae spp. Cc + +

TRICHOPTERA

Hydrobiosidae

Ptychobiosis nigrita (Banks) +

Hydroptilidae

Hellyethira ?simplex (Mosely) +

Calocidae

Calocidae sp.

Leptoceridae

Leptorussa darlingtoni (Banks)

Oecetis sp.

Lectrides varians Mosely Cc

Triplectides similis Mosely

T. truncatus Mosely +

Tiny leptocerids*

aaaanaa

Total number of taxa 91 15 6 23 4 27 24,0 8

% of total number of taxa 16.5 6.6 747 25.3 44 297 26.4 8.8

boat

ko]

50 ALT BOWLION

Boulton & Lake 1988), However, physiological and

behavioural adaptations employed by animals of

common heritage that share gross morphological

similarities and that are subjected io similar

environmental selective pressures are likely to

converge upon. a restricied miimber of solutions

(parallel evolyition sensu Mayr 1963).

In the Werribee and Lerderiderg Rivers, several

commim laxa (Nousia sp, some geipoprerygid

sloneflies) apparently hatched from desiccation.

resistant eges (ef. Leboikuh! 1971; Snellen & Stewart

1979; Malicky 1982) and were among the fist

invertebrates ta appear when Mow resumed, Similar

finslings have heen reponed in other intermittenc

streams (Harrison 1966, Churrer 1968; Hynes 1975;

Ladle & Bass 1975; Williams & Hynes 1976, 1977;

Abell 1984; Towns 1985}, Although simutiids

(Diptera) were also common shortly after Mow

resumed in the Werribee and Lerderderg Rivers.

none emerged from the drv substrata flooded in the

laboratory, implying chat epgs are laid by adults

flying in when flow starts (ef. Hynes 1975; Abell

(984).

Mictucrustacea (ostracods, cyclopoid and

harpactivaid copepods) emerged within hours of

flooding dry substrata in the laboratory and

harpacticoid copepods Were obscrved mating a day

later. Morton & Bayly (F977) recovered ovigerous

harpacticoid ferales only 24 hours after flooding

some dricd mud from a temporary pool at Clayton,

Victoria, Suypesting thal some species diapatise at

an advanced stage of development (ef. Cole 1953).

Another taxon, common shortly afler dry substrata

were inundated, was a neorhabdocce] tcntatively

identified as Mesostonta sp.. Bayly (1970) recorded

Mesastoma from a temporary saline Jake in south-

eastern Australia and observed thick-shelled exes

in the uteri of some specimens |hat are apparently

teleased when the animals die. A similar strategy

for desiccation-tolerance has been observed in

neorhabdocoels froma tentporary ditch m England

(Cox & Young 1974).

Gastropods in the Werribee abd Lerderdere

Rivers survive drought enter by secreting a

protective epiphmem (eg. Phvsasiza eibboso,

Ferrissia spp.) (cf. Kenk 1949; Eckbtad 1973; Leger

& Talin 1973) or by closing their operculum (eg,

Angrolia sp, and Glacidorbis hedlevi) (Boulton &

Smith 85), Most tended (o aestivate in moist

intcrohahitats under stumps, dry algal mats and leaf

litter (cf. Strandine 1941; Klekowski 1959; Casey &

Ladle 1976). Whe bivalve SpAgerium sosmenicum

probably minimizes water loss by closing its valves.

Other members of the Sphaeriidae are

oveyiviparous and broud their young while surface

wirter is absent (Heard 1977; Horobach ef af. 1980,

McKee & Mackie L981). Acstivating juveniles have

been found buried in the subsisaium (Way er ul.

1Y80) bul I did not recover any from my study sites.

Parastacid crayfish found in the Werribee River

over-summer in their burrows where water

temperatures remain quite constail, Surprisingly

few other invertebrates (pholeteros sensw Lake 1977)

appeur to use this refuge (ef. Creaser 1931; Williams

eral 1974; Williams & Hynes 1976; Wigeins er ai,

1980). The pholeteros in other Victorian Engeets

spp, burrows is also depauperate (Horwity ef ail,

1Y85), Nevertheless, this refuge seems to tw

important for the survival of janirid isopods in (he

Werribee River. Isopuds usually are absent from

femporaty warers because they lack desiccation:

resistant stages and are sedentary {Williams 1985).

‘The Water in burrows constructed by fish serves as

sources Of recolonists after droughts in some

sircams overseas (e.g. Tramer 1977; Glodck 1978}

but this refuge was not evident in the Werribee and

Lerderderg Rivers.

Few taxa were abundant in the hyporheos of the

study sites during summer. This paneity may reflect

the crude sampling methods because ayuptiputls,

janirid isopods, stoneflies, molluscs and

oligachaetes have been recorded Fram the hyporheas

of the intermittent Brachina River in South

Australia (W.D. Williams 1983), Overseas, the

hyporheus is. considered to be an important refuge

from both droughts and floods in sorte temporary

and permanent streams (Clifford 1966; Williams

& Hynes 1977; Williams 1977, 1984). However, in

desert streams whose beds ate mainly composed of

unstable sand, hyporheic dormant stapes are rare

because of high temperatures in the dry streambed

and severe scouring of the channel during flash

floods (Gray 1981, Fisher er a/. 1982).

One refuge not considered by Williams & Hynes

(1977), possibly uncommon in Ontario, is chat

provided by decomposing wood debris, abundant

ajong the banks and stream beds al many Avust-

falian intermittent streams, In my study, this refuge

harboured oligochaetes, amphipods, tipulids and

chironomids, More intensive sampling is likely ta

vield further taxa because other workers haye

recorded larze numbers of xylophitous taxa from

permanent streams in Oregon (Afiderson ef a/, 1978;

Dudley & Anderson 1978) und New Zealand

(Anderson 1982),

In the Werribee and Lerderderg Rivers, taxa

aestivared under racks and mats of algae (where

temperatures remained constant) as adults (e.g.

amphipods, dytiscids} or larvae (e.g, chironomids,

stratlomyids), Some latval stages seemed

surprisingly tolerant of desiccation; for exutmple, a

large water penny larva (Sclerocyphan stréatis),

collected from the exposed surface of a flat rock

on the bed of a riffle that had Wot Mowed for 19

OVER-SUMMERING REFUGES JN INTERMITTENT STREAMS 3)

weeks, resumed activity immediately after

ituimersion in water from 4 oearby pool. Sinllar

tolerance of desiccation by larvae of Scteracyphor

spp. has been described in Tasmania (Smith I98t)

and Queensland (Smith & Pearson 1985).

Adult aquatic Hemiptera {e¢. corixids,

nolonectids) and Coleoptera (e.3, dyjiscids) prob-

ably Ply in from nearby permanent waters. Such

aerial recolonization of temporary pools and

streams by these groups is commonplace {ee

Fernando 1958, 1959; Fernando & Galbraith 1973:

Williams & Hynes 1976, 1977; Wiggins ef al, 1980;

Abell 1984; Williams 1985), | also found several

species of adult dytiscids over-suminering below

rocks and dry litter near the margins of pools at

SC and WPS, This strategy seems less well-known

for this group; only D.D. Witliams (1983),

Boumezzough (1983) and McKaige (1980)' fin a

temporary pond near Colac, western Viclonia) have

reparted similar observations.

Although the receding pools harboured most of

the taxa thal over-summer at the study sites, they

appeared to be the most physivochemically “harsh”

refuge that [ sampled. The pools experienced a

considerable diel range in water temperature, oxygen

levels were Frequently below 20% saluralion and pH

levels fell to Jess than 5. Conductivity: rises as the

water evaporates and the pools are ofien stained

dark brown with eucalypt leachate. Similar

canditions have been observed in receding pools in

Other intermittent streams in Australia (Towns 1983,

1985, Smith & Pearson 1987) and North America

(Slack 1955; Larimore er af 1959; Clifford 1966;

Harrel & Dorris 1968),

The invertebrates that over-summer in hese pools

have yarious adaptations that allow then to tolerate

such physicochemical extremes, For example. larval

dyliscids come up to the surface of the pool to

oftain air through the terminal abdominal spiracle

whereas adull Uytiscids store air beneath their elytra

(Brition 1970), Some chironomid larvae (eg.

Chironomus Wi febrwarivs) are particularly

abiundanl in the receding pools and use

haemoglobin to facilitate oxygen uptake (Colless &

McAlpine 1970). MayNy nymphs (Asalophilehia sp.),

also vommmna in this refuge, have large ills that are

constantly oscillated to enhance respiration

(Boulton & Lake 1988). Terrestrial oviposition by

two species af leptacérid caddisfly common in the

Lerderderz River (Leptorussa derlingioni and

Lecirufes verians) may also be an adaptation to low

or unpredictably fluctuating oxygen levels (Towns

1983}, Development of vulnerable juvenile stages

' MeKaige, MLE. (1980) Emergence and development of

a@atatic invertebrate communities from dried awd after

flooding. ASe Hens thesis, Department of Zovlogy,

Manosh University, (Unpubl.).

In a physicochemically harsh environment may be

avoided or accelerated by ovovivipariry exemphilied

by the stonefly Austrocerce tavmaniza, also

recorded by Towns (1985) in a South Australian

Intermittent stream.

Although there is little known about the thermal

tolerances of aquatic macrpinvertebrates in

Australian intermittent strearms, it appears that

many taxa can cope with short-term exposure to

exiremes of warer teniperature Ibis likely that they

remain near the bottom of the pool where the

temperatures may be as much as 15°C cooler than

those of |he surface water exemplified by the study

pool at WPS. Such stratification is untcommon jn

shallow pons {@2. Byars 1960; Butler 1963; Moore

1970; Hartland-Rowe 1972). However, Eriksen

(1966) recorded surface-botiom differences of

9-16°C jn temporary turbid puddles less than SNe

deep, and a shallow (10 em) rockpool fn a stream

in the Pyrences had’a surface temperature of 29,09C

while the bottom was 19°C (Chodorowska &

Chadorowski 1966), Less marked stratification in

pools in North American intermittent streams has

been reported by Neel (1951) and Slack (1955),

Isolated pools remaining in the stream bed are

important oyer-sumimeting refuges for aquatic

macroinvertebrates in intermittent streams elsewhere

(e.g, Mack 1955; Paloumpis 1958; Larimore ef af

1959; Williams & Hynes 1976; Abell 1984), In

temporary streams in Ontario, they also provide

excellent breeding environments due to the ease with

which they warn up and che abundanr plant food

ihat develops within them, and they enable species

with long-lived aquatic staves to complete their fife-

cycles (Williams & Hynes 1976), Sintilarly, in the

Werribee and Lerderderg Rivers, Ihese puuls support

arich fauna stthough environmental conditons are

harsh and predators are numerous (Boulton & Suter

1986).

Taxa whose aquatic life-spans are brief enough

to be completed while water 6 presen! are under

less selective pressure to adop! Ufese stralewes than

specles whose aquatic development takes longer

than the period that water persists. Unfortunately,

we lack information on the duration of aquatic

stages Of many Australian macroinvertebrates and

few generalizations may be drawn. Most crustaceans

and moliuses listed tn Table 2 probably live longer

than a year and this may account for their abilipy

to survive in refuges other than the pools, Alchough

many other raxa (eg. chironomids, culicids) can

coinplete their aquatic stages in a matter of weeks.

their survival in intermittent streams depends upon

when their eggs hatch and how long water persists

afterwards, Interpretation of the stenifieayce of

many of these refuges will be possible when more

information on the life historles of

32 AJ, BOULTON

macroinvertebrates in Australian intermittent

sireamns i4 available.

In summiary, there appear to be five major over-

simimerlog strategies enpploved by the fauna of

these two rivers;

i) rolerating extreme and variable enviroumen-

tal conditions in the remaining pools,

ii) surviving in moist microhabitats below stones,

stuinps and mats of dried algae and leaf litter, and

in rotting wood,

iii) over-summering. jf flerohabitats where

environmental conditions are relalively mild and

comstan! such as in the hyporheos or the burrow

water of crayfish,

i+) surviving as desiccation-resistam! stitges Lit the

dry subsira(um, and,

v) living in nearby permanent water-bodies and

flying in and ovipositing when flow resumes.

The relative contributions from each of these

tefuges reflects the nature of the substratum (ex,

Clifford 1966; Gray 1981), the amount and pattern

of discharge during the previous spring, and the

severity of the summer, This last was illustrated by

the 1982 drought when all the pools in the upper

reaches of the Werribee River dried :completely,

extinguishing several Commion taxa (eg. the shrimp

Peratya oustratiensis) that | never recorded the

following year. Differential survival of fauna in

these various refuges undoubtedly influences the

commurity composition of the stream and has a

profound effect upon ils ecological succession

during, the ensuing period of flow.

Acknowledgments

Itis a pleasure to acknowledge the enthusiastic

eevouragement aud guidance fram De Sam Lake,

Cenrre for Stream Ecology, Monush University

where 1 did this research supported by i

Commonwealth Postgraduate Research Award.

Many taxonomists identi tied my

macroinvertebrates; | thank Dr M. Harvey

(Hydravarina), Dr C.MLS. Watts (Dytiscidae), Mr

T. Weir (Heiminthidae), Dr J, Davis (Psephenidae},

Dr Ci. Theischinger and Ms C, Yule (Plecoptera),

Dr P. Suler (Ephemerupiers), Dr A. Wells

(Hydroptilidac), Ms R. St Clair (Platyhelminthes,

Lepluceridac), Dr P. Horwitz (Parastacidac), Dr].

Lansbury (Hemiptera), Drs W. Pander and 8. Smith

(Gastropoda) and Dr A. Sokol (Anura}, Dr. R.

Marchant, Mr L. Metzeling and Mr, PL Lilywhite

allowed me access to laboratory facilities and the

vaucher collection at the National. Museum of

Victoria,

T was helped in the field by Leon Barmuta, Mark

Harvey, Pierre Horwitz, Paul Huniplirles, Sabine

Schreiber, Anthony Sokol, Gaye Spanguro, Alistair

Suren and Cathy Yule and the publican at Spargo

Creek deserves special mentun for his provision ol’

an over-sutimering refuge for semi-aquatic

vertebrates, Mr Devo Mortun kimdly supplied me

with aquaria and other equipment,

Lam indebted to Drs Margaret Davies, Sam Lake

and Keith Walker, Prof. Bill Williams and two

telerees for constructive comments on the

manuscript,

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_____ (1985) Biotic adaptions in temporary lentic waters,

with special reference to those semi-arid and arid

regions. Hydrobiologia 125, §5~110.

THE LARVA AND NYMPH INSTARS OF ODONTACARUS (LEOGONIUS)

ADELAIDEAE (WOMERSLEY) (ACARINA: TROMBICULIDAE:

LEEUWENHOEKIINAE)

BY R. V. SOUTHCOTT

Summary

The larva of Odontacarus (Leogonius) adelaideae (Womersley, 1944) is redescribed, and a

lectotype designated. Larvae have been collected repeatedly on domestic cats in suburban Adelaide,

South Australia, and reared experimentally to protonymphs and deutonymphs. The larva,

protonymph and deutonymph are described. Experimental transfer of larvae from cats to man is

described. A key is given to the known deutonymphs of Leogonius, and revisional notes on the

classification of the larvae.

KEY WORDS: Taxonomy, Odontacarus, Leogonius, South Australia, larva, protonymph,

deutonymph, rearing, cat.

THE LARVA AND NYMPH INSTARS OF ODONTACARUS (LEOGONIUS)

ADELAIDEAE (WOMERSLEY) (ACARINA: TROMBICULIDAE:

LEEUWENHOEKIINAE)

by R, V. SouTHcotrT*

Summary

Sourncor, R, V, (1989) The larva and nymph instars:of Odontacarus (Leogontus) adelaideae (Wamersley)

(Acarina: Trombiculidae: Leeuwenhoekiinue). Trams. R. Sac. S. Aust. 113(00), 35-45, 3) May, 1989,

The larva of Odontacarus (Leogonius) adeloideue (Womersley, 1944) is redescribed, and a lectolype

designated, Larvae have been collected repeatedly on domestic cats in suburhan Adelaide, South Australis,

and reared experimentally to protonymphs and deutonymphs. The larva, prolonymph and deutonytnph

are described, Experimental transfer of larvac from cats to man is described. A key is given tothe known

deutonymphs of Leogerins, and revisional notes on the classification of the larvae,

Key Worns: Taxonomy, Qdontecurus, Leogonius, South Australia, larva, protonyayph,

deutonymph, rearing, cat.

Introduction

Womersley (1944) described Leenwenhoekia

adelaideae from several larval trombidioid mites,

collected from domestic cats in southern suburbs

of Adelaide in 1931 and 1941, and three collected

from rats, in Cairns, Queensland, in 1939, He had

earlier (Womersley 1934) referred to some of the

same (1931) larvae as “Lewwenhoekia (sic)

australiense (sic) Hirst 1929" (sic), ie. as

Leeuwenhoekia australiensis Hirst, 1925, originally

described from larvae collected at Ashfield, Sydney,

New South Wales, bul subsequently recorded more

widely in Australia and the south-west Pacific area

{Gunther 1952; Wharton & Fuller 1952),

Following revisions of the taxonomy of the

leeuwenhoekiine larval mites, (hose known from

Australia, New Guinea and south-east Asia

{previously referred to Leeuwenhoekia Oudemans,

1911 and Acomatacarus Ewing, 1942) are referred

ty Odontacarus Ewing, 1929, subgenus Leogoriis

Vercammen-Grandjcan, 1968 (see Southcott 1986a),

In the present paper the larva of Oduntacarus

adelnideae is redescribed, The first atternpt fo rear

larvae was in 1941, but the protonymphs and

deuronymphs described below were nol obtained

until 1950-1952.

A key is given for deutonymphs of Leogonius.

The successful transfers of partly-fed larvae from

cats (0 feeding on man ts described,

Materials and Methods

Shde-mounted mites in the South Australian

Museum Adelaide (SAM) referred to O. adeluideae

und collected by D. C. Swan in 193] and R. V.

Southcott in 1941 were examined, They were

Originally mounted in gum-chloval media, bar

* 2 Taylors Rosd, Mitchum, S. Aust, 2062,

showed evidence of extensive remounting to gum-

chloral or polyvinyl alcohol-based media, with

associated damage. Some specimens labelled as

adelaideae from NSW. were excluded, The

specimens recorded by Womersley (1944) from

Cairns, Qld, 1939 were not located,

Further larvae were collected topotypically in

194] and between 1946-1952 from the ears of

domestic cats. Some were mounted in water-soluble

media (generally gum-chloral based), whilst

attempts were made to rear others to later instars.

In 1941 larvae freshly collected from cats. were

confined under a watch-glass attached to my

forearm with adhesive strapping. Although the

mites re-attached and increased in size, no successful

transformation occurred,

Mites were again collected from the ears of

domestic cals [rom Ure same Site, during summer-

autumn of 1946-1952, and placed on strips of damp

blotting paper in sealed tubes at ambient

temperatures. In une case a nearly-intact

protonymph was obtained, In several cases

transformation to deutonymph occurred. All

specimens were then preserved in lactic acid, or dry,

until studied, generally 35-40 years later. The

devtonymphs and residual pieces of larval of

protonymphal pelts were slide-mounted, through

lactic acid to Hoyer's gum chloral medium (Baker

& Wharton 1952),

Microscopy was by a Leitz Orthelux microscope

with phasc-contrast and polarizing facilities; its

camera lucida was used to make the line drawings,

Terminology and abbreviations are as in

Southeort (1986a,b). All measurements are in um

unless otherwise stated. Prefixes: ACB to aurhur’s

registration numbers, N to those of SAM.

Odoatacarus Ewing

Odontacarus Ewing, 1929, p, 188

(for synonymy sce Southcotl, 19868, p, 171, aid

contained references).

36 R. V. SGUTHCOTTE

Fig. | Odontacerus adelaideae (Womerslcy), larva. A Dorsal view of lectotype, legs on left omitted heyond trovchanters:

legs on right completed from specimens in same postures a5 lectotype, leg [ from specimen ACB520BA2, leg 11

from ACB270A, leg IIT from ACH270B. B Gnathosoma, dorsal scutuny and adjacent structures from paralectotype

NIORR333 (ACR941), © Same for specimen ACBS20BA2. D. Dorsal idiosomal seta. F Ventral ihosomal seta, F

Tibia 1 and tarus I, dorsal aspect, of paralectotype N1988333, (D, BE ww scale on right; otber figures to seale on left.

THE LARVA. AND NYMPH INSTARS OF ODONTACARUS (LEUGON/US) ADELAIDEAE 37

Definition of larva; Trombidioidea (as defined by

Southeott 1987) with anterior median dorsal

seulum, with nasus, Wo anteromedian setae, two

anterolateral setae, two posterolateral setae, and two

sensilla, cach with non-expanded sensillary seta. Leg

scemental formula 6 6, 4, Eyes 2 + 2, sessile,

Defiatlion af deutornymph and wails:

Trombidigidea with anterior median crista,

expanded anteriorly to a blunt arrow-head-like

shape (sagilla), bearing two nurmal setae, with two

§énsilla towards ils posterior end, each bearing a

slender, setulose sensillary seta. Leg segmental

formula 7, 7, 7, Deutonymph with two pairs of

suckers (acetabula) along each lateral penital yalye,

adult with three pairs.. Dorsal idiosomal setae not

leal-like, nor divided, bul may be present im two

distinct forms. Eyes absent..

Type species: Trombicuia dentata Ewing, 1925, p.

257,

Subgenis Leogonius Vercammen-Grandjcan, 1968

Definition of larva: as in Gulf (1979, p. 143).

Type species: Leeuwenhoekia australiensis Hirst,

1925, p, 150,

Odonracarus adelaideae (Womersley)

FIGS |A-E, 3-6

Leuwentoekia (sic) austrriense: (sic) Womersley.

1934, p, 217 (partint).

Leeiwenhvekia austrifiense; (sie) Womersley, 1937,

p. 82 (purtiin).

Leeuwenhoekia qusiraliensiss Womersley & Heaslip,

4943, p. 141 (partim); Thor & Willmann, 1947, p.

323 (partim).

Leeuwenhoekia adelaidewe Womersley, 1944, p, 105;

Gill, Moule & Riek, 1945, p, 29; Gill & Parrish,

1945, Plate 3; Taylor, 1946, p. 228.

Acomutecnms adelaideae: Womersley, 1945, pp. 98,

ILI; Southecott, 1957, p, 149,

Acomatacarus (Acontatacerws) adelatdeuwe>

Wharton & Fuller, 1952. p. 97.

Odontacarus adelaldeae: Southcott, 1973, pp. 46,

103; 1976, p, 139; 1978, p. 16 (unnamed); Lee &

Southeott, 979, p. 35; 1980, p. 7.

Ordontacarus (Leogonius) adefaidese: Southcott,

(96a, p. 180,

Descripuon af larvar Lectatype: Colour in life

orange red. Length of idiosuma (mounted on slide)

370, width 285; total length from tip of cheliceral

fangs to posterior pole of idiosonia 463.

Dorsal scutum wider than long (nasus in¢luded);

nasus well developed, tongue-like, slightly pointed

anteriorly, slightly waisted, meeting body of scunum

ai approximately tight angles; anterolateral angles

slightly obtuse, rounded; lateral borders slightly

corvex; posterolateral angles obtuse, rounded;

posterolateral borders sinuvus; posterior angle

obtuse, rounded, Scutalac narrow, tapering, slightly

bhutied at dip, with pointed, auistanding setules,

Sensillary setae with 8-10 setules in distal half,

Sensillary sockets level With PL seutalae bases and

set slightly obliquely, Shield lightly porase, wilh Lwo

larger pits near each PL angle.

Standard and other data of scutum and legs of

the lype series as in Table J.

Eyes oval, curijoined, posterolateral to dorsal

sculum; anterior eye with maximum diameter 16,

posterior WW.

Dorsal idiosomalae normal, liglily tapering,

slightly blunted at tip, moderately setulose, arranged

2 (“humerats"), then 6, then th vague rows across

dorsum; total abour 35,

Ventral surface of idiosoma with a pair of

pointed, setulose setae, 33 Jong, with centres of

bases 34 apan, between coxae LIL. Behind coxae 11

Opisihovoma wih 42 setae, curved, setulose,

pointed, but the more posterior setae longer, slightly

blunted, resembling posterior dorsal jdiosomalae,

Anus 24 long by 13 across; 16 setae anterior 1 level

of middle of anus, and 26 posterior, Urstigma well

chitinized, oval, 25 long by 16 wide.

Coxalae 2, 1, 1, long, well setulose, tapering,

pointed. Lateral coxala 1 44 long, medial coxala |

68, coxala I 53, coxala II) 46.

Dorsal tracheal Opening present between

posterolateral edge of gnathosoma and anterior

border of coxa | (or overlying latter).

Dorsal tracheal opening present between

posterolateral edge of gnathosoma and anterior

border of coxa I (or overlying latter).

Gnathosuma normal, Combined chelicerae bases

ce. 73 across, by 9L long from tip of cheliceral fangs

to posterior edge of cheliccral bases. Fangs stout,

curved, blunt-pointed, with 3-5 strong retrorse teeth

along concave (flexor, dorsal) ede, and 5-7 blunted

slenticles along convex (extensor, ventral) edec.

Galeala 26 long, simple.

Palpi normal, palpal seta! formula B, B, BNN,

So + 7Ik Palpal tibial claw three-pronged, No

supracoxala to gnathosoma or legs.

Description of lees of larva (trom ACBS20BA2 and

ACB270A,B); Leg lengths (including coxae and

claws}: | 425, 11 355, II] 425. Scobalar formulae:

trochanters 1, 1, 1; femora 6, 5, 4; genua 4, 4, 4;

tibiae 8, 6, 6 (including 2 mastalae on uibia IN); tarsi

26, 16, U3 [including one mastala on Ul}. leg

specialized seraé as folluws: Leg | {from

ACBS20BA2y SaGel,38ad(21 long), VsCiel.66pul4),

SeGel.7ipd(22), SoTil.59d(16), VsTil-88d¢2),

Sofil.9Sad(10) (i.e slightly distal to Vs), Leg [1 (from

ACB270A4): SoGell.3dpd(20), VsGell,64pd(),

Solill 42pd(16), SaTiULS9padilo). Leg Wd (from

ACB270B); SoGell.26pd(21), SoFilll.SOpd(27).

38 R. V. SOUTHCOTT

Tig. 2 Odentacarus adelaideac (Womerstey), larva. Ventral view of lectotype, tegs on Jet

omitted beyond trochanters, legs on right completed as in Fig. 1-

THE LARVA AND NYMPH I[NSTARS OF ODONTACARUS (LEOGONIUS) ADELAIDEAE 9

TABLE |. Metric data for type series of Odontacarus adelaideae (Womersiey) larvae

N1988331

(ACB67)

Character Lectotype n

AW 66 5

PW 84 5

3B 24 5

ASB 50 5

PSE 28 5

L 78 5

LA 16 5

LB 62 5

LN 27 5

Ww 96 §

AP 28 5

AM 36 5

AL 36 5

PL 49 4

PL/AL 1.36 4

AMB li 5

Sens 55 3

PW/LB 1.35 5

DS 28-55 4

Hum! 55 4

MDS 28-31 5

PDS 34-39 5

Gel 57 4

Til 60 3

Tal(L) 94 2

Tal(H} 27 2

Gell 44 5

Till 50 4

Tall(L) 78 5

Tall(H) 27 5

Gelll 51 2

Till 68 2

Talil() 95 2

Talii(H) 24 2

AW/AP 2.36 5

AW/TULT 0,97 2

PW/Tilll 1,24 2

PSB/SB 1.17 5

) Humeral seta length

@ For maxintum values

Both tarsi I and II bear a large central dorsal

solenoidala: SoTal.43d(15) and FaTal.4Gad(3) (from

ACB270A); on tarsus IL (from ACB270B) 1s

Sofall.40d(15), and FaTall.37ad(4) (ie. slightly

proximal and anterior to Sofall (in usual convention

of legs being considered stretched out horizontally

at right angles to main longitudinal axis of mite),

Pretarsal formula 1, 1, 0. Tarsal claws normal,

slender, falciform, simple, neomedian longer than

anterior and posterior, each with a fringe of minute

onychotrichs,

Description of Protonymph: (Fig. 3) from mounted

specimen ACB522BA5) Colour in life not observed,

presumably orange-red. Shape irregularly

spheroidal, 630 long, maximum width 430, Skin

without setae, thin, more-or less smooth but with

many minor creases, possibly artefacts of mounting.

mean s.d Observed

Tange

70,80 £3.56 (66-74)

87.80 +2.77 (84~91)

28.20 43.03 (24-32)

51,00 +1.73 (30-54)

29.80 +1.79 (28-32)

80.80 +3.38 (78-86)

16.80 +110 (16-18)

64,00 +2.83 (62-68)

25.40 +2.30 (22-28)

95.60 +0.55 (95-96)

30,40 +1.82 (28-32)

37.20 +1.64 (36-40)

35.80 £0.45 (33-36)

53,25 +5.44 (49-61)

1,478 +015 (1.36—1.69)

10.60 +1.67 (9-13)

53.67 +3.21 (50-54)

1.374 +0,08 (1.28-1.47)

55.502 +1.00 (55-57)

55.502 +1.00 (55-57 }e

32.802 +1.30 (31-34)2

40.602 $3.13 (38—46)2

56.75 $1.26 (55-58)

62.00 +2.00 (60-64)

95,50 42,12 (94-97)

29,00 +2.83 (27-31)

47.40 +2.30 (44-50)

54.00 $2.35 (50-56)

84.00 +3.46 (78-86)

27.00 +0.71 (26-28)

53,00 +2.83 (51-55)

70,50 $3.54 (68-73)

101.50 $9.19 (95-108)

23.00 +1.41 (22-24)

2.334 £0.14 (2,13-2.52)

0.985 +002 (0.97-1.00)

1.215 +0.04 (1.19--1.24)

1.066 +0.12 (0.93-1,19)

Developing limbs represented by finger-like

processes; those of chelicerae and palpi still adnate

to main mass of protonymph, those of legs

becoming free.

A few setae and a few fragments of larval

tracheae adhering to protonymphal skin.

Description of Deutonymph: (Figs 4-6) (from slide-

mounted specimen ACB522AA/DI, supplemented

by other specimens) Colour in life orange. Idiosoma

ovoid, more pointed anteriorly and posteriorly, not

waisted, flattened below; no division between

propodosoma and metapodosoma. Propodosoma

more or less conical; posterior pole of idiosoma

rounded, Length of idiosoma (from tip of sagitta)

512, width 370; over-all length from chelicerae tips

to posterior pole of idiosoma 634,

40 R, ¥. SOUTHCOTT

Fig. 3 Odontacanss udelaideae (Womersley). Protonymph,

entire, specimen ACBS22BAS_

Propodosoma bearing dorsally a short crista 151

long, with sagitta about I5 across, carrying [wo

almost nude pointed setae 41 long, Sensillary area

46 across, wilh two large sensilla, each with a

filiform sensillary seta 100 long, with faint setules

in distal half. Crista produced posteriorad beyond

sensilla, tapering to a blunt point 47 behind level

of sensillary centres.

Crista surrounded by normal setae similar to

those of whole of dorsum of idiosoma. Dorsal

idiosomal setae unexpanded, pointed, lightly

setutose, curved, 18-46 long, the posterior setae the

longer. There is no clear Separation into

morphologically different types of setae in any arca

of the dorsum.

Ventral surface of idiosoma thickly covered with

pointed, setulose setae, 15-34 long; setules more

prominent than for dorsal setae; setae less serulose

but longer towards posterior pole, where they ate

similar to the posterior dorsal idiosomalac.

Genital aperture 86 long, by 50 wide; valves

lightly chitinized; cach media! valve with two or

three setae 13-18 long at about mid-area, slender

and Jess serulose than surrounding setac. Anus 49

long by 31 wide, with 2 or 3 curved setulose setae

along each lateral valve, similar 10 surrounding

setae.

Fig, 4 Odontavarus adelaideae (Womersicy), Deutonymph,

entire, dorsal aspect, specimen ACBS21AA/Di.

Legs Jong, fairly thin; femoral to tibial segments

more of less cylindrical, Lengths (including coxae

and claws) I 960, I] $70, IIL 895, IV 925, Leg

scobalae pointed, setulose; interspersed among

them are many pointed sensory sctac (spinalae). A

small vestigiala, 5-6 long, present on each of tibia

I] and genua | and Ji, distally and dorsolaterally.

Tarsus I outline and elongate oval, other tarsi more

cylindrical (see fig, 4).

Other morphometric data as in Table 2.

Palpi normal, with generally lightly setulose setae

(many appeating almost nudc), but along dorsal

(extensor) edges of segments a number of

unilaterally setulose setae (pectinalac), Palpal tibial

claw stout, blunted, with four accessory thickened

setae on dorso-medial aspect. Palpal tarsus clavate,

ovoid, with several setulose setac, but carrying also

a cluster of terminal smooth sensory sctae.

Cheliceral fangs fairly robust, curved, blunt-

pointed, with a row of about 10 minute denticles

along flexor (concave) edge.

THE LARVA AND NYMPH INSTARS OI’ ODONTACARUS (LEOGONIUS) ADELAIDEAE. 4l

Fig. 5 Odontacarus adelaideae (Womersley). Deutonymph: gnathosoma and adjacent structures, dorsal aspect (partly

in transparency), from specimen ACBS21AA/DI1,

42 R, V. SOUTHCOTT

TABLE 2. Morphometric detq far deutonymphks of Odontacarus adelaideae (Womrersley).

Character Specimen n

ACBS22AA/D1L

cL 104 8

5B 25 9

PDS 33-46 9

Gel 135 9

Til 187 9

Tal(L)* 244 9

‘Tal(H) 67 9

Gell 75 i]

Till 110 9

TalI(L.)4 124 9

TalI (FL) 29 9

Gelll 77 7

Tull 118 8

TalL(L)2 117 R

Tall(H) 31 8

GelV 146 8

TiIV 198 8

Tal V(L)3 175 R

Tal V(H) 36 8

Tal(L)/Til 1,30 9

mean sd. Observed

Tange

115,38 +12.71 (95-135)

26.56 + 3.61 (20-31)

47,78 + 3.837 (44-456)

141.11 +14.00 (118-160)

206.1! $20.33 (178-226)

246.67 £18.26 (218-273)

68,56 + 4.45 (60-75)

79.63 + 5.58 (72-86)

113.00 +10,51 (96-128)

12.78 +13.27 (111-148)

30,00 + 343 (24-35)

86.86 + 811 (77-100)

129,88 $13.66 (1LI-146)

127,38 +11.33 (114-143

29.38 + 3.78 (25-34)

154.50 +11.65 (137-172)

219.63 +2423 (188-253)

182.00 +14,79 (162-205)

34.50 + 4.17 (29-41)

1.23 + 0.056 (1,15-1.30)

) CL = cristal Jength, measured from anterior lip of sagitta to level of midpoints of sensillary sockets

® For maximum yalues

4 Omitting claws and pedicle.

Identification of larvae of Odontacarus (Leogonius)

in Australia

Comments

The recording of about 42 ventral opisthosomal

setae in O. adelaideae larva invalidates caption

number 12 of my earlier key to the larvae (Southcott

1986a pp. 179-180), which was based largely on

previously published descriptions.

Accordingly, captions 12~14 should be replaced

by the follawing:

12 More than 45 ventral opisthosomal setae prESen

fobs ew ghte pupelem att ber ec dale ta Be ELE 4 a pene 1

Fess than 45 (about 42) ventral opisthosamal setae

present, AL seutalac in range 33-40 pm long. PAS

34-46 um long........ O. adelaideae (Womersicy)

13 More than 75 dorsal idiosomal setae present, ....

O. austratiensis (Hirst) (including QO. pirsti{ Womersley,

1944)4)

Less than 70 dorsal idiosomal setae present,.... 14

14 PL/AL <1.200000 200. ee eee O. novaguined

(Womersley, 1944)? (including O, /ougipes (Womersley,

1945)4)

PLVAL S20). epeitpa tte ole Sete 15

1 From Goff (1079),

2 Synonymized by Domrow (1956) and Golf (1979),

4 These two species (as larvaé) were synonymized by

Goff (1979), However, see the comment of

Southcott (1986a, p. 191).

Key to deutonymph instars af Leogonius

(Australia-SW Pacific area)

1 Dorsal idiosomalaé arising from small platelety and

about 25 pm long!

9305 0000 0 QO. navaguinea (Womersley, 1944),

Dorsal idiosomalae without the above combination of

CHATACHETS~ y/-'.. ote/eteb aw edt tice mesetelits orice oe 2

2 Posterior dorsal idiosomalae over 5() zm Jong, Tal/Til

mean 1,54 (tarsus | 210-228 ym long, tibia 1 130-154

1) a ar ©. australiensis (Hirst, 1925)

(syn. O. Airsté (Womersley, 1944)?,3

Posterior dorsal idiosomalae generally loss then 50 ain

POT sen sgee alo wf et ura aces - eh eo a h tetettht net 3

3 Tarsus I longer than 300 um {tarsus 1450 pm, tibia

1440 um, Tal/Til = 1.00, dorsal idiosomalae to 30

um longyl......... OL longipes (Womersley, 1945)

Tarsus I less than 300 pm long.............. 205 4

4 Posterior dorsal idiosomalae 15-20 xm lony, ‘lal/‘Til

1.36 (tarsus T 185 pm tong, tibia I 136 um)4

Mey emery es pan pedictis ¥ O. undyi (Radford, 1946)

Posterior dorsal idiosomalae generally to about 48 pm

long. Tal(L)/Til 1.15-1,30 (tarsus 1. 218-273 ym long,

tibia |! 178-226 pm............005. O. adelaideae

(Wometsley, 1944)

1 Womersley (1945)

2 Domrow (1956)

4 Goff (1979)

4 Nadchatram (1963)

‘THE LARVA AND NYMPH INSTARS OF OUDUNTACARUS (LEOGONIUS) ADELAIDEAE 3

Fig. f Odontacarus adelaideae (Womersicy),. Deutonyniph,

taternal genitaha aud anus,

from specimen

SCBS21AA/DI..

Metlerial examined of Odontacarus adelaideae: Type

series: South Australia: Glen Osmond, Nov, 1931,

D.C, Swan, from ears of cat, 3 paralectotypes

(N1988333-N1983335, ACBK9$1-943). Unley Park

{not Unley, as stated) by Womersley 1944), 171,194],

RK V. Sguthvott. from ears of cat, one lectorype

(NI988331. ACB67), one paralectotype (NI988332,

ACE 66}.

The lectotype was designated as N1988331

because it has not been remounted since 1943

{indicated by my. notes) and is, therefore, the most

intact. Also it is in the same attitude as Womersley’s

(1944) figure.

I have excluded from the type series the three

specimens recorded in the orginal description, from

“Cairns, Queensland, 1939 W. G. H”[easlip], “from

rats," as they can no longer be found in the

collection of the South Australian Museum, and

were presumably excluded by Womersley at some

time after his 1944 paper. He did not reler to Chern

in 1945. in his last publication on Odanfocerus.

Other material: 5, Aust. Unlev Park (all trom ears

of cats of topotype arva, cullewial RVG.) LiLIM4L,

one specimen, ACH68; 9.1), 1941, several specimens,

ACB72, 74, 928; LiitI941, 3, ACBT5-77; LLG.1941,

4, ACB 78,79 (S.ii1-3941, 2, ACBBO, 8); 29.711, 1941,

lL, ACBS2; 30.11.1941, 1, ACBB3; 4iy.194], 1,

ACBS4; 6,i¥.1941, 1, ACBRS; [1.[v.1941, 3, ACB86

BCE; I4ivi941, 1, ACHE7; 16,x7,1941, 3,

ACBIAIH-102; 23.x7, 1941, 5, ACBIO3: 29.xi.1941, 5,

ACBI04; 22. xi1.1946, several, ACB270; 16.41.1947,

several, ACB319; 31-xiL.1950, several, ACB520;

SLxii.1950, two batches of several larvae, from

which deutonymphs were reared, ACB521, 922;

25.i1.19Si, several, ACB523; 1iii1951, several,

ACBS24; 9.710951, 2, ACBS25; 61.1952, several,

ACBS560 (deutonymphs rewred).

Successful larva to nymph rearing experiements

Lurvae were handled jn the 1950-]982 series of

experiments as stated above. One protunymph

(ACB522BA5) Was obtained, but the transformation

was not observed, and discovered only when jhe

tube was examined on 29.1195]. Earlier examination

on 14.2.1951 revealed nothing unusual (however in

these wet tube studies observation is at times

difficult),

Efeven deutonymphs were obtained from the

1980-1952 experiments. Nine of these came from

experiments ACBS2) aid $22, with larvae collected

On 31.xi/.1950, and nymphs having emereed over

15-29,1.1951, With batch ACBSG6O, collected on

6.11952, two newly emerged deutonymphs were

found on 27.01.1952

The finding of an intact protonymph was

lortunale, as this instar is generally unrecognizable

alter the emergence of the deutonymphs

From the above experiments, the larva to

dewonymph {ransformations took 15-24 days

Domprow (1956) recorded that in QO. qusiraliensis

deutonymphs took 25-28 days to cmerge in his

laboratory in Brisbane.

Transfer of larvae of O. adclaidese from cats to malt

On the afternoon of [1 April 194] | transferred

seven attached larvae (batch ACB86) From the ears

of @ cat to the skin of the dorsum of my forearm,

under a watchglass kept in place by sticking plaster,

On removing the watchglass 2.5 hr later ] Found two

small larvae attached to my skin, Removal of the

watchglass periodically was necessary, as

condensation made detailed observations

impossible,

Next morning the underlying skin was sore, Three

maculopapules with attached mites were observed

at different sites; these maculopapules were 446 mim

wide, and were itchy. That afternoon the area of

erythema around the attachinent sites had grown

to 10-11 mm wide, these had become conjoined

(only two mites were now attached). Sinall while

specks were seen at the summits of the papules (four

bencath one mite, three beneath the other),

44 R, V. SOU PHCOTT

presumably mite faeces, Small vesicles were present

at the summit of each papule al (he mile attachment

siles (20-22 hours after initial attachments (codes

as § 4+ 20-22 hr).

The tiching, redness and vesiculation increased,

The attached mites increased slowly in size to “half-

grown”, Ac‘S + 39 hr the whole area under the

watchglass was itching, diffusely swollen and

somewhat indurated; this area was 45 mm across,

and by this stage the mites were considered ful)

gmwn (as compared with the larvae that had been

observed on the cat’s ears). The papules had become

sO prominent as to press upon the covering

watchglass. The mites continued to make amall

faecal pellets beneath rhem.

By S + ¢c, &) hr twa vesicles had broken and were

coving serum, No lymphangitis or lymph-

adenopathy had developed,

After the miles had detached the swelling reduced

rapidly, and scabs developed over the broken skin

surfaces,

By S + 8 days the papules had retrogressed to

being almost Hat, and the erythema was fading. No

general symptoms attributable Lo (he mite bitings

or thelr lesions develuped,

Unfortunalely, this attempt ro obtain mite

dentonymphs by human feedings was \insuiccessful,

A summarized account of this experiment was

recorded earlier (Southcott, 1973),

Acknowledgmeits

[thank the South Australian Museum far access

to material in their collection,

This work was supported by the National Health

and Medical Research Council and the Austrafian

Biological Resources Study,

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introduction to acarology.” {The Macniillan Campans,

New York).

Dewtrow, R, (1956) Three new Australian chigger nymphs

{Acarina, Trombicutidae), Proc, Linn, Soc. NSW RI

42), 144-152,

Ewing, H. B. (1925) A contributian to our Knowledge of

the taxonomy of chiggers meluding the descriptions of

&@ New genus, six new species aod a new Variely, 4 eer

JA trop, Med, 5 (A), 251-265.

(1929) A manual of external parasites. (Charles

C, Thomas, Springheld, Mlinvis & Baltimore,

Maryland}

(1942) Remarks on the taxanomy of same

American chiggers. (Tronthiculinae}, including the

descriptions of new genera and species... Parasiral. 28

16), 455-499.

Gi, DA, Moure. G. RK, & Rien, Ro EF (1945)

Trombidiosis in sheep in Queenstand, Aust, ver, 21

12), 22-31.

_, & Pawrisy, E. (1945) Microphotography as an

aid to the identification of trombiculine larvae. J,

Coune, Sek Ind, Res, V8 (4), 298-300,

Gorr, M. L. (L979) The genus Odontacarys LAcati:

Trombiculidwe> in New Guinea, with descriptions of

four new species. nied, Entomerl. 1S (2), 144-154,

GunTiter, C.F. ML (1982) A check list of the crombiculid

lurvate Of Asin and Australasia. Proc. Linn Soc NSH,

77 (1-2), 1-60.

Hikgst. S (1928) On a harvest bug (Leen wenhvekia

oustraliensis, sp. tt.) attacking humun heings a1 Sydney,

New South Wales. frams & Sar, frop, Mea, & Ave,

19 (3), 150-152.

Lite. DC, & Souvucort, R, V, (1979) Spiders and ather

arachnids of South Australia, Pp. 29-43, i Leonard,

B. E. (ed.) “South Australian Year Book. No. 14: 1979"

(Australian Bureau of Statistics, South Australian

Office), (Reprinted 1980 as separate, pp I-15. 4South

Australian Muscym, Adclaide)),

NabcHarrkAM, M, (1963) The latva and nymph of

Celantacaras audyi (Radtord) (Acarina; Trombiculidae).

Pacific lasecis 3 (3), 535-440.

OVOPMANS, AL. (IVI) Acarclogische aunteekeningen

NAXVL, Ent. Ber, Amst, S458) 137-134,

Ranrora ©. &. (1946) New species of larval mites

(Acarina: Trombiculidae) from Manipur State, India,

Proc Zool, Soe Londen 116 (2), 247-2165,

Soutucomr R. V. (1957) The genus Acomiaracaries

(Acarina: Trombiculidac. . Deseription of three new

species fram Triniry Ray, orth Queensland, Trans, R.

Soe, 9. Aus, BU, 146-155.

— —— (1973) Survey of Injuries to Man by Australian

Terrestrial Arthropods. {R. ¥. Souihcott, Mitcham,

Suuth Australia

(1976) Arachnid and allied syndromes in the

Sustain region, Ree, Adel. Childs. dfosp. 1 (1),

97-187,

——__ (1978) Australian harmful arachnids and their

alhes, 4R. ¥, Southeott, Mitcham, South Australia).

(1986a) The genus Odontacarws (Acarina;

Trombiculidite}, 1, Observations on the fife history and

morphology of Odontacarus swan n. sp., and related

forms. Ree. S. Awsi. Mus, (9 (12), 149-200,

(1986b) Descripnon of Odantucuras veitoni sp.

nov(Acarina: [rombiculidac). fbid. £9 (14), 213-217.

______ (1987) The classification of the mite families

Trombellidge and Johnstonianidae and related groups,

with 2 deseriptian of a new larva (Acarina:

Frombcitidac: Notirotrombidium) from Nanh

America, Trams. R. Soc. 5, Aust, Wt (1h, 2542,

Tavion, FH. (6946) Section |. Description, pp. 7-234.

(m Taylor, FH, & Murray, R. E. (Eds.) ‘Spiders, ticks

and miles including the species harmful wm man in

Australia and New Guinea" (Serviee Publication No.

6) School of Public Healt & Tropica) Medicine,

Uuiversity of Sydney,

Toor, S & Wittmann, Cy (1947) liombidiidae. Das

Vierreich Tho, xxix-xxxvi & L&7-S41.

VERCAMMEN-Ghas DEAN, Po A. (1968) “Phe (higgec

Mites of the Far Basr (Avarjna: ‘Jrombivulidac &

Lecuwenhoeklidae). At illustrated key and a synopsis;

some new tribes, gener and subgenera" 4U, 5S. Army

Med. Research and Development Command,

Washingtan, DC.)

Warton. G. W., & FULLER PL S (1952) A manual of

he cluggers, The biolugy, classifiedtion, distriburcien,

and importance to man of the larvae of the family

Drombicriaas;Agetinad. Mem Ent. Sac. Washingion

4, [-185.

THE LARVA AND NYMPH INSTARS OF ODONTACARUS (LEOGONIUS) ADELAIDEAE 45

WomeErRSLEY, H. (1934) A revision of the trombid and Australia and New Guinea. Trans. R. Soc. S. Aust. 68

erythraeid mites of Australia with descriptions of new (1), 82-112.

genera and species. Rec. S$. Aust. Mus. 5 (2), 179-254. ______ (1945) Acarina of Australia and New Guinea. The

_____ (1937) A revision of the Australian Trombidiidae family Leeuwenhoekiidae. Ibid. 69 (1), 96-113.

(Acarina). /bid. 6 (1), 75-100. , & HEasip, W. G. (1943) The Trombiculinae

_____—(1944) ~Notes on and additions to the (Acarina) or itch-mites of the Austro-Malayan and

Trombiculinae and Leeuwenhoekiinae (Acarina) of Oriental Regions. /bid. 67 (1), 68-142.

THE COMPOSITION AND DISTRIBUTION OF THE VEGETATION OF

NORTH-WEST EYRE PENINSULA

BY F. TIVER, A. D. SPARROW & R. T. LANGE*

Summary

The vegetation of inland north-western Eyre Peninsula, South Australia, is sampled for

phytosociological analysis by transect quadratting. Using a combination of numerical classification

(Bray-Curtis dissimilarity with average-linkage clustering) and ordination (detrended

correspondence analysis), seven main vegetation associations are identified: Eucalyptus dumosa-

Westringia rigida, Eucalyptus gracilis-Stipa, Eucalyptus gracilis-Eucalyptus oleosa, Eucalyptus

oleosa-Melaleuca pauperiflora, Eucalyptus oleosa-Enchylaena tomentosa, Eucalyptus brachycalyx

and Eucalyptus yumbarrana-Triodia irritans. The last occurs on the siliceous Moornaba sands, and

the other six on the calcareous Woorinen Formation. Environmental parameters correlate only

weakly with a floristic gradation over the calcareous soils, suggesting that historical influences may

be important in determining the distributions of the vegetation types.

KEY WORDS: Eyre Peninsula, vegetation survey, vegetation analysis, classification, ordination,

plant associations.

THE COMPOSITION AND DISTRIBUTION OF THE VEGETATION OF NORTH-WEST

EYRE PENINSULA

by F. Trver, A. D. SPARROW & R, T, LANGE*

Summary

Tiver, F,, SparRow, A. D. & Lance, R. T. (1989) The composition and distribution of the vewetation of

north-west Eyre Peninsula. Trans. R. Soc. S. Aust. 113, 47-61, 31 May, 1989.

The vegetation of inland north-western Eyre Peninsula, South Australia, is sampled for phytosociological

analysis by transect quadratting. Using a combination of numerical classification (Bray-Curtis dissimilarity

with average-linkage clustering) and ordination (detrended correspondence analysis), seven main vegetation

associations are identified: Eucalyptus dumosa—Westringia rigida, Eucalypris gracilis~Stipa, Bucalyptus

gracilis-Eucalypius oleasa, Eucalyptus oleosu—Melaleuca pauperiflora, Eucalpytus oleosa-Enchkylaena

tomentosa, Eucalyptus brachycalyx and Eucalyptus yumbarrana-Triodia irritants. The last occurs on the

siliceous Moornaba sands, and the other six on the calcareous Woorinen Formation. Environmental

parameters correlate only weakly with a floristic gradation over the calcareous soils, suggesting that historical

influences may be important in determining the distributions of the vegetation types.

Key Woros: Fyre Peninsula, vegetation survey, vegetation analysis, classification, ordination,, plant

associations,

Introduction

Until zecently, the vegetation of north-western

Eyre Peninsula remained relatively unknown, The

first detailed study was that of Crocker (1946).

Unfortunately the accompanying map only extends

northwest as far as Poochera, where the vegetation

is described as Eucalyptus aleesa ~ E.gracilis -

Edumosa edaphic complex. Specht (1972) also

included only the southern-most portion of the area

in his maps, describing the vegetation south of the

Eyre Highway as Aucelypius socialis — E. gracilis

open scrub and that north of it as Eucalyptus

inerassata — Melaleuca uncinata open scrub.

Boomsma & Lewis (1980) described 16

communities, defined by the tall dominants, which

occur on north-west Eyre Peninsula. Their

distribution maps are not detailed enough to be able

to extract the relative importance of any of these

communities.

Descriptive surveys such as those mentioned

above are not always capable of repetition by others

(Nilsson 1986). For this reason, most modern

Surveys use quantitative data of the whole flora, and

various forms of computer-assisied multivariate

analysis, which result in a higher degree of

objectivity and repeatability, A mumerical

classification of the vegetation of western Eyre

Peninsula, one of the few such studies in South

Australia, was undertaken by Margules & Nicholls

(1987). Using complete species data from 104 plots

each of 0.1 ha, cluster analysis was used to identify

six vegetation communilies on western Eyre

Peninsula. This survey did not extend northwards

* Dept of Botany, University of Adelaide, GPO Bux 498,

Adelaide, \S. Aust. S001

onto the sand dune formations, Statistical models

were constructed to relate the occurrence of the six

communities to three environmental variables:

depth of carbonate layer, distance from the coast

and latitude. Although allowing prediction of likely

vegetation types in other remnant patches, these

factors do not adequately describe the

environmental factors important in determining the

vegetation pattern.

The aims of this study were to classify (he

yegetation of north-western Eyre Peninsula (Fig. 1)

using repeatable, numerical methods, and to provide

some explanation of the main factors controlling

the vegetation pattern,

Fare r

Geduna

Yor) finale

£ SP pa

— ae

100 ?cO

Kilomactes

Big. §. Showing the location of the study area on Torth-

western Eyre Peninsula, S.Aust.

48 F. TIVER, A, D. SPARROW & R. T, LANGE

Geology, Landfarm and Soils of the Study Area

Most of the surface geological formations of

western Eyre Peninsula (see Fig. 2) are derived from

calcareous sands, It has been suggested that these

sands were blown in from the exposed marine

continental shelf during periods of low sea level and

aridity, which occurred during the Pleistocene ice

ages (Crocker 1946; Twidale & Campbell 1985).

Another possibiliry is that the sands represent

palaco-dune systems that were formed by wave and

wind action during periods of high relative sea level

(Short ez al, 1986).

The Bridgewater Limestone is composed of

calearenite, grains of calcareous sand cemented

together with calcite crystals. In the maritime zone

itis exposed as coastal. cliffs but is otherwise covered

by unconsolidated, deep, calcareous shelly sand of

low water-holding capacity (Wright 1985), The

Bakara Calcrete consists of calcarenite in which the

shell fragments have been progressively dissalved

and replaced by nodular and platy calcrete. To

reduce the ecological complexity of the survey, these

coastal limestone formations were excluded from

the study.

The Woorinen Formation closely corresponds to

the Chandada Plain landform in which the dunes

have consolidated to form an undulating plain with

MOLINEAUX

SANDS

Maltee

no surface drainage (Twidale & Campbell 1985).

Calcrete is usually present as a bed of nodules

beneath the surface. The soils are calcareous

throughout, and range in texture from sands to clay

loams. They are usually shallow and reddish brown,

but sandier grey varieties and redder varieties with

higher clay content have been reported; they have

been extensively cleared for agriculture (Wright

1985). Areas of the calcareous sands normally

associated with the Bridgewater Limestone also

occur on the Woorinen Formation. Near the coast

these sands are coarse and shelly, but further inland

they are firmer with less-apparent shell fragments,

Wright (1985) reports deficiencies in a wide range

of minerals in these sands,

The Molineaux or Moornaba Sand is of

extremely recent origin (c, 10.000 years bp. onwards)

and composed of sand dunes aligned in a NW-SE

direction. Bleached white siliceous sands with a

yellowish B horizon occur on the dunes. The

underlying scils of the inter-dune corridors contain

calcareous carths typical of the Woorinen

Formation.

Methods

Sampling Strategy

A system of transects. was chosen as the best

compromise between equal coyerage of the region

‘

eee AUX SANDS

Kilometres

™,

Nunjikompita

™.

ae,

*Pimbaacla

WOORINEN

FORMATION §Wirrulla

BaKARA

GCALCRETE

Fig. 2 The study area, showing major surface geological farmations (after Blisset 1969).

VEGETATION OF NORTH-WEST EYRE PENINSULA 49

fo 14 20 25

Kilometres

Nunjikompita ©

. s

° S

my Pimbaacts

Wirtulla «&

Fig. 3 The study area, showing location of sites along seven iransecis, Shading represetuts uncleared vegetation.

and efficiency of time spent on data collection.

Transects were placed in suitable areas of native

vegetation, running north and south from Ceduna,

Maltee, Mudamuckla, Nunjikompita, Pimbaacla,

Wirrulla and Cungena (Fig. 3). Transects were

extended southwards only as far as the boundaries

of the Bridgewater formation and its overlying

coastal sands, and northwards as far as. was

accessible by four-wheel-drive vehicle. In the

northern part of the study area, sites were placed

1 km apart, but in the southern part of the area,

which is widely cleared for agriculture, native

vegetation was often restricted to roadside

reninants, and some sites had to be located as much

as 2 km apart.

Field experience showed that quadrats of 0,1 ha

as recommended by Whittaker (1978) and used by

Margules & Nicholls (1987) were an appropriate size

to achieve a good representation of species present

without overlapping different vegetation types. One

$0 x20 m rectangular quadrat was measured at each

Site,

vegetation Datu

In each of the 162. sites sampled, all seed-plant

Species were recorded, although only native

perennials were used for analysis. Annual species

were excluded because their occurrence depends on

recent seasonal events; their inclusion would

therefore lower the repeatability of the results.

Introduced species were also excluded, since they

do not form part of the natural vegetation pattern,

Total cover was estimated for cach species

present, and expressed as a semi-quantitative score,

an adaption of the cover scale as Braun-Blanquet

(1932); see Table }.

Environmental Data

Antivipating some influence on floristic

composition by various disturbance factors.

disturbance by erosion, grazing, animal tracks,

undergrowth clearance and carthworks was visually

assessed on a scale ranging from O (nil) to 4 (Severe).

Sixty-two sites rated 2 (mild) or greater were

excluded from analyses for definition of vegetation

types,

Landtorm was assessed in the field as dune-crest,

dune-flank, rise or plain (including wide interdune

corridors). A soil core was dug in the centre of each

quadrat and the depth to limestone recorded, in

intervals of 10 cm, Texture was determined using

the field method described by Northcote (1971).

Existing rainfall maps were not detailed enough for

the purposes of this study, so a rainfall map was

50 F, TIVER, A. D. SPARROW & R. T. LANGE

constructed, using complete Bureau of Meteorology

rainfall records for 24 recording stations within or

near the study area. This enabled the average

rainfall at each site to be estimated and classed into

12 mm intervals,

Vegetation Analysis

The need to classify vegetation raises the question

of whether vegetation types occur as discrete entities

with distinct boundaries, or whether classifications

are merely abstractions imposed by ecologists onto

a continuously variable vegetation (Greig-Smith

1983). If vegetation consisted of random

combinations of species a continuum would result

and classification into discrete groups would be an

entirely arbitrary process. If, on the other hand,

variation was always discontinuous and no

intergradation occurred, there would be a finite

number of vegetation types in existence, and

classification would be a simple matter. The actual

case lies between these two extremes, ranging from

the near discontinuous and distinct vegetation types

to almost continuous ecotones (Webb 1954).

GROUP DESIGNATION NUMAECR DF SITES

iN GROUP

165

Consequently, a combination of two

complementary analysis techniques is recommended

when investigating vegetation patterns; a

classification to define vegetation types, and an

ordination to examine the relationships between the

groups and the extent of ecotonal behaviour

between them (Gauch 1982).

For classification purposes, the dissimilarity

between all pairs of sites was calculated with the

Bray-Curtis distance coefficient, and then sites

TABLE 1. Semi-quantifative scores expressing percentage

cover (adapted from Braun-Blanquet 1932).

Cover Score Percentage Cover

absent

0-1

1-2

2-10

10 — 50

50 -— 100

UhwWheS o&

BRAY-GURTIS GISTANGE COEFFICIENT

+7Q +75 50 285 -90

| Eucalyptus dumgeg 5

Wastringie riglda

W Eucalyptus gracilis 6

Stipa sp+

GUNGENA 293 |

tH Eucalyptus gigogg 30

Eucalyptus grucilig

Iv Eucalyptus vleoua 17

Melaleuca paupariflora

Vv Eucalypius plegsa 15

Enchylaens tomentosa

VI Eucalyptus brachycalys it

Vil Eucatyptus yumbarrang 4

Triodia inritans

T a a T art 1

———

Fig. 4 Summary of dendrogram trom clustering analysis showing arrangement of groups at 0.65 distance level.

VEGETATION OF NORTH-WEST EYRE PENINSULA iJ

grouped from the least to the most dissimilar by

the unweighted pair group method using arthmetic

averages (UPGMA; Legendre & Legendre 1983) to

produce a hierarchical dendrogram, the branches

of which represént vegetation types. The

complementary ordination method chosen was

Wetrended correspondence analysis, DCA (Gauch

1982), Ordination methods such as DCA rank sites

along a series of axes which describe the major

floristic variation. The results are displayed as a

scattergram from which continuity and disjunction

of vegelation types can be determined and with

which environmental factors can be correlated. The

numerical taxonomy computer software package

NTP (Belbin é7 a/, 1984) was used for both analyses.

For the purpose of preparing vegetation maps,

the sixty-two disturbed sites were returned to the

data set, artd a further cluster analysis. performed.

By the proximity of these sites on the dendrogram

to sites of known vegetation type, most of the

remaining sites were identified as belonging to one

of the major vegetation types.

Resuiis

The classification dendrogram is summarised in

Fig. 4. Field notes indicated that the groupings

defined at the 0.65 distance level were the most

readily interpreted, a higher level producing groups

with large internal variation, and a lower level

producing too many groups for consideration.

Seven major vegetation types were identified at the

0.65 level, one of which, ne Vl, was widely

AMIS = (Variation Explained 21 Gb?

dissimilar from the other groups, only fusing with

them at 0.90 level, A minor vegetation type was alsa

identified. Represénted by only one site, Cungena

29, this site was dominated by Callitris preissii,

rather than a eucalypt species.

The relationships between the vegetation types

can be seen from the ordination; a plot of the first

two ordination axes is given in Fig. 5. The first axis

explained 50% of the variation in the data set, and

corresponds to the first major branching of the

dendrogram, separating vegetation type VII from

the others, The separation between the remaining

six communities is largely supported by the

ordifiation, although there is a certain degree of

overlap, most marked between types [ and II, and

types V and VI, The relationship of these six

Vegetation types to each other obviously tends

towards the “ecotone” rather than the distinct

“association” end of the scale in vegetation pattern

as described by Webb (1954).

The species consistent in each of the vegetation

types are presented in Table 2 and Fig. 6 summarises

the associated landforms, soils and rainfall. Table

3 gives a full listing of species found in rhe seven

vegetation types. Understorey species are equally

important as shrubs and trees in delineating

vegetation type.

The vegetation types, their distributions (Fig, 7)

and their environmental correlates are discussed

below. The types have been named “associations”

according to usual ecological practice, even though

they do not fully fit the formal definition af

“association” given by Braun-Blanquet (1932).

AMIS § (Vatlatien Exptained SOM)

Tig. 5 DCA ordination showing grelauioriships between the seven major vegetation types. Symbols denoting vegetation

type are as follows: O , |;

4, 1h 4, 15 @, IV, @, Ve 0, Vis VW, VOL.

52 F. TIVER,, A, D, SPARROW & R. T. LANGE

TABLE 2, Plant species characteristic of the seven major vegetation types, Marked species had u cover score of two

or greaier when present, and could be considered daminants,

VEGETATION CHARACTERISTIC PLANT SPECIES

TYPE Key Species (75-100%) of sites) Usually Presenr (50-74% of sires)

f Cussia nemophila Eucalyptus gracilis ®

var. nemophila © Eucalyptus oleosa ¢

Eucalyptus dumosa* Dianella revaluia

Geijera linearifalia ® Exocurpos aphyltus.

Hestringia rigida * Pittosporum phylliraeoides

Acacia colletioides Rhagodia preissti

Eremophila glabra

Melaleuea lanceolata

Zygophyllum apiculaturm

ul Eucalyptus gracilis * Acacia colletioides *

Stipa sp. © Eucalyptus oleosa ®

Enchyluena totnentosa Eremophila glahra

Geijera lineorifolia Melaleuca lancedlata

Sclerolgena obliquicuspis

a Eucalyptus gracilis © Eremophila seoparia *®

Eucalyptus oleosa * Triodia irritans *

Westringta rigida ¢ Maireana eriaclade

Zygophyllum aurantiacum Rhagodia crassifolia

TV Eucalyptus oleosu * Rhagodia crassifolia

Melalexca pauperiflora @

V Eucalyptus oleosa © Maireana erioclada ®

Enchylaena tomeritosa * Rhagodia crassifolia *

Geijera linearifolia Scleroleana diacantha

Zygophyllum aurantiacum Sclerolaena obliquicuspis

Threlkeldia diffusa

Vi Fucalpptus brachycalyx * Melaleuca pauiperifiora «

Maireana erioclada Enchvlaena tomentosa

Rhegodia crassifolia Sclerolaena diacaniha

Threlkeldia diffusa

Zygophyllum aurantiacum

Vil Eucalyptus yumbartana © Dianella revoluta

Melaleuca eleuthrosiachya *

Trivdia irritans *

The Vegetation Associations

Association f Eucalyptus dumosa-Westringia rigida

The closest counterpart in the literature is the

Melaleuca lanceolata association of Boomsma &

Lewis (1972), which is said to occur with a number

of species as co-dominants, including Eucalyptus

dumosa and &. eracilis. Their classification inclhides

a much wider range of vegetation associations than

is here defined. It occupies a somewhat. irregular

distribution throughout the study area, restricted

to rising ground on shallaw calcrete, and the least

sandy soils (sandy loams and Ioams).

Association IT; Eucalyptus gracilis-Stipa

This vegetation association could be described

as a depauperate variation of Association I.

Although some of the same species are present,

Eucalyptus gracilis is the most important tall species

uniling the group. The lowér diversity understorey

Eremophila crassifolia

Podolepis capillaris

often includes a high percentage of perennial grass

(Stipa), The relationship of Associations | and II

is indicated by their proximity on the dendrogram

(Fig. 4) and overlap on the ordination (Fig. 5).

The association has not been previously recorded

on western Eyre Peninsula, and &. gracilis has only

ever been reported as a co-dominant with other

mallee species for the whole statc (Boomsma &

Lewis. 1980). lts distribution is also irregular,

occurring on the dry central to northern section of

the Cungena transect, but also scattered through

the southern Pimbaacta and Wirrulla transects at

higher rainfall, Very shallow, light soils on low lying

ground are consistent with its occurrence. The

resulting harsh moisture conditions probably reduce

the number of understorey species present to. those

which can tolerate them, such as the shallow-rnoted

Stipa and Sclerolaena spp,

VEGETATION OF NORTH-WEST EYRE PENINSULA 53

Association {i> Eucalyptus gracilis-oleosa

Association UIT probably forms the core of the

Eucabprs oleasa / £. gracilis association identified

Ixy Crocker (1946) and Specht (1972), although

Specht appears to misidentify £. ofeasa as &.

Sociaifs. lt is the most common vegetation

association in the northern part of the study area,

but was not recorded by Margules & Nichatls (L987),

Its ovcurrence in Lire southern section of the

Nunjikompita transect at slightly higher rainfall

seems to sugpest that its distritution is more

restricied by soil than climatic factors. [t is not

present in che south-western part of the study ares

because the calcareous sands there favour other

vegetation associations such as [VY and V1, or on

the south-eastern section where calerete outcrops

favour Associations | and JI.

The occasional occurrence of Association JL) on

dune flanks may be explained hy historical factors.

[tis the most conimon inter-dune vegetation of the

iorthern part of the study area, occurring on

medium depth to shallow soils. Since the Molineaux

sands have been deposited over only the lyst few

thousand years, they are relatively unstable. In some

instances sand from these dunes has drifted on to

calcareous earths in the anterdunes already occupied

iw Association 11 vegetation, and the long-lived

species such as E. ofeos@ and & gracilis persist un

a soil which would perhaps not at present favour

their establishment. The understorey in these cases

tends to include species less typical of the

association, such as Triorlia irrifans,

Association [¥> Lucalyptus cleosa-Melaleuca

pauperiflora

Association 1V contains a very species-poor and

sparse understorey of chenopods such as Rhagedia

crassifolia and Muireana erioclada, A known

yegetation association including Eucalyptus aleosa,

Rhegodia crassifolia, Zygophyllum aurantiacurn,

Selerolevne diccantha and Melaleuca lanceolaio

(Margules & Nicholls 1987), is probably equivalent

to it, their Afelafeuca /anceoleta a possible

misidentification of Af. pauperif{lora.

Association LV occurs mainly in the

southernwesiern part of the study area, on shelly

calcareous sands, with scattered occurrences in the

north, for cxample, on the Nunjikompita lransect,

where Quilbers of this soil (ype occur on rises in the

inler-dunes, These outlicrs are probably remnants

of the old coastal dune environment which would

have been present before the Molineaux sands werc

deposited. Although the vegetation association

appears to occur al slightly higher rainfall, this may

simply be a result of the soil type which favours

its occurrence being characteristic of the southern

coastal higher rainfall areas. The low uiversity of

tie association may be an indication that only a

few understorey species are able lo persist an the

coarse shelly sands,

Association VO Eucalyptus oleosa-Enchylaena

lomentoga

Vegetation falling within Association V ts very

variable and a large number of disturbed roadside

siles were roost closely allied ro u, indicating rhat

it may not bea naturally occurring association, but

a result of disturbance. Mathy of the species are

unpalatable, such as Geijere linearifolia, or tow

growing chenopods, which suggests removal of

more palatable understorey species by grazing.

Further investigation would be required to ascertain

if any of the sites represent @ naturally occurring

Association V,

Assuciation Vie Eucalyptus brachycalyx

Vegetation Association V1 includes Eucalyptus

brachyeniyx as the important upper-slorey species,

together with a nange of low chenopods similar to

that of Association V. The floristic similarity results

in considerabje overlap between these two

associations on the ordination (Pig. 5), [fis must

common towards the coast, occurring in the

southern central and western part of the study area

on shallower soils than Association LV,

Axsocivfien¥i; Eucalyptus yumbarrana-Triodia

irtitans

It is probable that kvcalppius yunebarrane was

previousty recorded as Eucalyptus socials, and that

the £. socutis/Melaieuca uncinata sandhill

association of Specht (1972) corresponds to

Association VII, as does Margules & Nicholls’

Community [af their Melalence adnvta is re-

identified as M. eleuthernstachpa.

Vegetation Association. VII is restricted to the

sand-dunes of the northern, low rainfall part of the

study area. The specles characterising it are entirely

different from those found in the other vegetation

associations, and include many species with

distributions in higher rainfall parts of the state,

such as Lomendra leucocephala and Guhriia

lanigera (Jessop & Toelken 1986). This major

Flonstic dasjunctinn corresponds to a similar

observation in north-western Victoria, where the

most marked discomtinuiy of mallee vegetation was

between a Aucelypius incrossata/Albbertiad

Lepitospermym association ‘with “southern

temperate” affinities, and all the other associations,

m which semi-succulent herbs, especially

chenopods, were prominenr (Noy-Meir 1971).

The disjunction observed is related to Le geology,

since the heavier soils of the exposed Woorinen

Formation present in the infer-dune corridors

lypically support vegetation of Associations 4, 11

or I, The combined effect of coarse texture and

54 F, TIVER, A. D, SPARROW & R. T. LANGE

TOPOGRAPHY RAINFALL

SOIL DEPTH SOM TEXTURE

VEGETATION TYPE

FREQUENCY

PERCENT AGE

Vil

Fig.6. Histograms summarising observed environmental patterns for each of the vegetation types. For topography:

P, plain; R, rise; F, dune flank; S, sandhill, For soil texture: L, loam; SL, sandy loam; LS, loamy sand; S, sand,

increased depth of the sands of the dunes

presumably creates a more favourable water

relations environment, which allows the presence

of a suite of species normally requiring higher

rainfall, There is a higher percolation rate and

deeper penetration of rainfall into sands than into

clays, while evaporation dries the surface soil to a

similar depth regardless of texture, so that at the

end of along dry period, there would be more water

available in the root zone of sandy soils than in clays

(Walter & Stadelmann 1974). Clays also have a

higher water holding capacity due to their smaller

pore size (Ball 1986). This means that a larger

amount of rainfall is required To bring air-dry clay

to a range where water is available to plants than

for sands.

Discussion

There is a clear vegetational discontinuity

between the two predominating geological

formations of north-western Eyre Penmsula, the

siliceous Moornaba sand dunes and the caleareous

Woorinen soils. Although it is possible to recognise

plant associations within the vegetation of the

Woorinen Formation and to relate their

distributions to environmental features in a general

way, sharp discontinuities do not exist. The strong

gradation of associations is typical of the mallee

on Woorinen Formation and its equivalents across

the whole of S. Aust. (Sparrow unpublished data),

There are two possible interpretations of these

observations, Firstly, since there is no strong

environmental discontinuity within the Woorinen

Formation, the gradation may reflect combinations

of species with independent environmental

responses, which, further confounded by

disturbance, results in almost continuous ecotones.

Alternatively, the gradation of mallee associations

could be caused by some degree of independence

of the mallee and understorey strata, The mallees,

VEGETATION OF NORTH-WEST EYRE PENINSULA 55

Fig. 7 Distribution maps of the seven major vegetation types. Shading represents uncleared vegetation and the broken

line marks the Eyre Highway.

56 F. TIVER, A. D, SPARROW & R. T. LANGE

TaBLe 3: Full species list (nomenclature following Jessop & Toelken 1986) showing occurrence in the seven major

vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped in the final

column. Introduced species are indicated with an asterisk.

Species Vegetation Type

i IL Ill IV Vv VI VII“ Other

Acacia calamifolia + + + - - = + 7

Sweet ex Lindley

Acacia colletioides + + + _ + + + os:

Benth.

Acacia hakeoides + - + = + + - ~

Cunn. ex Benth.

Acacia ligulata + = = - _ = + +

Cunn, ex Benth.

Acacia merralli + + + + + + + -

F. Muell,

Acacia notabilis + = = = 2 _ _ .

F. Muell.

Acacia oswaldii + + + = + = = 7

F. Muell.

Acacia rigens ~ = = - = _ 4 _

Cunn, ex Don

Acacia sclerophylla . + + = + = _ =

Lindley

Acacia spinescens = - - - + = = =

Benth.

Alyxia buxifolia = = - _ + _ _ _

R. Br,

Amyema melaleucae ~ — = pa ve “3 = +

(Miq.) Tieghem

Atriplex acutibractea + + + + 4 + + +

R. Anderson

Atriplex stipitata = + = e + 2 = =

Benth.

Atriplex vesicaria = + 4 + + A _ a

Heward ex Benth.

Beyeria lechenauttii + - = = = _ _ e

(DC.) Baillon

Beyeria opaca = - = + + _ rs =

F. Muell.

Billardiera cymosa = a _ + = 2s = _

F. Muell,

Boronia coerulescens = = = = cs - = _

F. Muell.

Callitris preissii 3 + = + = = -

ssp. preissit

Mig.

Callitris preissi ~ = - - { - - n

SSp. verrucosa

(Cunn, ex Endl.) J. Garden

Carpobrotus rossti - - = + + + > =

(Haw.) Schwantes

*Carrichtera annua - + - + + + = +

{L.) DC.

Cassia nemophila var coriacea + + a: 7 + _ “= &

{Benth.) Symon

Cassia nemophila var nemophila + + = + - _ 4 >

Cunn. ex J. Vogel

Cassia nemophila var platypoda = + = + fr = + +

(R. Br.) Benth.

Cassytha glabella + = = 4 J _ in! *

R. Br.

Cassytha melantha + = + + + + + _

R, Br.

Casuarina cristata = = = L + = = +

Mig.

Chenopodium desertorum — = = = = = _ _

(J. Black) J. Black

Cratystylis conocephala + a ; “ 4 , : P

(F. Muell.) S. Moore

VEGETATION OF NORTH-WEST EYRE PENINSULA 37

TABLE 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven

major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped

in the final column. Introduced species are indicated with an asterisk.

Species Vegetation Type

I Il lt lV Vv val Vil Other

Crypiandra leucophracta = rs -_ = _ _ + _

Schldl.

Cryptandra tomentosa = = 7 = 2 s. + i.

Lindley

Danthonia spp. 4 + L. aS _ = 1 7

Lam, & DC,

Daviesia asperula - = = = - - + _

Crisp

Daviesia benthamii = : = 2 = - + =

Meissner

Dianella revoluta + + + + + + + ¥

R. Br.

Dillwynia uncinata = = = = = = + _§

(Turez.) J, Black

Dodonaea bursariifolia + - > ~ 2 5 + .

F. Muell,

Dodonaea stenozyga - 2 + * = = = +

F. Muell,

Dodonaea viscosa + = = = 7 5 rn =

ssp. angustissima

{DC.) J.G. West

Enchylaena tomentosa + i + + + + + +

R. Br.

Eremophila crassifolia — - + + — _ + _

(F. Muell.} F. Mucll.

Eremophila deserti + = is = + 23 = +

(Cunn. ex Benth.) Chinn

Eremophila glabra + + + + — + + _

(R. Br.) Ostenf.

Eremophila scoparia + ef + + + + - 3

(R. Br) F. Muell.

Eremoaphila weldii + “h + + i‘ + + _

F. Muell.

Eucalyptus brachycalyx/concinna + + + + + + + +

Blakely/Maiden & Blakely

Eucalyptus dumosa + ' + 4s + + + +

Cunn. ex Schauer

Eucalyptus foecunda = = _ = _ - + _

Schauer

Eucalyptus gracilis + + + ef + 4 < +

F. Muell.

Eucalyptus incrassate _ + = = = s + =

Labill.

Eucalyptus oleosa 4 + + + + + + 4

F. Muell. ex Mig.

Eucalyptus socialis an — 4 = _ _ . _

F. Muell. ex Miq

Eucalyptus striaticalyx ss + a _ - mn _

W. Fitzg.

Eucalyptus yumbarrana s = + = - s + +

Boomsma

Eutaxia microphylla _ = = — ” = + -

(R Br. in WT. Aiton) J. Black

Execarpos aphyllus + 4 + + ' 4 + +

R. Br.

Exocarpos sparteus = = = - - - + =

R. Br.

Gehnia lanigera - - \ = _ = + _

(R. Br.) Benth.

Geijera linearifolia + + + + + 4 + +

(DC.) J. Black

Goodenia varia — = = = = = 4 -

R. Br.

F. TIVER, A. D. SPARROW & R, T. LANGE

Tae_t 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven

major vegetation types. Occurrence in sites that could not be assigned to a major vegetation type are grouped

in the final column. Introduced species are indicated with an asterisk.

Species

Grammosolen truncatus

(Ising) Haegi

Grevillea huegelii

Meissner

Hakea francisiana

Maconachie

flalgania cyaneu

Lindley

Halgania lavandulacea

Endl.

Helichrysum bilobum

Wakef,

Hybanthus floribundus

(Lindley) F, Muell.

Txiolaena pluriseta

Haegi

Lasiopetalum behrii

F. Muell.

Lepidosperma laterale

R. Br.

Leptospermum coriaceum

(F. Muell.) Cheel

Leucopogon cordifolius

Lindley

Logania nuda

F. Muell.

Lomandra collina

(R. Br.) Ewart

Lomandra effusa

(Lindley) Ewart

Lomandra leucocephala

(R. Br.) Ewart

*Lycium ferocissimum

Miers

Maireana appressa

Paul G, Wilson

Matreana brevifolia

(R. Br.) Paul G. Wilson

Maireana enchylaenoides

(F. Muell.) Paul G. Wilson

Maireana erioclada

{Benth.) Paul G, Wilson

Maitreana pentairopis

(Tate) Paul G. Wilson

Maireana sedifolia

(F. Muell.) Paul G, Wilson

Maireana trichoptera

(J, Black) Paul G. Wilson

Melaleuca acuminata

F. Muell.

Meluleuca eleutherostachya

F. Muell.

Melaleuca lanceolata

Otto

Melaleuca pauperiflora

. Meull.

Microcybe muttiflora

Turez.

Muehlenbeckia gunnii

(Hook. f.) Walp.)

Myoporum platycarpum

R. Br.

Olearia brachyphylla

(F. Muell. ex Sonder) Wakef.

Vegetation Type

af Il Ill IV Vv VI VIL

VEGETATION OF NORTH-WEST EYRE PENINSULA

TABLE 3 cont. Full species list (nomenclature following Jessop & Toelken 1986) showing occurence in the seven

major vegetation types. Occurrence in sites that could not be assigned to a major vegelation type are grouped

in the final column, Introduced species are indicated with an asterisk,

Species

Olearia floribunda

(Hook. f.) Benth.

Olearia magniflora

Vegetation Type

I II Ill IV

(F, Muell.) F. Muell, ex Benth.)

Olearia muelleri

(Sonder) Benth,

Olearia pimetioides

(DC.) Benth.

Pimelea microcephala

R. Br.

Pittosporum phylliraeoides

DC.

Podolepis capillaris

(Steetz) Diels

Ptilotus obovatus

(Gaudich.) F. Muell,

Ptilotus spathulatus

(R. Br.) Poiret

Pultenaea elachista

(F. Muell.) Crisp

Rhagodia crussifolia

R. Br.

Rkagodia parabolica

R. Br.

Rhagodia preissii

Mig.

Rhagodia spinescens

var. deltophylla

F. Muell.

*Salsola kali

Santalum acuminatum

(R. Br.) A. DC.

Scaevola bursariifolia

J. Black

Scaevala spinescens

R. Br.

Schoenus subaphyllus

Kiik

Sclerolaena diacantha

(Nees) Benth.

Sclerolaena obliquicuspis

(R. Anderson) Ulbr.

Sclerolaena uniflora

. Br.

Sida corrugata

Lindley

“Solanum coactiliferum

J. Black

Stelligera endecaspinis

A. J. Scott

Stipa spp.

Templetonia battii

F. Muell.

Threlkeldia diffusa

R. Br.

Triodia irritans

R. Br.

Velleia connata

F. Muell

Vittadinia dissecta

(Benth.) N. Burb.

+ + + + +

= - - = +

+ + + = +

mJ - 4 a 4,

~ _- — — _

+ + - = +

= = 2 b =

+ + + + +

- - - - +

+ + = + +

+ + = - +

+ - - + +

+ + + + +

—- - - + +

+ + + - +

+ + a + +

+ + + + +

= + 4 = _

+ = = =

it =. + + =

+ + + + +

_ — — +

+ d + + +

+ - + - —

VI VII Other

+ + =

+ = =

+ + -

= 4 =

+ + +

s 4 #

“ + Pa

+ + +

F + +

- - +

+ + -

= - +

a + s

+ - +

- - +

= +

+ - =

+ + +

+ = «=

+ ¢ +

_ + =

60 J_ TIVER, A. D. SPARROW & RK. 1, LANGE

TABLE 3 cout, Full species list (nomenclature following Jessop & Toclken 1986) showing oveurettce in the sever

mgjor vegelatton types. Occurrence in sites that could not he assigned fo a mu/jor veperation type are grouped

in the final coluinn, Intraduced species are indicated with an asterisk,

Species

Hestringia rigida =

R '

. Br.

Zygophylluny anienaphitum

F. Muell,

Zyguphvllum apiculatum ‘

F Muell.

Zypophyllun) aurantiacuin/ovatunt 4

(Lindley) F. Muell./Ewart & ). White

Zynophyllunt glaucum +

F, Muell, ~

Verctation Type

I i

nt 1v V vi VIL Other

' ' ‘ +

+ - \ - -

' = - ca ~

+ ‘ -

_ + 4 + ‘ + =

being very Jong-lived plants, may be indicating same

historical influence, while the shorter-lived

understorey species reflect current environmental

conditions, Lange & Nicolson (1982) report mallees

tecotding former extensions of a palacosol, and

TWidale & Campbell (1985) and Short, ef al. (1986)

provide ample evidence of dynamic changes in the

surface geology of north-western Kyre Peninsula

during the Holocene.

A further issue is the occurrence of uncommon

yepelalion associations in the area. One example is

the Callitris preissii association represented by a

single site on the dendrogram; another, towards the

northern end ol the Mudamuckla transect and

removed fram analysis due to disturbance, ~was

unique lor the presence of Casuarina cristata. A

regular sampling strategy inevitably Ieads 1a the

omission of rare vegetation types, su there are

undoubtedly other uncommon associations in the

area,

Acknowledgments

We would like to rhank the staf? of the Vegetation

Retention Branch of the S. Aust, Department of

Environinent and Planning for practical assistance

during the course of the study, The work was

carried out as: part of the Middleback Ficld Centre

programme for 1987,

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TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 113, PART 2

TIMING, EXTENT AND CHARACTER OF LATE, CAINOZOIC FAULTING

ON THE EASTERN MARGIN OF THE MT LOFTY RANGES,

SOUTH AUSTRALIA.

BY R. P, BOURMAN* & J, M. LINDSAY}

Summary

Stream erosion on the eastern flank of the Mt Lofty Ranges has exposed a reverse fault near

Cambrai on the Milendella escarpment, indicating a compressive component to the uplift of the

ranges. Cambrian rocks have been thrust over Pleistocene fanglomerates and Miocene limestone

has been dragged up along the fault zone to an elevation of 160 m asl. This fault may indicate the

reactivation of a compressive Palaeozoic fault during the Cainozoic. Diagnostic foraminifera have

been identified in samples of the limestone, which is of the Early Miocene Mannum Formation

(about 20 Ma), adding support to the unpublished view that the position of the limestone on the

escarpment is due to tectonic uplift of 60-90 m since the Miocene.

KEY WORDS: Tertiary limestone, foraminifera, reverse faulting, Mt Lofty Ranges, South

Australia.

TIMING, EXTENT AND CHARACTER OF LATE, CAINOZOIC FAULTING ON

THE EASTERN MARGIN OF THE MT LOFTY RANGES, SOUTH AUSTRALIA,

by R. P, BouRMAN* & J, M, LINDSAYT

Summary

Bouwmas, R. P. & Lainpsay, J. M. (1989) Timing, extent and character of late Cainozow faulting on the

ewslern wnarein of the Mt Lofty Ranyes, South Australia. Trans. RX. Soc. 8. Aust. 13 ), 63-67, 31 May, 1989.

Stream erosion on the eastern flank of the Mt Lofty Ranges has exposed a reverse fault near Cambrai

an the Milendella escarpment, indivating «compressive component to the uplift of the ranges. Cumbrian

rocks have been thrusi over Pleistocene fanglomerates and Miocene limestone has been dragged. up along

ure fault zome to an elevation of 160m asl, This full may indicate the reactivalion of a compressive Palaeozoic

fault during the Cainozoic., Diagnostic foraminifera have been identified in samples af the limestone, which

is of che Early Miocene Mannom Formation (about 20 Ma), adding support to the unpublished view chal

the positica of the limestone on the escarpment is due to tectonic uplift of 60-90 m since the Miacen&

Key Worp® Tertiary limestone, foramunifera, reverse faulting, Mi Lofty Ranges, South Australia,

Introduction

The fault origin of the ML Lolty Ranges hus long

been discussed (Benson 1911; Fenner 1930; Sprise

1945) and the majorivy of these workers considered

that the faults were normal types. Glaessner (1953)

presented a model for the tectonic evolution of the

Mc Lofty Ranges, which involved basement complex

flexuring, that resulted in reverse faulting on the

margins of the ranges and norma! faulting within

them, Campana (1955) preferred to explain the

origin of rhe M1 Lofty Ranges by compressional

doming, which led to minor fault disruption on

their margins, Gibson (1963) noted that the

Clarendon-Ochre Cove Fault displays a reverse habit

where intersected by tunnelling operations during

the construction of the Clarendon-Happy Valley

pipeline, Recently Wellman & Greenhalgh (1988)

favourtd the view thal compressive forces have heen

important in the formation of the Mt Lofty Ranges,

A re-evaluation of exposed fault contacts is required

to resolve the nature of the faulting responsible for

the uplift of the Mr Lofty Ranges.

The Milendella Fault

There is clear evidence. of reverse lauliing on the

eastern escarpment of the Mt Lofty Ranges as first

reported by Mills (1965)2, and it may indicate the

* Gibson. A. A. (1963) Final geological report on tye

Happy Valley Tunnels Project. S. Aust. Dept of Mines

report 37/92 (unpubl)

2 Mills, K. J. (0965) “The stnictural peredlogy of ihe

Milendella area of South Australia’. Ph.D. Thesis,

University of Adelaide (unpubl)

* Depr of Soi) Science, Waite Insitute, University of

Adelaide, Urrbrac, S. Aust, 5064,

| Cieolowival Survey, Soil Austvatian Departinent of

Mines & Bnergy, Box 151, Eastwood Px. §. Aust, $063,

re-activation of Palaeozoic ihrusi faulis, Stream

exposures of the Milendella Fault occur west of

Cambrai on ithe escarpment (Angaston MR

390/630) at 160 m asl (Fig. 1). The fault here |s of

reverse type with brecciated Cambrian schists of the

Kanmantoo Group of metasedimentary rocks thrust

over Pleistocene fanglomerates. Two pods of

Tertiary limestone have been dragged up at a high

angle in the fault zone, which dips to the west aL

45° (Fig. 2). The Pleistocene fanglomerates affected

hy the faulting are similar to Pleistocene

fanglomerates tilted into a vertical position by

movemen| along the Willinua Fault as Sellicks

Beach (May & Bourman 1984), The sediments al

Sellicks Beach were related to the Ochre Cove

Formation, which May & Bourman (1984) regarded

as of Middle Pleistocene age. If rhe uwo deposits

are correlatives then faulting on the Milendella

Scarp probably accurred until the late Middle

Picistocene or the early Late Pleisiocene. No

unequivocal evidence of dislocation of Late

Pleistocéne sediments in the area was noted.

It is suggested that during faulting, the

Pleistocene fanglomerates were folded, and

breeciated basemenr rocks broke from the hanging

wall to tuinble on top of the fanglomerates. Calcrete

mantles the convoluted fanglomerates, dying out

al the laulr zone, so that the culerete may have

derived fram solution and reprecipilation of the

Tertiary limestones dragged to the surface along the

fault zone and draped over the top of the

fanglomerates. Calcareous pisoliths in this calerere

are lypically laminaled around votes of locally

derived bedrock. This supports the interpretation

of in sine pisolith formation from the redistribuuon

of Tertiary limestone, Alternatively, if the calerete

has stratigraphic significance (eg. the upper Middle

Pleistocene Bakara calerete of Firman 1967) then

R. P. BOURMAN & J. M. LINDSAY

\ : ; E

= ts 5 a

z 5 z =

Ww = re Oo

=; ie < [oa

a; = fs)

Ww =

= oO Cambrai

= 4,

e @

N < a)

Qa A I e

\ Ss / ©) Walker Flat

& i

fa a

\ yu

DELAIDE ‘\. o

6 a Ms 3 MANNUM Teale Fiat

SY, vo fe)

35° § & Ay , Caloote >

& © ; =

ri Sy aS J <

Sr Sh. &

s </ + ~ .\ OCMURRAY BRIDGE

= Oo” Ef os "

= A SK

. v/ &

A wf O

% 2 v OTAILEM BEND

« >

Sy x

ed WESTERN MARGIN OF, FS

MURRAY BASIN Ke Lake Alexandrina >

, =

« VICTOR -% x,

yy" HARBOR 97 CE

we SS

Lake

, 10 20 30 40 50 Albert

KILOMETRES

Fig. | Darnton map showing major faulp zones; | site of fault exposure near Cambrai and © Lepidoeyellna-bearing

limestone in the Bremer valley.

the relationships may indicate fault disruption of

the calcrete in the Late Pleistocene. It is not possible

to determine the amount of offset accomplished

during this postulated Pleistocene phase of faulting

without detailed drilling. However, Mills (1965)?

shows a thickness of some 30 m of Pleistocene

sediments in section at the scarp (Fig. 3), arid as

the limestone must have been dragged up trom

below this level, this suggests an offset of less than

30 m. Dislocation of Early Pleistocene marine

sediments across the Willunga Fault was reported

by May & Bourman (1984) illustrating similar post-

Farly Pleistocene faulting of about 40 m on the

opposite side of the ranges. It is interesting to nole

that although there is clear evidence of geologically

recent faulting on the eastern margin of the Mt

Lofty Ranges, there is no pronounced fault scarp

associated with it, which may suggest thal the scarp

formed largely in brecciated rock and rapidly

degraded.

Mills (1965)* suggested that displacement on the

Milendella Fault since “pre-Tertiary peneplanation”

was approximately 335-366 m, and he attributed

250 m of this to the Early Tertiary to Miocene, and

60-90 m to post-Miocene faulting. He noted

Miocene sediments (?Morgan Limestone) at

elevations up to 14() m on the scarp, correlated them

with Miocene strata under the Murray Surface and

LATE CAINOZOIC FAULTING IN THE MOUNT LOFTY RANGES 6

Fig. 2. Fault exposure near Cambrai. View south along the eastern escarpment of the Mt Lofty Ranges. Cambrian

schists (K) of the Kanmantoo Group of metasedimentary rocks have been thrust over (?) Middle Pleistocene

fanglomerates (F). Light coloured calcrete (C) mantles the fanglomerates and two pods of Miocene limestone (L)

occur in the fault zone (arrowed), which dips to the west at approximately 45°

attributed the difference in elevation to tectonic

dislocation. Twidale & Bourne (1975) did not deny

recent fault activity but preferred to interpret the

perched limestone as essentially due to Tertiary

eustatic influences, with the limestone under the

Murray Basin being the older, lower Mannum

Formation and that on the scarp as the younger and

higher Morgan Limestone. Thus they discounted the

view that there had been 60-90 m of dislocation

on the Milendella Fault since the Miocene and

argued for an erosional origin for the lower part

of the scarp. The resolution of this conflict may be

achieved only by an accurate age determination of

the Miocene limestones on the Milendella scarp to

compare with those from the Murray Basin.

Palaeontological work reported here demonstrates

that the limestone on the scarp, at least in this

locality, is from the lower part of the Mannum

Formation.

Age and facies of the Tertiary limestone in the fault

zone

Four samples have been examined from the two

pods of Tertiary limestone dragged up in the fault

zone.

Portions were crushed gently in a pestle and

mortar to disaggregate partially, boiled in a dilute

solution of sodium bicarbonate with a drop of

detergent, washed free of mud through a very fine

sieve, dried, and picked for foraminifera under a

binocular microscope. Slides with the foraminiferal

microfaunas are stored in the Biostratigraphy

Branch collections of the S. Aust. Department of

Mines and Energy.

Lithology

All four samples comprise quartzose fossiliferous

sandy limestone (biogenic calcarenite), The rocks

are cream to brown, hard to friable, somewhat

recrystallised and, in part, leached. Quartz and

minor lithic sand content is considerable and poorly

sorted, ranging from very fine grained to very coarse

grained. Quartz and lithic grains up to 10 mm are

present. There are occasional pale green glauconite

infillings, somewhat oxidised.

Barnacle plates are frequent and notable;

echinoid, molluscan and bryozoal fragments are

frequent to common; foraminifera are common but

mostly recrystallised; ostracods, Ditrupa tubes, fish

and decapod fragments, and algal oncolites are rare.

66 R. P. BOURMAN & J. M. LINDSAY

EVANS TRIG.

age <- 397m

300

METAMORPHIC ROCKS

Height in metres

Pleistocene alluvium

100 . .

Miocene limestone

200 Displaced Miocene limestone blocks

Recent alluvium

Eocene lignitic sands

SEA LEVEL

Kilometres

Fig. 3. Cross section through the eastern margin of the Mt Lofty Ranges to the Murray Basin near Cambrai (after

Mills 19652).

Discussion of foraminifera, age and correlation

The four samples have much in common and the

microfaunas will be considered together (Table 1).

Numerically, all the microfaunas are dominated by

Elphidium chapmani and Notorotalia cf.

miocenica, but these are not the most

biostratigraphically significant.

The overlap of the top of the range of

Sherbornina atkinsoni and the bottom of the range

of Operculina victoriensis in S. Aust. indicates an

early Longfordian, Early Miocene age (Wade 1964;

Lindsay 1985). For the Murray Basin proper,

Ludbrook (1961) only overlapped these ranges in

lower Mannum Formation, and this has been borne

out by subsequent experience. Other foraminifera

present in the four samples are generally consistent

with Longfordian Stage and Early Miocene age.

Notorotalia miocenica has been scarcely recorded

previously from S. Aust., so the slightly earlier age

here of a related form, compared with the range

in Victoria (Carter 1964) is not regarded as

incongruous. The solitary and small planktonic

specimen identified as Globigerina sp. is

non-diagnostic.

TABLE 1, Foraminifera: Table of occurrences of selected species.

Foraminifera Samples

F23/83 F24/83 F2/88 F3/88

Discorbis sp. from group D. balcombensis

Chapman, Parr & Collins -D. cycloclypeus

Howchin & Parr R R F F

Ammonia sp. - Vv - R

Pararotalia verriculata (Howchin & Parr) Vv Vv - Vv

Sherbornina atkinsoni Chapman - - - Vv

Elphidium chapmani Cushman ce, Cc A A

E. crassatum Cushman F R Cc F

E. parri Cushman = - - Vv

Notorotalia cf. miocenica (Cushman) F F Cc A

Operculina victoriensis (Chapman & Parr) Vv R Vv R?

Globigerina sp. - - - Vv

Amphistegina sp. _~ — R

Cibicides vortex Dorreen - - Vv -

Planolinderina plana (Heron-Allen & Earland) - - Vv -

(Abundances: V = very rare; only one or two specimens recovered in thorough picking. R = rare; 3-5. F = frequent;

6-10. C = common; 11-25. A = abundant; more than 25).

LATF CATNOZOIC FAULTING IN THE MOUNT LOFTY RANGES 67

The lithology of the saniples is consistent with,

but not unique to, Mannum Formation as described

by Ludbrook (1961) and observed subsequently, This

Early Miocene Mannum Formation in the fault

fone is distinctly older than the clevated,

Lepidecyelina-bearing, Morgan limestone reported

from the Bremer valley east of Kanmantoo (Lindsay

1986"); about 20 Ma compared with 16 Ma (Lindsay

1988),

Environment af deposition

The variety and nature of the fauna indicate

marine conditions. ‘hree lines of evidence suggest

& shallow-marine, littoral-nerific environment: the

notable frequency of barnacle remains, the

restricted Alphidium-rich foraminiferal facies

almost lacking planktonic forms, and the quartzose

and lithic clastic component of the limestone, with

gravel-sived grains, This corroborates the

sedimentological evidence of Mills (L965)*, who

proposed a near-shore environment for the

deposition of the limestone,

Conclusion

The evidence presented here demonstrates that

most Of the elevational difference between the

3 Lindsay, J. M, (1986). New record and tectonic

implications of a high level Lepivocyedinabearing

linestorig, Bremer Valley, casrer) Mount Loft y Ranges.

S.A. Dept of Mines and Energy cepore 86/66 (anpubl,),

Muannum Formation limestones on (he escarpment.

near Canibrai, and under the Murray Basin can be

allributed to tectonic dislocation in the past 20 Mu

as originally suggested by Mills (1965)2,

Furthermore, it has also been demonstrated thal

there has been at least 100 sm (and prabably much

more) of tectoni¢ offsel of the early Middle

Miocene / epidocyclina-bearing Morgan Limestone

that occurs in the Bremer Valley (lindsay 1986)4

in the past 16 Ma, Consequently, evidence of

considerable tectonic dislocation of Tertiary

limestones on the eastern side of the Mr Lofty

Ranges, separated by 50 km, together with faulted

Pleistocene sediments provides strong support for

pronounced tectonism since the Middle Miocene.

Without denying the evidence for eustalic influences

in the Tertiary (eg. Vail & Hardenbol 1979) iris clear

thal post Middle Miocene faulting is far more

significant in the evolution of the east Mt Lofty

Ranges than suggested by Twidale & Bourne (1975),

Furthermore, the clear exposure of a reverse fault

on the eastern margin of the Mt Lofty Ranges

supports the notion of a compressive component

to the forces involved in the Tertiary uplift of the

eastern Ranges

Ackauwledgments

This paper is publisted with permission of the

Director-General of Mines and Energy, South

Australia,

References

Besson, WN, (1911) Notes descriptive of a stereoursm

at rene tale Lofty Ranges, Trans. R. Soc. S. Aust. 35,

103-111,

CAMPANA, B. (1955) The structure of the Eastern South

Austrahan Ranges; the Mount Lofty-Olury Ane. Gen.

Soe. Aust, 2, 47-61.

tArtrR, A, N. (1964) ‘Tertiary loraminilera trom

Gippsland, Vicloria, and their stratizraphical

significance. Mea. geol Surv. ier. 23.

Penner, C, (3930) The major structural and

physiographic features of South Australia. Trans. R.

Soe. 8. Aust. 54, 1-36.

Firman, J.B. (1967) Stratigraphy of Lare Cainozoic

deposits in South Australia, 7bid, 91, 165-178,

Gy) aessvpr, M, Fo (1953) Some problems of Tertiary

geology in southern Australia, Jaun Proc. R. Sec,

NSM RT, 31-45,

Linpsay, MA (W985) Aspeers vot South Awatratian

‘Tertiary foraminiferal biostraligraphy, with emphasis on

studies of Mussilina and Subbatina. Spee. Publ. S,

Aust, Dept Mines and Energy, §, 187-232,

LuprRook. 8. A. (1961) Stratigraphy of (he Murray Basin

in South Australia. Bull, zeod Surv, S. Aust, 36,

May, R, 1, & BouRMAn, R. P (1984) Coastal

landslumping in Pleistocene sediments at Sellicks Beach,

South Austratia, Trans KR. Soe. S dust. 108, 85-94

Spricic, KR, ©, (1945) Some aspects af the geomorphalaay

of portion of the Mount Lofty Ranges. Jhid. 64,

277-302.

TWibale, C. Re & Rovrnwe, FA, (1975)

Cieomorphological eyolution of part of the eastern

Mount Lofty Ranges, South Australia, shied. 99,

197-209,

Van, PR, & Harpenren, J, (1979) Sea-level changes

during the Tertiary. Oceanus, 22, 71-79.

Want, M. (1964) Application of the lijeage concept ta

biostratigraphic yoning based on planktonic

foraminifera, Adieropyleontalagy WW, 273-290,

Wertman, Po & GREENHALON, S. A. (1988) Flinders

Mount Lolly Ranges, South Australia, theic uplift,

crosion aud relationship te ccustal structure. Zraets, &

Soc, S. elust. 192, W-19.

INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF,

SOUTH AUSTRALIA

BY R. C. AINSLIE, D. A. JOHNSTON & E. W. OFFLER*

Summary

The fauna of the intertidal mudflats of northern Spencer Gulf was monitored between 1982 and

1986. Two main habitats were identified, the mid intertidal zone, and the seagrass fringing low

intertidal zone. These habitats supported quantitatively differing faunas.

There was no evidence of the species impoverishment reported elsewhere for the epizoic fauna of

this region. Comparison of the characteristics of the fauna with that of another negative estuary, the

Port River in Gulf St Vincent, has provided the basis of a monitoring programme to assess potential

environmental stress imposed by power station development in northern Spencer Gulf.

KEY WORDS: Intertidal habitats, benthic fauna, seagrass, Spencer Gulf.

INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF, SOUTH

AUSTRALIA

by R.C AINSLIE. Dh A. JOHNSTON & EL W. OFFLER*

Summary

AtNsLip, RUC. JoHnstTon, DA, & OFFLeR, E, W, (1989) Intertidal communities of northern Spencer

Goll, South Australia. Trans, R. Soc. S, Aust, 113( ), 69-83, 31 May, 1989.

The fauna of the intertidal mudflats of northern Spencer Gulf-was moniored between 1982 and 1986.

Two main hahitats were identified, the mid intertidal zone, and rhe seagrass fringing low intertidal zone:

These habitats supported quantitatively differing faunas.

There was no evidence of the species impoverishment reported elsewhere for the epiauic fuuna of thin

region. Cormparisan of the characteristics of the fauna with that of another megarive estuary, the Port River

in Gulf St Vincent, has provided the basis of a monitoring programme to assess potential environmental

stress imposed by power station development in northero Spencer Gulf,

Key Worps: Intertidal habitals, benthic fauna, seagrass, Spencer Gulf.

Intraduction

Northern Spencer Gulf can be defined as thar

portion of Spencer Gulf, South Australia, north of

the line from Ward Spit to Lowly Point, i.e. north

of 33°S (Stefanson 1977). Tt is a unique, narrow

body of water, projecting inland some 200km from

the Southern Qcean coast (Fig. 0),

The waters of northern Spencer Gulf are

characterised by high summer salinities, up to 48

(Nones & Lennon 1986) and high summer

temperatures with wide seasonal fluctuations,

11-25°C (Johnson 19811), Due to bigh evaporation,

and low fresh water inflow to the northern Gulf,

there 15 a longitudinal salinity gradient of about four

in 20kin. The region has been described as a

negative estuary,

A limited number of studies has been carried out

on the marine communities of the “middle Gulf”

(SEA 19812; Ward & Young, 1982, §3) and concur-

Johnson, J,E. (1981) Hydrological Data for Upper

Spencer Gulf 1975=1978. Fisheries Res. Paper No. 3,

1-30, (Dept. fisheries, Adelaide), unpubl.

Social and Ecological Assessment (J9R1) Vrati

Environmental Impacr Statement for port and terminal

facilities at Stony Point, South Australia. Prepared for

SANTOS, unpubl.

Kinhill (1987) Lipper Spencer Gulf Inlertidiat Survey,

Final Repert, May 1987. unpubl.

* Electricity Trust of South Australia (1977) Northero

Power Station Enyironmental Impaet Statement. July:

1977, unpubl.

Electricity Trust of South Australia (L985) Northern

Power Station Environmental Impact Statement, Aupust

1985. Prepared by Kinhill Steartis, unpubl.

Environmental Sciemes an Tnineering Branch,

Electricity Trust of Sourh Ausiralia, PO. Box. fi,

Eastwood &, Aust. S063

rently with this study, Kinhill (1987)9 have examined

intertidal communities wt Ward Spit and Lowly

Point. Shepherd (1983) provides the only published

description of benthic communities of the narrow

reaches of the northern Gulf. Shepherd found a

marked reduction in species number in the epizaic

‘communities on Pinna bicolor in northern Spencer

Gulf compared to that of the same habitat in Gulf

St Vincent, raising the possibility of a general

species impoverishment in the hyper-saline waters

of northern Spencer Crulf, He emphasised rhe need

for further detailed study of other “component

parts” of the biological system to determine its

capacity fo accept additional stresses of industrial

wastes and discharges,

Singe the mid 1950's Playford Power Station las

been operating on the castern shore of the northern

Gulf, discharging warmed cooling water to the Gulf.

The discharge from Playford Power Station,

operaling at it’s Maximum nominal capacity of

330MW, resulted in surface water temperatures

ubout 6°C wbove ambient near the power stalion,

with only occasional incursions of warmed, watet

(about 3° above ambient) to nearby intertidal

zones (ETSA 19774). In 1977 and 1985 rhe

Electricity Trust of South Australia issued

Environmental Impact Statements for three further

power station units, with a total gencrating capacily

of TSOMW, to be built in this area, This

Uevelopment forms the Northern Power Station.

Thermal discharges were predicted (a more than

uouble {ETSA 19774, 19855),

The effects of warm water discharge fram power

stations on benthic fauna have been the subject at

a Tiumber of reviews (Coutant & Talmage 1975;

Talmage & Coulant 1980; Craven e/ af 1983;

Langford 1983). Effects vary from site to sile asa

aL A.C. ATNSLIE, DD A. JOHNSTON & E, Wo OFMLER

function af the climatic, hydrological and biolugicul

features (Crema & Paglial 1980}. Greatest effects

are Ghserved jiear the outfall, ac the hotcest times

of the year (Langford 1983) and include changes

in the biological characteristics of the benthic

tummunilies within the zone of influence of the

warm water plume (Robinson e7 af 1983; Robinson

1985; ‘Thomas ef a/ (986), In particular, Barber

& Spenver (1984) and Thomas ef af (1986) have

described the decline in populations of particular

bivalve mullusce species and the establishment of

dense popwations of opportunistic worm species

as characteristic of \hermal disturbance in soft

substrates,

The present study began in 1982, three years prior

tu the commencement of operation of the new

power station. Intermittent operation of the first

250M W unit commenced in the suntmer of [9RS,

with SOMW operating by summer 1986. The aim

of the study was lo investigate spatial and Lemporatl

nallerns of the benthic communities in the negative

estuary of nocthern Spencer Gulf. It was anticipated

that detailed descriptions of these communities

would identify whether or not there were any

biplogical impacts from the existing power station

development (and ¢arly operation of the new

development), and establish bascline data for long

Lerin assessment of the eflects of increased thermal

discharge; This study concentrated on the intertidal

and shallow sublittoral fringe, a5 the ouly other

published South Ausiralian study on the impacts

uf thermal discharge on the marine environment

had deseribed well defined paticrns of change in

the communities af this yone (Thomas ef a! 1986),

The thfluéice on communihes of sediment Lype,

degree of emersion, and presence of scavrass are

investigated in this study.

Stalion focniiens, elevatians, and sedintents.

Near Port Augusta the narrow channel basin of

northern Spencer Gull is bordered by shallow

sublittoral seagrass meadows and extensive intertidal

mudflats backed by stands of the grey nangeove,

Avicennia marina var restniferd (Fig. 1),

Seven transects were established along the eastern

shore-of the gulf, bath adjacent to and mmote from,

exisling and proposed power station developments

(Fig. 1). Three stations were established at dilferent

intertidal zones un each transect, Hach station hud

an area of about 25m?, On each transect, station

1 was located 30m scaward from the mangrove

fringe, station 3 was loyaled al the seaprass verge,

(emergent at extreme low tide} and station 2 was

located midway between | and 3. The locations and

elevations af each station were confirmed relative

to gn esmblished Porc Aneusta Power Station

darum, using an BDM theodnlite.

Sediment samples were collected from cach

siation on all transects and analysed for pruin size

according to the Weni(wurth scale (Folk 1968) with

injnor modifications (Thomas ef af 1986). Graphic

means, Mz, (Folk 1968) were determined for the

samples. The Bray-Curtis classitication technique

was used Lo determine homogeneity groups bases

on the percentage distribution of particular

sediment size classes. in each sample (Miedeke &

Stephenson 1977; Thomas ef af 1986).

Werer ane oir temperature.

For the study period 1982-1986, a continuous

record of Gulf water temperature was made at the

cooling water intakes of Playford Power Station.

A continuous three hourly record of ambient uit

fremperature was mnade at rhe SA Weather Buredi’s

Port Augusts weather station No, (19066.

Fauna

Each station was sampled using a hand operat:

ed voret collecting 3 sediment sample of JAG cme,

surface area 80 ern? 10a depth of 20 em. The sam-

ples were taken during daylight hours.

The pallern of species accumulaiion with repent-

ed tandom coring within a station area was deter-

mined. By subsample 20, about 90% of the spe-

cies recorded in 40 subsamples had been found, and

those species which occurred after subsample 20)

comprised an extremely small number of the total

number af individuals sampled. The decision was

made to proceed using 20 subsamples per station,

exceeuing the species-area requitements proposed

by Cain & Castro (1959) and mecting the stringent

propusaly of Wemberg (1978), Each Station was

sampled twice yearly (winter and summer) From

June 1982 10 Sune 1986. Samples were treated as

desyribed in Thomas er .a/(1986). Faunal relation-

ships (berween stations, transects, tidal zones, aid

samipling mes) were interpreted using Bray-Curtis

classificu(iun aller rool-reot transformation of the

data, and dendrograms were constructed with

group dverage sorting (Swariz 1978; Bield. ed al

1982),

Species constancy, abundance rank scores, aid

average rank scores were assigned us in Hitilstone

(1976) for 160 spectes totalling 34,312 individuals.

Species associations were detentined using Bray

Curtis classification analysis an non-transformed

data, considering the 40 top ranked species for the

period 1982-1986.

Results

Seation elevations and sediments

Tidal Puciuations relative to station zones result

in dilferent periods of emergence for statins 1, 2

and 3 om all 1ransects. Considcring February 1984,

Fig. 1. Sampling station locations. Northern S

denoted.

INTERT(DAL COMMUNITIES OF NORTHERN SPENCER GULF

3Guik

ausStAacia

SOUTHE AN

OCEAN

INSHORE

7] SEAGRASS

FRINGE

PA= Posidonis

PS- Petigania stnuona

AA—Ampnibalis antarctica

Caaliing Water

Ourret

Channa

PORT

PATERSON

pencer Gulf, South Australia. Inshore fringing seagrass species also

72 RB. C. AINSLIE, DA. JOHNSTON & E, W, OFFLER

the station | zone is emergent for over 200

hours/mionth, station 2 for about 80 hours/month,

and station 3 for about 5 hours/month, or less than

LY of the time, In winter even shorter penods of

emergence oveur at the low intertidal zone. Local

meteorological canditions, such as wind velocity,

also affect the degrce of exposure on low spring

Tides, Thus stations ) and 2 are clearly mid interti-

dal stations while station 3 is in the low intertidal

zone.

The mid intertidal stations ate devoid of seagrass

cover. The seagrass species al stations 3 include

bath Posidonia australis and Posidonia sinuosa,

without clear cui zonution in the shallow water,

Some Amphibalus antarctica is present at transect

The hierarchical classification of sediments from

all stations gives two major groups ol stations,

proups (a) and (b) Fig. 2. The sediments of statiuns

forming group (a) have Mz values Which gencrally

range from medium sand (1.50) to coarse sand

(0,250). The exceptions are station 1.1 which is clas-

sifted ax fine sand (Mz = 2.5}, and the subgroup

formed by sediments of stations 6,1 and 6.2. These

last two stations have Mz values between 0 ancl

— 25 (ie. at the lower end of the very coarse - sand

scale),

Fhe second major group of sediments (b) have

Mz values ranging from very coarse sand [—259)

hough to granules (1.7), with stations 7.1 and

7.2 having the coarsest sediments recorded int the

survey, Although sediments of station 7,3 have un

Mr? value of 1,256 (medium sand), this station

forms its own subgroup on the dendrogram because

of the restricted distribution in grain size around

the median, unlike all other samples which had

mire OF less even ranges of grain sizes.

Aur and water temperatures

Air and water temperatures 1982, 1984 and I98S

show that seasonal water temperatures follow air

temperatures, Ambient average nonthhy water tem-

peratures range from [1-12,5°C in winter to

24-28°C in summer (Fig. 3).

here is no indication of change in the overall

umbient cemperature from 1982-1985, as a result

of intermittent cooling water discharge fram the

existing pawer station development.

Fauna

A total of 160 species were recorded ; polychaetes

were the most Numerous with 52 species, includ-

inw several new records for S, Aust. (Table 1). Mol-

iuses: with 50 species and crustaceans, 40 species,

were next i atuindance, with 18species from var

ious ofher taxa (lable L),

“0

= eo

= 70

A i is io, i a a

a ee ee

a STeTiom b

Fig. 2. Dewdfograincl northern Spencer Gull stations by

sediment prades.

Fig, 4 presents the station faunal homogeneity

dendrograms for the first two seasonal surveys

(winter 1982, summer 1983), and for the last two

surveys (summer and winter 1986). [0 all cases, at

about the 25-30% level the homogeneity dendro-

grams consistently show two major collection

clusters with high intra-group homogeneily (Pig.

4 (1-4), Groups A and B). Generally the faunas

of stations Land 2 on all transects comprise Group

Aon each dendrogram, and station 3 faunas lorm

Group B. In two of the dendrograms presented

(winter 1982 and simmer 1986) there are exceptions

to this generalisation, but there is no seasonal or

spatial paticrn to. the exceptions, and the major

trend persists, There is no evidence of consistent

subgrouping within the major observed associations

{Group A and Group B, Fig. 4). This tendency for

faunas of mid intertidal stations | and 2 to group

more closely together than to those of station 3 has

been consistent throughout the our year study peri-

od and is also reflected in the the tidal zone analy-

sis, (Fig. 5) for each survey 1982-1986.

The species homogeneity dendrogram cleter

mined using the 40 most common species.is.present-

ed in Pig. 6. Accepting the arbitrary cut-off level

of 25% similarity, this dendrogram shows two

Major groups of species (Al and Bl, Fig. 6).

Although all but two of the forty most common

species occur over all stations, those which form

Group Al always contnbute a greater proportion

of the total number of individuals in the samples

of stations | ard 2 and those of Group BI are al-

ways more abundant in the station 3. faunas,

Together these analyses (Figs. 4, 5 & 6) indicate,

thar the two habitats represented by the mid inter

lidal region (stations | and 2) and the low interti-

INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF 73

TABLE 1. Species recorded throughout the study, New records for South Australia are indicated by an asterisk.

eee ee en ee EE PEE a

Exclusively Exclusively

mid intertidal low intertidal

POLYCHAETES

Olganereis edmondsi (Hartman)

Neanthes vaalii Kinberg

Neanthes cricognatha (Ehlers)

Ceratonereis transversa Hutchings & Turvey +

Ceratonereis mirabilis Kinberg

Perinereis nuntia (Grube)

Perinereis amblyodonta (Schmarda) +

Perinereis sp

Nereis trianguluris Hutchings & Turvey

Nereis sp 4

Nematonereis unicornis (Grube)* +

Marphysa sp*

Arabellidae 1* +

Arabellidae 2* +

Lysidice sp

Eunice sp +

Schistomeringas sp

Lumbrinereis sp

Nephtys austratiensis Fauchald

Glycera americana Leidy 7

Phylo sp* +

Scoloplos cylindrifer Ehlers

Leitoscoloplos normalis Day*

Leitoscoloplos sp

Naineris grubei australis Hartman

Phyllodocidae 1* +

Phyllodocidae 2* +

Phyllodocidae 3* +

Cirriformia filigera (Grube)

Cirratulidae sp

Boccardia chilensis Blake & Kudenoy

Harmothoe sp 14

Harmothoe sp 2*

Harmoathoe sp 3*

Lysilla apheles Hutchings*

Liomia medusa (Savigny)*

Eupolymnia nebulosa (Montague)

Streblosoma sp*

Terebellinae 1

Terebellinae 2 +

Terebellinae 3

Terebellinae 4

Polycirrinae*

+++

A - species seperated on basis of anterior elytrae.

Barantolla lepte Hutchings

Notomastus sp +

Paraonidae

Pherusa sp ae

Armandia sp

Galeolaria caespitosa (Savigny)

Scalibregma sp*

Maldanidae*

Maeelona sp*

NEMERTEAN

Nemertean spp +

SIPUNCULID

Golfingia margaritacea adelaidensis Edmonds

74 R. C, AINSLIE, D. A. JOHNSTON & E. W, GFFLER

BIVALVES

Tellina deltoidalis Lamarck

Laternula recta (Reeve)

Katelysia peronii Lamarck

Katelysia scalarina (Lamarck)

Brachidontes erosus (Lamarck)

Xenostrobus inconstans (Lamarck)

Ostrea angasi Sowerby

Pinna bicotor Gmelin

Malleus meridianus Cotton

Anomia trigonopsis Hutton

Circe scripta Cotton

Electroma georgiana (Quoy & Gaimard)

Trichomya hirsuta Lamarck

Chlamys bifrons Lamarck

Musculus paulucciae Crosse

Corbula flindersi Cotton

Lucilinidae

GASTROPODS

Salinator fragilis (Lamarck)

Nassarius burchardi (Philippa)

Nassarius pauperatus (Lamarck)

Monodonta constricta Lamarck

Bembicium melanostomum (Gmelin)

Batillaria estuarina (Tate)

Batillaria diemenensis (Quoy & Gaimard)

Bedeva paivae (Crosse)

Cominella erburnea (Reeve)

Cominella sp

Austrotiotia densilineata (Tate)

Diata lauta A. Adams

Quibulla tenuissima Sowerby

Lepsiella vinosa Lamarck

Conus anemone Lamarck

Cantharidus irisodontes (Quoy & Gaimard)

Clanculus plebejus (Philippi)

Clanculus weedingi (Cotton)

Phasianella australis (Gmelin)

Hydrabia sp

Microcolus sp

Sophismalepas nigrita (Sowerby)

Pterynotus triformis (Reeve)

Nudibranchiata

Opisthobranchia

Netomella sp

Elegidion occiduus Cotton

Asteracniea sp |

Asteracmea sp 2

Stomatella auricula (Lamarck)

CHITONS

ischnochiton variegatus Adams & Angus

Ischnochiton contractus Reeve

Heterozona sp

CEPHALOPODS

Hapalochlaena maculosa Quoy & Gaimard

Exclusively

mid intertidal

Exclusively

low intertidal

+ ++

+444

+++

INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF

Exclusively

mid intertidal

Exclusively

low intertidal

DECAPODS

Penaeus latisuicatus (Kishinonye)

Processa gracilis (Baker)

Leander serenus (Heller)

Leander intermedius (Stimpson)

Paguristes frontalis (M. Edwards)

Paguristes brevirostris (Baker)

Callianassa aequimana (Baker)

Callianassa ceramica (Fulton & Grant)

Crangon socialis (Heller)

Pilumnus fissifrons (Stimpson)

Philyra laevis (Bell)

Cryptocnemus vincentianus Hale

Polyonyx transversus (Haswell)

Gomeza bicornis (Gray)

Halicareinus ovatus (Stimpson)

Litocheira bispinosa (Kinahan)

Portunus pelagicus (Linnaeus)

Helograpsus haswellianus (Whitelegge)

Ceratoplax punctata (Baker)

AMPHIPODS

Maera mastersii (Haswell)

Birubius panamunus Barnard & Drummond

Elasmopus bampo Barnard

Allorchestes compressa (Dana)

Ceradocus serratus (Bate)

Cymadusa sp

Gammaridae |

Gammaridae 2

Gammaridae 3

ISOPODS

Cymodoce longicaudata (Baker)

Zuzara venosa (Stebbing)

Exosphaeroma sp

Cirolana woodjonesi (Hale)

Cymothoidae

Anthuridae 1

Anthuridae 2

TANAIDS

Apsuedes australis (Haswell)

Paratanais ignotus (Chilton)

Tanaidac

CIRRIPEDS

Batlanus amphitrite Darwin

Elminius modestus Darwin

CNIDARIANS

Anthothoe albocincta (Stuckey)

Cricophorus nutrix (Stuckey)

PISCES

Nesogobius hinshyi (McCulloch & Ogilby)

Gymnapistes marmoratus (Cuvier & Valenciennes)

Plotosidae

+ +4444

76 R_. AINSLIE, D, A. JOHNSTON & GB. W. OFFLER

ECHINODERMS

Leplosynapta dolabrifera (Stimpson)

Taeniogyrus roehucki (Joshua)

Amblypreustes sp

Trachodotu sp

Ophiuroidea |

Ophiuroidea 2

Dendrochirotida

Evhinoidea

ASCIDIANS

Ascidia sp

DIPTERA

Dolichopodidue

Exclusively

mid intertidal

Exclusively

low intertidal

dal fringe (stations 3) can be considered to have

quantitatively differing faunas.

fig. 7 compares mean nuniber of species/m?,

and mean numbers of individuals/m? for these two

hahitats over the period 1982-1986, Significanily

more spectes/m? are found in the low intertidal sta-

ions. Reference to the species list for all surveys

reveals that of the 160 species collected, 86 (54%)

oceur over the whole tidal range. This 55% includes

37 of the 40 most common species, Fifty-nine spe-

cies, or 38% of the total, occur exclusively in the

low intertidal zone, and only 8% occur exclusively

in the mid intertidal zone occupied by stations 1

and 2 (Table 1), In contrast to species abundance,

the communities of mid intertidal stations | and

2 consistently have significantly higher mean num-

bers of individuals (Fig. 7) than those of the

scagrass fringing stations 3, Nine of the 10 most

common species are More abundant at stations |

and 2 (Table 2, Fig. 6); conversely, of the 61 spe-

cies exclusive to the station 3 community, only two

appewr in the “top 40” list (Table 2). Nevertheless

the less common, but exclusively station 3 species,

contribute to the consisient separation on the sta-

tion and tidal zone faunal homogeneity dendro-

grams (Figs. 4 and 5).

Fig. 8 shows {he intersurvey time dendrograms

comparing whole survey collections for each of the

two communities for the period 1982-1986,

Discussion

Community-habiat relationships of the mid inter-

tidal and low intertidal zones.

In many studies scdiment characteristics have

been shown to be a significant factor in determin-

ing, benthic community patterns (Gray 1974;

Saenger ez uf 1982" Jones e¢ af 1986). In intertidal

regions particularly, there is the potential for wave

action to grade sediments, resulting in coarser par-

ticles on the upper shore line. Such a graded series

of sediments is potentially reflected im the nature

of the benthic communities (Gage 1974; Robinson

1985). On the. low energy shore of northern Spencer

Gulf there is no sediment gradation, with random

groupings of station sediments irrespective of the

zones in Which the stations occur. There is a high

overall similarity in the sediments, the two main

groups discerned (medium to coarse sand, and

coarse sand to granules) showing about 70%

similarity on the homogeneity dendrogram (Fig, 2)-

In common with other studies of the South Aus-

tralian Gulfs where a limited range of sediments

is found (Kinhill 19874; Thomas er af 1986) there

is no evidence that this factor influences benthic

community distributions in the northern Spencer

Gulf,

The mid intertidal zone is characterised by spe-

vies which, although more common in this zone,

oveur over the whole tidal range (Table 1), The

most abundant species is the mussel, Bruchidontes

erosus, Which forms dense mats in this zone. Such

mussel mats are characterislic of many low energy

sand and mud flat substrata in 8, Aust. (Womers-

ley & Thomas 1976; Thomas ef a/ 1986), Other

bivalves, although less abundant thai the mussels,

are also characteristic of the mid intertidal zone.

These include Je/ling deltoidalis, a species belong-

ing to a genus known to have a behavioural

response to light direction which contributes to tts

zone maintenance (Newell 1979), Katelysia scala-

rina, and Laternula recta. These bivalves contrib-

ute to the abundance of many other species as they

provide the substratum for the settlement of the

barnacles Elminius modextus and Balanus amphi-

INTERTIDAL. COMMUNITIES OF NORTHERN SPENCER GULF 77

PORT AUGUST& POWER STATIONS

AIR-SESWATER TEMPERATURE COMPARISONS

welee Temri me rose MEeSURED AT BLaT FORO Baste Cur pce?

v 36

w 25

~ 29

< i

wood

# s

= WoMFHLe AW, OST

w 19 ~~" wake Thue,

montatr ay,

5 TT waren Teme

wom Fee WAN APR dey UR 4uL AUG Bte SCH Woy Dec

aay FER MAP APR WAY GUN 4UL 4UG §8® CT WEY bec

—_

eS FER be BPR mk) Gu Jul 4uG sae pot Mey cer

Fig. 3. Monthly averages of daily maximum air

temperatures recorded at SA Weather Bureau Station

No 019066, Playford Power Station, Pt Augusta (Fig

1), Monthly average ambient Gulf water temperature

derived from vontinuous hourly records at the

ciyculating water intakes, Playtord Power Station.

trite, and the sea anemone Anthothoe albocincta,

There also appears to be a close relationship be-

tween the large bivalve species and the omnivorous

worm species Nephiys australiensis, Neanthes

vaalit, Olganereis edmondsi, and Nereis trian.

gularis, The last species seems generally to be as-

sociated with mussel beds (Hutchings & Murray

1984). Several species of scavenging and car-

hivorous snails (Nassarius pauperatus, Bedeva pai-

vat, Batillaria estuarina, and Batillaria diemenen-

sis) also Occur in the community dominated by the

mussel beds, as does the scavenging erab PAilyra

faevis, characteristic of the mid intertidal zone on

low energy coastlines of S, Aust, (Womersley &

Thomas 1976). The herbivorous top shell,

Monodonta consiricia is common among the dense

mussel beds which provide a substrate for algal

growth, particularly Enteromorpha spp.

In the extreme weather conditions of the north-

erm Spencer Gulf it is clear that a number of ubig-

uitous intertidal species are restricted to the low in-

tertidal zone. These include the polychaetes

Nematonereis unicornis, Ceratonereis transyversa,

Barantolla lepte, Cirriformia filipera and

Leltoscoloplos normalis, for which a range of

habitats has been documented (Hutchings & Mut-

ray 1984) and the gastropods Nassarius burchardi,

and Ausfrolioiia densilineaia, also found elsewhere

over broad tidal ranges (Thomas er al 1986), Can-

versely, there are a number of species which are

generally characteristic of the subtidal zone. These

include such species as the hammer oyster Malleus

meridianus, the scallop Chlamys bifrons, the blue

swimmer crab Portunus pelagicus, and several

other species primarily, or only, found in associa-

uon with seagrass (Electroma georgiana (Ludbrock

1984), Golfingia margaritacea adelaidensis (Ea-

monds 1980) and Amblypneustes sp. (Shepherd &

Sprigg 1976)). The major environmental factor

which appears to separate the habitats described

in this study is the “degree of exposure”. This is

a function not only of the length of the periods of

emersion (in a region of high summer air tempera-

tures) but also the protection from environmental

extremes provided by the presence of seagrass in

the low intertidal zone,

Comparisons with other areos,

Are the intertidal communities of norchern

Spencer Gulf impoverished as evidence may sug-

yest for epizoic fauna on Pinna bicelor? There are

inherent difficulties in comparing coramunities of

various marine and estuarine areas. Generalisalions

from one area to another are often of limited value

beyond the broad observation that marine domi-

nated sites generally have more specics than estua-

rine sites (Collett et a/ 1984), Accepting this timi-

tation it can nevertheless be seen that the total nurn-

ber of species found in the present study (160) is

within the range of specles number recorded in

several studies of eastern Austratian “estuarine”

regions (Rainer & Fitzhardinge 1981; Jones e7 a/

1986), and is comparable 10 the number recorded

in # recent study of the intertidal mudflat fauna

of the Port River estuary, Gulf St Vincent (Tha-

mas er a! 1986), About one and a half times as

% SIMILARITY

R. C, AINSLIE, DB. A, JOHNSTON & E. W. OFFLER

Fig. 4. Station faunal homogeneity dendrograms, (1) winter 1982, (2) summer 1989, (3) summer 1986, (4) winter 1986.

many species are recorded in the present study as

were recorded by Kinhill (1987)8 for intertidal mud-

flats about 50 km further south in Spencer Gulf

Where salinities are four to five lower (Nunes &

Lennon 1986). Kinhill (1987)4 did nal sample at the

seagrass fringe (D Evans pers. comm.). Disregarding

the 38% of the total number of specics which are

exclusive to the seagrass (ringe in the present study

leaves a species number very similar to that found

by Kinhill 987)". Therefore this study provides no

evidence of species impoverishment in the intertidal

zone, related to high salinity or other environmental

factors characteristic of the area.

Natural vartations in the intertidal communities of

northern. Spencer Gulf

For.a baseline study to be of use in post develop-

ment monitoring an. understanding of natural var-

iations in the species matrix is necessary, The faunal

homogeneity dendrograms for both stations and

tidal zones demonstrate the consistent relationship

between the mid intertidal and tow intertidal com-

munities over several seasons. Between 1982 and

1986 the common core species characterising these

two habitats remained constant (Table 2). Perhaps

surprisingly no seasonal pattern emerged from the

intersurvey comparisons. The over-riding pattern

INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF 79

which showed for both habitats was the trend for

sequential subgrouping ol surveys, with survey fau-

nas within each of the two habitats showing high

similarity, grouping at the 60%-70% similarity ley-

el in each case, Sequential grouping of survey fau-

nas does not appear to be unusual. Saenger et al

(1980) interpreted such a pattern as demonstrating

progressive recolonisation following environmen-

tal disturbance, Buchanan ef a/ (1974, 1978),

however, observed similar sequential subgrouping

of survey faunas from undisturbed marine environ-

ments and interpreted this pattern as indicating a

more or less continual change in the presence and

abundance of occasional or rare species whilc the

core species remain relatively constant; Given the

overali similarity in survey faunas over time this

seems the most likely explanation here. The slightly

lower intersurvey similarities of the low intertidal

communities, compared ro those of the mid inter-

tidal communities, isa consequence of the fact that

the former are characterised by more rare or occa-

sional species.

Monitoring Jor changes associated with environ-

menial disturbance

Changes in estuarine faunas in response to or-

ganic or thermal disturbance have been well

documented (Gray 1976; Parker 1980; Bamber &

Spencer 1984; Thomas er a/ 1986). In particular,

in the negative estuary of the Port River, Gulf St

TABLE 2. Cornman species of the intertidal fauna-of Northern Spencer Gulf, determined by reference to constancy,

and abundance rank scores (Hailstone 1976). Note that the top 40 species ranked in this table includes all 21 species®

which would be selected on Field e7 ai’s (1982) arbitrary cutoff for dominant species, namely species which contribute

4% or more to the total papulation of any one survey,

% ABUNDANCE % CONSTANCY

RANK CODE SPECIES

(Refer Fig 6)

ia | Wi Oleanereis edmondsi

>} wi Nephtvs australiensis

°3 B7 Brachidoniés erasus

4 BI Tellina deltoidalis

oF RIS Nassarius pauperalas

*g, 6 C4 Paguristes frontalis

7 Bia Batillaria diemenensis

°8 B4 Katelvsia scalurina

9g C7 Elminius modestus

"10 W6 Scatoplas ¢ylindrifer

IT w2 Neanthes vaalii

12 BY Monodonta constricta

"12 C2 Callianassa ceraniica

14 B3 Salinator fragilis

14% w7 Cirriformia filigera

°16 CR2 Balanus amphitrite

°17* Wwi0 Baraniolla lepte

F1RY CIR Apsuedes australis

19* W30 Nemeriean spp

20 Wil Nereis (riangitlaris

20° BS Nassarius burchardt

°22" Wi15 Maldanidse

23 wo Marphysa sp

24 wig Terebellinae 1

24 W22 Lysilla apheles

226, cl4 Crangon sovialiy

27 co Philyra laevis

28 B2 Laternula recta

29 B26 Asleracniea sp 1

50) Bll Batilluria estuarina

3H Bi2 Bedeva paivae

32 X5 Anthothoe albacineta

o33% W33 Eupolymnia nebulosa

°33 W20: Streblosoma sp

35 BI4 Katelysia peronii

aA* W38 Golfingia m udelaidensis

°37* B35 Electroma georgiana

33” B20 “Austroliotia densilineata

3R* Wa Schistomeringes sp

aut Wwi2 Leitoscoloplos normatis

10.2 73.5

7 95,1

36,8 65,9

3.0 73.0

22 67.6

23 62.2

39 47.0

2.0 56.7

4,3 33.0

1.7 49.2

hg 45.9

0,95 34.0)

0.8 53.5

3.7 23.8

1.95 44,0

46 20.5

0.9 28.1

0.9 23.8

0,35 34.6

0.55 24.3

0.5 28.1

OS 26.5

0,37 28.1

0.6 17.8

0.5 22.2

0.28 23.8

0,26 23.8

0.21 23.8

0.3 25

0.7 9.7

0,17 17.8

0.2 15.)

0.4 97

0.24 13.0

0.14 15.7

O14 13,5

0,2 10.8

0.5 7,6

0.12 15,1

ONS 10,8

* Most common species in the low intertidal zone Group BI Fig. 6.

80 R, C, AINSLIE, D. A. TOHNSTON & E. W. OFFLER

WINTER SIMMER WINTER SUMMER WINTER SUMMER WINTER SUMMER WINTER

Wu2 1989 1863 1964 1964 1985 1985 1966 1086

GQ Simi, aarry

ate %

LOW INTERTIDAL STAT

INTERTIDAL

Fig. 5, Faunal homogeneity dendrograms with respect to tidal zones for all surveys, 1982-1986.

wi eo WZ a7 mt s2 we c7 B19 W22 812 xs 82 WIO Ca wih Ge cl4 w3e e20

as wit #4 wa cz we 0811 814 CS s26 wi? c32 W7 W30 W344 82 S35 W339 W20 wi?

60-

% S!MILARITY

Al SPECIES Bt

Fig. 6. Species homogeneity dendrogram determined on non transformed data for the 40 mast common species far

the period 1982-1986 as determined by average rank scores assigned as in Hailstone (1976).

INTERTIDAL COMMUNITIES OF NORTHERN SPENCER GULF BL

—— Species

——— Individuals

7oa

600

SY ly

20 i 500

th 4000

a 15 { a

E | bd

o 3005

bas t a

10 I >

a 1 Qa

* \ 2002

a |

= 9

2 6 z

a 1004

= { =

{

Stations

142

Station

Fig, 7. Comparison of numbers of species/m® and

numbers of individuals/m? for mid intertidal stations

1 and 2, and for law intertidal stations 3's,

Vincent, Thomas et a/ (1986) describe responses of

intertidal communities to progressive increase of

thermal discharge from Torrens Island Power

Station. These include suppression or elimination

of populations of bivalve mollusc and worm species

characteristic of undisturbed regions, and increases

in populations of opportunistic worm species. OF

the 120 species recorded from the Port River estuary

about 50 species (40%) also contribute to the 160

species of the present study, These include several

bivalve and worm species which had a negative

response to thermal discharges, and which played

a significant part in defining thermal perturbation

in the Port River, namely the bivalves Te/lina del-

toidalis, Katelysia scalarina and K, peronu, and the

worns Néphtys australiénsis, Neanthes: vaulii and

Scoloplos cvlindrifer. All of these species are

common in. the communities of the mudflats of

northern Spencer Gulf, particularly in the mid

intertidal zone (Table 2), The presence of these

common indicator species provides the basis of a

monitoring programme to detect potential thermal

perturbation in northern Spencer Gulf. Any

changes in abundance of these species in areas near

thermal discharges will be reflected in recognisa-

ble changes to the stable faunal homogeneity rela-

(a)

% SIMILARITY

SURVEY

% SIMILARITY

SURVEY

Fig. 8. Intersurvey faunal homogeneity dendrograms. (a)

stations | and 2, (b) stations 3. Survey number 1 —winter

1982, 2—summer 1983, 3 — winter 1983, 4 - summer

1984, 5— winter 1984, 6 — summer 1985, 7 — winter 1985,

# — summer 1986, 9 — winter 1986.

tionships described in this study over the period

1982-1986. Such changes in faunal homogeneity

dendrogram patierns have been clearly demonstrat-

ed by Thomas ef a/ (1986) in studies of changes in

the fauna of the Port River estuary with progres-

sive increase in thermal discharge to that region.

The cirratulid worm Cirriformia filigera is common

in the low intertidal.zone (Tables 1-2), Although

the life-history of this particular species is not well

known, other members of this genus have been

shown to have typical opportunistic characteristics,

capable of rapid colonisation of disturbed regions

82 KOO. AINSLIE, DAL IGHNSTON AE W OFFLER

(George 1964 a,b; Thomas e7 a/ L986}. ‘Nhe ar-

cherypal opporiunistic worm species Capirella

cupttata (Grassle & Grasse 1974; Gray 1976; Tsut-

sumi 1987) has not deen recorded fn the present

study. (Cas, however, present in inferlidal commu-

nities 30 kin further south (kimhill 19874), and it

is nol unusual for opportunistic species in disturbed

regions to be imnmugrants from other regions (Tho.

mas ef «! 1986), As with changes in the indicator

species discussed above, any changes in populations

of opportunistic worm species near the thermal dis-

charge can be expected to noticeably alrer che pat-

terns Of the faunal homogeneity dendragrams,

warning of potentially significant environmental

perturbation, ‘To 1986 foe impact of the gecusion-

| incursions of warmed water [rom Thomas Play-

ford Power Siitiun to the intertidal zone, and the

initial discharge from the Nerchern Power Station

(from summer 1985) can be discerned from com-

munity relationships described in the present study.

Avknowledpments

This work was funded and supported by the

Electricity. Trust of South Australia as part of its

Ongoing programme of covironmental research.

We are grateful to those who identified organisms,

espectally Jr P. Hutchings (polychaetes) and hdr Wy,

Zeidler (amphipods) and to the South Australian

Museum for access to their relerence collections,

We also thank Dr M, Geddes for his suggestions

and comments,

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ROTIFERA FROM AUSTRALIAN INLAND WATERS.

HI. EUCHLANIDAE, MYTILINIDAE AND TRICHOTRIIDAE

(ROTIFERA:MONOGONONTA)

BY W. KOSTE* & R. J. SHIEL}

Summary

Diagnostic keys are given to the Australian representatives of the Rotifera: Monogononta in the

families Euchlanidae (Manfredium, Diplois, Dipleuchlanis, Tripleuchlanis, Euchlanis), Mytilinidae

(Mytilina, Lophocharis) and Trichotriidae (Wolga, Macrochaetus, Trichotria). All species known

from Australian inland waters are described and figured, as are some widely distributed taxa not yet

recorded from the continent. Distribution data and ecological information also are given.

KEY WORDS: Rotifera, Euchlanidae, Mytilinidae, Trichotriidae, Australia, taxonomic revision.

ROTIFERA FROM AUSTRALIAN INLAND WATERS.

lil, EUCHLANIDAE, MYTLLINIDAE AND TRICHOTRIIDAE

(ROTIFERA:MONDGONONTA)

by W, Koste* & R. J. SHIELi

Summary

Kostr, W. & Sxies, R. J. (1989) Rotifera from Australian Inland Waters, If}. Euchlanidae, Mytilinidae and

Trichotrijdae (Rotifera:Monogononta). Trans, R. Soc. Aust, 113,85 14, 31 May, 1989.

Diagnostic keys are given to the Australian representatives of the Rotifera:Monogenonta in the families

Euchlanidae. (Mun/redium, Diplois, Dipleuchlanis, Tripleuchlanis, Euchlanis), Mytilinidae (Mytilina,

Lophocharis) and Trichotriidac (Wolga, Macrochaetus, Trickarria) &\| species known from Australian

inland waters are described and figured, as are some widely distributed laas not yet recorded from the

continent. Distribution dala and ecological information also are given.

Key Worobs: Rotiféra, Euchlanidae, Mytilinidae, Trichotridas, Australia, laxonomic revision,

Introduction

The families of Rotifera:Monogononta

considered in this part of our review of the

Australian species (see Kosle & Shiel 1984b, 1987h)

commonly are encountered in the shallow littoral

zones of billabongs or vegetated lake margins {cf.

Koste 1981, Tait et af. 1984, Koste & Shiel 19862,

1987a, Koste e¢ a/, 1988). Some species, ee.

Euchtanis dilatata and related fortns, are known ta

become planktonic in eutrophic waters during

cyanobacterial btooms (Ruttner-Kolisko 1974). ‘I'his

species Occurs seasonally in the lower R. Murry,

S. Aust, (Shiel ef a/. 1982), Otherwise, species of

the Euchlanidae (Manfredium, Diptuis,

Dipleuchlanis, Tripleuchlanis, Euchilanis),

Mytilinidae (Mysilina, Lophocharis) and

Trichotriidae (Moleag, Macrochaetts, Trchotrin)

generally are benthic or epiphytic in habit, and

o¢eur in open water of lakes, reservoirs or rivers

only as incursion species during seasonal flushing

flaws,

Mast of the 3S taxa recorded in the three families

were collected as such incidental incursions into the

limnetic region of billabongs. It is likely that more

taxa remain to be described, particularly endemic

species, because our initial sampling program was

directed at the planktonic communiry and the

sampling bias has “undersampled” the prelerred

habitar of these Jittoral rotifers, Only four endemic

species are known in the three families, one

mytilinid (Lopheecharis curvata) and three

trichotriids (Tricholria pseudocurta and T, buchneri

from Tasmania, and Meerochaetus darineeli, from

ihe N-T.), In contrast, about 25% of the taxa in the

Epiphanidae and Brachionidae (Koste & Shiel

* Rontgenstr. 1, 2903 Bad, Zwischenatin, Fed. Republic

of Cermany

tMurray-Darling Freshwater Research Centre, POL Bow

821, Albury, NSW, 2640

1987b), true limmetic species (and therefore more

intensively sanipled), apparently are endemic,

All known Australian representatives of the three

families ave described and figured, and diagnostic

keys are provided to enable identification to species.

Widely-distributed taxa moc yet recorded from

Australia, but which are likely to be found here, are

included in some genera. The format follows thar

of carlier papers; dichotomous keys are followed

by individual species’ descriptions and known

Australian records. Relative abundance from > 2000

collections is indicated by: “rare” (fewer than ten

tecords, “uncommon” (10-30 localities) and

“comnban" Gnore than 30 widely dispersed

localities), Brief ecological data are included where

available, generally in the sequence: temperature

°C), pH, dissolved oxygen (DO, mg 1°‘),

conductivity (Kya, pS om ') and turbidity

{nephelometric turbidity units {NTU)).

To minimise the citation of previous references

to the rotifer families reviewed here, early Australian

references which are included in Shiel & Koste (1979)

are not repeated, nor are the majority of European

references, which are included, for example, in the

reviews of Hudson & Gosse (1886, 1889), Weber

(1358), Voigt (1957), Bartos (1959), Rudescu (1960),

Kutikova (£970) and Koste (1978), A more detailed

treatment of the Rotifera outside Australia is given

by these juthors,

Family Buchlanidae Bartos

The following genera Were integrated hy Remane

(1929-1933) in the subfamily Brachionidae. A

special family was erected by. Bartos (1959), later

accepled by Kutikova (1970) and Koste (1/978). With

the exception of Manfredium the genera are

characterized by a lorica with plates which are

connected with sulci, a segmented foot, more or less

elongated toes, a corona of the same type

(Euchlanis-type, Fig3d) and malieate trophi

(Fig, 2:2b,3).

Bh W. KOSTE & Ro J. SIIEL

Key tu genera of the Family Kuchlanidae

1 Lariea thin, without lateral sulci, long toes

-Maafrediumn Gallagher

Lorica tigiel, With dorsal and sometimes yentral plates

ee ee ee

W1), Lorica composed of three plates, dorsal divided,

separated by sulci. . b ..Diplois Gosse

Dorsal plave not divided. 2... 3

3(2).. Dorsal plate arched_ concave, narrower than arched

Ventral plate......-- _Dipleuchlanis

Reauchainp

Dorsal plate arched, convex o7 triangular... .. 4

4(3). Dorsal and ventral plates nearly of same. size,

conneeled by «lateral double longitudinal membrane

within 4 stiff projection. ... Trpleuchlanis Myers

With and without lateral longitudinal sulei

vate alia ae Exchlanis Ehreribere

Genus Mantredium Gallagher

piciaeite Manfredi, 1927, p. 58 (non Walsingham.

1911).

Boauchanipiella Remane, 1929, p, 107,

Eudactylota Manfredi, 1927 is a junior hornonym

of Exdactylota Walsingham, 1911 (Lepidoptera).

Recuchampiella Remane, 1929 is a nomen audem

under articles 12 and 16 of the International Code

of Zoological Nomenclajure (name published

without description, definition). Manfredium was

Proposed by Gallagher (1957) to replace Eudactylora

Manfredi, There is a single species, Man/reditern

eudactplotunt.

Manfredium eudactylotum (Gosse)

FIG, |

Sedridiun exdacivlotunt Gosse In Hudsan & Guese

1886, p. 74, Fig, 2h.

Eudae -rylota ecigersiont (Gosse) Manlredi, L927, p. 58.

Heauchamsiella euductylota (Gosse) Remane, (929, D,

Marfrediuim eudactylotmm (Ciosse) alter Gallagher,

1957, p. 182.

Djagnosis: Body pear-shaped with soft,

transparent cuticle; head stall; foot with two longer

segments; toes. long, rod-shaped; striated muscles

visible in caudal part of body and foot: masrax rei-

lobed, trophi malleate (Mig. 1:1); oesophagus. with

small paired glands; large ted cerebral eve; resting

egg with fine hairs (Fig. le.

Lenguh; 420-760 um, four S0-110 ym, toes

275-320 pm, trophi 25 am, longest uncus tooth 12

wo, manubrium 18am.

Oustribuzion: Cosmopolitan, panconitinental in

Australia, rare. Warntstenotherm, 11,5-17,0%, pH

4.8-6.5, conductivity to 1600 «S crim. The rotifer

may swim) betwee walter plants, also jumps by

Vicking the toes.

Literature: Koste 1978, 198): Ridder LYBI; Kose &

Roberison 1983; Koste & Shiel 1987a

Genus Diplo Gosse

EXplois Gosse Jv Hudson & Gosse, 1886, Vol. 2, a. 86.

Diplois daviesiae Gasse

FIG, 2:1

Diplors davesiae Goxse In Hudson and Cinsse, (886,

Vol, 2, j. 87. Pig. 24:3.

D, phlerrava Vroso, 1910, p. 301,

D. seulpturara Yaday, 1897, p. 135 (from New Guines).

Diagnosis: Lorica with three plates; arched dorsal

plate divided longitudinally by sulcus; lareral sulci

small; foot Lhree-segmented; toes thin; lorica surface

smooth, flexible; head shart, broad; eye with lens:

oesophagus short; stomach compact; gastric lands

large.

Length: (6 500 pin, width 225 ym, toes 100 jum.

Ostribation; May be cosmopolitan in the benthic

of freshwater Sphagnum pools. Not yet recorded

from the Neotropics. Not recorded in this srudy, bul

there is an early record from Old (Thorpe 1887).

Genus Diplenchlanis Beauchamp

Diplenchlaniy Beauchamp, 1910, p, 122,

Dorsal plate narrower than ventral plate; foot

three-segemen(ed; tocs more or less long, rod-

shaped with tapering points occasionally swollen

terminally; rami points with minute comb (Fig,

2:2b); unci with 7-10-teeth; gastric glands long in

younger individuals, broader and lobed in adult;

red cerebral eye. Two variants of # single species 12

propatula) are known trom Australia; individuals

can be discriminated by toe structure. DB. propatula

has toes without a swelling belore the points,

whereas, D. propaiula macrodactyla has toes with

a swelling,

Dipleuchlanis prapatula propatuta (Gaosse}

FIG. 2:2a—f

Diplois Ppropatula Goss /n Hudson & Gasse, ENMG, fi

ig Dinlewctanis propaiuia (Gosse) alter Beauchamp.

Diagnosis; Caudal part of dorsal plate sometimes

stretched to. rounded point znd reaches nearly te

end of ventral plate; tees tapering to start paint.

Length; 338-508 ~m, lorica length to 500 pm;

width to 200; toes 70-110 pm.

Distribution: Cosmapolitan in littoral, sometimes

occurs in plankton.of ponds or billabongs, Rare,

Vic, NT. Warm stenutherm, pH 4,5-6.4, also in O=

poor waters on the suriace af mud.

Literature: Koste 1978, (981.

Dipleuchlanis propatula macredactyla (Hauer)

FIG, 2:22

Dipleuchlarus. maervdacipla Hauer, 1963, p. 351,

Dipleuchlanis Prange f, mucrodactyte ( Haner) alter

Koste, 1978,

Diagnosis: Bava part of dorsal lorica with shore

paired cuticular projections; tocs long.

Length; 124-244 pm, widil) 69-212 pm, toes

112-168 am,

ROTIFERA FROM AUSTRALIAN INLAND WATERS

Fig 1: Manfredium eudactylotum (Gosse

Scales lines, top

(e) resting egg; (f) trophi;

Wulfert (1940); e after Koste (1978).

‘0 pm (e); lower left LO wm (f-i).

): (a) lateral; (b) ventral; (c) dorsal; (d) lateral;

Fig. la-d, f-i after

(g) opened rami; (h) uncus: (i) fulerum.

100 pm (a-d); lower right

88 W. KOSTE & R. J. SHIEL

5a 5b 5c 4h

Fig, 2: 1, Diplois daviesiae Gosse, dorsal. 2, Dipleuchlanis propatula propatula (Gosse): (a) dorsal; (b) trophi; (c)

yentral, swimming; (d) toe; (e) unci; (f) cross-section; (g) D. propatula macrodactyla (Hauer), dorsal. 3, D. elegans

(Wierzejski): (a) dorsal; (b) lorica cross-section (cl, D. propatula). 4, Tripleuchianis plicata plicata (Levander):

(a) lateral; (b) traphi; (c) dorsal, swimming; (d) cross-section; (e) dorsal; (f) ventral; (¢) Uophi, apical; (h) lateral.

5, T. plicata razelni Rodewald: (a) lateral, (b) dorsal, swimming; (c) ventral. Fig. 2:1, after Weber (1898); 2a, b,

f, 3a-d, alter Myers (1930); 2c, d, 3, after Fadcew (1924); 2g, 3e-g, after Hauet (1965); 5a—c, after Radewald-Rudeseu

(1960). Scale lines Fig. 2:1-4, 100um (adult lorica each number group); 5,50 zm; 2b, 4b, g, 10 pm.

ROTIFERA FROM AUSTRALIAN INLAND WATERS 89

Distritution: Panjropical, N7., Vie. Water ta 32°C,

pH 5.4-65.

‘iferatures Myets 1930; Haver 1965; Koste 1974,

1981; Berzins 1982.

Doubtiul or insufficiently described species:

Dipleuchtanis elegans (Wierzejski, 1893) = Ewchlanis

orapatula elegans {Fig. 2:3) according to Beauchamp,

1910, p. 122,

Bipleuchlams canradi Evens, 1947, p, 179, Fie 5.

Dipleuchtanis paludosa Haver, 1936, p. 139, Fiz. 1:12.

Cienus Tripleuchlanis Myers

Tripleucklants Myers, 1930 p. 379-

Lorica ovoid in shape, truncate in front,

smoothly-rounded caudally; dorsal and ventral plate

hearly of same size, connected by pair of lateral

longitudinal sulci. Longitudinal flange of stiffened

cuticle extending for entire length between each,

giving bellows-like appearance to cross-section. Foot

three-jolated, guarded by shield-like process

extending downwards from median longitudinal

Nange; tocs short ending in abrupt paints; mastax

of modified malleate type (Fig. 2:4h); six club-

shaped teeth on each uncus; rami triangular with

minute denticulate combs at inside of tip; double

cerebral eye, A single species, 7. pifeate, with a

doubtful variant (7 plicata f. rezelmé (Big. 2:5))

frou Romania distinguished on the basis of au

elongated dorsal plate. caudally with a rounded

notch (Rodewald 1940;88).

Tripleuchlanis plicata (Levander)

FIG, 2:4

Euchlanis plicata Levander, [RM p. 48,

spe neural plicata (Levander) after Myers, 1930, p.

Diagnosis: Lorica stout; foot glands and reservoirs

elongated; retrocerebral organ small,

Length: 250-270 pm, dorsal plare 90~I20 ~m,

ventral plate 100-115 pm, toes 25-37 ym.

Disiribution: Cosmopolitan; rare in fresh-, more in

brackish and marine waters, also (11 warm springs.

Qld, Vic

literature: Hauer 1925, Koste 1978.

Genus &ywohlanis Ehrenberg

evenianis Blirenberg, 1832, p. 13).

Loriva transparent with oval or ovate outline;

dorsal plate arched, convex, sometimes with median

keel or lateral wings, larger than ventral plate,

variable in height and shape; ventral plate caudally

somewhal indented, nearly flai. Two plates

connected by thin, Flexible cuticle forming

longitudinal sulci. Foot with two or three joints; toes

long or short, sword-shaped oF parallel-sided, with

sham tips, Paired long selae situated on dorso-distal

portion of foot-segments (Fig. 3a:ss). Corona of

family type (Fig. 3d); long sensory bristles and cilia

in tufts atrd solitary on apical field, Trophi (Fig.

3e, !) moditied malleate; fulcrum (Fig, 3ffr} broad,

pointed rami (ra) with and withoul minute comb;

uncus with main and variable number of accessory

teeth, Brain with large cerebral eye (Fig. 3b ey)

behind large mastax, Large retrocerebral organ

present (Fig, 3b:re), with subcerebral glands. Dorsal

antenna large; lateral antenna (Fig. 3b:la) with

sensillae tufts on small tubules. Stomach sometimes

with sacculi but not constant between species.

Excretory organ with large contractile bladder (Fig.

3a,b:b1), Vitellarium with eight large nuclei, Male

thinly loricate (Fig 3i:m) with reduced alimentary

tract; protonephridia present. Foot with twa to three

joints; two long sensory bristles on terminal joint.

Toes short. Dark resting eggs may be atlached to

plants (Fig. 3h,j),

éuchlanis species ave littoral rotifers living among

aquatic plants, byt in the pelagial they occur in

eutrophic lakes and ponds at Cyanophyta maxima,

especially of Gloeutrichia, the main food of

Evchlants dilatata (Ruttner-Kolisko 1974), Other

foods consists of diatoms, desmids, other alyae and

Getricus.

The taxonomy of the cifferent species is difficult:

even within the same population lorica shape and

cross-section is variable: The shape of the anterior

margin of both plates is of little value in jhe

determination. Most useful is trophi structure,

which is species-specific. Fucklanis should pot be

pressed by a coverslip. See for cxample & meretr

cross-sections and the different forms of £. diletera

Key ta Species of the Genus Fuchlanis

1, Cross section of lorica more or less arched to

High keel... .- Teer

wie wy wee a

2(1). Posterior edge of dorsal lorica with disiinet nolch

Venter membranous ar rudimentary ...,,-.-..5

413), Ventral plate ca. 44 dorsal plate width; wing-like

bic} .€. alata Voronkay

Ventral plate ca. 44 dorsal plate width, constricted

AL posterior end; fiw expanded margins (Fig. 6:3)

res -E. tera Hudson

5(3). Lateral constrictions in medial dorsal luriea; flanged

Jateral margins (Pip, 8:3. ..£. pyrifarmis Gusse

Dorszl forica not constricted, lateral margin not

flanged (Figs 4:3, d:d).... .-£, deflexa Gasse

6l2), Cuticular shield-like process just below caudal part

of dorsal plate (Fig. 7:1). --.. 8 mene Myers

Shield-like process lacking... .....-2.... 2. 7

AG) Ventral plate rodimentary; lungitudinal sulci abserit;

toes very Jong (>100 jan), slender (Fig, 4:25

vee ttiiineet vere eee. elpidia Myers

Ventrml plate well developed... ocr 4

i ia eras

0 W. KOSTE & R. J. SHIEL

Fig, 3: Auehianis dilatate dilatatq Ehrenberg: (a) ventral; (b) lateral; (c) cross-section; (d) apical field; (e) traphi;

(1) tYophi, lateral; (z) &. triquetra, amictic egg; (h) mictic epg; (i) EB. dilalafa, mictic egg: (j) subitaneous egg) (k)

snerm; (1) male; (m) £. deffeva, male. (Key to a,b: bl = bladder; dpl = dorsal plate of lorica; ey = cye; gg =

gastric glands; la = Jaleral-antenna; iv = mouth; me = membrane; ov = ovary; re = retrocerchral sag; sl =

stomach; 55 = sensorybristles; vi = vitcllarium; ypl = ventral plate of lorica. Key to f: ma = manubnium; fr

= fulcrum; ta = ramus; u = uneus). Fig. 3a, ¢, j, after Koste (1978); b, after Beauchamp (1965); d, afrer Remane

(1933); e, alter Hauer (1930); fafter Voigr (1957); g, h, afler Wesenberg-Lund (1929); i, alter Kavar (1914); k, after

Ruttner-Kolisko (1972); 1, alter Wullert (1956); 1n, after Leissling (1924). Scales lines, top, a-d, g-m, SQum; ¢, f 10 nm.

8(7). Dorsal loriga with winglike lateral extensions toes dilated distally for 44 length and constrict to

deflected ventrally al tips (Fig. 8!2)2. phryne Myers acute points (Fig, 8:1)......,,.8. orapha Gosse

No lateral extensions of dorsal lorica.....----. 9 10(1)Mentral plate completely developed, lateral sulci

1 present (Pip. @:1)..... sow stete's etotaa E. tacisa Carlin

9(8). Foat and tocs slender; toes 4 length of dorsal plate, Ventral plate reduced to thin hyaline membrane; wide

blade-like, fusiform (Fiz. SabjE. dilatata Ehrenberg dorsal plate flanges (Fig. 8:le-e..... E. trigitetra

Foot and toes stout; toes. 14 length of dorsal plate Lhrenberg

ROPIFERA FROM AUSTRALIAN INLAND WATERS ut

Euchlanis alata Voronkov

HG. 41

fuehlanis alerd Voronkoy, 1912, p, 210, Figs 2,3.

Diegrasis: Shape of body ovoid: dorsal plate

without terminal totch or embayment, but may

have lateral wing-like extensions (see Fig, 4:tb, d)-

Wingless individuals casily mistaken for & tyre or

£. deflexa. Intermediate forms common, Ventral

plate elongate, relatively marrow, but without

constricuon before end as in E. fvre. Foot twoe

segmented and robust; toes stout and fusiform. Tips

of tami incurved with pair of finely denticulate

combs; four stout teeth in each uncus with several

accessory teeth.

Length of dorsal plate 260-360 um: ventral plate

width 102-156 xm; toes 70-98 ym; toes width 12-20

am,

Distribution: Nearctic and Palearctic, in acid boreal

waters, littoral and psammal, Not vet recorded from

Australia.

Liferarnre; Pejler 1962; Kaste 1978; Chengalarh &

Koste 1983.

Euchlanis calpidia Myers

FIG, 4:2

Fuchlanis calpidia Myers, 1930, p. 371, Pl. 20, Figs La

Diagnosis, Dorsal plate round, or highly domed

(“obscure triradiate", ic. bhelmet-like) in cross

section. Middle portions of lateral edges straight

or pinched, extending downward below venter:

ventral plate rudimentary {cl. #, defleva and E:

porYormis), radiments connected wah dorsal plate

by flexible membrane; lateral sulci absent; foot iwo-

jointed, slender; toes very long, parallel-sided with

abrupt point. Trophi characterized by five slender

patred tecth in unci with Iwo ur three accessory teeth

beside smallest, Tips of rami withour minute

denticulate comb. Trophi similar to those of é&

deflexa and &. pyriformis, E, calpidia distinguished

by presence of distinct posterior netch at end of

dorsal plate, and by relatively long toes.

Length of dorsal plate 280-400 ym: lorica width

220-397 pmy.toes 12U-155 pm; toe width to 18 xm.

Distribution: Nearctic Palearctic (Europe to bar

Basi). Single record from a Goulburi R. billabong

near Seymour, Vie. 22°C, pH 7,3, DO 8,0 mg 1-3,

Literature: Hauer 1936; Wang 196{; Kutikova 1970):

Kasie 1978.

Evechlanis deflexa Gosse

FIG, 4:3

Fuchlanis deflexa Gosse, 1851, p. 200.

en deflexa (Gosselin Myers 1940, p. 3689, PL. 21,

"Danidit = Euchlents afier Carlen, W354, p, 16)

Dirgnosis: Rady resembles are of circle i) cross-

section, dorsal plate oveld, without distince

Posterior notch, sometimes wilh shallow

emargination, Lateral edges of dorsal plate

connected by flexible membrane somewhat

thickened in position occupied by ventral plate in

other Zuch/anis. Posterior portion of this arca more

hardened as rudimentary ventral plate No

longitudinal sulci present. Foot stow! and awo-

jointed, long setae on dorsal end of first joint:

trophi with five long teeth on each uncus, clubbed

at tips, with 2—3 accessory tveth; rami drawn oul

ta long tips, minute inside combs lacking (Fiz. 4:31).

Length of dorsal plate 190-350 wm, width

140-240 wm, toes 55-100 um, toes ta 15 wm wide,

subltanzous egg !90x89 pm.

Disiribution: Cosmopolitan in (he littoral,

ovcasionally in the pelagial. Rare; N.SW., Tas., Vic,

W.A. There may be local yanants, eg. the

Tasmanian form closely resembles & deflexa lore

(Fig, 4:4), described from Lake Balkasch, Siberia

(Kutikova 1959), [4.0-22PC, pH §,0-7,7, DO

8.0-8.6 me I-1, <70 wS cmb

Citerature: Evans 1951; Koste 1978; Shicl & Koste

1979.

Euchlaniy dilatata Ehrenberg

FIGS 3, S:1

Evrhlenis iifatata Ehrenberg, 1832, p 131, Fig, 4:3.

Diagnosis General body shape ovoid, rruncate

anteriorly, rounded posteriorly; dorsal! plate variable

in height and shape of cross-section, varying from

low ate of circle to high triangle: ventral plate

almost as large as dorsal; longitudinal sulci nor deep

and narrow (Fig. 5;lc), Posterior dorsa! plate

divided by deep elongate natch of variable form,

ef. f. lucksiana (Fig, 5:4), f. unisetata (Fig, 5:3) or

fF, farga (Fig, 5:5), Foot slender and two-jointed:

paired setae on distal margin of first segment: toes

mostly parallel-sided, tapering terminally to sharp

tip, Unci of trophi with four opposing teeth: vetitral

large teeth each have rudimentary touth; tips of

rami with inner minute denticulate comb:

retrocerebral sac large (Vig, 3b:re, cf. also Fig. S:4ik:

brain with red eye on dorsal side

Intraspecific forms are described exclusively

according to differences in the shape of the lorica

and cross sections, however research by Parise (1963)

suggests that different morpliological types are

correlated with ecological differences, This alsa may

apply to some distinguished species; & parva

Rousselet, tor example, resembles &. difatate and

is distinguished from it by smaller size and long

slender toes. All known measurements fall within

the range of variation of &. dilutata; it is seen as

un ecotype of £. dilatuta,

Lorica leneth 140-320 ym, dorsal plate width

100-253 am, ventral plate 95-172 am, toes 50-100

am, posterior potch 12-61 win Weep; male 115-148

W. KOSTE & R, J. SHIEL

ROTIFERA FROM AUSTRALIAN INLAND WATERS 93

Fig 5: 1, Euchianis dilatata dilaratad Ehrenberg: (a) dorsal; (b) toe; (c) cross-section. , E, dilatata macrura (Ehrenberg):

ventral. 3, F: dilatata t, unisetata (Leydig): (a) dorsal; (b) cross-section; (c) toe, 4, E. dilatata f. lucksiana (Hauer):

(a) cross-section; (b) dorsal; (c) ventral; (d) trophi, apical; (e) swimming; ({-g, k) cross-sections; (h, j) different

lorica forms; (i) @. difatata, lateral, swimming. 5, £. dilatata larga Kutikova: (a) dorsal; (b) toe. Fig: lta—c, 2, 3a-c,

5a, b, after Kutikova (1970); 4a-d, j, k, afler Hauer (1930); 4e—1, after Myers (1930). Scale lines 100 zm.

Fig. 4: 1, Euchlanis alata Voronkov: (a) without wings, ventral; (b) with wings, ventral; (¢) toes (d) ventral: (c) habit,

lateral; ({) cross-section. 2, &. calpidia (Myers): (a) dursal, swimming: (b) trophi, apical; (c-f) crdss-sections; (2)

lorica, ventral. 3, E. deflexa Gosse; (a) dorsal; (b) lorica, ventral; (c, 2, j), cross-sections; (d) trophi, apical; (e)

tog, lateral; (Ff) other trophi; (h) dorsal; (i) resting egg; (k) lorica, broad form, ventral; (!) toe. 4, E. deflexa larga

(Kutikova): (a) lorica, ventral; (b) toe. Fig. 4:1a, d-f, afier Yamamoto (1953); 1b, ¢, 3j, |, 4a, b, after Kutikova (1970);

2, Ja-e after Myers (1930); 3f-h, after Donner (1964), Scales lines 100um (adults lorica in each number group),

ud W. KOSTE & R, J, SHIEL

jem, toes 34-240 ym. [E, parva Rousselet, 1892;

dorsal plate length 140 pm, width 100 «m, Yentral

plate length 125 yin, width 75 zr, tes 70-yin, depth

al posterior notch 50 pr).

Pistriburion: Cosmopolitan in fresh and brackish

waler! pH 4-10, Cl 140-4 1-1 (Koste 1978). The

Most common and widely tolerant euchlanid in

intaid waters; pancominental, including Tasmania,

8.0-25.9°C, pH 35.44-7,9, DO 3.6-13.8 mg I-|,

23-1600 4S cm-1, 2-235 NTU.

“iteratures Myers 1930; Mauer 1935; Kutikova 1970;

Ridder 1972; Kaste 197%: Shiel & Koste 1979 Koste

& Shiel 1980.

Evettants tnctse Carlin

FIG. 6:1

kuchlenis tncisa Carlin, 1999, p. 17.

Euchlanis inguetra Hudson & Gasse, 1389, PT. 23, Fig.

Diagnosis Body ovoid in shape, truncate in front

with variable anterior notch; dorsal plate triradiate

in cross-section with median keel extending from

neck to angle of posterior y-shaped notch. Ventral

plate completely developed, Longitudinal deep sulci

conmect bath plates; foot rwo-yointed, setae on first

foot joint; toes slender, fusiform; trophi (Fig, 6:le,

hy) with five teeth on cach uncus, First and last (Fig.

6:1h) have accessory shorter tooth; minute combs

on rami tips. Male (Fig. 6:11) with normal anatomy

for genus (cf. Fig. 3:3),

Length of dorsal plate 210-270 ym, ventral plate

180-240 pm, lorica width to 240 wm, toes 70-85 wm;

male 145 pm, toes 26 pum.

Listribation: Cosmopolitan in the littoral,

occasionally in the pelagial of shallow ponds and

billubongs Rare, N.T., Tas., Vie: 13.5-29.4°C, pH

5.3-7.4, DO §.45-8.2 mg 171, 23,0-80.2 pS cem-1,

2.0 NTU.

Comment A variant, & triguetree mucronata (=

& incisa mtucronate) (Vig. 6:2a, b} described by

Ahlstrom (1934) from the Neotropics may be an

ecotype; it resembles & ineiva and may co-occur.

The dorsul plate has an elongated keel. Length of

dorsal plate-to 320 am, loriva width ty 252 pm,

height to 120 amy; toes lo 120 pn. Not yet recorded

trem Australia.

Literature; Mycrs 1930; Wulfert 1976; Kessie 1974,

197h, 1981; Shicl & Koste 1979; Berzins 1982.

Buenlanis lyra Hudson

FIG, 6;3a~j

fMuchlarus lyre Hudson In Hudson & Gasse, 1886,

ao Fig, 23:1

£ myersi Kutikova, 1959, ys, 224,

Oieunosis: Body clongate and ovoid: cross-sevtion

resembles are of circle; lateral sulci present; dorsal

plate without posterior notch; ventral plate totally

developed, somewhat constricted caudally; foot

slender, two-jointed; paired long sctae on first Foor

joint; ophi with five paired unequal tceth, with

{wo or three accessories; rami tips with minute

inside comb, Length of toes variable.

Length of dorsal plate to 335 jm, ventral plate

302 um, lorica width 110-180 pm, toes 76-90 xm,

toe width 14-16 pm.

Distribution: Cosmopolitan in inundation areas,

also in periphyton, Single record from L. Mulbwala,

Vie. 12.0°C, pH 7.5, DO 10.7 nig I-71, 46 pS emn,

17.5 NTU, Variants ure known in che northern

hemisphere, ez. 2 dyravryersi (Kutikova 1959; 223)

(Fig. 6:4) a Palearctic subspecies with long slender

toes (86-107 pani long, 7-8 pm wide) and Lf. lyr

larga (Fig. 6:3k, 1) (Kutikova 1959) from N. Siberia.

Variants aré Hol recorded from Australia.

Literature; Myers 1930; Pejler 1962; Koste 1978;

Shiel & Koste 1979.

Fuchlanis meneta Mycrs

FIGS 7:1,2

Euchlants menera Myers, 1930, p, 378, Pl19, Fizs 5-8

buchianls evophe Lucks, 1912, p. JOS, Fig. 31

Euchlanis proxima Myers, $930, p 377, Pl. 19, Figs bo.

Diagnasis> Shape nearly ovoid; well-developed

ventral plate almost as large as dorsal plate; cross-

section seiiicircular arched bul alsi) roughly

tritradiate; dorso-ventral width of Iateral sulet

(relatively) uniformly wider than wm any other

congener; cuticular shield-like process bencath

posterior notch above first foot joint; posteriar

notch of dorsal plale very deep and often large (Fig,

Tk), foot rwo-joinled; toes very lang and slender,

sometimes with minute claw at lps. Maye

colourless, with short toes; dorsal and ventral plate

still conspicuous; brain long with red eye; two

Sensory hairs on Uny Lubules in apical lield, as in

female. Trophi (Fig. Tle) with four main teeth,

aecessorics not Vistble; minute denticulate combs

on inside of rami tips (Figs Tle, 2d),

Leneth of dorsal plate 10S-473-hm, width 80-140

yim, depth of posterior notel 35-45 am, width of

ventral plale 50-90 pm, toes 60-75 pm, width of

toos 3.5 xm, Male total Jengrhy 170 um, toes 25 um

Distribucow: Cosmopolitan, mostly in aeidie

walers; rare in submerged Spaagiuwm,; N, America,

W. Germany, NSW. NLT, Vie WA. 10,0-29,99C

pH 5,.3-7.4, DO $.45-10.1 mel 3, 20-523 aS cm |

22 NTU, alk, 2.6-2,7 mg L |

Literature: Myers 1930; Hauer 1935 (yrraxinia);

Wullert 1991; Pejler 1962; Berzins 1963, 1982; Koste

1978, 1981; Koste & Shicl 1980,

Exenlanis oropka Gosse

rig. Bl

huchlanls oropid Liosse, TSR7, p. 3, Pig. 2:16,

non Luchlanis vravhy Lucks, 1912, p. 10S, Fig. 31.

ROTIFERA FRQM AUSTRALIAN INLAND WATERS 95

Diagnosis; Resembles EF. ditatata but usually

smaller, with stouter foot and more robust,

differently-shaped fusiform toes; lateral antennae

suluated in cavities beside somewhat keeled end of

dorsal plate; (rophi construction as &. dilatare (et.

Figs 3e, 5:4d). Comb-like processes on inside of

each ramus tip; four apposed club-shaped

functional teeth,

Length of dorsal plate 164-266 pm, width

127-172 xm, ventral plale length 123-225 wm, toes

49-78 pm, width 1-14 pm, posterior notch 24-29

Distribution, Cosmeopalian. NT, Qld, Vie.

8,0-18.5°C, pH 48-84, Dt) B4-10,0 me b-),

67-400 pS em |, 45-160 NTU.

Luerature: Myers 1930, Kutikova 1959; Koste 1978;

Shiel & Koste 1979; Berzins 1982,

Euchlanis pocameneia Berzins

FIG. 7:3

Euchlanis perameneie Werzins, 1973, p, 126, Figs. 4, 5,

Diagnosis; Resembles E, meneta (see above). ‘Toes

with pseudoctaws; unterier margin af dorsal plate

with two short keels.

Length of dorsal Jorica 95-104 pm; ventral! plate

70-72 am; width, dorsal 72-75 pm, ventral 50-55

ym; anterior margin width 35 pm, caudal opening

26-30 xm long x 18-25 pm wide; toes 45-50 p.m.

Distribution: Recorded from New Zealand, not yet

known trom Australia,

Luchlanis phryne Myers

FIG. 8:2

Euchlanis pkryne Myers, 1930, p. 372, Pl, 14, Fig. 4s

Pl. 15, Pies |-4

Diagnosis: Dorsal lorica oval, with slight

constriction of median lateral margin; deep inverted

L-shaped caudal notch almost a$ long as toes:

sciftened yentral plate connected to dorsal plate by

longitudinal sulel; toes short, fusiform, ca, '4 length

of dorsal plate; trophi with four club-shaped teeth

on each ramus; rami with fan-shaped denticulate

combs on inner distal margins.

A superficial resemblance to E, pyrifermis and &.

ealpidia was noted by Myers (1930), and Koste

(1978) considered E. phryne 4 possible variant of

£, calpidia or E. dilutate. Comparison of lorica

morphology of the species above, particularly cros¢-

sections, and differences in trophi structure,

indigates that E, phryne is a distinct species.

Length of dorsal plate 225 wim, ventral plate 210

am, dorsal width 180 pm, ventral plate width 150

pm, toes. 65 am,

Distribution: North Americ# (Maine), single

lueality, a billabong of the Goulburn R. near

Sevmour, Vic, in spring 1976 and again in summer

1973, 17,0-22.0°C, pH 7,1-7.4, DO 8.7-9,8 me b 1,

Comnrent: The above anomalous distribution is

noteworthy; £. phryne is one of sevetal rotifer and

microcrustacean species recorded from Goulburn

River billabongs with similar disjunct distributions

(Shicl 1976 and unpublished dara; Koste 1979). The

importation of heavy dam-building machinery by

the Utah Construction Company in the mid-1950"s

(during the consituction of Eildon Dam) ts

implicated a3 a possible source of transport uf

resting eggs/ephippia. Populations have become

established ity sheltered billabongs downstream of

the dam. site,

Euchlanis pyriformis Gosse

HIG) 8:3

Evchlaris pyriformis Gasse, 1851, p. 201

Dapidia pyrrifermis Myers, 1930, p. 370, Fig. 15:5-7.

Didenosix Shape nearly circular, may have shelit

emaremations instead of caudal noteh; lareral

margins turned downwards, occasionally pinched

in the middle; ventral plate rudimentary, only

developed posteriorly; no lateral sulci; foot twe-

jointed, pwo long pairs of setaé on dorsal side of

the first Foot segment; toes relatively short; trophi

resembles that of &. deflexa (cf. Fig 4:3F).

Length of dorsal plate 285-320 pm, width

275=<315 zm, tocs 80-85 pm, toe width 10-13 um.

Distribuiion> Cosmopolitan in the littoral,

occasionally pelagial. Vic, Tas.

Literature: Kaste 1978; Berzins 1982; Stel & Kose

LORS.

Euchlanis triquetra Ehrenbere

FiG. 9)

Euchilanis triquetre Ehtenberp, 1833, 0, 461, Pig. $7:8.

E, pellucida Harring, 1921, p. 6, Fis. 2,

E, langubordica Manfredi, 1927, p, 24, Pin. 7h

Panidia carinata Cartin-Nilssun, 1934) pW. 6, hig. 2.

0, lata Carlin, 1939, p. 14,

aa eae iriquerta plerigaides Grese, 1455, nh &\, cigs

non-F. (riquerr after Ruttier-Koliska, (972, p, 182, Fig.

Diggnesis: Body in dorsal view circular, sometimes

somewhat elongated (Fig, 9:1); triangular with a

high keel in cross-section; median keel extends fron

neck 10 caudal region; dorsal plate laterally with

wide Flanges (Fig. 9:te-e); no ventral plate, na

posterior notch, no lateral longitudinal sulci; some

specimens With semicircular ventral line over base

of toes; foot obscurely two-jainted; toes long,

slender, nearly straight, ending in abrupt points

most loricas very transparent with coloured inner

organs (retrocerebral sac dark, mastax yellow,

stomach olive green, yellow or brownish), Male (Fig.

33n0) with cuticular plates and dorsal keel, Trephi

ef. Figs 9:lj—n) tps ef rami with minute verbs (Fig.

S1j-k).

W, KOSTE & R. J. SUTEL

ROTIFERA FROM AUSTRALIAN INLAND WATERS 97

Lorica length 300-710 am, width to 540 pam, toes

90-15 wm, lorica height! to 300 pm, trophi: rami 40

am, folerum 32 ym, marubria 30 pm.

Disirihution: Cosmopolitan in the littoral,

sametimes pelagic in shallow waters, NSW, Qid,

NG, Vie.

Eiteralure: Pejler 1962; Kosice 1978, 1981; Shick &

Kaste 1979: Berzins |982,

The following species are not considered here:

£. arenoya Myers, 1936, known. only fram N. America

and ?Volga;

£. bioculita Issel, 190} (nosmen nude: name published

without description, and apparently tater synonymized

with &. plicata Levander by Issel (1906). Incorrectly cited

in Voigt (1957) and Koste (1978),

& cullysta Myers, 1930 (Fig. 9:4) is possibly a juvenile

form of & triquetra, Only known from the Nearctic.

Length of dorsal plate 170 xm, lorica width 80 pm, toes

40 pm. (Myers, 1930; Kosre, 1978);

£. callimorpha Berzins, 1957 (Fig. 9:2) resembles, £,

fncisa, only known fronv Gambia, W, Africas

£. contorta Wolter, 1939. known only from central

Germany and the Volga estary;

£ dapidula Parise, 1966, see Koste 1978 (misquared as

Parise, 1963). Doubtful species;

E_ hyphidactyla Pavise, 1963, see Koste (9738, Doubtful

species.

£. Hieulata Kutikova & Vassileva, 1982, endemic in Lake

Baikal;

&. mamorokuensis Berzins, 1973 (Fig. 9:3), recorded

from Madagascar;

E. mikropous, Koch-Althaus, 1962

ceniral Germany;

£, pannonica Bartsch, 1877, known only from Hungary,

single record;

£&. perpusilla Ridder

Caribbean;

E turfoysa (Rodewald,

Romania.

5 recorded only fron

1977, known only from the

1940), knawn only from

Family Mytilinidae Bartos

Loricate rotifers; cross-sections of lorica mostly

triangular or nearly rhombic; ventral plate and

dorso-lateral plates firmly fused: long dorsum with

or without sulcus, latter common with double keel;

three or less foot sections; toes pointed, straight or

shehtly curved ventralwards; malleate trophi, AU

species littoral and benthic, occasionally (but rarely)

in the plankton. Two genera (see Bartos 1959,

Kutikova 1970, Koste 1978),

Key jo penera of Mytilinidae

1. oriea thin or rigiet with dorsal sulcus and double keel;

lofida Wnornamented; tocs fong

Myltilina Bory de St Viticent

Lorica without dorsal sulcus, one strong keel; lorica

ornamented wilh dislinet pattern ard cavities; toes short

J uphtcharis Ehrenberg

Genus Afytitina Bory de St Vingent

Mytilina Bory de Si Vincent, 1826, p. 87 ¢=Selpina

Ehrenberg, 1831), p, 46 = Diplux Gosse, 1851, 9, 201 =

Diplacidium Lauterborn, 1913, p, 483).

Two recognized form-series (“Formenkreise"),

One series strongly loricate, usually with variable

stiff anterior and posterior spines; anterior margin

of lorica generally granulated; toes stiff and sword.

shaped, Second group thinly loricate without

anterior spines (except ML acanthophera, (Fig. 10.1,

2) anterios matgin not granulate, toes mostly long

and slightly curved, flexible.

Key Species of the Genus Mytilina

L. Lorica stout, anterior margin granulated, wes

sword-shaped sy...) oc eee ee ee eee

Lorica thin, anterior margin not Hrartulated, toes

long and curved. q

21), Antetior lorica margin with two short dorsal and

two short ventral spines, Mf. rnvcronata Muller

Anterior lorica margin with only two ventral spines

-.. M. ventralis (Ehrenberg)

31). Ventral margin af ‘lovica with variable projections

M, acunthophora Hauer

Ventral anterior margin without projections. ..4

AM), Toes with claws...., 202002.

Toes without claws, ...-- re 6

3(4), Head part lpricate; 3 foot segments

.M. crassipes (Lucks}

! foot scaments

-M. ungripes (Lucks)

A(4). Toes relatively shorl; ‘anterior fel in of lorica with

Lf. CS eee eee WsWleata Lucks)

Tocs long, straight, anterior Pack without falds

(iesembles Miurcuathophara).M. trigona (Gass)

phage

Head part iNarivaie; 2

Mytilina ocanthephora Hauer

FIGS 10:1,2

Moetifivg acunthophora Hauer, 1948, p, $50, Figs 73a-c.

Diagnosis: Dorsal lorica strongly arched,

granulated; ventral lociva fattened, with depression

beneath head opening; triangular lorica projections

flank medioventral head aperture (Fig, 101b);

shield-like projection over two-segmented foot; toes

Tong, pointed,

Length: 195-253 jum (total); to ISO ym (lorica):

height to 92 wm; width to 64 um; toes HO-94 yr,

An unusually large specimen from Lago Grande,

Amuzonia had the following measurements

(sequence as above); 342, 198, 125, 137, 140-144 win

(Koste unpubl).

Distribution: Probably pantropical in polysaprobic

waters. Not yel recorded from Africa, Occasiarially

Pie. fi: 1, Euchlanss incisa Carttin: (a, 6) lorica cross-sections; (c) yeniral, (d) Tateral, swimming; fe} trophi, apical;

(1) dorsal; (uw) toe; (fh) uncus; (i) male. 2. & incisa mucronata (Ahlstrom); {a) dorsal: (b) Jateral, 3, &. dene Hudson:

(a) dorsat; (b) veniral; (c) trophi, apical; (d, c) cross-sections; (f) dorsal, swimming; (g, h) Jorica, ventral; (i) toe;

(j) tae, (kK) E dvrw darga (Kutikova), vencral; (1) toe 4, E. Gira myersi (Kutikova): (a) yeniralt (bh) another form,

ventral: (c) trapihi, Spical, (d)} cross-section; (¢) toe. Fig. 6: la-t, 3a, b, da-d, Myers (1930); if, uy Joel, de, after

Kutikava (1970); 2a, b,

, alter Koste (1974), Svales lines 100 am (acule jortea in each number group).

98 W. KOSTE & R. J. SHIES,

Fig. 7; 1, Buchlanis meneta Myers: (a) lateral; (b) dorsal; (c) ventral; (d) lorica cross-section; (e) trophi apical; (ft)

lorica ventral; (g) cross-section; (h) lateral, swimming; (i) lorica dorsal; (j} toe; (k) dorsal, swimming; (1) dorsal;

(m) cross-section; (n) male; (0) male, lateral. 2, E. proxima Myers; (a) lateral, swimming; (b) cross-section; (c) ventral;

(d} trophi, apical; (e) dorsal; (f) ventral: (g) dorsal. 3, & purameneta Berzins: (a) dorsal; (b) ventral. Fig. 7:la-e,

2e-g, after Hauer (1935); Lf, g, i, j, after Kutikava (1970); Ik, 0, after Wulfert (1960); 3a, b, after Berzins (1973);

2a-d after Myers (1930). Scale lines, top, 100 xm (Fig 7:1, 2), battam, 50 wm (3a, b).

ROTIFERA FROM ALSTRALIAN INLAND WATERS 99

in plankton sansples. Qld (Townsville, Mr Isa).

Literature: Koste 1978; Shiel & Kuste 1985,

Murlling bisyleata (Lucks)

FIG, 10:3

phx bisuleara \acks, 1912p. 95, Figs. 18a-c.

M. trigune alter Harring, 1913 p. 75.

Diagnesis: Lorica with folds on anterior margin;

three weak dorsal keels in cross-section; sulcus

indistinct, toes shorter than in M. tertguipes: with

Jong tips (cf M. wnguipes (Fig, 1232).

Length t30-180 jam, toes 60-70 pm, tips 8-14 wm.

Distribution: In peat bogs, areas of decomposition.

ACE, Vic.

Literatures Hauer 1936; Koste 1978: Koste & Shiel

1980; Berzins 1982; Tait ev al 1984.

Mynilina crassipes (Lucks)

FIG. 10:4

Diplax crassipes Lucks, 1912, p, 96, Fig. 30,

Mytilina orassipes after Carlin-Nilsson, 1934, p. IL.

Piagnosis: Lorica laterally compressed; head with

two lateral rounded plates; trophi small, uncus with

six teeth; mouth opening with palpar organs;

stomach and gastric glands large; two black spots

at “forehead” region (eyes),

Length 220-280 am (total); lorica 138 pum (oes

57-75 pln; claw 9 jam; (rophi 21-30 wm,

Aistribulion: Cosmopoatiian in polysaprobic waters,

occasionally in shallow-water plankton. N.T.

(Magela Ck).

literature; Kuste 1978, |98l; Koste & Shicl 1980;

Tait er al, 1984.

Mytitina mucronata (Muller)

FIG. 11

Grachionus mucronatig Miiller, 1773, p, 194.

Maitilina otucronata alter Holsten, 1908, p. 54.

See Koste (1978) for extensive synonymy.

Diagnosis; Shape of lorica variable. of Little

taxonomic valuc (eg. MM. mucronata var. Spirigera

(Ehrenberg, 1832)); corona as in Evchlanis; lorica

granulated anteriorly: dorsally curved spines visible

in lateral view beside foot opening; keels eud in

short hook; foot three-segmented; large cerebral eye

present (Pig, Wila eve lateral anrennae indisriner:

dorsal antenna (Fig Lh:ta da} visible; trophi of

malleate type, unci with five teeth; mastax (Fig. Ula

mx) with salivary glands; stomach cellular with

small gastric glands (Fig. Il:fa gy), Male

approximately half female size, also loticate with

cerebral eye (ct. (Fig. 3:1, m)).

Length: 170-250 am, height 96-100 um, toes

33-60 pm, subitanedus egg 80 xm, male to 135 zm.

Oistribution; Cosmopolitan in eutrophic waters,

also on the surface of decomposition areas between

witler plants and in brackish waters. Occasionally

in the plankton of shallow ponds and billabongs.

Vic, 13.5-22,0°C, pH 7.1-7.4, DO 44-102 mg [-1,

240 pS om-), § NTL.

Literature: Evans 1951; Koste 1978,

Myciling trizena (Gosse)

FIG, {2:1

Diplax trigama Gosse, 1851, p. 201.

Mytiling trigona after Harring, 1913, p, 75.

Diagnosis: Surface of lorica slightly datted;

antcrior margin with rounded edge ventrally (ef M.

deanthophora Vig, 101, 2; cress section almost

triangular; narrow double keel; toes with long tips.

Tolal length: 245-245 an, lorica length 150-160

aM, Toes 68-83 pum.

Distribution: Cosmopolitan in decomposi-

tion/inundation areas, Qld.

Literature: Koste 1978; Shiel & Koste 1979; Berzins

1982,

Mytilina unguipes (Lucks)

FIG. 12:2

Diplay wnguipes lacks, 112 p. 96.

Movtiting bisuleara > unguipes (Lucks) after Carlin

Nilsson, 1934, p. UL

ae ie unguipes (Lucks) afler Wiszniewski, 1983, p,

Diagnosis: Anterior margin of lorica without folds,

laterally compressed; foot with two joints; toes long,

slightly curved with distinct, inflexible claw,

Doubtful species, possibly identical with MW.

hisulcuta,

Length 193 ym, height 94 wm, toes 58 xm, claw

1} pm,

Distridition: European peat bog, nor yet recortled

from Australiz.

LEiferarure: Kutikava 1970; Koste 1978

Mytitina vertiralis (Ehrenberg)

PIGS 1:3, 4,5

Salpinia verrratis Ehrenberg, 1832, p. (33, Pl.4, Fig. 7,

yf ana veatralis (Ehrenberg) after Harring, 1913. p.

"For extensive synonymy sce Harring oe. cit.) and

Kunrikova (1970:519-520),

Diagnosis: Lorica margin granulated; shape and size

of lorica and spings very variable; only ventral

anterior spines, Short-spined form (var, brevispina,

176-215 pm) shown in Tig, (:3a-c; long-spined

form (var, macracantha, 250-350 um) shown in Fig,

11:4 Pig. 15 shows rare form described by Wulfert

(1965) (M. ventrutiy f, longidactyla) with short

anterior but long median-and shorter caudal venual

hooks, Lorica (90-115 pm) without spines, median

caudal spine 62 zm, ventral caudal spines 48 ani,

tues 72 wm. These forms may be ecorypes,

Distribution: Cosmopolitan. Probably

pancontinentul, not yet recorded from S, Aust. Cf.

typ. from N.SW., N.T., Qld, Tas., View brevispina

from N.T., Qld, Vic; maecrecantha from NT, Gld,

W. KOSTE & R, J, SHIEL

ROTIFERW "ROM AUSTRALIAN INLAND WATERS tal

W.A. 135,.5-28,5°C, pH $.44-7.9, DO 5.8-11.6 me

1-1, 23-1500 pS om>!, 4,8+67 NTU, alkalinity 27

me th

Not yet recorded from Australia:

M. bicarinata (Perty, 1850)

M, macrocéra (Jennings, (894)

M. mutica (Perty, 1849)

MM. videns (Levander, 1894)

faee Kosre 1978 for details),

Genus Lophocharis Ehrenberg

Lophockaris Ehrenberg 1838, p. 458.

Strong lorica, surface soft, granulated, but may

have honeycomb structure; cross section nearly

rhomboid but specimens with flexible integument

nearly oval or compressed; dorsum always arched;

more or Jess distinct keel with and without

transverse folds; ventral plate with longitudinal and

transverse ribs; apical margin with and without

servation; ventral aperture rounded, dorsal more

triangular; foot-opening ventral; foot with three

juris: toes with sharp points; trophi malleate (Fig.

L3:Id); cerebral eyes only visible in living animals.

Littoral and benthic in habi(, seldom found in the

plinkion, Seven species are Known, four have been

recorded from Australia,

Key to Speeies of the Genus Lophocharis

L Lonca >120 um long with distinct dorial heel

producing, Wianaular cross-section. a"

Lorica 100 pm lone, kel indistinct, crass- -Keehion

more elliptical, . --.. 4) curvaia Berzins

20) Anterior margin ‘at lonce sironely serrated, dorsal

keel with transverse folusL. selpina (1s hrenberw)

Anterior margin smooth or lizhtly serrated, dorsal

keel without falds.......-...--..-.--.-2-- 3

(2). Lorica sinooth, dorsal noteh of head aperture V-

shaped, ventral a brouder L-shape; fine striae on

both sides of dorsal keel..... “, euias Wulferi

Lorica sculpiured as 1. sulpina; head aperture

notches V-shaped: in striae beside keel... 2.

oxpsiernan (Gossey

Lophocharis curvata Berzins

FIG, (4:3

Lophocharis curvata Berzins, 1&2, p. 12, Fig. '5a—c.

Diagnosis: Small species, lorica clongated U-shape

without typical dorsal keel, cross section more

dorso-ventrally flattened than in congeners; head

aperture margin with dorsal und ventral sinuses,

ventral slightly broader; foot aperture broadest

posteriorly; tacs thin, curved outwards.

Lorica length 95 ym, width 55 pm, toes 18 pm,

foot apetture width 32 xm,

Distributions Endemic; single record from King

Parrot Ck, Kinglake, Vic. 18.x,53,

Comment: Although the original description and

figures of this rotifer do not meet the requirements

of the Code, we feel that the distinctive dorsum,

foot.aperture and small size warrant retention of

spectfic status until material can be examined.

Lophocharis naias Wulfert

FIG, t4

Laphochoris naias Wulfert, 1942, p, 188, Fig. tak.

Diagnosis: Dorsal lorica with shallow keel and

lateral longitudinal striations, surface morphology

Otherwisesmooth, unornamented; apical margin of

lorica finely serrated; uncus with 7 teeth) inner

margin of rami with Fine denticles.

Lorica length to 170 wm, widith to 105 wm, toes ta

21 um.

Distribution; Europe, Single record, from Coongie

Lakes, S. Aust.,, coll, Jane Roberts, Botany Dept,

Univ. of Adelaide, 06.21.86.

Literature: Koste 1978,

Lephocharis oxysternan (Gasse)

FIG, 13:1

Metopidia oxysternon Gosse, 1851, p, 201,

Lophocharis oxysternon (Ciosse) after Harring, 1916,

). 564, Fig, 976-13,

Diagnosis: Surface of integument soft or weakly

pusiulated; ventral plate with deep depression over

foot-opening. Many intermediate forms to L.

salpina, e.g. with slightly serrated apical margin,

Trophi (Fig. 13:1d-c) with six uncus teeth.

Lonica length 120-200 pm, toes 24 pm.

Distribution: Cosmopolitan, benthic in fresh and

brackish waters, occasionally in plankton. Vic.

Literature: Koste 1978, Berzins 1982.

Lophochuaris salpina (Ehrenberg)

FIG, 13:2

Lepadetla saliping Ehrenberg, 1834, p. 205

Metypidia salpina after Hudson & Gosse, 1889, p\ 46,

Fig, 34:4;

bs 7 el saline (Ehrenberg) afler Marring, 1916,

For extensive synonymy see Koste (1978:153)

Diagnosis: Axterior margin always serrated; keel

with transverse folds mostly aver caudal part of

loriva; variable, intermediate forms to /,

oxysternon, traphi also with six uncus teeth: foor

with three joints; tocs sharply pointed, ventrally

curved.

Length 175-224 pm, borica 120-135 wm; wiklth

Fig 8: |, Euchlants aropha Gosse: {a) dorsal; (4b) dorsal; (c} cross-section; {d) alternate ane loricas; {e) Lroplii,

apical, 2, £, phryne Myers; (a) dorsal; (b) ventral; (¢) trophi, apical; (d) crass-seerion, 3

»& pyelformis Gosre; (ay

VeEriyal) (by) larica cross-sections (¢) rrophi. 4, &. ewtlyste Myers: (a) lateral; (b) eross-stctian: (c) trophi; (d) ventral

Fig. S:la, after Kosiv (1978); led, after Donner (1964); le, 2a-d, 3a-c, 4a-d, after Myers (1930). Seale lines 100

um (Pig. 81-3); 50 pm (44, b, d),

102 W. KOSTE & R. J. SHIEL

3b 3c

Fig. 9: 1, Euchlanis triquetra Ehrenberg, (a) lorica, ventral; (b) lateral; (c—c) different cross-sections; (f) dorsal, swimming;

(g) toe; (h) male, contracted; (i) male, lateral, swimming; (j) trophi (fulcrum and rami); (k) ramus; (1) manubrium;

(m) fulcrum; (n) uncus. 2, &. callimorpha Berzins, dorsal. 3, &. mamorokaensis Berzins: (a) dorsal; (b) cross-section;

Wulfert (1956); 2, 3a—c, after Berzins (1973); 4, after Ridder (1977). Scale lines 100 zm (adult lorica in each number

group).

ROTIFERA FROM AUSTRALIAN INLAND WATERS 103

Af 4g 4h

Fig. 10: 1, Mytilina acanthophora Hauer from Magela Ck, N.T.: (a) ventral; (b) lateral; (c) cross-section; (d) lateral

(cf. 3a, b). 4, M. crassipes (Lucks): (a) lateral; (b) dorsal; (c) lateral; (d) dorsal; (e) lorica, contracted; (f) trophi;

(g) trophi, lateral; (h) uncus; (i) mouth with palpar organs (m = mouth). Fig. 10:3, after Hauer (1936); 4, after

Wulfert (1939). Scale lines [00 wm (adult lorica in cach number group).

104 W. KOSTE & R. J. SHIEL

3c 3b

Fig, 11: 1, Mytiling mucronata (QO, F. Miller): (a) swimming, laleral [b/ = bladder, br = brain, de = dorsal antenna,

eve = cerebral eye, f= foot, 2g = gastric gland, / = intestine, mx = mastax, st = stomach, ¢= toe, vi = vitellarium);

(b) male, swimming, lateral; (c) lorica, lateral; (d) lorica, dorsal; 2, Myzilina sp. cross-section. 3, M. ventralis

(Ehrenberg): (a) lorica, lateral; (b) ventral; (c) male, dorsal. 4, M4. ventralis macracantha (Gosse) lorica, dorsal.

5, M. ventralis longidactyla (Wulfert), lateral, from E. Africa. Fig. 11:la, 2, 3a, b, 4, after Koste (1978); 1b, c, after

Feiler te jd, 3c, after Weber (1898); 5, after Wulfert (1965), Scale lines 100 um. Top Fig. 1l:lb-d; left, 3, 4;

ottom, 5.

ROTIFERA FROM AUSTRALIAN INLAND WATERS 105

Fig, 12: 1, Mytilina trigona (Gosse): (a) lateral; (b) ventral;

(f) ventral. 2..M. zguipes (Lucks): (a) lateral: (b) cross-

section. Fig. 12:1, after Donner (1954); 2, after Hauer

(1936). Scale lines 100 zm,

80-95 um, toes 25-40 pm.

Distribution; Cosmopolitan, benthic in fresh and

brackish waters, also in polysaprobic pools and

ponds. Pancontinental, 10,5-30.4°C, pH 5.6-8.4,

DO 3,7-11.0 mg 1-1, 28-3330 «S cm~-1, 5~92 NTU,

alkalinity 4.0 mg 1 1

Literature: Evans 1951; Shiel & Koste 1979; Koste

1981; Berzins 1982.

Not yet recorded from Austratia;

Lopkocharis hutchinsoni Edmondson,, 1935

L, naias f. ambidenta De Ridder, 1960

L. parva Rudescu, 1960

L, rubens Wulfert, 1939

(see Koste 1978 for details).

Family Trichotriidae Bartos 1959

Three genera with loricate head and body; surface

with facets, mostly granulated and with spicules or

spines, sec. on dorsum; foot freely movable or with

stiff joints; trophi malleate. Generally occur

between aquatic macrophytes and in periphyton. In

plankton only as migrants.

Vig. 13; 1, Lophocharis oxysternon (Gosse): (a) dorsal; (b) ventral; (v) lateral; (d) trophi, apical; (e) trophi, lateral.

2, L. sulpina (Ehrenberg), (a) swimming, lateral; (b) lorica, lateral: (c) lateral; (d) another lorica, lateral; (e) cross-

section. 3, L. curvata Berzins, (a) lorica, dorsal; (b) cross section; (c) lorica, ventral. 4, L. naias Wulfert, (a) torica,

dorsal, (b) lorica ventral. Fig, 13:1 a-e, 2b, after Harring (1916); 2a, aller Wulfert (1956); Id, 2c, after Haver (1925);

2d, after Klement (1960); 3 after Berzins (1982); Scale lines, centre, 100 pm (la—c, 2-4); lop right, 10 pm (d, e).

106 W KOSTF & R. 1. SHIFI

Fiv. 14) Wolea spinifera (Western): (a) dorsal, swiinining;

(b) lateral; (c) ventral; (d) cross-sectian a-c. aller

Western (1894); d, after Vranovsky (1257), Seale line,

$0 yarn,

Key tu penera of Trichotriidae

1. Lorica with anal segment... ....-.. a ,

Lorica without anal segment, ,., Wo/gu Skorik

211), Lorica with distinct elongated spines an dorsum

ya sonn fie toe 90rd Macrachaelus Perry

peeesuy Erichelria Bory de St Vincem

=p ib eee

Genus Holga Skorikuv

Wolgd Skorikay, 1903,, p. 37,

Monospecific, Holem spinifera.

Holga spinifera (Western)

FIG. 14

Distvlu spinifera Western, [894, p. 427, Fig. 241-4.

Cuthypna spinifera after Murray, 1913, p, 545-54.

Lecane spinifera afier Harring, 1913, p. 62.

Mala spintferu (Wentern) after Skorikov, 1903, p, 37,

Diagnosis: Dorsal Jorica divided by ribs into panels

or facets; foot not loricate, freely movable; ventral

plalé with oval foot-opening undsy transverse

distinct line; toes shart, slender, painted. Cross-

section of body somewhat compressed (Fig. 14d).

Male unknown.

Total length 20)-255 um, lorica 95-145 jum, width

70-90. ym, spicules 5-6 pm.

Distribution: Cosmopolitan in fresh and brackish

water, Tately in the pelagic. Twa localities, probably

more widespread, but tare. N,SW., Tas, 10.5-13.0°C,

pH 6.7-8.4, DO 10.0 mg! 7.

Literature: Koste 1978, Kaste & Shiel [980,

Genus Trichotria Bory de St Vincent

Trrehtointa Bory de St vincenl, 1827, p, 752.

Head, body and three foot segments heavily

loricate; cross-section hexagonal (Fig. 15:1f) (see

comment with 7) buchneri for exception); Most

lorica borders with spicules, grearer on “shoulders”

if head contracted (Fig. 15:3, 4a); lorica with facets,

granulated; head lorica composed of several plates;

second foot segment with relatively long, spines (Fig.

\S:la Jsp); toes long; corona of Euchlanis type?

trophi (Fig. 15;ld-e) maleate; red cerebral eye (Fig.

15:]4-eye). Male loricate (Fig. 15:1b-c), Food algae

and detritus, diatoms preferred. Trichotria accurs

tarely in the plankton (as migrants). Seven species

recognized, with many variants; five known from

Australia.

Key to Species of the Genus Trichotria

|. Cross-sectian hexagonal, three foot segments. .2

projecting beyond posterior lorica margin; lorica

clongated U-shape...) pseudecuria Koste er al,

All three, of at least lwo, foal segments and toes

projecting beyond lorica margin.........--. - i

3(2), Last foot joint with dorsal minute spine between

the toe bases, 6.0.4, 6% poeiiun (Miller)

No minute spine between the tye bitses.. os... 2

4(3). Lateral part of dorsal Jorica conspicuously large;

marginal spicules of lorica directed forward

wey s ot TIM, dates Ob T. truncata Whitelegee

Lateral lorica not very expanded, spicules if present

not directed forward, ,.. 7) fetractis (Ehrenberg)

Trichetria buchneri Koste, Shiel & Tan

FIG. 14:1

Trichotria buchneri Koste, Shicl & fan, 1988, pp.

120-12, Figs 3-5.

Diagnosis: Lorica of triangular cross section with

median keel ending in elongated caudal spine;

dorsal lorica with longitudinal rows of denticles to

keel (appears granulated by light microscopy);

anterior dorsal margin deep U-shaped aperture

between two lateral cusps; ventral margin with

shallower U-shaped aperture projects beyond dorsal

margin to level of lateral cusps (or slightly beyond

lips); rectangular plales cover head, when

contracted a pyramidal projecrian extends to or past

anterior lorica margins; foor two-segmenred,

without spines; Loes long, ngid, with acute paints,

Length 160-182 pm; width to 125 wm; height to

84 ym; proximal foor segment 24 wm, distal 15 ym;

toes 50 um.

Distribution: Known only from marginal reedbeds

of acid dune lakes north af Strahan, on West coast

of Tasmania (1, Garela, L, Ashwood and

surrounding dune pools), 17,.0-20.0°C, pH 3.1-5.15,

80.6-188.4 nS em 1, 0.9 NTU.

Comment: The mutphological differences between

this species und congeners were noted by Koste e/

a, (1988), particularly the domed and keeled cross-

section, caudal spine and two-segmented foor, The

ROTIFERA FROM AUSTRALIAN INLAND WATERS 107

Fig, 14: 1, Trichotria buchneri Koste et al. a. dorsal, b. lateral, ¢. ventral; 2. Trichotria pseudocurta Koste et al. a.

dorsal; b. ventral. After Koste et af. (1988). Scale line 50 yam.

armoured head is peculiarly trichotriid amongst the type locality in Jan, 1989, and additional sites for

Rotifera, but the distinctive features of T buchneri the rotifer recorded. A detailed SEM study of this

suggest that generic status may be appropriate. species will appear at a later date (Shicl & Koste,

Accordingly, additional collections were taken atthe in prep.).

108 W. KOSTE & R. J. SHIEL

¢ os

ey it

—*

ROTIFERA FROM AUSTRALIAN INLAND WATERS 1c)

Trichoiria pocillum (Miller)

FIG, 16:2

Trichoda pocilhum Miller, 1776, po 281.

Dinocharis pocillum aller Ehrenberg, 1830, p. 47.

Trichotria pocitlum Bory de St Vinecnt, 1827, p. 752.

yen extensive synonymy see Flarring 1919) and Koste

Diggnesis: Length of font segments and spine on

seoond foot segment variable; lorica spindle-shaped,

surface divided into granulated facets by ridges,

corona simple, can be retracted in neck and lateral

head plates; male Ioricate. Rate in plankton.

Length 200-300 pm (total), lorica length 110-140

pm, foot spine 55-66 xm, median minute spine on

last foot segment 20-28 pm, toes $3-145 um.

Distribution; Cosmopolitan in the littoral of fresh

and brackish waters. N.S.W., Old_

Literature: Shiel & Koste 1979,

Trichotria pséeudocurta Koste, Shiel & Tan

PIG, 15:2

Trichotria pseudocurta Koste et al, 1988, pp, 121-3, Fig.

Diagnosis: Lorica small, U-shaped in outline with

both surfaces granulated; triangular cusps ar

external angles; two raised ribs on ventral surface

terminate anterior to raiset| tim of foot opening:

similar ribs form twin keel on dorsal surface,

commence either side of median notch on dorsal

anterior margin, fuse to a single keel on posterior

lorica; foot 3-segniented, without spines; toes with

short claws,

Length to 125 pm; width to 74 pm; ioes to 30 pm,

Distribution: Known only trom humic waters of

Lake Pedder (White Spur Inlet), Tasmania. 21°C,

PH 4.2, 32.6 nS em-1, 0.5 NTU, Resembles 7.

curta (Skorikow) from the Volga R., however the

latter is smaller (110 gm tong) with nore angular

morphology, and lacks the frontal cusps.

Wriehotria tetractis (Elwenherg)

FIG. l6é:1, 3

Dinacherty tetractis Ghrenberg, 1830, p. 47,

Trichatria rerractis (Fhvenberg) after Carlin. 193, p, 40.

For extensive synonymy see Koste (19781157).

Diagnosis: All morphological characteristics

variable; loricu spinulated or not; measurements of

lorica, foot segments, dorsal and ventral lorica

ornamentation difler between individuals, with

morphological series between T. ruacate and 7,

fetractis similis (latter has. very lang toes; 120-190

ui) (Fig. 17))) caudal part of lorica with one or

patred hooks (7) tetractis cvudate; spinules on

second foot segment may be very short (7 sepractis

Paupera).

Length typical form; 218-380 um, toes to 160 win;

T. tetractis similis (Stenroos, 1898): 330-40) jum,

second foot segment 42-48 pm, loes 120-190 ym;

T. letractis caudatar 230-260 pm, caudal hook of

lorica 22-28 «im, toes 50-72 um.

Distribution: Cosmopolitan in macrophytes,

littoral, Commonly in shallow waters of billabongs

in N.SW,, Qld, Vie. (typical form) and N.T.

(similis), 13.5-24,5°C, pH 6.3-8.0, DO 5.8-9.2 mg

1°), 59-400 «2S cm> 1. 2-160 NTU.

Literature: Koste 1978, 1981, 1984; Shiel & Kaste

1979; Green IY]; Rerzins 1982.

Trichotria truncata Whitelegge

FIG. |6:4a—b

Dinacharis truncarunr Whrelogee, TBP px. 315.

Dinacharts intermedia Bergendal, 1892, p. 107, Vig.

6:33.

Trickotria cornuta Myers, 1938, } tH.

Trichotria tetractis var. fruncata (Whitelegge) in Koste,

1978, p. 157.

Diagnosis: Margin of loricu with spicules, also on

foot segments, all directed forward; most with

shorter toes as in 7 tefractis, but many intermediate

forms; variable facetration on lorica.

Length 95-146 jan, greatest width 28 ym, toes

HOMME em,

Distribution: Acidophil, ii SpAcgnve, mainly

between plants, seldom in open water. N.SW., NOT,

Vie, 25,0-28.5°C, pH $.44-7.6, DO 6.0-4.2 mg Ir),

23-145 wS emi-).

Literature: Koste 1978, 198), 1984,

Comments Although morphologically variable, 77

Jrencafum is readily distinguished from T. retracrés

in material we have examined from both Austrahian

and Canadian collections (cf, Chengulath & Koste

1983), and we consider it a distinet species,

Not yer recorded from Australia

perracits caudata (Lacks, 2); cf. Koste 1984, p. L16,

i fetractis puupera (Ehrenberg, 1830);

T. curta (Skorikov, 1914);

T, eukesmrrra Myers, 1934,

(See Koste, (978, pp. 187-8 for details).

Pig. 16:l, Trickutria tetractis (Ehrenberg); (a) temale, lareral, swimming; [a = anus, Al = bladder, br = brain, al

= dorsal lorica, da = dorsafantertna, eve = cerebral eye, fe = toot gland, fj = foot. scement, fsp = foot spines,

8a = aasiric gland, fa = lateral antenna, oe = ocsophagus, sz = sultvary gland, s¢ = stomach, vi = vitellarium,

vf = ventral lorical; (b) male [dd = dorsal antenna, eve = cerebral eve, f = foot, p = penis, pe = prostate gland,

ss = (estis, vy = vestigial gut], (c) male, Jateral [ef = cihared fields, corona, eve = cerebral eve, p = peiis, ss

= pestis, vg = vestigial put); (d) teophi [ra = rainus,# = uncus ma = wanubrium); (¢) uncus, lateral; (f) lorica,

wross-section, 2, T pocidian (Miler), fermate, dorsal, swimming; 3, 7) ¢etractis, contracted dorsal, 4, 7) tranceta

(Whitelegwe): (a) dorsal, from Canada; (b) from Romania TT, trvacara f, langisping after Rodewald 1935] Fw.

15:la, 1, 2. 3, 4a dher Keste (6984): 4b, after Rodewaldl (1935). Seale Lines, leh 50 jut (Pig. 15da-c); lower cighe

VOX) pret (2-4),

110

W, KOSTE & R. J. SHIEL

er &), wt

‘A Ta

ROTIFERA FROM AUSTRALIAN INLAND WATERS 11

Genus Macrochaetus Perty

Lorica flat, hexagonal cross-section, head part

retractile; lorica granulated with minute spinules;

more or less developed anal section; two loricate

foot segments; toes generally spindle-shaped; paired

symmetrically inserted spines (Fig. 18:la); lorica

margin with spicules, shoulder hooks larger (Fig.

18:la, 7a—b); integument opaque; corona simple,

with central buccal field and longer lateral sensory

hairs (Fig. 18:3b); dorsal and lateral antennae on

visible papillae; one cerebral eye; mastax with

malleate trophi; eggs and males unknown. All

species live between or on water plants and are rare

in open water.

Key to Species of the Genus Macrochaetus

[refer to FIG. 18:1a]

1. Only one pair of rudimentary anteriosubmedian

spines present (2a, b).M. danneeli Koste & Shiel

More spines present..............ccc cece ees 2

2(1). Two anterolateral spines, two anteriomedian spines,

two posteriomedian spines present, central dorsal

spine pair absent (4a, b)....M. collinsi (Gosse)

Central dorsal spine pair present............. 3

3(2). Two anteriolateral- , two anteriomedian- , two

central median- and two anteriosubmedian spines

present ay ace caennns M. altamirai (Arevalo)

Two anteriolateral-, two anteriomedian, two

anteriosubmedian- and two posteriomedian spines

PIESENL. 5 ji rae Seles eed M. subquadratus (Perty)

Macrochaetus altamirai (Arévalo)

FIG. 18:3a, b

Polychaetus altamirai Arévalo, 1918, p. 1-47.

Macrochaetus esthonicus Riikoja, 1925, p. 7, Fig. 2).

Macrochaetus altamirai (Arévalo) after Selga, 1952.

M. australiensis Berzins, 1982, p. 18, Figs. 17, 38.

Diagnosis: Shape, size and spicules at lorica margin

variable; distinguished from M. collinsi only by

paired central dorsal spines (see key to species).

Lorica length 80-115 xm, width 76-108 ym, toes

20-28 pm.

Distribution: Cosmopolitan warm stenotherm, rare.

Probably more widely distributed in Australia than

a single record from Victoria would indicate (as M.

australiensis by Berzins 1982).

Literature: Koste 1978; Shiel & Koste 1985.

Macrochaetus collinsi (Gosse)

FIG, 18:2

Dinocharis collinsi Gosse, 1867, p. 269, Figs. 1-4.

Macrochaetus collinsi (Gosse) after Harring, 1913, p. 67.

For extensive synonymy see Koste, 1978, p. 161.

Fig. 17: Trichotria tetractis similis, lateral (toes 190 um).

After Koste (1984).

Diagnosis; Shape, size, length and number of

spinules on margin variable; number of dorsal

spines constant; lorica brownish,

Total length 94-250 ym, lorica 53-112 wm, width

62-112 ym, second foot segment 20-25 pm, toes

16-20 ym, dorsal spines 48-66 um, posteromedian

spines 24-48 wm, anal segment spines 32-35 ym.

Distribution: Pantropical and subtropical, warm

stenotherm, benthic. N.T., Qld, Tas. 28.5-29.9°C,

PH 5.44-S5.47, DO 5.45-6.2mg 1-1, 23-44 pS

cm~', alkalinity 2.6-2.7 mg 1-1,

Literature: Koste 1978; Shiel & Koste 1979; Tait er

al. 1984,

Macrochaetus danneeli Koste & Shiel

FIG. 18:5

Macrochaetus danneeli Koste & Shiel, 1983, p. 116, Figs.

Ja-d.

Diagnosis: Lorica granulated/pustulated; dorsal

lorica with terraced sides to blunt keel; margin of

keel base with longer spinules; uppermost keel

dorsum covered with large cuticular ledges and

beading; anal segment spineless; shoulder spines

(Fig. 18:la, 7a), posterolateral- (Sa, b),

TT - —————— Sn— —— SSFSSSSSSSSSSSSSSSSSSSSSFFMsesF

Fig. 18: 1 (a) points of insertion of spines and setules in Macrochaetus Perty spp., dorsal lorica and foot segments

[/a, 6 = anteriolateral spines, 24, b = anteriosubmedian spines, 3a, b = anteriomedian spines, 4a, b = central

dorsal spine pair, 5a, b = posteriolateral spines, 6a, b = posteriomedian spines, 7a, b = shoulder hooks, 8a, b

= anal segment spines, 9a,b = foot segment]; (b) Macrochaetus lorica, cross section. 2, M. collinsi (Gosse): (a)

dorsal, swimming; (b) lorica, ventral. 3, M. altamirai (Arevalo): (a) dorsal; (b) ventral. 4, M. subquadratus Perty:

(a) dorsal; (b) dorsal; (c) ventral; (d) lateral. 5, M. danneeli Koste & Shiel: (a) dorsal; (b) ventral. Fig. 18:la, b,

2a, b, 4a, 5a, b, after Koste & Shiel (1983); 3a, b, after Rodewald (1940); 4a-d, after Wulfert (1964). Scale lines

50 zm each number group.

112 W. KOSTE & R. J, SHIEL

posteromedian- (6a, b) and only one pair of

rudimentary anterosubmedian spines present.

Lorica length 125-130 wm, width 130-137 pm,

foot segment 20 pm, toes 16-18 «pm.

Distribution: Single locality known: Buffalo

Billabong, Magela Creek, N-T. 30.9°C, pH 5.38, DO

4.15 mg 1-1, 29 pS cm-!.

Macrochaetus subquadratus Perty

FIG, 18:4a-c

Macrochaetus subquadratus Perty, 1850, p. 22.

Polychaetus subquadratus Perty, 1852, p, 45, Fig. 1:6.

Diagnosis: Lorica generally hexagonal (Fig. 18:4a);

ten dorsal spines of variable size and shape;

shoulder spines most conspicuously large.

Lorica length 80-135 ym, width 73-140 um,

second foot segment 8-16 zm, toes 15-22 pn.

Distribution: Cosmopolitan in periphyton of

aquatic macrophytes. Rare migrant in plankton,

more common in billabongs. Thermophile.

Literature: Koste 1978; Shiel & Koste 1979; Green

1981.

Not yet recorded from Australia

M. hauerianus Wulfert 1964 (=M. collinsi var. Hauer

1938), p. 534 (Indonesia);

M. longipes Myers 1934, p. 13; see Koste 1978 p. 161;

M. multispinosus Myers 1934, p. 12; recorded from the

neotropics and S. Nearctic;

M. philopax Wulfert 1961, p. 87; single find in central

Germany;

M. sericus (Thorpe) 1893, p. 182; see Koste 1972, Koste

& Shiel 1983 p. 117 Fig. 8.

Acknowledgments

The support of the Australian Biological

Resources Study in grants to RJS, the use of the

facilities of the Botany Department, University of

Adelaide in manuscript preparation and long-term

loan of microscope facilities to WK by the

Deutschen Forschungsgemeinschaft are gratefully

acknowledged. The Department of Lands, Parks

and Wildlife, Hobart, supported fieldwork in the

World Heritage Area in Jan, 1989, Particular thanks

to the staff of the Peter Till Laboratory, Albury-

Wodonga Development Corporation (now Murray-

Darling Freshwater Research Centre) for providing

collections from billabongs in their study area, to

Professor D. G. Frey, Biology Department, Indiana

University for the donation of an extensive series

of rotifer publications from his Limnology Library,

and to an anonymous referee for critical comments

on an earlier draft. Our thanks also to Charles

Hussey (BM(NH)) for checking obscure references,

Anne Grant, CSIRO Publications Service, for

editorial expertise, and Lor Wai Tan for final

corrections.

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BRIEF COMMUNICATION

REDISCOVERY OF UPEROLEIA ORIENTALIS (PARKER)

(ANURA: MYOBATRACHINAE)

Glauertia orientalis was described from three male frogs

collected by W. H. Stalker in 1905 on Alexandria Station

(19°08" , 136° 43" vide!) on the Barkly Tableland, Northern

Territory, and a female frog collected by G, H. Wilkins

in 1925 from Groote Eylandt'.

The species was referred to G/auvertia Loveridge because

of its moderately extensive webbing on the toes (Fig. 1)

and its widely exposed frontoparietal fontanelle.

Fig. 1. Plantar view of the foot of Uperoleia orientalis

(Holotype, Alexandria Station),

Glauertia orientalis has been reported over a wide range

in the north of Australia2,3,4. The genus was referred to

the synonymy of Uperoleia® when it was shown that the

generic characters represented extremes in continua

expressed within Uperoleia. U. orientalis was redescribed

from the holotype and topotypic paratypes® but the Groote

Eylandt paratype was not examined at that time. Its

conspecificity was called into question because of

distribution and other data provided in the generic

revision®. No other referrable material was available to

these authors.

Later collecting ® failed to produce further specimens

of U orientalis at the type locality, and further studies

on Groote Eylandt resulted in the female paratype being

referred to U. inundata Tyler, Davies & Martin.’

Uperoleia orientalis thus remained known only from

the type locality and from the holotype and two topotypic

paratypes.

Whilst examining a collection of Uperoleia from the

Northern Territory Museum (NTM), I found a specimen

referrable to U. orientalis.

The frog (NTM RA88) was collected by Mike Gillam

during the Cox River Survey on 2.7.1977 adjacent to the

Arnold River (15° 43’, 134° 37°), at night in paperbark

woodland (Melaleuca spp) in a spring-fed swamp. The

species was abundant at the locality9.

The specimen is small (probably sub-adult) with a snout-

vent length of 17.6mm. Toe webbing is moderately

extensive (characteristic of the species) and there are very

prominent subarticular and supernumerary palmar

tubercles on the hands. The head is small in proportion

to the body and the parotoid glands are prominent.

Inguinal and coccygeal glands are extensive and coalesce.

No mid-vertebral stripe is apparent.

The site of collection represents a range extension of

about 500 km to the north north-west.

Parker, H. W. (1940) Novit. Zool. 42, 1-106.

Barker, J. & Grigg, G. C (1977) “A field guide to

Australian frogs” (Rigby, Adelaide).

Cogger, H. G. (1975) “Reptiles and amphibians of

Australia” (Reed, Sydney).

Tyler, M. J. (1976) “Frogs” (Collins, Sydney & London),

Tyler, M. J., Davies, M. & Martin, A. A. (1981) Aust. J,

Zool. suppl. Ser. 79, 1-64.

Tyler, M. J., Davies M. & Martin, A.A, (1983) Trans. R,

Soc, S. Aust, 107, 237-242.

Tyler, M.J., Davies M. & Watson, G. F. (1986) Zool. J.

Linn. Soc. 88, 91-101.

Tyler, M, J. & Davies M. (1986) “Frogs of the Northern

Territory” (Conservation Commission of the Northern

Territory, Alice Springs).

Gillam, M. Jn litt.

MARGARET DAVIES, Department of Zoology,

University of Adelaide, G.P.O.Box 498, Adelaide,

S, Aust, 5001,

THE OCCURANCE OF PLANT PARASITIC NEMATODES IN THE ARID

REGION OF SOUTH AUSTRALIA

BY JACQUELINE M. NOBBS

Summary

BRIEF COMMUNICATION

ul?

‘THE OCCURRENCE OF PLANT PARASITIC NEMATODES IN THE ARID REGION

OF SOUTH AUSTRALIA

There is Jittle information on the occurrence of plant

parasitic nematodes in arid South Australig. Twenty-four

genera of plant nematodes from the area encompassing

the Northern Flinders Ranges to Cape Jervis have been

identified tovether with a réelanonship between the

ihistribuuon of plant species and the distribution of plant

nematodes 7. The distribution of plant species was related

to Yainifall. Following on from this work, a survey of the

arid region (mean annual rainfall less than 250mm) was

undertaken over a four year period (1983-1986).

The sorl was sampled from under native vegetation, the

nematodes extracted using a modified Bacrmann’s funnel

technique 2 and where possible the nematodes identified

to species level. At many sites few mature female specimens

were extracted, so identification to genus only was. possible

(Table 1).

There were several genera of fungal feeding tylenchids

identified which included Aphelenchus avenae,

Aphelenchotdes sp, Coslenchus c.f, costatus,

Neotylenchus s.1. Dlenchus sl, and Dinvlenchus sp.. These

are not included with the plant nematodes in Table 1.

‘Jwelve genera of plant nematodés were identified from

the arid region ay well as juveniles and migles of the

Heteroderidae which were not identifiable because of the

absence of females or cysts.

The most widely distributed plant nematode was

Tilencharhynchus tebari (Sauer & Aanells, 1981), Of the

374 sites sampled, 63% contained 7 tobari. All other

species occurred at less than 8% of the sites. More than

Ohe taxon was present in 26% of the sites and a similar

percent of sites had no plant nematodes present. The arid

regions are an adverse environment for moxt organisiny.

To survive, organisms musr be drought resistant and also

respond rapidly ta the stimulus of rainfall. Some

nematodes are ideally suited to the arid region as they car

form an anhydrobiatic state in which metabolic respiration

is greatly reduced bur are quick to reyive with addilign

of water 3 and begin to reproduce rapidly after revival,

T, tobari (Sauer & Annells, 1981) is a widely distrituited

nematode within the region and must be well adapted to

the environment, The species was first described from a

site near Wentworth, N.SW, * The species was. cultured

by planting seeds of certain species of Chenopodiacexe

(a common family in arid Australia) into soil known to

contain Tf tohari.

Studies of host/parasite relationships aad covironmental

effects on nematode population growth will be presented

ia detail in another paper,

| meee & Wallace, H. R. (198!) Nematologica 27,

329,

2 Schindler, A. KF. (1961) Plant Discase Reporter 45,

747-748.

> Freckman, D, W., Whitford, W. G. & Steinberger, ¥.

(1987) Biology and Fertility of Soils 3, 3-10.

4 rhe i R. & Aunells, CM, (198) Nematologics 27,

JACQUELINE M. NOBBS, CABI Institute of Parasitology, St Albans, Herts,, UK. AL4 OXU.

TABLE |. Nuytiber und percent of sites Jrom whieh plant nematodes (PP) were extracted in the years 1983 t0 1986

rarciusive.

Genus and Species

Trlenchorhpncdus tobari Saver & Annelis, 1981

Pratyleachus sp.

Morulaimus sp.

Helicotylenchus sp.

Seutellonenia sp,

Radopholus sp.

tulenchorhynchus velatus Sauer & Annells, 1981

TWienchorhynchiss sf.

Reivlenchys sp,

Heaplolatinus sp.

Morulaimus geniculatus Sauer, 1966

Puralongidorus sp-

Heteroderidae males aud juvenites

Séutellanema laeviflexum Phillips, (971

Trlenchorhynchus unmiulgies (Cassidy, 1930) Golden, 1974

Scutellonema minutum Sher, 1964

florlenchus hastulaius (Golden, 1960) Siddiqi, 1963

Melicotylenchus yariebilis Phillips, 1971

Radopholus crenatus Colbran, {971

Paratylenchus sp.

Hemicycliophora sp.

foral number of sites sampled

Sites with >1 genera af PP

Sites with PP

Sirés without PP

Sites Percent of

Sampled Total Sites

237 63.3

24 6.4

16 43

16 4.3

ity 4.3

14 37

12 3.2

? 1.9

7 1

§ 13

4 tl

3 0.8

4 0.8

2 05

1 0.3

1 03

“4 0.3

i 0.3

374 100.0)

07 25.9

276 Ti8

98 26.2

VOL. 113, PARTS 3 & 4

30 NOVEMBER, 1989

Transactions of the

Royal Society of South

Australia

Incorporated

Contents.

Koste, W. & Shiel, R. J. Rotifera from Australian inland waters IV. Colurellidae (Rotifera:

Monogononta) - - 2 : : = : = :

Short, A. D., Buckley, R. C,, & Fotheringham, D. G. Preliminary investigations of beach

ridge progradation on Eyre Peninsula and Kangaroo Island -

Pledge, N. S. The occurrence of a diprotodontid marsupial referable to Euowenia grata

De Vis in western New South Wales - - = = =

Austin, A. D. & Allen, G. R. Parasitoids of Uraba lugens Walker (Lepidoptera:Noctuidae)

in South Australia, with description of two new species of

Braconidae - = = a * = p ey

Barker, S. Contributions to the taxonomy of Australian Buprestidae (Coleoptera): New

species of Astraeus and Stigmodera (Castiarina) and a key to Astraeus

CS ee

Jenkins, J. F. & Hasenohr, P. Trilobites and their trails in a Black Shale: Early Cambrian

of The Fleurieu Peninsula, South Australia - - - =

Gowlett-Holmes, K. L. & Holmes, N. J. C. Rediscovery of Primovula (P) heleneae Cate

and description of a new species of Crenavolva from South Australia

(Mollusca: Gastropoda: Ovulidae) - - = = =

Davies, M. Developmental biology of the Australopapuan hylid frog Litoria eucnemis

(Anura: Hylidae) - - - = Ee a 4 z

Brief Communication:

Beardsell, G. R. Hybridisation of Litoria chloris and L. xanthomera, (Anura:

Hylidae) - - - © : J , = : ¥

119

145

163

169

185

195

205

215

221

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

VOL. 113, PART 3

ROTIFERA FROM AUSTRALIAN INLAND WATERS IV. COLURELLIDAE

(ROTIFERA:MONOGONONTA)

BY W. KOSTE*, & R. J. SHIELT

Summary

Diagnostic keys are given to genera and species of the rotifer family Colurellidae (Colurella,

Squatinella, Lepadella, Heterolepadella) recorded from Australian inland waters. All species are

figured and available distribution data and ecological information given. Some widely-distributed

taxa not yet recorded from the continent are included.

KEY WORDS: Rotifera, taxonomy, Australia, known species, Colurellidae

Transeenions OF he Ravel Suctety af §.

Atyt, (S989), 10303) 19-145

ROTIFERA FROM ALSTRALIAN INLAND WATERS [V, COLURELLIDAE

{ROTIFERA:MONOGONONTA)

by W, Koste* & R. J. SHIELT

Summary

Kosta, W., & SHlel,

R. J, (1989) Ronfera fram Australian inland waters, CY. Colurellidae (Kocifera:

Menogononta), Trans, R. Sov, 8, Alust, 113(3), 119-143, 30 Noveinber, 1989.

Diagnostic keys are given Lo genera and species of the rotifer family Colurellidae (Coluredia, Squatinalla,

Lepadella, Heteralepadella} recorded from Australian infand waters. All species are figured and available

distribution data and ecological informavion given, Sonte widely-distesbuted taxa nor yet fcoordéed trom

the continent are included.

Key Worps: Rotifers, iaxdnomy, Austraba, known species, Colurellidae

Intraduction

The monogonont rotiter families considered lu

ate in our revision of the Australian Rotitera (Koste

& Shiel 1986b, 1987b, 1989) have included the mosr

common planktonic taxa in Australian inland

waters. Here we review the Australian represen-

(atives of the family Colurellidae, several genera of

which: have widely-distribured species in plankton

and littoral collections in Australian waters and are

considered heleoplankters (semsw Hutchinson 1967)

rather than accidental ineursions, i.e.

pseudoplankton (= fychoplankton af Ruttner-

Kolisko 1974) (eg, Lepadella patella). In general,

however, the Australian representatives of the

Colurellidue (Colurella, Squutinella, Lepudella,

Hererolepadella) are littoral (epiphytic or

epibenthic) in habit. They are encountered

cammonly in the shallow vegetated areas of

billabongs and other werlands, where diverse

mImgermphyte associations effectively partition the

habitat, providing abundant microniches.

Diagnostic keys, brief descriptions and figures of

all known Australian representatives of the family

are provided to enable identification to species,

Some widely distributed taxa not yet recorded from

Australia are included, The formar follows that of

eurher parts; dichotomous keys are followed by

individual species” descriphions and koown

Australian records, Relative abundance is indicated

bys “conmmoan” (more than 30 widely dispensed

records), “uncommoan" (10-30 localities), “rare”

{<10 records). Brief ecological dats are included

Where available. Early Australian records cited in

Shiel & Koste {1979} are nol repeated here, nor are

the majority of northern hemisphere citations given

by Koste (1978), which contains a detailed treatment

* 5, Ludwip-Brill-Strasse, Quakenbrtick 9-4570, Fecera)

Repubtic of Germany

} Murray-Darbng frestiwwier Research Contec, PO. Box

92), Albury, 6 SW 2640)

of the Rortifera outside Australia, Keys to families

are included in Kostc & Shicl (19876).

Family Colurelidae Bartos

Loricate rolifers, the vorena in all genera with

4 membraneous cap or semicircular shield

(retractible in all except Squatinel/a); no lateral sulct;

Colurella and Paraceluretia with ventral sulcus;

(rophi malleate; ventral plate with caudal foat-

Opening or enlarged Jor basal toot segment; toes

slender, sharply pointed, Four genera; Colurella,

Lepadella, Squatinella, Paravoluretla.

Key to genera of the Family Colurcllidae

}. Lortea laterally compressed, wilt venwal or sori

and ventral apertures (Fig. 2),....-...-...... 2

Lorica dorso-yentrally flattened, without such

apertures -.--...-.--- ----- - 3

2(l), Lorica a single structure, with ventral aperture

(Pig. U.....---..Calurella Bory de St Vincent

Lorica two plates, with. dorsal and ventral apertures

(Fig, 2:8)....... Puracolaretia Myers (not known

from Australia).

3(1). Hood or head-sineld large, non-retractible

(Figg) oo... Squatinelia Bory de St Vincent

Hood small, retractible (Fig. 5)

poems Lepadella Bory dv St Vincent

Gentis Calyrela Bory de St Vincent

Colunmila Bory de St Vincent, 1824, 17.203,

One piece loriva, in lateral view oval, egg- or

hatchet-shaped (outline of lorica differs according

to degree of contraction, particularly ventral

margin); anterior lorica margin more or Jess

(rancated or rounded: posterior margin rounded,

gradually merging into blunt, straight or curved

proj-tions, in some cases sharply pointed, offset

win ike extensions of posterior margin (Fig. Ic}:

lan W, KOSTE & RK. J, SHITE)

ventral cuticle between mussel-like lorica borders

delicate, membrancous, may be distoricd in

preservative; corona of the Euchlunis type (cf, Koste

& Shiel 19876); loot with three or four segments,

basal segment mostly pliable and often of indistinct

structure; distal foot segment (before toe) with

dorsal sensory pore; toes of variable length, species

with long toes more compressed laterally; toes

separate, however may appear fused as frequently

are stuck together; gastric glands mononucleate;

trophi malleate; eyeless or with two. eyespots, which

may or may not be red-pigmented; lateral antennae

very conspicuous; male little known and

insufficiently described. Resting eggs retained in

lorica of parent an deuth (cl. Fig. 2:3).

Variability in the described species requires

investigation, particulatly whether the morphology

of different races (“var”) or forms represents

thes

Pig. tl,

ecotypic variation [“Millienfaktoren” (Hauer 1924)],

e.g. the variability in the lorica ends in C uncinata

or © adriatica, Eeotypic variation is common in

the Australian brachionids (Koste & Shicl 1987b),

and variants are known in the five species of

Colurella recorded from Australia. It is likely that

other species occur here; in view of their small size

(most <100 ym) colurellids are easily overlooked

in detnitus-rich samples.

Key to species of the Genus Coturefla

{, Lorica surface smooth. 2.00. 00.2-- 02.00.0008, 2

Lorica surface with longitudinal furrows or facets

icht it pew. tr CG. fesselala (Glascolt)

2(1). Loriva "Vale (lateral) slender, posteriorly with

Watiable apices,,.. 2.0.0... e ee ee ees

Lorica valve rounded, Pliiied Faocek here ses a

Colurella adriativa (Ebrepberg): (a) taterat (an = anus; du — eyes er > subilaneous eg, ax = mustilx;

vi = vitellarivint); (b) lateral, contracted; (e) ventral: (dd) posterior lorica showing foot and we (DS. = dorsal sensory

mit}; (e-1) lateral views showing Variation in lonca morphology, 2

lateral: (dg) male, lateral. 7, ¢*

Corda (1834); 1h, 3d, alter Carlin (1939): le,

(1953): Va-c, atter Hauer (2924). Scale lines 20 pm,

WC. colurus (Ehrenberg): (a) dorsal: ¢b) ventral: (c)

vitusy (Coasse), (2) dorsal: (b) ventral: (a Vater, (d) anorher morph, lateral. la, afrer

d, wtier Waulfert (1980); fest, Zuse, aller Hauer (1928); 20) aller Wiszmewsky

ROTIFERA FROM AUSTRALIAN INLAND WATERS Pal

3(2).. Posterior end of lorica with raised lines terminating

in varnible tupering, downward-curviny (oveasion-

ally lightly upward-curving) spines; lorica high

(eneth/height rario <t.8); tows short, ...

qr tS eens C uncinuta (Niitler)

Posterior end of loriga broadly rounded, or blunt-

spied or tapering, loriga low (ratio >1.8); toes long

Sein Lorica Valve postenarly rounded, or at most

extended tes blint apex (Fig. 1;2c)...,, 128906

-C ecalurus (Ehrenberg)

Loriva valve posteriorly ‘with curved spines (Fig. Hc)

Baalebne'a''s g & adriatica Bhrenhere

ColureHa adrartica Ehtenberg

FIG, V4

Coluretla adctatiea Ehrenberg, 1831, Fig. 3:3.

Piapnosis: Extremely variable lorica size and

phenotype, particularly points at posterior end of

lorica. These are the only features separating C

adriatica from C calurus, Pejler (1962) noted

transitional forms, Different ecotypes were des-

eribed by Hauer (1924), with increased loricu size

and Woe length correlated with increasing salinity,

The measurements below encompass the range of

ecolypes (Koste $978),

Length 85-113 ~m; width 29-48 pm: height 44—66

pam; toe length 28-48 xm.

Disiribution: Probably cosmopolitan, Euryhaline.

Rare: fresh and slightly saline waters of southeast

S. Aust, Tas. and western Vic. Berzins (1982)

recorded C udriatica asx widely distributed in eastern

Australia. 5.0-30.0°C, pH 5.5-10.5, dissolved

oxygen (DO) 6.0-11.2 mg | |, conductivity (K,,)

§7-13,940 uS cm‘ turbidity 21-67 nephelometric

turbidity units (NTU).

Literature: Ridder 1972; Kusle 1978

Colurella colurus (Etrenberg)

FIG. b:2

Meno colurus Ehrenberg, 1430, p, 44,

Colurella colurus after Harriny, 1913, p, 29,

Diagnosis: Distinguished trom C adriatica only by

the rounded or blunt-spined posterior lorica,

Resting eve and male known (Fig. 12d).

Length 71-110 um; width 25-37 wm; heieht 39-54

xm; toe length 25-40 ym.

Distribution: Cosmopolitan, widely tolerant, oecuc-

ting in fresh, arhallasic saline and coastal waters.

Huropean temperature 1,0-28,0°C and pti 4.9-10,0,

Ridder (1972) records ©. colurus from 15,035 2 Cl

Ll . Rare: Tas, Vie 1.8-16.0°C, pH 3.7-6.4,

495-2130 eS cm.

Lilerature: Koxte |978,

Colurella vhtusw (Gosse)

FIG, t.3

Colurus obsusus Cisse fr Hudsen & Gidsse, 1886, p. 103,

Fig. 26:3,

Colurella abtusa after Holsien, 1909, (x. 84.

Diegnasis; Lonca outline plump; margin of bead-

uperture directed obliquely from ventral lorica;

ventral lines variable; head width approx. halt Jorica

length, or smaller; eyespots with or withour

pigment.

Length 60 pm; width 28 pm; toes 17 am,

Distribution: Cosmopolitan between aquatic mac-

rophytes; pancontinental, uncommon, 8,.0-29,0°C,

pH 3.4-7.5, DO 11.2. mg 1", 36.4-1600.p5 cm !,

{5 NTU.

Literature: Eyans 1951; Koyte 1978, 1981; Shicl &

Koste 1979; Green 1981: Koste ef of, 1988.

Colurella lesselata (Glascat)

FIG, 2:1

Calurus tesselatus Glascott, 1893, p, 74,

Colurella tessélata atier Hofsten, 1909, p. 84.

Diagnosis, Loriea with dorsal keel; facet-like pattern

with ribs. on lorica surface; toes relatively short,

Length 55-64 am; width 24-31 pin; height 32-41

puny, toes 18-27 wm.

Distribution: IC -osmopohtan (nat recorded from

N.Z,), sporadic in oligotrophic waters, mostly in

moss, also in psammion, Single record from Coghills

Creek, Tourello, Vie.

Literature: Koste 1978; Berzins. 1982,

Colureila uncinata uncinala (Miller)

FIG, 2:2

Brachionus uncinatiy Muller, 1773, pp, 134,

ColureHa uncinata after Having, 1913, p, 30,

Diwgnasis: Lorica posterior end gradually merges

into downward-curving short spines; head-opening

smaller than greatest lorica height.

Length 77-106 jem; width 30 pm; height to 60am:

toes 18-23 wm; male 50-85 «wm,

Distribution: Cosmopolitan, eurvhaline, Common-

pancontinental. &.5-24.5°C, pH 4.8-8.0, DO

7.8-11,2 mg | ', 20-1600 ws cm~', 3-160 NTU,

‘jtérature; Koste 1978,

Comments; Two. morphologically distinguishable

taxa described originally as distinct species, but

regarded by Koste (1978) as “forin” yariants of C

uncinata also até known from Australia.

Colurella uncinata bicuspidara (Ehrenberg)

FIG, 2:3

Coluras bicuspdaias Ehrenberg, 1832, pm 123,

Cohtrella necinatd C. bieuspidata alter Mewche, 1939, px

W. KOSTE & R. J. SHIEL

ROTITERA FROM AUSTRALIAN INL ANI? WATERS 121

Diegnosis: Loriva width up to twice that of

noininate species; highest extension of torica al

head-opening; lorica ends short, most cume

downwards; toes relatively short; resting exe with

punctiform cavities,

Lengih 60-108 «mj; width 53-62 pm; height to

A um; toes 13-27 pm.

Distribution: Cosmopolitan in littoral of fresh and

athallasic saline waters, alsa marine Littoral.

Uncomnion: N.SW,, Qld, 8. Ause, Tas. Vic.

10,0-23.5°C, pH 4.5-7.5, 93-927 pS am |

Literature: Koste 1978, Stuel & Koste 1979; Green

IYRI; Berzins 1982,

Calurella unciaala deflecd (Ehrenberg)

FIG. 2:4

Cotarus defleris Ehrenberg, 1834, p. 203.

Celurelia uncinata \, deflexa alter Meuche, (939, p, 400},

Diagnosis: Lorica end robust, clongated, turned

outward, most directed ventrally. Very variable.

Length 60-100 am; width to 55 yin; height 35-55

um; toes. 17-25 pny,

Distriburion: Cosmopolitan, sporadic, euryhaling.

Rare. Qld, Tas, 9.5-20,0°C, pH 6.5.

Hiterature: Shiel & Koste (979; Koste & Shiel 1987a.

Not recorded trom Australias

CO. denticauda Carlin, 1939. Scandinavia

dicentra (Gosse, (887), N. Eurype, tance

| hindenburai (Steinecke, 1917)(Fig. 2:6), Probably

cosmopolitan

©. geophila Donner, 1951. Burape

©. mariqavi Althaus, 1957, Black Sea, mesopsammal

Aan

C. mucronulatea Atitstrom, (938, U.S.A,

C. oxycauda Carlin, 1939, Scandinavia

C. paliidasa Carlin, 1939, USSR,

C satiaa Althaus, $957 (Fig. 2:7). Europe, N.4.

C. sinisira Carlin, 1939, Furope

C. suleata (Sienroos, 198K Fyr. 2:5), Probably

cosmopolitan, warm <tenotherm, neidophil.

C. umieauda Eriksen. (968. N, Europe.

Genus Squatinella Bory de St Vincent

Sguctinella Bury de St Vinognt, 1826, po 87,

Body cylindrical te ovoid: head and corona under

hyaline semicircular shield (Fig. 39), which is

pontect apically in some species, also with

triangular side plates (“ears”); dorsal lorica smooth,

also with occasional symmetric longitudinal ribs:

ventral lori¢a smooth, with allached. shveléd-tke

plate, with one or two scales, or also with keel;

posterior lorica rounded, truncated or tapered! to

variable points; some spogies with lonz dorsal spine;

short spine in inedian line of posterior lorica may

be present; foot 2- or 3- segmented with long

pointed toes; small thin spine may be present at base

of tocs (Fig. Sule); syeninetric malleate trophi (Fie.

3:6); gut without peculiarities, dorsal and ventral

antennae small; two widely separated distinct

eyespots with crystalline bodies, Male and resting

ege not yet described, Ecology of Squarinella little

known. Koste (/978:174) notes enus restricted to

littoral, with some spp. acidophils, Four species

recorded from Australia. For details of other species

and variants, see Koste (1978:175-179),

key to species of the genus Squatinelia

| With dorsal spine, , ---- ' a 2

Withour dorsal spine -- Vries itsse reid

2(1), Foot 3-xegmented; short spine al base of roes

whe bee teense fomaispinata (Tatem

Poet 2-segmented) spline absent... . 022... tt

A. _..-..§. devas? (Zacharias)

41) With & short spine abeve the base of the foes

S. rostrum (Schmarda)

Withaul spine above toe bases. oe

oa, S. mutica (Firenberp)

Squatinella longispinata (Talem)

FIG, 3:3

Stephanops longispinarus Tatem, 1867, p. 252, Pl, 10, Fie

1-3,

Squatinedla lomgispinora atiey Warsing, 113, p. Yb,

Diagnasis: Characicristic tong, thin and curved

dorsal spine. Pejler (1962) noted two xhart spires

flanking dorsal spine (Fig.3:3a). Ventral side

sometimes wrinkled. Ratio dorsal spine:body. length

0,8. Foot three-segimented.

Length 82-160 pm; dorsal spine 96-116 am; toes

3-10 wm, spine(s) at the base of the roes 4—5 pm,

Distrifation: Widespread but rare, mostly in moor

waters beiween SpAaenunt (Koste 1978), We have

not cnecountered this species in > 2000 collections,

however it Was recorded by Whirelegge (1889) from

N.SW,, also by Evans (1951) front Victorian waters.

1 iterature; Pejler 1962; Kaste 1978.

Fig. 2, Coluretfa tesselata ({Glaseott); fa) dorsal; (bu) lateral. 2, C vnemnara uncinare (Miter): (a) lateral: (b) dorsal

te) male, ventral; (d) male, lateral. 3, C wactnala bicwspidaty (Fhcenbers): (a) tatecals db) another moeph, lateral:

fe) ventral (d) ventral, with resting egy. 4, CL uavinare defieve (shrenberg): (a) literal; (b) hubiv, dorsal. 5, C. stefeutes

(Stcnrnos), lateral 6, C. hiadendured Steineekes (a) dorsal; (b) literal. 7, CL valine Alnaus: (a) laterals (b) ventral.

4, Paravalirefie lagna Myers: (a) lateral; (b) ventral, Ja, 6, 2b, 46, afer Haier (1924)5 lo, 2a, Ra-c, da. 3, alter Curlin

(1940); Ze, U, alter Weber (1898); 3d, Sat. b, aller Kasic (1978); 6a, by alter Steineeke (YLT), Fa, by after Alulmuy (19571

Seale lives SO pan.

124 W. KOSTE & R. J. SHIEL

KOTIFERA FROM AUSTRALIAN INLAND WATERS 124

Squarinella devdlell lougiseta (Pourriot)

FIG, 34

Squadinetla teveigiiT, fangisera Pouiriot, [Y71, pp, 99-100,

ripe 4s

Lnagnosis: Robust body; dorsal spine longer than

body; two foot segments; no spite on distal end of

terminal segment (at base of toes) Much larger

animal than §. /ongispinate.

Body length 210-235 am; dorsal spine 270-378

pm; ratio spinethody length 1.6; tocs to 37 pm.

Distribution: Rare ino submerged Sphagnum,

Myriophytium, etc, Single record trom @ pool near

L. Gareia, western Tas, 17.0°C, pH 3.1, 80.648

cm 7,

Comment: The nominate S. feydigil is not recorded

from. Australia, [t is distinguished from S.

longispinata by foot morphology and from the ssp,

longiseta by its markedly smaller dimensions (body

length 10-130 um, dorsal spine 90-110 jm:

spine:body ratio 0.8-0.95; toes Lym).

Literature: Koste 1978,

Squatinella mutice (Ehtenbere)

FIG. 3:2

Stephanopyomuticus Ehvenberg, 1832. p. 138.

Squaetinel/a mutica after Harring, (YI, ps. 97,

Diagnosis: Caudal mann uf dorsal lorica semi-

cirenlay, tongue-shaped, three-lobed, with three

short spines or truncate (Fig, 3:2d-g), All forms may’

occur in same popwation, Foot three-seemenied,

Body length 100-226 pm; headshield width 60

ym; toes 10 30 pun.

DEiribyhon: Europe, N.Z. Rare, from billahongs,

nonds or dams: N.SMW,, NW, Tas., Vic. 16.5-24,5°C,

pH 3.8-5.3, 45.5-46.2 pS um |.

Literature: Russell 1961; Koste 1978; Berzins 1Y82:

Koste & Shiel 1987.

Squatinella Mmuricn tridertteata (Fresenius)

PCH, 3:5

Srepharnans tricdendius Fresenius, 1858, p. 28, PL 1d, Fig,

Sauatineda maticu tridentata afler Wisaniewsh), 1953, p.

401.

Diagnosis: Posterior lorica oval, terminating in three

pointed, clonguted spines of variable length.

Resembles §. rostrum, but lacks spine at base ul

toes, Headshield semicircular.

Body tength 134-190 pm; toes 20-30 jatn,

Distribution: Ponds and pools, often abundant in

Sphagnum. pH 4.8-6.2, Rare, Qld, N.SW., Vie.

Literature: Evans 1951; Koste J978.

Squatinella rosirum (Schmarda)

FIG, 3:1

Lisirion rostran Schimarda, 1646, p 20, Pl 2. Pig. t.

Syuatinella rostrum after Carlin-Nilson, 1949, p. 3

Pragnosis; Caudal lorica with three long or short

pointed processes which arise from narrow bases,

Lateral antennae sunken in dorsal lorica margins.

Length 150-217 pm; toes 24-28 pm; headshield

to 76 pm; caudal processes fo 36 fm; spine at base

of tes ta 23 pin.

Distribution; In Aulwuchs (i.¢, periphytun attached

to submergent stems), Pwo records only, Brisbane,

Qld and from Creswick Ck. Clunes, Vie.

Literature: Thorpe 1887; Koste 1978; Berzins 1982.

Not recorded from Australia:

S. biferca (Bolion, 1884) in Hudson & Gosse (L886)

Kurope, lecland.

S. mivredactyla (Miicray, 1906), Europe, Newtoundland,

in SpAagerum,

Genus Lepadella Bory de St Vincent

Lepadella Bory de St Vittcerit 1R26, p &6,

Outling-oval, pear-shaped or circular, more or less

dorso-ventrally Hattened; dorsal and ventral lorica

plates firmly joined: deep notch for head and wor;

ocvipital margin of lorica often with granulated

collar; head with cap-like plate, Dorsal lori¢a

domed, smooth, with or without keel development,

also with several ledges or fillers; lorica margins ih

some species drawn out into winglike extensions.

Head and foot apertures of different Shape;

differences useful in taxonomy of group, Foor of

three flexible segments, one inflexible basal

segment, Toes short and pointed or long and nvedle-

like, similar or dissimilar in length, inserted laterally

or dorsal-ventrally, in some species completely or

partly fused. Corona single ciliary wreath with

lateral ciliary tufts and buccal field. Two lateral eve

spots, occasionally with separate pigment granules

Fig. Mb Squutnedts rostram (Sehmarda): (a) headshield, dorsal; (6) headshicld, ventral; (c) dersal td) lateral amenna:

(ep ventral: (1) lateral, 2; So inuticg (Ehrenberg): (a) ventral; (by lateral: (ce) dorsal (d-e) variations in vaudal

morpholowy, 4.8. ompispinata (Tatem): (a) taveral; (b) dorsal; (c) foor, lateral. 4, 5. /evdieit tonetsera Pourriol.

SS. nitien tridentate (Fresenius): (a) dersals (b) lateral, lH, by afler Webou (1898), Je-t, 2a, fy, da, alter Wultert

(1999); 2c-z, ather Koste (1978); 3a, after Pejler (1962); 3b, atler Slenroos (1898), Ju alter Mycrs (942). dh, ater

Gillard (1948), 8, after Pourtie) G971). Serle lines $0 yam,

126

nearby. Dorsal and lateral antennae present, lattct

generally dorsally situated at level of upper border

of fuot-opening, Last foot segment usually with

caudal antenna. Trophi maleate, uncus with five

teeth. Gut with large, sometimes lobed gastric

glands. Vitellarium large with eight nucleii.

Proionephridia with distinct terminal ces and large

bladder, Litthe known of reproductive cyele, Some

males (90-120 pin) wre Known, Resting ees, will

prickly outer shell, retained in lorica of parent on

death.

JTepadella species. are benthic, commen in

billabongs and ponds, but occasionally collected as

incursion species in the plankton of reservoirs and

rivers, og. L. patelhy (Shiel ef al, 1982) Further

details wre given by Koste (1978), who notes thar

all Spevies can be identified trom preserved nvaterial,

although caution must be takeb with preseruition

artelucts, particulady uo the case of formalin-

preserved specimens, Several of the 45 tasa ot

Lepetelia identified from Australia probably are

misidentified for this reason. These are considered

in the texc. Nevertheless, about half of the known

species upper to be indigenous; Lhe genus is thus

second only wo Brachionys (Kost & Shiel 1987b)

in ats vonsplerment of endermiv taxa.

Three subgenera are recognized on the basis of toc

morphology;

i) fepedella (Woes of equal length, entirely

separated);

(ii) Hererolepadella (toes af unequal lenethy;

(iii) Nenolepadella {toes complelely or partly Tused).

Por convenience, the Australian species of the three

subgenera are considered separately in the following

keys.

Key to species of the genus Lepadelia (s. str)

1, Dersa) lorica without keelor ribs (may be lightly

suieiate, Hur otherwise wnomamented)....2

Doroal Jones with keet declan longuudinal ribs

or Other ornamentation... . ccs peeee esl?

20). Lorea tength > 3s lorica width, wlmast cylinitrival

(rig, 4:4) ody anensra Berzins

Loviea length <3adocica with, ovoid, circulay,

trapezoidal Or rhormboidal,... 2.2.2... 26. 3

Dorsallonica cross-section 1/3 of eirele ta highly-

domed hemisphere (Pig. $:2d)......----.-- 6

Dorsal lonica a stiatlow dome, or darsoventrally

Wattenud -- é a!

Loriva oval to circular; head aperture notehed

both Roof und ventrally (Fig, t:3a).......

4. evalis (Muller)

312).

Lori iva 2x ‘wide in posterior third Man anterior,

dorsal murgin of head aperture straielit, ventral

mitt only weakly y-shaped (fig. @:9b)

4 5

eee oe -

W. KOSTE & R.

3(4).

(3).

7(6).

YS),

L(A),

HIG),

1211).

13(12).

14(13).

15¢14)

1612),

(7d).

107)

W177.

JUSHIEI

Ventral lorica with twer lateral spines directed

forward: (ues striuglue (Pig, O82). 000.00 ee,

penta corey be tana Koste & Shiel

Ventral Jorica Apineles a4 ae curved,

ee bee 13 netmlsst Bervins

Devipit al maint ol head ape ee protruding (Fig,

SOMEED Heenan on a): : _..

Qecipiial margin of head Aperture concuvely

noiched (Fin. S4a)......, pecininiciibic! Lopes = 7

Lorica outline elongate oval to cireutay (Fig. 5;7a)

Lutica outline rhombaidal (Pig, 3: da},

se 1. vhensalathi Kosic

Posterion margin ot Joriva rounded oF trungaie,

never pointed (Pig, SFad. vere ee

Posrerion maraih of loriga tapers lo a pointed

projection extending wes bitse OF roes (hig. 9:7)

wilhatis) Kose & Shiel

Head! aperture vential sinus with parallel cuticular

ribs; dorsally with a koe prantlared collar

ellipned Willert

Head apeitiise- Wittiout rs or dorsal collar

, patella (M*iler)

Ventral forica with long pleaty.......

trifa Myers

Ventral tories “umpleared sda ome ue heath

Cross-section with Sie bead- like foteral

margink. sc... voceyedy daerplisera (Stencons)

Crossesection lateral margins produced lo aqgute

en ere it

BIC Wee ete sc veatalle ai leletene tn 12

Toot-opening excised dorsally. .-...., 5 ao TES

Foot-opening nat excised dorsally... 0.0, iy

Extremely wide foot-opening, tonger than wide;

corners With ourwardly-cHived points... ._. i4

Foot openine wider dhan long, no points on

57 tt TR a L. henxamow Warring

Head ajentuce ventrally with lateral biftreare

pointed extensions extending beyond dorsal

Margins... 0. -L. cornuti (Koste)

Nor balecal poly oo head aperture, ,,, 15

Head aperture with convex dorsal margin last

foot seument <b) foe leneth. oc cece eee

ied renin: site (dhe naps L. fatusinus (Aigendort)

Head apetiure with concave dorsal mucins last

foot segment Se toe kingthy.occee

. eer S271 Shephard

Head aperture: Greate cabaie L. apsida Warring

Llead aperture with deen yeniral sinus,

erase aie scoln' Ui fyi leeln etn bp 1. ratwnhlrad (Lacks)

Loricu ourline trapezmidal; loniua ornamented

(appears punctate oO. granulated under 1.)

ante te te semtel ao a lelecs sok ot latices idesetalh fete anes 14 ni!

Lorica curling cireular of ovale. 00.0.0... 19

Lateral extensions (“horns”) from posterior loriva;

rows of finy rodlike structures covet lorcet (Fig,

Artay ... + --1, dinar’ Koste

Tauteral horns absent; puncitorm ormmamentition

of dorsal und vermral loriea (hig. 9:44). , .

: of, tert Roste & ‘Shiel

horica outline circular; Jivad aperture circulur,

venually placed: lorica cross-section an trverted

nducer (Pig. m2)... 4 Ly findaui Koste

Lorica onthe ovoid, eal aperture not circular;

rate

ee he

ROTIFERA FROM AUSTRALIAN INLAND WATERS 127

Fig. 4:1, Lepadella acuminata (Ehrenberg); (a) dorsal; (b) caudal inorphology, ventral; (¢) dorsal: (d) caudal morphology,

ventral; (2) ventral: (1) lateral; (g) cross-section; (h) ventral; (i) L. acuminata sexcosiatu, dorsal: (4) L. chorea, ventral:

(k) £. chorea lateral. 2, L. amphitropis Harring: (a) ventral; (b) cross-section; (c) dorsal; (d) lateral. 3, 2. angusia

Berzins: (a) ventral; (b) dorsal; (c) float, distal segment, and 16es, dorsal; (d) toes, lateral; (e) cross-section; (1) anterior,

lateral, la-d, h, after Kosice & Shicl (1980); la, [, g, i. k, afler Berzins (1982); 2, alter Harring (1916); 3. aller Berzins

(1960). Scale lines 50 pm,

128 W.. KOSTE & &

Jorica ¢ross-section lightly donied, vibbeu Gr

townpgwar — - ete Hs inti toe att 20

Posterior lorica lapers bo ait acute poly, which

may be ridged dorsally... ......, 0.2.20

2 ieienape 1 1. aeuntmata (Ehrenberg)

Posterior lorica rounded, indented, a7 coneavely

norched . Reig Selintel es eo ee

. Dorsal torica with “median ‘keel sstutiyg Vi 2d

Dorsal lorica without median keel. yy ie

. Dorsal loriga with open-squared pattern (three

rows); ovcipital margin stright... yee

Ly decora Berzins

Dorsal tora unttered, with {4 short ribs over

foot-opening .......- aoe 23

1-7 short ribs at posterior end over rr oot -OPeninys,

bross-section rhombic “ig

Nees etele ale elope tants L. umphitropis Harring

4 parallel shart tibs over foot-opening; cross-

section a shallow dome With lateral concavities

of ventral margin (Pig. 8:2C).....

a , quadrivarinata (Stenrops)

Median heel of dorsal lone without side ribs

ieee RE ph dts eet opel eo sehen Bee ...28

Median keel with 2-3 pairs of side ribs... ..-

wh cehaseewnd tts L. quinguecosiuta CLueks)

Median keel law and wide, hordered by lateral

grooves; granulated vera

L. phoultivictes (Gose)

Median keel high, triangular, arising from ourrow

basu; no granulated. collar. Some farms have fine

sirined/pustululed lorica surface. .

4 a sea talal sta L. iriptera (Ehrenberg)

2019).

24(2N),

25124)

NE. Some European forms of LZ. triptera are noted by

Kasie (1978) to be rhombodal, Only owl forms are known

from Maistritia,

Lepadella acuminata (Rhrenhberg)

Vite, 4:4

Metogidia acununata Lhrenderg, 1834, p. 210.

Lepadela acuminate after Dujardin, Wal, p. 634.

Lepadelta chorea Berzins, 1982, p, 18, Fiz. 36.

Diagnosis: Dorsal lorica with (wo laleral keels;

caudal dorsal lorica variable, with small notch or

strong elanguled spikes, which may show short

sharp ar blunt ridge dorsally. Nominate sp. with

points of dorsal lorica slightly offset laterally.

Length 64-110 xis width 42-72 am; toe length

[8-35 parm.

Distribution: Cosmopolitan between water plants,

in littoral of [resh standing and running waters,

tolerates slight salinity increases, however prefers

acid walers (Koste 1978), Uncommon: pancon-

tinental, 10,0-29.8°C, pH 4.3-7.8, DO 6.1 ing! *',

28-1020 pS em 7, 8 NTU, Alk, 1,9 mp t-'.

Comenents: Several variant forms are known, On

present evidence these are not considered of specific

status. In ZL. eevmingta yexcosiuta Bartos (Fig. 4)

the dorsal lofica has six ribs, which run medially

1 SHIEI

ta form an elongated ridge to the end of the lorica.

i acuminate septemecostata Berzitis probably ty an

ecotypic variant. L. chorea Berzins (Tig. 4:1), k) is

inadequately figured, and appears io be a

preservation artefact of #. aeumiinuta.

Literatures Bartos 1955; Russell 196]; Koste 1978;

Shiel & Koste 1979; Green 1981; Berzins 1982; Koste

er ul, 1983,

Lepadella amphitropis Harring

bIG, 4:2

4. amphitropis Varring, 1916, p, 543

Loamphitropis vierorleasis Berzins, 1982, p, 10, Fig, 28.

Diaerosis: Lorica ecross-seclion rhombic duc to

curvature of the ventral lorica; lorica endl with small

notch; dorsal lorica end with one to [hive short ribs.

Length 69-76 yin; width 54-56 yom; heighi to 29

pen; Lows 20-24 pm.

Distribution: In Sphagnum, middle kusepe and Nth

America, Rare; N.SW,, Tas, Vic. 9.5-17.0°C, pH

3.1-5.7, 80.6-98.3 oS em -

J ierulure; Haver (958; Koste 19622 Koste & Shiel

1987a.

Comment: A population from the Tarwin R., Vic,

designated Lepadella amphitropis victoriensis by

Berzins (1982) on the basis-ot the deep sinus on the

ventral forica does not differ in this feature from

Ihe nominate species. The measurements of the ssp.

are, however, appreeiably different: lengity 86 pm,

width 60 pm, head-apertire 20 wm, dorsal sinus

depth 8-12 ym, ventral sinus depth t4 pm, foot-

opening 18=20%13-16 ym, to¢ length 14-18 pm. We

Tetain subspecific status for this taxon until the

extent of ecotypic variation can he determined,

Lepudella ungusta Berzins

FIG, 4:3

L. angusia Berzins 1960, pp. 85-86, Figs 7-12,

Diagnosis: Lorica elongated, ventrally flat, dorsally

convex; occipital margin with deep U-shaped dorsi!

sinus, deeper V-shaped ventral sinus; dorsal lorica

sparsely plinetate; dorsal pores unusually far

forward, in anterior Vsrd of dorsal lorica: foot

groove broad, distally not reaching rounded

pusienor margin of lorica; foot long, with distal

scemeor jonger than two proximal segments

vombined; terminal segment with two small dorsal

projections (Fig, +43c, d); toes long, thin, acute,

wider proximally.

Lengih WS gm (loriga 105 gm), width 46 0m,

Jorsal sinus 34x18 pm, ventral sinus 34423 pn, foot

groove 25x18 zm, foot and tues 65 jun (terminal

seginent 16 pm, toes 41 ym),

Disrributien: Apparently endemie Collected in

ROTIFERA FROM AUSTRALIAN INLAND WATERS 129

summer from the Plenty R., Morang, Vic, No other

locality known. Ecology unknown. Superficially

tesemtbles 1. el/intica (Fig. 5:7).

Lepadella apsida Harring

FIG, 511

J. apsida Harring 1916, p. 536, Fig. 891-23,

Diagnosis: Lorica almost circular; hed aperture

round, with posterior border drawn downward) toes

very short,

Leneth 70 pin, Width 60 pm; toes 12 pm.

Disiribution: \n riparian vegetation. N. America,

Asia. Rare: billabongs (NT), 1. Euramoao (crater

lake} (Qld). 28.5°C, pH 5.4, DO 6.2 mg 1-', 23 wS

cnmi -.

Literature: Green 1981, Koste 1981.

Lepadetta benjamini Harring

FIG, 5:2

L. benjamin Harring 1916, pp. 548-549, Fig, 93-8.

Diagnosis: Lorica owline broadly oval, head

aperture more or less tubular, narrow, with

ventrally-diregted margins; lateral lorica margins in

cross-section with bulging, beaded crim, or

downturned; foot aperture wider than long: toes

long, gently curved ventrally.

Length Hl2-Il4 pm; width §5-100 jum) height

44-52 amy head aperture 32-40 am; foot aperture

24x30 pm; toes 45=48 pm.

Distribution: N. America, 8. America (1. braziliensiy

Koste 1972). Rare: L.. Boort, Vic, N.T. billabongs.

20,0-24,5°C, pH 63-74, DO 51-88 me |,

42-1500 nS om |,

Literature; Koste 1981

Lepadella chengalathi Koste & Stel

PIG, 3:4

L. chengalatht koste & Shicl, 1980, p. 138, Figs 7 a-d,

Diagnosis: Rhomboidal lorica »with rounded

corners, Widest in first third; head opening stall,

nol-deflected ventrally; foat-opening semicircular

al widest point of caudal boundary of dorsal torica:

dorsal lorica weakly arched towards interior; three

Nexible foot-articulations, distalmost slender and

elongated; toes symmetrical, long, needle-like,

straight (Tig, 3:46); lorica cross-section shows highly

vaulted dorsal torica with slender downward-drawn

lateral wings; ventral plate with delicate double

convex keel (Fig. 5:4d),

Length 135-140 ym (loriva 96 um); Width 92 pm;

height 48 pin; head-opening width 28 ym: foor-

opening 28x28 pum; distal foor sezment 19-20 pm:

iovs 33-36 pm.

Distribution: Endemic, only known locality Li

Boort, Vic. 23.5°C, pH 7.9, PO6,0 me] |, 42048

em~, 67 NTU,

Lepadelle coraute nom. nov.

FIG, 5:3

Lepadetla latusturs f miuecronaa Koste, 1981, p, 119, Vip.

I4a, b,

non 4, macranate Schmarda, 1854 9.57, Pl 13, Fig, 20.

non Meropidia nueronata Daday, 1908, p. 30,

Diagnosis; General posterior outline of lorica

resembles 1. dafusinus, with extemely wide [oot-

opening; in 4, cornuta opening. wider in proportion

to lorica width, 1.2. body slightly more elongate;

caudal niargin lightly concave dorsally, ventrally

with v-shaped sinus flanked by distinctive pointed

lateral projections extending beyond dorsal margin:

points are bifurcare it lateral view (Fig, 5:3 inset},

with upper longer tips reflexed ventrally over shorter

tooth.

Lorica length 110 am; width 62 pam; foot-opening

width 40 «my; toe length 32 pm.

Distribution: Endemic, only record from Nakeen

Billabong, Magela Creek, NT. 29.C:, pH 5.47, DO

545 mel ', 44 pS cem~!, ulkal. 24 mg]

Lepadella dactsliseta (Stentuos)

FIG, 3:5

Metopicia dactvliseta Stenroos, 1898, p. 165, Vig. 3:1,

Lepadelia dactyliseta after Harring, 1916, p. 547, Figs

921-3,

Dlasnosis; Caudal margin of dorsal lorica flat, even

or convex; ventral margin with variable sinus (ef.

¢. fatusinus), loriea cross-section always shows very

pronounced dome; toes relatively. short,

Lenyth 93-100 pm; width 60-66 ymy height 40

pm, (oes 2432 um,

Distribution: Cosmopolitan in algal-rich standing

waters, #lso lake litorals. Rare, Vie. 8.5-27.0°C, pH

7.0=7.6, DO 7.0-11.2 mg 1 ', to 575 pS em ',

21-52 NTU.

J.ierature: Koste & Shiel 1980: Berzins 1982.

Lepadetli decora Berzins

FIG. 3:6

L, devora Berzins 1982, po 1, Figs Ja, h.

Diagnosis: Oceipilal margins nearly straight; dorsal

lonica with open-squared pattern, arranyed in three

rows; robust pores on last third of dorsal loricay

ventra) Jorica slight; foot-opening wider than long,

occupies Whole posterior margin toes. straight,

sharp,

Length 70 wm; width $2-54 ym; head-opentng 34

W. KOSTE & R. J. SHIEL

130

ROTIPFERA FROM AUSTRALIAN INLAND WATERS Ra]

vm; fooropening 12x3-32 pm; locs 24-26 am.

Oistrebution: Only known locality Laddon R,,

Victoria. Ecology unknown.

Cuereture: Berzins 1982.

Cormment: inadequately figured in Berzins (1982),

We have reproduced Berzins figure, which does not

show the patterning or pores described above. We

have wot encountered the species, however on the

basis of the above description réganl ir as valid.

Jepadella ellipiica Wulfert

FIG 5:7

L elliptica Wuliert 1939, p. 609, Fig. 21.

Diagnosis; Elongated lorica, hemispherical in crass-

section (cf. L. angusta), Wead aperture with shallow

U-shaped dorsal! sinus, deep V-shaped ventral sinus,

later with accompanying parallel-hordered lateral

cuticular pleats; distal foot-seement conspicuously

Jong, tapering slightly to base of (ocs: toes relatively

long, curved ventrally.

Length 1OK-H15 po width 70 zm; distal foor-

segment 28 yum! toes 20-22 ym,

Distribution: Sporadic in littoral decomposition

zone Known only from a billabong of the Magela

Ck, Jabiluka, NT. 255°C, pH 6.2, DO 2.9 mgt 4

fl ns om |,

Literuture: Kosice 1981.

Lepadella lathsintus (Hilgendert)

PIG, 38

Metonidia solidus larusnias W\laendort, 1899, p. 131, Fig.

Wy1sa,

Lepadelia latusinus atier Warring, 1913, p, 63,

Diagnosis: Lorica broadest in middle. tapers slighely

to cach end: cress-sectim vaulted hemisphere;

occipital margin Cowl-like, protruding; head

aperture relatively small) extremely wide {por

opening, small spikes at marains hooked dorsally

upwards,

Luriva length 80-92 pm; width to 64 xm: height

35 jm; head aperture width 24 zm; coes 24-28 pin.

Distribution: \n subtropical and tropical shallow

waters (bul recorded from Mt Cook, N.Z.), warm

stenotherm, Rare: N.t. 24.0-27.0°C, pH 4.5-6,3,

Ds) met |, 42 pS cm '.

Corurent’ & variant described by Koste (1981) from

Nankeen Billahong, NT as Z. fucesires cl, visenda

(Vie. 6:la) had features of L. lotusinus and L.

visenda Myers (Vig. 6;1b), Only one Individual yas

encountered, insufficient maternal for further

analysis.

Literature: Myers 1934; Russell 1Y45: Kost 1978,

Lepadella lindaui Koste

FIG, :2

L. lindaui Koste 1981, pp. t09=U10, ig 2

Diagnosis: Wide egg-shaped Jorica, ovcasiunally

with almost circular outline; head aperture ventral,

nearly circular, at umes bordered around lower purt

by cuticular ribs; foot opening paraholic (o

elliptical; caudal margin rounded or Jighths

indented. Lorica lat, Dorsal lorica in vross section

slightly arched with wide raised hump, which in

posterior ‘rd, accompanied by two converging

pleats, carries flat keel, terminating in fine poinr

reaching caudal margin, Lateral antennae papillae

small on end of oblique lotica fold (Fiy. @:2a)-

Sensory pit present on cistal foor segment,

Loriea length 80pm; width 68 zm; head aperture

22 pm; lorica height 20 um; foot aperture 23 pm:

toes 12 am.

Distribution: Only known lovalilies. Mombasa,

Kenya and Winnmrra billabong, Magelu Creck pear

Jabiluka, NT 285°C, pH 5.4. DO 6.2 mgt *, 23

esem | alkal. 2.7 mel ',

Lepadella minarut Rosle

FIG, 6:4

L. minor’ Koste, t98l, pp. HU-113, Fie. 3,

Diagnosis; Lorica Outline in upper part trapezoidat

with wings (ductitke, terminally concave (Fig. 6'4a,

b)); loriea tapers symmetrically from wings, ending,

in pointed protrusion covering foot-opening; head

aperture (ventral) smoothly indented, dorsally with

median notch; thin keel runs from noteh to

posterior forica margin; ventral plate flat, with

robust bead-like structures (uapatterned); darsal

loriea covered wilh 3-6 pm long rods in tows

paruflel fo marzins outside, and with keel om inner

rows, Rous sit on circular bases. Larcral antennae

dorsal, loweted approximately level with anterio;

margin of foot-opening, with pistil-like papillae:

foot 3-seamented; toes Jigtily curved ventrally.

pointed,

Lorica length 72-80 yom; width 76-80 um; height

48 pm; toe Jength 24 gm; head aperture width 24

am, depth [2 pm; foot-opening width 16 am,

Viv Sit, Fepadedas apvida Harring: (i) ventral; (b) lateral: (c) erosssecuan. 2. 7. benjqrntin Hartinus (a) vennal,

(6) dorsah Ce) lateral; (el) ehosseseonon, 3, 2, curnete (hoste), ventral, o£. chengalathl Koste & Stel: day ayeninals

Ab) chstil segment of foot and lags: (2) peeipital muruin, dorsal, (c) cross-section. 3, 7.

lect viiseia (Stents): (a)

veriral, (b) fareraly (eo) vross-soehiono(, 4, deeura Berzins: (a) darsal; (b) ventral 7%, effiotica Walhert. (ay dovsal,

(b) serial, be) orgos-seution, Ke d furasdias (ilgeneonty: (ay ventral, (O) luteal: le) cross-section, bo S Raley

Harring (910); do arfrer Komi Qa aL after Rosie ie Shieh (O80), 6 alter Berane (W8abe 7 otrer Wallen (939),

Seale lines 40 yan.

W. KOSTE & R. J. SHIEL

132

ROTIFERA FROM AUSTRALIAN INLAND WATERS 133

Dixtrbudion: Endemic, Leichhardt aid Winmurra

billabongs. Magela Creck near labiluka, NT,

28.5-30,1°C, pH 54-55, DO 5,5-6.3 me |-',

23-58 zS cm ‘, alkal. 2,7-2.9 mg |‘.

Lepadella neboiss? Berzins

FIG, 63

L. aehotss Herzins 1960, p. 83, Flies I-J.

Diggnasiz Outline of body broadly ovate, greatest

width slightly posterior to middle of burly; Jorica

strongly dorso-ventrally flattened, without ridges;

anterior dorsal margin straight, vent’al margin with

wide V-shaped sinus and short fold present.on each

side. lorica rounded posteriorly, Sinus and fold

slightly behind the middle of body, on each side

of lorica, with corresponding pores on dorsal side;

foul groove nearly as long as wide; foot rather short,

terminal segment longer than first and second

seginents Combined; toes relatively long, divergent,

pointed at apex.

Total length 105-108 am (lorica 78-81 pm); width

of anterior lorica 63-66 xm; ventral sinus 11-13 ym

deep! foot groove 18-19x!2 ym; foot 14-16 wmy toes

mM) wm.

Distribytion: Rndemic, King Parrot Creck. near

Kinglake West, Vic.

Commeni: Resembles L. monodi Berzins from

Senegal, Aftica, but differs in the form of the lateral

sinus and the pores for lateral antennae on the

maypin.

Lepadelia ovalis (Muller)

FIG, 6:5

Brachronys aviclis Muller. 1786, (. $45, Fig. 49:]-3,

Lenvielia ovelis after Ehrenberg, 1830, p, 45, bie, 7:4

non Metypidia avalis Anderson & Shephard, (892, p. 78

(sec, virrea).

Divenosis: Lorica outline aval lo circular; wide

space between internal organs and loriva margins;

ventral lorica nearly flat; lorica at rim thinly

tapered; narrow striated area at outer margin from

head aperture to foot-opening; outline of toot

opening variable; gestric glands ollen lobed and

long-stalked. Juvenile unimals sometimes have

caudal notch dorsally over foot-opening. Males

known, Resting egg shell covered with robust, lightly

curved spines.

Lorica Jength 90-}70 ym; width 70-140 jm;

height 25-30 ym; toe length 22-36 pan; foot opening

to 48 pm deep, 32 um wide,

Distribudion; Cosmopolitan, between macrophytes

in benthos of fresh and saline waters, Uncomnien:

pancontinental, Ecotypes described from Australian

waters include a form from the Magely Creck, NJ

(Fig, &5d) and a minute form from L, Dulyerton,

Tas (Fig. 6:Se). 8.2-24,0°C, pH 5.8-7,7, 273-3330

as an '

Literature: Koste 1978, 1981; Shiel & Koste |O84;

Kosie & Shiel |986a,.

Lepadella patella (Muller)

FIG, 7:1

Brachionas patella Miiller, 1786, p. 34), Fiz. 48:15-19.

LepadeHa patella after Bory de St Vincent, 126, p 8A.

Diagnosis; Lorica outline ovoid. to circular: dorsal

lorica domed, without wings; ventral lorica fat,

occasionally with somewhat overhanging margins;

wide lorica forms more domed in cross-section than

narrower forms (Wulfert (1960) described Mat and

domed forms from Sphagnum); variable foot

opening (Hauer (1962) found caudal corners of

foot-opening extended into spines, while one

specimen described by Wulfert fac. cif, had almost

circular foot-opening); occasionally, dorsal lorica

adjacent lo foot-opening has weakly distinguishable

folds, Male known.

Length 120-145 pm; lorica length 70-110 pm;

width 65-90 ym; leneth:width ralio mostly < 1.2;

toe length 20-35ym; male 110-120 pm.

Distribution: Cosmopatitan, euryopic in fresh and

suline water's. Common: panesntinental. Ecotypic

variants are known from NUT. (Fig. 7:le) and Tas.

(Fig. 7;lg). 10,0-24.5°C, pH 3,9-8,8, BO 7.8-11.6

meg !-', 92-6600 uS cm-', 3,0-262 N'TU.

Literature: Koste 1978, 1981: Shicl & Kaste 1985,

Comment: Several recognized subspecific variants

of L. patelia are recorded from Australian waters,

The status of these taxa may change with detailed

systematic work;

Lepadella patella bitoha Hauer

FIG. 7;2

Lepadetla patella biloha Hauer, 1958, pp. 27-24, Vig. 214.

4, patella |. bilaba (Hauer) by Koste, 1978, p, 185:

Diagnosis; Lorjca outline resembles L. patella,

however differs from f. typ. in three features: lateral

corners of toot-opening reflexed dorsally (Fig. 7:2);

Fig. 21, (a) Lepodelly latusinus cf. visenda; ventral; (bh), visendi Myers, ventral; 2, /. Jindgul Kostey (a) NLT. form,

dorsal; (bt ventral; (ey Mombasa form, ventral; (cl) dorsaly (2) (rass-section, 3, 1, #eboissi Berzins: (a) dorsal: (b)

ventral (e) cross-seclion, 4, 2. wrimaras Koste: (a) dorsal: (by ventail, (c) cross-section. 5, 2..dveis (Muller)! (a) ventral;

UD} laterals (c) cross-section, (UW) NT, form; (2) Tas, form la. 2. 4, Sd, after Koste (1981); 1b, after Myers (1934); 3,

after Berzins (1960); Sa-c, after Harring (1916); Se, avier Shiol & Koste (1985), Seale lines 50 jr.

W. KOSTE & R. J, SHIEL

134

Fig. 7:l, Lepadetla patella (Miiller): (a) dorsal; (b) ventral; (c) lateral; (d) cross-section; (e) N.T. form, ventral; (f)

cross-sectian; (z) Tas. form, ventral. 2, £. patella bifoba Hauer: (a) dorsal; (b) cross-section; (¢) ventral; (d) lateral;

(e) Tas, form, ventral. 3, L. patella oblunga (Ehrenberg): (a) ventral; (b) ventral; (c) cross-section; (d-f) variations

in foot-opening morphology. 4, L. patella similis (Lucks): (a) ventral; (b-d) different loriea forms, ventral: (@) cross-

section: (fg) fool-opening, ventral; (hb) L. buengensis Berzins, ventral. la-d, afler Harring (1916): le, f, after Kasre

& Shiel (1980); lz, 2e, after Koste & Shiel (1986); 2a-d, 4b-g, after Hauer (1958); 3a, ¢, after Wulfert (1939); 3b, d-f

alter Bjorklund (1972); 4h, alter Berzins (1982). Seale lines 50 ym,

ROTIFERA LROM AUSTRALIAN INLAND WATERS mS

ventral lorica with lateral concavilies, side margins

deflected downwards (Fig, 7:2b); nwo stort cuticular

folds of caudal dorsal Jorica.

Length $0-107 pm; width 59-64 am; height 36

pm; toes 19-25 um,

Distribution: Europe. Rare: Tas., Vic., southwest

WA. 11,8=24,0°C, pH 3.5-7,.9, 183-2130 ecm |,

( Herature: Koste ef a/, 1983: Shiel & Koste 1985.

Lepadella pulella ablonga (Ehrenberg)

FIG. 7:3

Syuaniuia oblonga thienberg, (834, p. 220,

Lepadelld Oblonwa (Ehrenberg) alter Harring, 1913, ). 64,

Lepadelty pulella t, oblonga atter Wulfert, 1960, p, 285,

Rig. 24,

Diagnosis) Distinguished from £. patella only by

relatively low lorica height and narrow head

aperture. Lorica outline mostly elliptical,

Lorica length 80-110 pm; width 50-70 wm;

length:width ratio >1.5; toes 20-25 yrn..

Mistribution, In Tresh and brackish water pools,

often sympatric with the nominate species and L,

patella similis. Bidrklund (1972) demonstrated

blometrically distinguishable populations, however

infergrades with 4. patella typ. occur,

Only Australian record from Sheepwash Billabong,

near Yeu, Vic, 17iv.76(Shiel unpubl.). 19.9°C, pH

7.2, DO 8S mye) , 85 pS em |, Secchi trans-

parency RO cm,

Giterature: Koste 197i

Lepadelta patella similis (ducks)

FIG. 74

Metopidia similiv Lucks, 1912. p. WY, Pie, 34,

Lenadelta similix (Lucks) afler Hauer, 1925, p. 17

t patella vay, similis atten Remuine, 1924, 9, Fig. VAA-E.

“. budngensis Bereiny, 182, p, 1 Me. 9.

Diagnosis, Lorca outline ellipucal: taay-openiig

with teinforced uppec arch (Fig. 7:4 f, 2}, mustly

hexagonal with angled edges and base straight to

indentud. Not reliable taxonomic character because

oF vanability in foot-opening. Distinguished fram

L. guitella (s,.sir,) by lorica lengih:width ratio (> (2)

and overall smatler habit.

Length 68=85 pm; width 49-60 xm; wes 17-21

pin,

Dixiribution: Bewween macrophytes, detrivus, in

psamnmon and rock pools; fresh, brackish and salt;

pH to 10 (Koste 1978). Single record from Australia

(as £. buangensis Berzins), but recognizable as a

nreservation artefact of L, patella similis, Mt Donna

Buang, Vie

Lepadella plilota Berzins

FIG. 834

C. pliléta Berzins, 1960, pp. 83-85, Figs 4-6.

Diagnosis; Body ovoid in outlines venira) surface

slightly medially convex, dorsal surface very

strongly 50; dorsal median line with sharp, uneven

longitudinal ridge; frontal margin dorsally convex,

with blunt projection in the middle; venually with

rounded sinus; laterally, lorica enlarged with three

pairs of thin, transparent tamellary projections;

lorica prolonged posteriorly (to rounded Jobe: crass

section somewhat triangular,

Length 72~75 wm width 60-67 pm; ventral sinus

depth 8 xm; length of foot groove (2-15 ym; toes

12 yarn,

Distribution. Endemic. Only known locality

Creswick Creek near Clunes, Victoria, Ecology

unknown,

Comment Resembles J. ¢riprera (Ehrenberg) but

differs in (he form of the trantal margin of the

Torica, the sharp dorsal ridge, and the presence of

lateral cuticular projections (termed "flappers” mn

the original description althaugh they have ne

muscular contact],

Lepadella quadricurinata (Stenroos)

FIG, 8:2

Melopidia quadricarinaia Stenroos, 1R98, 5. 165, big. 3-2,

Diagnosis: Dorsal lorica with four short ribs over

foot-opening; posterior end tapering to short

tongue; ventral lorica flat; foot opening variable.

Lorica length 81-92 ym; width 61-72 wm; height

1o 34 um; toe length 20-26 jam.

Distribution: Littoral, between macrophytes, pr

6.2-10.2 (Kaste 1978), Only record that of Berzins

(1982) trom Coghill’s Creek neat Tourello, Vic. /,

q. Sexcarinata (Klement), recorded fram the same

localily, muy be an ecotype,

Literature: Koste 1978,

Lepadella yuinquecostata (Lucks)

FIG, 8:3

Metopidia quinquecestata Tucks, 1912. p. 126, Pig. 47.

Diagnosis: Lorica pyriform, narrowing towards

head; dorsal lorica with median low keel which

hegins as witely separated ridges behind callar.of

head aperture; two or three pairs of side ribs:

distinctive cross-seclion (Fig. &:3c): Jorica end

variable, sometimes with short ridges foot-opening

variahle,

Lorica length 92-112 ym; width 60-70 wm; tne

lenyth 22-30 yin,

Distributions Cosmopolitan in tirural, efter

W. KOSTE & R, J. SHIEL

136

ROTIFERA PROM AUSTRALIAN INLAND WATERS ity

collected in Sphagnuen. pH 4.$-6.7, temperature to

PC (Koste 1978). Single record: dam near

Chillagoe, Qld.

Lepudelta rhomboides (Gosse}

FIG. 84

Metopidivrhamboides Giosse in Hudson & Classe, 1886,

p. 18, big. 25:70,

Lepudela rhamboides alter Harring, 1913, p. 65.

Diagnosis; Dorsal lorica. with moderately high,

mare or less arched median keel, bounded by lateral

grooves; anterior margin Width 4 lorica length;

dorsal sinus broadly U-shaped; ventral sinus ¥-

shaped; dorsal aperture of head-opening with collar,

whieh can be bordered by cuticular thickening; foot

opening narrow, U-shaped with parallel sides: distal

foot seament long (cl. L, elfipticah; toes short (<4

body length) slightly decurved, tapering to points.

Lorica length }10-120 pm; width 35-88 yam; wwe

length 20-28 pm,

Disicibution: Cosmopelitan im periphyton, pH

Tolerant, Rate: N.S.W., NLT, Tas,, Vie. 10.5-30,7°C,

pR 4.7-7,.9, DO 4,6-6.0emg 1 ', 23-750 pS om |

to 67 NTU, alkal, 2.7~2.9 me L-'.

Comunent: Two variants are kitown From Australia,

described elsewhere as ssp, form or var: df.

rhamboides hauer{ (Wulfert) (Fig-8:5) from a stack

vam south of Burnie, Tas. (168°C, pl 4.7, 42 uS

em and L. /homboides caringta (Danner) (Fig.

8:6} from the Uroken R. at Benalla, Vie. (27,0°C,

mil 7.6, DO 7.0 mp t-', $2 pS em '),

Literature: Kutikova 1970; Koste 1978: Koste er al.

IDS; Koste & Shiel 198ha,

Lepadella rarrenhursi (Lucks)

FIG. Yt

Metapidia redtentyied Lacks, (912, p. 127, Pip. 48ay 0,

(.rovenburei atler Harring 1913, p, 65:

Digenosis: Side margins of lorieu cross-section

earend to pointed tips below level of median ventral

bulge; head aperture dorso-ventrally deep, only:

ventral sinus; dorsal margin straight, without sinus:

fool-opening variable. Resembles £. dactylisera,

Can be-contused with Colitrella spp. in lateral view,

Lorica length 75 am; width 50 pm; toes 25-28 pm,

Oistribution: Isolated occurrences in Sphagnum and.

peat swamps. Only record Riddel’s Creek, Sunbury

Vic.

Liferature: Koste 1978, Berzins |82.

Lepadella tana Koste & Shiel

FIG. 9:2

Lepadedle tara Koste & Shiel, 1987, p. 102, Fig. 19.

Diagnosis: Loviva hraadly ovoid, ventral loriea Mat,

dorsal micdially conver tn median cross-section:

dorsal margin of head-opening almost straight.

ventral margin with weak V-shaped sinus: fold

behind middle of bedy on ventral surface; with

pointed, slightly curved spine on either side, directed

apically; foot-opening broad; toes relatively long,

straight, sharply pointed,

Lorica length 79 xm; width (medially) 61 yam;

anterior width 29 am; foot-opening 18xl8 pm; toes

29 wm} fateral spine length 18 am.

Distribution; Eudemic. only known lucality L,

Pedder, Tasmania. 165°C. pH $.3, 46,2 aS emo.

Comment: Resembles L. nedoissi, a Victorian

endemie (Fig. 6:31, however |he latter has ventral

folds (no spines). and differs in foot and toe

morphology,

Lepadella triba Myers

FIG, 9:3

Lepadella triba Myets, 1934, pp. 4S. Figs 10-12.

Diagnosis: Lorica elongated; crostseytion shallow,

evenly arched dorsally; dorsal margin of head

aperture almost straight, ventrally broadly V-

shaped; no stippled collar; ventral lorica with

distinctive Euch/anis-like slightly clevated mid-

section (Fig. 9:3c), lateral margins run to head

Aperture or diverge io side apices; foot opening

Maring: distal foot segment approximately 4 toe-

length; toes long, tapering to slender drawn-out tips.

Lorica length 71-80 am; width 42-45 jim; ventral

sinus depth 12 jan; foot opening depth 20-any; foot

length 26 wm}; distal segment 14 jam; toes 30-33 pn.

Distribution: Noth Ameriga, Single record from

Old (Berzins 1982) needs verifieution.

Lepadella Sriptera (Ehrenberg)

FIG. &:4

Metopitia triptera Ehrenberg, 1830, pp. 74-83,

Lepadelta triptcra Ehrenberg, 1832, p: 72,

Fig. Rl, Lepedelia ptilota Berzins: (a) dorsal; (h) ventral; (ec) crass section. 2) 2. quadricarinate (Stenroos):; (a) dorsal:

(h) ventral: (¢) cross-seetion: (0) dorsal: (ey ventral; (f) dorsals (8) cros#section. 3, 2, guinguecostate (Lacks): (a)

veniral; (b) laccrak; (ev) crass-seetion; (d) dorsal. 4, 2. rhermdbeices (Gosse): (a) ventral: (b) lateral: (c) dorsal: (a) cross

secuion. $5, 2. rhamboides hauert (Wulterl): (a) SCT. form, dorsal; (b) Luropean form, ventral: (c) utass-secrion. 6,

{ rhomboides Corina (Dunner): (a) ventrals (b) averal (c) venieal: GH cross section Spee. dh CO} Choss-secuiOn spec,

11) loriea stinfaee morphology. L, alter Berzins (1960); Jaw, 1 Shu, alter Bartos (1959): 2d, fy after Pejler (1982);

Te alter Biciblugd (972). 4, 4, ater Harring (916) 3, aller Koso 98th after Donen (943). Seale lings 30.j0i,

138 W. KOSTE & R, J, SHIEL

ROTIFERA PROM AUSTRAI LAN INLANIY WATERS {39

Diagrosis; Lorica outline vatiable: circular ta

chombic according to Koste (1978). Dorsal lorica

vaulted, with high, somewhat protruding median

keel; head aperture with shallow U-shaped sinus

ventrally, usually slight median notch dorsally:

forica surface may be Vinely striped (Fig, 9:41) wirh

small founded proirnsions, mast, however, are

smueth,

Lorica length 58-65 ym: width 30-57 pm, toes

13-22 wm,

Oistelution: Cosmopolitan in fresh and brackish

Waters, and on muddy litrorals. Rares NSW. NT,

Old, Tas, Vio 12.4-20.0°C, H 48-77, )7-3330)

Soom '.

Literature: Shriel & Koate 1979; Koste (981+ Berzins

1982,

Lepodella tyleri Koste & Shil

FIG, 9:5

Lepudella tyler] Koste & Shiel, W8Fa, p. 37, Pigs 3, 4.

Diogaosis: Rigid loricu of trapesoidal outline; cross-

section nearly triangular; dorsal plate with median

keel} dorsal and yentral plates finely pranulated;

head aperture lightly convex dorsally, deep V-shaped

sinus ventrally, with lateral lamella; caudal Jorica

with slight constriction aboye blunt lateral corners;

fool-opening broadly flared; three foot segments,

distal with caudal antenna: toes very thin,

elongated, pointed.

Total length 123-125 pm; lorica length 78-SD pm;

greatest width 58 vm) height 36 pm; head aperture

width 21 wm, depth 22 pm; foot opening 18x18 um:

distal foot segment 12 4m; toes 32 pm,

Distribution: Endemic; only known locality a stock

dam south of Swansea, east voast of Tasmania,

134°C, pH 6.0, 390 nS cm ', 82 NTU,

Lepudetla vitrea (Shephard)

FIG. 9:6

Metopidieavatis Shephard in Anderson & Shephard, TRYD,

p. 78, Fig. 66a, Won ML valis (Muller, 1786)

M, vitrea Shephard, Wil. p, 53

Lepadella wired alter Harring, L913, p. BS.

Diaynosis: Ovoid, tansparent Jorica, taperine to

cach end from widest part posterior to midline:

head aperture slightly convex dorsally, without

sinus, ventrally with deep, broadly elliptical sinus:

dorsal margin of foot opening convex between

lateral points of slightly outcurved caudal Jprica

projections; ventral foot apenine deep s-ellipse;

distal foot segment almost twice length of proximal

sépments, more than half length of toes, toes short,

Tapering to pointed fips.

Length 123 pam; width 36 wm, [Na other

measurements given in ihe original description,

however from Fig. 6 in Anderson & Shephard

(1852), on the basis of cheir lengrh/width

measurements, the following wre estimated: head

aperture 23 jun wide x 20 pm deep; foot openiny

20 am wide x 22 ym deep; distal tue joint 12 pm;

tors 17 ym]

Distributions Subsequent to the ormeinal Cine

LBrizghton, Victoria), L. vitree was recorded from

South Westland, N.Z. (Russell 1954), Dimensions

were slightly different (14 «m Jong, toes 20 ym),

with “a series of dots round the lateral edge of the

dorsal plate”. L. virrea is possibly more widespread-

Comment 1. vitrea was synonymised with LZ.

fu(usinus by later ceviewers (ee. Koste 1978),

however can be distinguished from che latter by

more elongated lorica (lengthiwidth ratio 2.2 vs

<15 in L. fitusinus), absence of cowl-like head

aperture, elongated distal foot-segment and

markedly shorier 10¢s,

Lepadella williams Koste & Shicl

FIG, 9:7

fepadetla williarns! Koste & Shiel, 1980, pp. 138-139, Fix

Olasnusiss Smoothly oval, highly vaulted lorica,

with ventrally-directed head-opening; dorsal lorica

tapers caudally to variable-length laterally curved

point, foot-opening narrow, ending.at base of dorsal

lorica projection; three foot segments af similar

length; long, dorso-laterally curved sharp toes,

Lorica tength 112-6 pm; width 60-64 pm;

height to 56 xm; head uperture 33-36x28-30 pm;

caudul lorica projections 19-28 p~m; foot opening

17-20x19-20 jam, distal fant segment 10 jm; toe

length 36-40 um,

Distributions Kuala Lumpur, Malaysia, and Jabj

Iuka, NT. only known localities, 25.5°C, pH 6.2,

DO 29 mel |, 62 pS ecm ') 5 NTU,

Fie. 4, Lepedella rotienburgi (Lucks): (a) ventnats (b) fatural: (¢) verttrals (el) cross-section 2, £. ane Aoste & Shivl,

venifl, 3, 2. frida Myers (a) dorsal; (b) ventral; (c) cross-section. 4, 2. edprera (Etirenberu): (a) dorsal: (b) ventral:

{(c) eross-Keurion, (d) cross-sectiom: (¢) lareral: ((-h) different Joriea forms, ventral; Gi) Tas. farm, ventral; (j) 2d ‘Tas,

form, wential 5) 4, Wert Koste & Shiel: (a) venenaly (4) dorsal (c) lateral QD) cross-section: 6. L.

Witrea (Shephard }:

(a) dorsal; (b) ventral. 7, 2. williams? Koste & Shiels (a, b) lwo morphs, ventral: (c) fateral: (d) cross-section. 1, b,

afler Caling (939); bo, d, after Bartos (1959), 2, 4), 5, ier Koste & Shret (ORG): J, alter Myers (1934): tit hy, ates

Having (1916), 5, ater Rosle A Shiel (9874); Git. orig, 6h, aber Shephard (90s 7, aller Koste Moshe (4880) Seale

lines 40 pm,

140 W. KOSTER & R. 1. SHEEL

Key to species of the genus Lepudelliz

(Heterolepadelia)

1 PR Pec (Gs Le 2) | nnn DO re eat

Leariea thombotdal.. 00.00... 2 eee ee Arar)

Ze). Head aperture with Veshapead dorsal ancl ventral

KIMUSOS, oo sere bh. HY, Aeteredavivia Tadeey

Head aperture with only a sential sinus, dorsal

Margin stright. ....... L. 7H.) upsicara Myers

AA), Larice wilh posteriori pore spurs, loot puoove

Wilh lateral pointed piujections... ,, -

; Cod. CAD @hrenberai (Peri)

Lateral wings of loriva.arid postenar margins af foot

groave rounded... .. L. UY) heteroxtvle Murray

poets

Lepitdella (Heteralepadella) apsicora Myers

ric. We

Fepadeia upsicare Myers, 1934, pp. 5, 7, Migs t6-18.

Diaenosis Loyica oval; crass section shallow, evenly

arched dorsally; dorsal anterior margin nearly

straight, ventral margin with deep V-shaped sinus;

stippled collar present; foot groove wide, flared

posteriorly; discal foot joint 2x length of combined

first and second joints; foot twisted, left toe lies

beneath rights (oes asymimetrie. right 24 fenuth of

left,

Lorca lengrh 80 jams width 65 am; ventral sinus

depth 18 prs anterior points 28 pm. foot groove

Zt pms foor 30 pny distal segment 20 yr; right foe

30 pms lett yam.

Oistethution: North America, Two records from

Attstrahia: L. Boort, central Victoria, and Winmurra

Billabong, Magela Creek, NUT. 23.5°C, pH 3.4-7.9,,

DO 60-6.2 me | 1, 23-750 pS cm

alkal, 2.7 mel '.

hiterufure: Koste & Shiel 1980; Koste 1981.

» 67 NTU,

LL. ¢H). ehrenhergi (Perty)

PVG, 10:2

Niutawania ehrenhersi Perty, 1850, p. 20.

Moerapidia clhrenbergi otter Jennings, 1894, py 26.

Lepudelia ehrenberge ater Harring, (913, p. 63

Lvagnosis: Lotica rhombic, with posterolateral

margins produced to triangular, dorsully ridged

spurs reflexed upwards in cross-section (Fig. 10:2¢);

smaller painted spurs on each side of foat granve;

stippled collar on ventral plate, with less obvious

Jornal stippling; foo. groove Vi lenegrh of lorica,

rounded anteriorly, flaring slightly posteriorly; foot

stout, 3 length of lorica; distal foot joint longest;

toes long, asymmetric, laper evenly,

Loriva tength 70-95 pin; with 70-90 pn; toe

length 27-32 wm and |9-27 yon.

Divtribution: Cosmopolitan in periphyton, Orie

laria and moss. Rare; in billabongs and vegetated

lake margins in N.SW,, NTL, Qld, Vic. 24.0-29.2°C,

pH 6.3-7.5, DO 5.1-8.0 mg 1-', 42-85 pS em |, 6

NTU.

Literature: Shiel & Koste 1979; Koste & Shiel 1980;

Greeiy 1981; Koste 1981.

L. (H.) heterodactvia Fadeev

FIG, 10;3

L. (F.) heterodacivla Fadeey, 1925, p. 73, Phot, big. 8,

Diagnosis; Lorica oval; dorsal and ventral mareins

of head aperture with deep, almost triangular

sinuses; last foot segment longer than preceding

segments.

Lorica length 106-113 pm; width 70-75 pro; distal

foot segment 22~23 pm; toes 33-30 and 20-25 jum.

Distribution: Rurope, S. Ameriva (Amazonia),

Single record, L. Boort, central Vie. 20.0 °C, pH

7.4, DO 8.0 ing Io!

Literature: Kutikova 1970; Kaste 1978: Koste & Shicl

1980,

1, (fh) heterustvia (Minrray)

IG, 10.4

Metopidia heterostvla Murray, (913, p. 489, Ph19, Pig.

6a-c.

Diagnosis: Lorica rhombic; lateral wings rounded

rather than pointed (cf. L, ekrenbergi), with tips

more or less deflected dorsally; ventral lorica wider

than dorsal in region of head sperture; stippled

collar present.

Distribution: Probably cosmopolitan in littoral

zones, stagnant waters, Rare: N.SW., Tas., Vic.

10.0-23.5°C, pH 5.8-7.9, DO 40-98 mg 1°,

80-750 pS cm |, 2-67 NTU.

Literature: Koste 1978; Shiel & Koste 1979: Koste

& Shiel Lgo.

Not revordéd from Australia:

L..(H.) eyrtopus (Harring), Eastern Europe, N. and

Central America.

A single free-living species of the subgenus

Lepadella (XNenalepadella) is tecorded from

Australia. In view of the preferred habitat ol most

oi the other known species of the genus, ie. the

branchial chambers of a range of crustaceans, it is

likely chat orher species oceur here but have heen

overlooked.

Lepudelle (Nenolepadella) mondactyla Berzins

FIG. 10:5

Lepadolla monodactyla Reveins, YU, pp. 5,6, Fizs B=,

ROTIPERA FROM AUSTRALIAN INLAND WATERS

Fig. 10:1, Heterolepadella upsicura Myers: (a) dorsal: (b) ventral; (e) cross-section; 2, 4. chrenberei (Perty): (a) dorsal:

(b) ventral; (c) cross-section; (d) lateral, 3, Af. heterodactyla Fadeev: (a) ventral; (b) toe detail. 4, /1. heterost via Murray:

(a) ventral; (b) dorsal; (¢) lateral; (d, e) different cross-seclions: 3, Xenvlepidella rronodaciyla Berzins: (a) dorsal;

(b) ventral; (¢) cross-section, |, 2, aller Harring (1916); 3, after Kudkova (1970); 4, after Harring (1916); 5, after Berzins

(1960), Scale lines 50 pm,

Diagnosis: Lorica outline elliptical; dorsal lorica

arched, punctate or lined; ventral plate with light

bulge medially, otherwise unstructured; head

aperture with V-shaped dorsal and ventral sinuses.

Total length 170 um; lorica length 127 wm; width

77 wm; foot-opening 31x10 um; toe 57 pm.

Distribution: Madagascar, variants known from

Brazil. Single record, billabong, Jabiluka, NT.

25.5°C, pH 6.2, DO 2.9 mg 1-!, 62 zS cm !.

Literature: Koste 1978; Koste & Shiel 1980,

Not recorded from Australia:

L. (X.) astacicola Harring, L. (X.) borealis

Harring, L, (X.) branchicola Haucr, L. (X.) haueri

Rodewald, (.. (X.) lata Wisniewski, L. (X.) parasitica

Hauer, £. (X.) pygmaea (Gosse). See Koste

1978:198-200 for detuils).

Acknowledgments

Collectors acknowledged in our earlier Papers

also contributed further material to this work, Their

assistance is appreciated. The Deutshcen For-

schungsgemeinschaft, Bonn-Bad Godesburg,

provided microscope and photographic facilities to

WK.

i2 WoRKOSTR & RoI, SHIP

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PRELIMINARY INVESTIGATIONS OF BEACH RIDGE PROGRADATION

ON EYRE PENINSULA AND KANGAROO ISLAND

BY A. D. SHORT*, R. C. BUCKLEY**, & D. G. FOTHERINGHAMT

Summary

Field investigations along 2150 kin of the South Australian coast including the Great Australian

Bight, Eyre Peninsula and Kangaroo Island revealed numerous sites with low wave energy beach

ridge and occasional chenier plains. Examination of 14 mainland sites and six sites on Kangaroo

Island indicate that all ridges are wash deposited during episodic wave events, probably over a

period of decades to centuries. The ridges overly intertidal shelly sandflats. The oldest innermost

tidal flats date at 6.3 ka and 5.9 ka BP, but the oldest ridge dated at 4 ka BP suggests a period of

tidal flat formation and progradation is required before a source of sediment and plat form for ridge

formation is available. All ridges and ridge sequences dated between 4 and 2 ka BP with no ridges

material younger than 1.8 ka BP. While this confirms a delayed start for ridge development it does

not necessarily imply no present ridge building as reworked, older shells may be incorporated in the

modern ridges. The age and elevation of the sites lend support to a published sea-level curve for

northern Spencer Gulf. The limited evidence available suggests that sea-level reached its present

level in the Bight by 6 ka BP with no conclusive evidence for changes since.

KEY WORDS: Beach ridges, sea-level, Eyre Peninsula, Kangaroo Island.

Hansuctions af the Raval Suctety of 5. odast, (1989), 1304) 144-16).

PRELIMINARY INVESTIGATIONS OF BEACH RIDGE PROGRADATION ON

FYRE PENINSULA AND KANGAROO ISLAND

bv A, D. SHOR’, RoC. Buckiey**, & D, G. FOTHERINGHAM/

Sutnmary

SHORE. A. DL, Boehtbh, ROCL & PorirkingHam, DOC. (1989) Preliminary investigations of heaeh ridge

mogradauion on Eyre Peninsula and Kangaroo Island. 7rans. R. Soc. §. Aust. 113(3), 145-161, 30 Novernber,

1989,

Meld nywstwutons dong 2150 km al the South Austrilian coast indudiiy the Great inant Breht,

Eyre Peninsiute apt Kangitrow [stand revealed mumeratts sites with los wave encres beach ridge and pecasiioal

ehemer pliitis EXAM Aron Ol 14 mainland sites aad sis sites on Kangaroo tykind indiguhe hat all mages

Ones wiih deposited during episadic wave events, probably over a period of devades Wy contunies Chie rides

overly inferiehil shelly samdflives, the oldest innermost sida) (has dare at 6.4 ku ane 29 ba WPS bur the

midest Hidhve daledal ka BP stiepests a pertod of tidal tid fornition and progradariny 1 tequired before

weenie OF scdiment and phiitform for ridge formarion is available, AN rilges ain ride scquences died

belween 4 and 2 ke BP with nev ridves manernal younger clan 8 ka BP, While this confirms a delayed

stint Ton fidve development itdoss, notnevessarily triply no present ridge buildin as reworked, older shells

Iidy he ineorpardied ithe modern ridges, The age and clevarion ob ute sites tend support toa published

sda-hvel burve lor dortherd Speaver Gull) The limited evideriee availible sugeests chat sea-level roagheal

is present level in the Bishi by 6 ka BP with no conelusive evidenee tor changes since.

Kru Words: Beach ritlyes, sea-level, Byre Peotnoula, Kangaroo [ylud

Introduction

The South Australian coast, west of Whyalla and

including Kangaroo Island and the 7& islands of the

Eyre Peninsula, represents 60%; of the state's

coasting. Phe cous! contains a variety of Flalacene

Shoreline types ranging from. tide dominated.

saudflats fo, Will iereasing eave energy, cheniers,

beach ridges ind regressive (lorechine ridges) iid

transgressive barrier-dune svsiems (Short f988a).

The Jatter include some of Australian's ogost extensive

voustal dune systems, including more than S00 kin

of clifftogy dunes (Shore t988b).

Whilst the 3700 kis, Aust. coast possesses a

wide range of Holovene coastal depositional

systenis, The nutune and evolidon of rhese systents

received Only cursory attention unit Chis decade.

Revional studies und detailed invesdauion have

only just begun to address (ie status of Lhe couse,

In the cull regions Hologene sedimentation and

Pluistovcne and Holocene sea-level history

wxsocraicd wath beach tides plains huve been

investivaled by Burne (1982), Belperio et at (1983),

Hails & Ciostin (1984) and Belperioa er af (988).

fails & Ciostin (984) and colleagues also presented

aseries of papers on the evolution af northern

Speneer Gull and its implications for modern

sedimentation. In St Vincent Gulf, Bowman &

Harvey (1986) conducred extensive coring and

= (ous) Studies Unit, Dental Cremgniphy, University

af Sydney, Sydiwwy, SSW. 200A.

sOAMPEL. 162% Miller St, O'Conmmen, ACT 2600,

1 Coast Managenrtent Brareh. DA. Oo Box 667,

Adtvlnide, So Aust S00),

cating of the Le Fevre Peninsula to reconstract the

evolution of Wiis imipertant beacheridge cand plex.

On the open coast three revenl regional stuaies

of the Seuth East, Kangaroo Island and Eye

Peninsula (Short & Hesp 1984; Shan &

Borhenmyehainn 1986; Short er al, 1846) provide the

first comprehensive assessment of the morpho

dynamtivs and Holocene evolution of these seerions

of coasiline. The aii of this paper is Co assess We

evojution of the low wave energy beach ridge and

chenier sites on Kangaroo Island and Eyre

end their implications for mid to late Holovene sen-

fevel in the region, Beueh vidges are tere defined

its Shore linear, low sundysstelly swash deposits,

separated by wider iitet-and suprasGdal sane Mats.

They are a product of periodic wave acon and

showld nor be contused with aeolian foredune

ridves, Wineh have also been called beach ridges (see

Hesp 1954). Cheniers have similar ridees but are

vader lain aid separated by finer sediments such as

minds, silts or fine muddy srids,

Reach ridges occupy only 4°) (66-km) of Evre

Peninsula and (44.1 30-kmn of Kangacoe stand. This

is Tt sucprising given the generally high wave

enerey together with high percentage Of racky east

(66 and 4") respectively), On Fyre Penttisulie beach

ridges ate common in thy ventral gull and oeeur

it several protveted bays on the Bivhe coast, while

on Kangaroo Island they ure restricied do the

proiccted north eastern caves (see Short et al JORG

and Short & Fotheringham 1986 for locwtion of all

stiles). These sttes are, however, very importint for

the mierpretation of hor Hotovene shoreline

evolution and sea-level history. parnenierly gevern

M46 A, D. SIIORT, R. C. BUCKLEY & 1 1G, FOTAHERINGHAM

the noise-or ubsence (due io erosion) of the more

duminant higher energy beach and dune deposits,

Bach Burne (1982) and Belperio eral. (1983, 8k)

use such deposits (0 interpret Holocene sea-level

history, particularly invorthern Spencer Gull, while

Burne (1982) and Bowman & Harvey (1986)

identified episodes of ridge building in northern

Spencer and Sc Vincent pulls. respectively, This

paper provides a preliminary assessment of the

initiation and timing of ridge building at 2) sites

together with some implications for mid to late

Holocene sea-level in these regions,

Study Area

All beach cridge-chenier sites were identified by

mapping the entire coastal geomorpholoey using

1:10,000 calour aerial photography. Twenty

representative sites were selected for field

investigations (Pig. 1). They represent about 300

al both regions beach ridge systems. Table 1 lists

environmental parameters of each site,

Field investigations involved surveying ul cross

ridge tratscety using a theodolite, and bath surface

and shallow (<1 m) subsurface sampling, the latter

using both shovel and hand auger. All samples were

collected within 10-cny Stratigreplic unin usually

from in situ tidal flat deposits, All shells were

collected within cach layer, For dating, only the mosi

intact valve or bivalve shells were collected assuming

that the more intact and fresh the appearance, the

less distance they have been transported and

therefore (he younger they ruight be, thereby

providing a more accurate Mininium age of the

deposit. ‘Lonless otherwise indicated, dated shells

were obtained trom the base of the anger-shovel

hole The locdhion of each sample is Tllustrated in

the accompanying figures.

All sites are chacavterised hy low breaker wave

corditions, with essentially zero modal occan wave

effects, Wave energy is delivered bath by greatly

attenvated (<1 im) extreme icean waves and, more

lreyuently, by local wind waves (<b my. Tides

range fram micro ¢~) ni) onthe open coast to 2.5

ron the mid Spencer Gulf (Tig) th) Orientation

lathe daminant west to south-west winds and sea

breezes is Important only m relation co wind wave

generation, since the heach ridges sites hitve lite

or no weohan cappuie.

Resulls

Spenver Cinll

The five Spencer Ciall sates are listed in Table |

und iusiraled in big. 2. Tidal range liereses from

Linat lulka Wellro 2.3 mar Qensea, Glensea ind

‘Tulka Well receive periodic low wind waves, while

Point Gene. The Knob and particularly Lipson

Cove receive more regular southerly wind waves.

Gilensea’ Beach Ridges

The eight swash depayiled beach ridges (site |,

Fig. 1; Fig. 2) near Glensea Homestead are part of

an extensive beach ridge plain which runs (km

from just south ol Whyalla ta Lucky Bay. Lie

ridges. are fronted by extensive intertidal sandflats

foraling 100 km-. Many of the ridges, such as

those ar Cilensea, imerdigitaie with partially

submerged Quaternary longitudinal dunes, with the

dunes forming low ‘headkinds’ and ridges occupying

the intervening ‘embayments’, Dating of five of the

tidyes (Viv. 3a and Table 2) suggests ridge

development commenced after 2.6 ka BP, The

mixjure of ages in ridges 6, 7 and 8 suggests that

the ouler ridges may be contaminated by older

shells. If so, their age may be younger than

indicated. Daring this time their carbonate content

increased from 2 ta 30-60% indicating dependence

on biogenic production for sediment supply, The

surface slope ol the tidges and particularly the

swales (Fig..3a) could be attributed either toa fall

in sea-level of 1.0 m during this penod, or a

reduction in incident wave energy because of

sandtlat development. Examination of the

subsurface morphostrangraphy will be required to

assess the causes,

Point Germein Beuch — Foredune Ridges

Point Germein (site 2, Fig. |) is part of the 13

kr Tong Windmill beach-barricr that has prograded

notrbward, thus partially blocking Pranktin

Hurbour, and subsequently seaward, The ave of the

innermost beach ridge at ihe transect Iseation only

indicates the arrival of the northward prograding

burrier there at 3220 © 80 yr BP (Pig, 3b; Table

2). Subsequently, the burner tas prograded 250 m

eastward into the Guilt with increasing aeolian

cupping producing a 3 om high foredune at the

present shoreline, The elevations of the three inner

beach ridges are all within the limit ol present day

swish action and also suggest rapid shoreline

Progmadalion compared to the higher outer ridges

(4-8, Fig. 3b),

The Knob — Shingle Ridges

The 11 exposed shingle ridges at The Rirab (site

3, Fig, 1; Mig. 3c) record siurm wave deposition of

coarse Tertiary conglomerate material. Whilst no

dares were obtained, the seaward deerense in ridge

elevation could be attributed to a relating fall an sea-

level of « [m, However (he evidence is inconclusive

withour subsurface stratigraphy and dares: all mdee

BEACH RIDGE PROGRADATION 147

Head

of Bidht

Panone

10 Km — a eneng

C ay

Streaky Ba

streaky Bay

Buti Bay Whyalla

Vopus Bay

Port Pirie

Anvinds Bug 59 Tu0 kin a), )

Cowell Port Brouabton

a \ ' ; CG g Son

| s ) es

ok getiton

Pt Whidbey +e Feure

~ Peninsula

i histle

a é

Rigascore vote

o the Spit

NEPEAN

Rwy

Cape Borla

ESE) Tidai Flats :

KANGAROO

EA samphire SCANT Willoughby

36S

Cane Cape

Du Couedic Ganiheaume

Fig. 1. Location map showing portion of the South Australian coast and jocation of the 20 sites (1-20) discussed

in text. See Table | for names of sites. Lower insert is of Western Cove (sites 15-19) on Kangaroo Island.

crests are also within the reach of contemporary Lipson Cove — Shingle Ridge

storm swash, Following progradation the ridges

were buried by dunc transgression, and later Shingle ridges back many of the sandy beaches

exhumed as a deflation surface, suggesting that along the western Spencer Gulf, particularly in the

shingle nidye formation preceded the deposition of vicinity of the Tertiary conglomerates. At Lipson

barrier sands, corresponding to the finding of Burne Cove (site 4, Fig. 1) a single shingle ridge (Fig. 3d)

(1982) in northern Spencer Gull, has been emplaced landward of the present beach,

148 A.D. SHORT, RL, BUCKLEY & BG. POTITERIBGITAM

Table 1. Site Locgtrun* and charucteris ics

Lacvetion Distanve’ —

WEST SPENCER GULF:

1 ‘Olunsew’ {579 km 8 beach rides

2, Point Germein 1628

3. The Kiab 1633

4. Lipson Cove 172]

5. ‘lulka Well [856

GREAT AUSTRALLTAN BIGH'L

a. Colin Bay

6 Morgan's banding 20KY

7 Salt Matterhiole M14 (

8. Yangiv Bay 2433 tidal flats

9, Mount Dulron Bay aNTz 3 beach ridges

b. Western Eyre Peninsula

10, Perluhie Beach (5) 2544

LL. Acraman Creck 2553

12. Fyre Istiand 2635

13. Laura Bay 2643 6 beach ridges

fa. Tourville Bay 2766

KANGAROO ISLANL):

Ls, Brownlow 26 kin 5 beaeh ridaes

6. Brenyrlow (Zoll voursey 38

17. Cygnet River a

1k. Merreson Beach a2 tidal flat

19. Morrison Beach 4B

20, Pennington Bay 156

Morphology

7 beach foredune ridges

I! shingle ridges

I shingle ridge

low beach ridges

4 beach odves

7 faredune ridees

beach-loredune ridges

THulliple beach-mdses - spits

TAulLple beach-ridees - spits

J cheiens wid mangroves

NW 4

7 beach sidees N&

2 chemer-beach ridges N &

N &

back barvier depression

shure plailyrm

Marine Froerey

southerly wind wayes

southerly wind waves

wocthevly winel waves

beoas. swell and N. wind waves

rMiet; preyieusly low swell

ne waves

low S wind waves only

W wind waves

Wo & S wind waves occas, swell

occas. swell & Woawind waves

oveas. swell & S wind waves

occas, swell

& F wing waves

E wind waves

& E wind waves

& BE wind waves

N & E wind waves

oper swell

(Mainland distanee based on CSIRO Australian Coastal Inventory kas () km al SA’ Vie harder with km increasing

to wes 10) 3277 km at SAYWA border (Galloway er ah 19d).

Distance on Kaneroo Island based oa clockwise measurement from Narnth Cape (Short & Poiheringham (986).

‘The location of the ridge suggests deposinon prior

fo sandy beach formation, while its erest elevation

of 4 m above ISLW is within the reach of

contemporary storm waves,

Tulka Well — Beach Ridges

Tulka Well (site 5, Fig. 1) is a protected low wave

energy site receiving only wind waves generated

across Port Lineoli proper by nerth-cast winds. The

small, isolated beach ridge plain consists af several

low ridges which have prograded approximately 130

m inte the bay. Dates obtained from three holes dug

in Joe of the innermost ridge indicate thar rhis was

tidal flat by at least 3510 «© 80 yr BP (sample CS

53) Table 2, Pig. 3c), The dates (WK 837 and CS

S30) indicate upward ieeretion of 24.cm over 0.4

ka, a rate of 0.04 mm ye'. Following the upward

accretion of the tidal flats, some ime subscquent

to 2.5 ka BP, the low ridges prograded across rhe

flats 150 an into the bay. The inner samphire — dal

Hat tidal tlat hint could be interpreted ay tying up

to 40 em abave the present ridal Mat limi! suggesting

a slight fall in sea-level, While De Deekker ef al

(1982) also invoked a progressive fall in seaevel over

the Jast 3-6 ka to explain decreased salinity in

nearby Pillic Lake, the present data were

Ineemiclusive.

The Great Australian Bight

(Western Eyre Peninsula)

The Great Australian Bight (the Bight) is exposed

to some of the world’s bichest and most persistent

waves (Chellon ef al 1980). However shelf

attenuation (Provis & Steedman 1988), coast

orientation and the presence of extensive nearshore

calearenite substantially reduces wave energy at

many sds. even orn tie open coast (Short er al.

1984). As a result low energy beach ridges and

cheniers are found in a number of locations

particularly an the farger bays — Coffin, Venus,

Baird, Streaky, Smuky, Decres und Tourville (Fig.

1). Nine sites were investigated. They are located in

Coftin (4), Streaky (2), Smoky (1), Decres (1) and

Tourville (1) bays, (Table 1, Figs 1, 4).

BEACH RIDGE PROGRADATION 149

Coffin Bay

Morgan’s Landing.

Morgan’s Landing (site 6, Fig. 1) contains four

beach ridges backed by a back barrier depression

dominated by a Melaleuca swamp (Fig. 4a). In

addition, shell-rich beachrock is exposed on the

beach in front of the ridges (Table 3). Three dates

were obtained. The oldest (WK 829), from the back-

barrier swamp, dates at 3690 + 190 yr BP, with the

rear of the innermost ridge dating at 2540 + 60

yr BP (WK 827) and shell in the beach calcarenite

at 2680 + 80 yr BP (CS 527). This suggests that

progradation of the three ridges occurred around

3-2.5 ka BP and that since then the shoreline has

been relatively stable. This stability is interrupted

by west to east moving pulses of sand (sandwaves)

that alternately prograde (such as when beach

calearenite was deposited) and erode the beach

(such as at present) as they pass.

Salt Walterhole.

A 1 km wide series of at least seven low foredunce

ridges has prograded north-eastward of the Salt

Waterhole (site 7, Fig. 1; Fig. 4b). The ridges

apparently prograded at a time when a tidal channel

existed between the Waterhole and the present Point

Longnose. The eastward growth of Point Longnose

subsequently closed the channel and terminated

ridge progradation. Thyee dates were obtained from

the back barrier depression and the swash deposited

base of the first (oldest) and fourth ridge. The back-

barrier sample (WK 829) dated at 3440 + 60 yr BP,

the inner ridge (WK 830) at 3930 + 60 yr BP, and

the fourth ridge (CS 528) at 3190 + 80 yr BP. These

dates suggest a period of ridge progradation

beginning about 4.0 Ka and continuing to some

time beyond 3,2 ka BP, probably around 3.0 ka BP,

At this time it is likely the channel was closed by

the eastward growth of Point Longnose causing the

ridges to be cut off from periodic ocean wave attack

and sediment sources.

Yangie Bay.

Yangie Bay (site 8, Fig. 1, Fig. 4c) is a subsidiary

embayment in the southern part of Coffin Bay. It

is a Zero wave energy environment where Holocene

sedimentation is dominated by tidal and biogenic

activity. One date (CS 529) was obtained from tidal

flats surrounding an outcrop of Pleistocene dune

calcarenite and should indicate the commencement

of tidal flat progradation in this part of the bay,

It dated at 2780 + 80 yr BP (Fig. 4d). The other

date (WK 832) was taken at 60 cm depth beneath

samphire flats extending from the calcarenite and

dated at 2080 + 60 yr BP. Both dates suggest that

substantial tidal flat progradation did not

commence until around 3.0 ka BP. Stratigraphic

examination of the site (Fig. 4d) also suggests that

there has been little if any change in sea-level since

that date.

Fig. 2. The beach ridge plain near Glensea is located amongst longitudinal Pleistocene dunes (D). The transet (Fiz.

3) extends from A across the ridges to B on the inner intertidal sand flats.

150

cssi0 a Ve)

2560 280 cg525

2101-80

Site 2; Point Germein

ra lh

G5526

3220/80

Site

Auger

Holes

METERS (ISLW)

A, D, SHORT, R. C. BUCKLEY & D, G, FOTHERINGHAM

Site 1: ‘Glensea’ beach ridges

cssia

Tidal Flats

2120'80 ——S=

Site 3: The knob

Erasion

Scarp

5. Tulka Wells

a WKS37 2900760

b.CS631

3510+ 80

cWKS36 2420160

100 200

DISTANCE (meters)

100 200 300 400

Fig. 3, Spencer Gulf field sites, Chronological ridge numbers indicated above some ridges.

Mount Dutton Bay.

On the north-western side of Mount Dutton Bay

(site 9, Fig. 1) is a number of very low energy low

beach ridge sites, One of these was surveyed and

sectioned (Fig. 4e). The innermost sample (WK 833)

dated at >40,000 yr BP indicating Pleistocene

material. A sample obtained from tidal flat deposits

below the base of the innermost ridge yielded a date

of 6330 4 90 yr BP (CS 530), the oldest Holocene

date recorded in this study. This date confirms the

arrival of the Holocene sea-level stillstand by at least

6.3 ka BP, in agreement with its arrival in Spencer

Gulf (Belperio e/ a/. 1984) and Australia in general

(Hop!ey & Thom 1983). This date also is supported

by the age of the first swale which dated at 4590

+ 80 yr BP, the third oldest date in this study. The

elevation of the samphire — beach ridge, and tidal

flat — samphire contact between ridge | and 2 (Fig.

4e) suggests a slight (~50 cm) fall in relative sea-

level. The timing of this apparent fall however

cannot be determinded from the existing data.

Streaky Bay

Perlubie Beach (south)

Perlubie Beach (site 10, Fig. 1) is located in south-

east Streaky Bay and consists of a 1 km wide plain

of Holocene beach and foredune ridges (Fig. 5a),

A single date was obtained from the shell-rich swash

zone immediately behind the innermost Holocene

ridge (CS 499). It dated at 3290 + 90 yr BP, again

Suggesting a time gap between the stillstand and the

onset of low energy beach ridge progradation.

Acraman Creek

The Acraman Creek (site 11, Fig. 1) region in

northern Streaky Bay consists of a 10 km section

of shoreline dominated by multiple episodes of

recurved spit development and bayward beach ridge

progradation (Fig. Sb). The shoreline at its

maximum has prograded up to 3 km seaward. One

date was obtained from the inner portion of the

back barrier depression. Whole shell samples (CS

BEACH RIDGE PROGRADATION 1st

TABLE 2.. Radiocarbon Dates

Sample Lab. Location CSIRO Environment Material Elevation Age BP Corrected Age

No, No. km (m MSLW) Yrs BP = la Yrs BP i da

a. Eyre Peninsula

See

Spencer Guil

j-2 cS 50 Glensea 157¥ Back Swale shell 3 3020 = 280 zsali = 280

3-6 CS dt Gilensea 1579 Beach Ridge shell 23 3580 = 80 220 + 40

4-| CS 324 Glensea 1579 Beach Ridge shell 3 250 + BO 2070 4 80

4-4 CS 525 Glensca 1579 Beach Ridge shell | 2470 + 80 2010 = 80

4-5 WK 826 Glensea 1579 Swale shell 2 mos 30 B30 4 50

a7 CS $26 Point Germein 1628 Beach Ridge shell 3 Fasu - 80 JI 2 BO

4-50 CS 531 Tulka Well 18560 Tidal Flats shell | 3970) + RO 3510 4 a0

AY. WK 837 ~~ Tulka Well 1856 = Tidal Flats shell 16 M60 4 AO 20) ~ 60

4-48 WK 836 = Tulka Well le56 Tidal Flats shell 12 ZAR) + 60 2420 + 60

Coffin Way

4-25 WE R28 Gate Waterhole 2089) inter Barrier shell ~| 4150 + 190 4690 4 190

4-24 Wh 827) Morgans Landing 2089 Back Swamp shell = 3000 = 60 2540 + 40

4-23 Cs 427 Morguns Landing 2089 BReavh Calearenite — shell inter lidal 3140 4 Bt MR + 80

4-26 WK §29 0 Salt Warerhale- 214 Back Swamp shell ~1 3000 + Al) W404 60

4-27 WR #300 (Collin Bay) 214 0 Swast Deposit shell ~2 4390 + 60 3930 + €D

4-28 Ch 528 (Coltin Bay) 214 -Swash Deposir shell -~2 3650 + SU Win + BO

4-37 CS §29 Yargzie Island 2193 Inier Tidal shell 07 3240 4 80 2780 = 80)

4-38 WK 832 -Yangic Island 2133 Inter Tidal shell ~08 2540) + 60 TORO = AN)

4-34 WE 8330 Mt Dutiog Hay 272 Unter Tidal shell 17 40,000 > 40,000

4-40 CS 330 Mr Dutton Bay 2172 Inter Tidal shell 15 679 - 90 6330 + 90

4-41 WK 834 ML Dutton Bay 2172 Inter Tidal shell 13 05 + 80 459) 4 ORO

Gteat Austratian Bight

3-103 CS 499 Perlutic (south) 2544 Inet Ueach Ridps shell tt 3750 4 0 324) a

3-54 CS us Avantman Cresk 2543) Innermast Swale shell ? 610 4 9G S630 + 96)

re ee ail f : ; _ ayy + 3000 yy yoy + F000

3-88 CS 469 Point Collinson 2611 Shell Rock shell 1-45 42,400 arin 42,300” sn00

ne athe : » AF0G + 1700

—* ¢ By 5 , . =) < a dV 4 4

3-4) CS 469 Point Collinson 2611 Shell Rock shell 1-1.4 17,500 1400 ARAM) © i400

3-10) CS 509 Evie Istarid 2033 Inter Tidal shell ~1 T8604 80 140) b RD

3-71 CS 508 Eyre Island 2033 Inter Vidal shell ~1 3300 = 90 2kd0 4 YO

3-71 CS 506 Fyre Lslanel 2635) Inter Vidal shell ~1 3680 + 4 3220) + 9

3-73 CS 308 Fyre Island 2635 Inter Tidal shell -) 3770 = 160 VO + 160

4-24 WK SIS Laura Bay 2683 Beach Ridge shell -l 6l4@ - 90 S6k + 90

4-45 CS S04 Latira Bay 2683 Beach Ridec shell 2 740 4 90 2280 4 00

3-46 WR A24 Laura Bay 2683 Beach Ridge shell 23 2340 1 70 WW»

3-07 C§ 302 laura Bay 2683 Beach Ridge shetl 2 3520 4 91) 260 + 90

3-98 CS 50) Laura Bay 2683 Reach Ridge shell 3.5 2760 + 90 2 + 90

3-99 Wh s25 Laura Bay 2683) Beach Ridge shell 24 3480 = 60 3020 + a

3100 CS SQ) Laura Biv 2681 Swale shell zs 4 - WA, YO

3-3) Wh 816 fourville Bay 2739 Bench Ridge shell Z 2050 | il ZL) + A)

4-34 WR RIT Davenport Creek 277h Chenier Ridge shell =2 2470 + © 90) 201 = Yd

4-35 WK RIB Davenport Creek 277 ~=Chenier Base mangrove ~I 2860 4 SO 2860 4 50

3-36 WK 819 Davenport Creek 2776 Chenier Base mangrove ~1 2370 4 Sal 2571 + 50

3-07 WK AIO | Davenport Creek 2776 Chenier Base nud ~1 2540. 5h 2340 + 30

3-38 Wk 821 Davenpori Creek 2776 = Intertidal mangrove =~) 269) » 50 260). FD

b, Kangaroo Island

nn een ee

kl 13 WK 839) Mutrion Beach 42 Reach Ridge Suale — shell mn S40 — 70 ask = Th

KL Su Wh K42 Cyenet River a She!l Ridge shell 25-3 30. AU 2A + AD

KL SI Wk 84] Cyenet River dM Shell Ridge shell 254 2750 + 90 D2 4 MI

1 ~ unditates uearveved, exiiete Only

2 Colrecvon Paton ef 466 years (sed, sue Henerera a Hlarvee (1983),

152 A DOSHORT, B,C. BUCKLEY & BD. G TOTTERINGHANM

503) gave uw dale ol! $850 ~ 90 vr BP the second

vldest a! the Holocene dates, This alsa supports

fheampirom age ot the Holocene stillstand in ibis

ienion a >6 ke BP.

Fyre dstunil

Fyre Island (site 12. Fig, 1). whieh partially blocks

Smoky Biv isthe only unconsolidated inland inthe

Right. teconsists of wamoltiple series Of onlapping

recurved spits. beagh ridges and low foredunes

whieh surround a core of samphire = mioigrove

Hats (Pig. 4). Ao transect was made across. [he

westernmost and oldest of the existing ridges (Fig,

Se). Phe date obtamed trom the three swiles and

behind the present beach indicate that Chis scetion

of the ishind bexan building around 3.8 ka BPoand

progracded to around L4 ku BP. This sechion is

presently crogding as sedimerms are reworked

eastward alone both sides ol the land fo progride

Iherecupved spit complexes, As the island may have

been reworked uastward prot to tka BP the dates

can only be used as a pide (o (he ape of the oldest

section of the existing istand, and donot nevessarily

indigate the commencement of iskted formation

laura Bay

Laura Bay (site 13, Tie. Dis a 2 kin wide bay

within Deeres Bay, Locontains a 1.2 km wide beach

ridge plain frontal by several huadred metre wide

intertidal sand flats, The bay receives easentially no

Dcean wave energy us attested by the growth of

mangroves on irs northern sand(lats. A senies of

Seven dates obtained from two swales and the base

of five ridges (Fig. Sd) suggests that progradation

commenced alter 4.0) ka BP (CS 500). A period of

slay progradation (100 m between 4-3 ka BP, ridges

| and 2), was foliowed by rapid progradation (S00

m) between 3-2 ka BP (Fig, 5d), All ridges ure

within the reach of present storm swash elevations

und do nol suggest any apparent change in sca-level.

Like the Glensea ridges, the 2.3 ka age for the outer

active ridge (6) may indicate age contamination by

older shells, and hence i younger ape may be

appropriate,

loweville Bas

Taurville Buy (Site M4, Big. )) isthe westernmat

enclosed bay in 8. Ausi. and contains the

wesleromiost mayigreve woodland am South and

southern Australia. A tora) of five dates were

obtained in the bay, lour lor Dayenport-Creck andl

ove from the northern side of ihe bay (Fie. Se).

Davenport Creck consists of a 10 kim’ memgrove

woodlind described by Butler eral C977) within

whiel He rhree disvaminuoas cheniens (Eig, 7). The

second of the three cheniers was seetiondd and

provided j sequence ob Three dates (WOR 820, 818,

819, Fig, 30), These dates confirm the enrplavenient

of this Second ehemer at between 2.5 and 2.0) ka

BP, and the development ial the underlying Heliatl

flats by 2.5 ka BP. Mangrove stumps on the south

side ol! Davenport Creck, presently being Huried by

dune transgression datecl af 2690) 4) 30 (Wh R21),

conteniporanous with the chenier hase mangroves,

The date on the nerthetn shore of Tourville Bay was

obtained fram the intertidal hase of the imnermest

beachoaidge (Wik $16). Its dare of 2190 © OO Ve BP

coincides with the development of the secund

chenier. these dates are confirmed by the rysulls

oF Belpeno eat (988) who present the results af

13 vibrocores across Tourvillé Bay, They lound the

basal Posidome seagrass facies bevan accutane

about 6800 vr BP, with the embayment taracty

infilled by 5000 vr BP Bench ridge formation

followed the mantling of (he seagrass banks by

intertidal and mangrove factes. Phere uinerniost arid

single teach ridge date of 1840 yr BP is in

aurcement with our furdings.

Kangaroo Island

Mostof the Kangaroo Island coast is (Wypitied by

high bedrock aod calcarendte city exposed to tush

wave and wind conditions, However the north east

leeward pare of che island contains rhree

embayments — the Buy of Shoals, Western and

Eastern Coves which offer varying degrees of low

cuerty Holocene sforeline progeddation, Westert

Cove bas experienced che most sobstanital

progradation with sediment beinw delivered

Alonyshore from ityeasiern und western arm, as well

as from the Cvener River and (4 sity depasits, Four

aves were Caamined around the shoreline (rigs 1, &).

Brownlow

A series of live. low, shelly, swash-deposiled

bench ridges was surveyed at Browlaw (site 15, Lig,

1; fig. Ba). Their uniform elevation suggests no

aupparcnl chive to seadevel Na dithes aye been

oblained,

srownlow wolf course

Sever low shelly ridges were Survevedt immediately

south of the Brownlow golfoourse (silt 10. Fig. 1

and avain their relatively uniform elevation gives

no incication af sea-level change (Pig. Sb),

Cyenen River

Aseres of low discouttueus beach ridge and

possibly eheriers were surveyed | km south ol phe

Cyonet Riverimourh (sive (7, Fiz. (i. The two ridges

A, D. SHORT, R. C. BUCKLFY & 1, G. FOTHERINGHAM 154

Site 7: Salt Waterhole

Site 6:Margans Landing

Holocene Beach 7 \. Coffin Bay. |

Ridges ge hae dress wae?

.< . Coffin Bay rier

i Sand Flats

ATS

; Samsnire, Bieta

a.WK828 3690-190

0 tkm b.WK827 2540-60

c.CSS27 2680+ 80

d.WK829 3440 “60

&.WK830 3930-60

1.CS528 3190-80

Site 8: Yangie Island

(m)

ee Tidal \eo 8

ye Channehy 2

transect

g.CS529 2780:80

h.WKB32 2080°60

pe ia lly Sand

7 5 ae

DISTANCE (m)

| WK833 -40,000

|. CS530 6330-30

k WK 834 4550 ° BO

= jis Or L] o>

F ee, ae

[Calesrentte pe te ge

ELEVATION (ISLW)

20 40 ~ 60 80 100

DISTANGE (m)

Fig. 4. Coffin Bay field sites, Sites 6, 7 and 8 were not surveved. Sketches based on 1:10,000 aerial photographs.

154 A. D. SHORT, R. C, BUCKLEY & D, G. FOTHFRINGHAM

Site 10:

Perlubie Beach

Streaky

Bay

Pigfuce Is-

Holocene Ridges

a G3499 3290:'90

Site T2: Eyre Island

©. 68505 3310160

d.CS506 3220°90

8.68508 2840-90

E5509 }400180

200 400 *

DISTANCE (m)

Mangroves

6 pay

(

Southern

PiJaines

Ocean

ikm| Site Td: Acraman Creek

b CH50393 5650: 90

Mangroves

Se x) Acraman Ck

Beach Ridges “Su )/

PL. Lindsay

Streaky Bay

Site 13: Laura Bay

060-80

770'70

OSS00SN

Oe mutA 0

Shelly Sand

Calcarenite

4 L L

z00 400 1000 1200 1400

400 300

DISTANCE (m)

Site 14: Davenport Creek

Chenier Section

Organic

Shelly Sand

Manygrave Facies

nWKE16 2190 "60

owWK B17 2010'90

bp WKET8 28860.°50

qg-WK81S 2570 °50

1WKS820 2540'50

B.WRKB21 2690'50

Fig. & Western Eyre Peninsula field sites. Sites 10, {1 ard 14 not surveyed. Sketches based on 1:10,000 aerial photographs.

~ Site 12 survey elevations suspect, distances correct. Chronological heach ridve numbers given for sites 12 and 13.

yielded dates of 2650 . G0 and 2290) + 50 yr RP

(WK 840. WK 841). The ridges are now frotited by

300 m of densely vegetated samphire flats.(Fig, 8c).

Their distance from the present shorefine and the

elevation of the samphire flats could have resulted

Irom a slight fall in sea-level subsequent ta their

formation. However this cannot be confirmed from

the present data.

Morrison Beach

Two sites were examined at Morrison Beach (sites

Tk and 19, Fig. , Ina waterhole al the eastern end

of the heach (Site 18) the elevation of intertidal

shingle exposed in the waterhole is ~0.5 or higher

than the present beach shingle (Fig. 8d), and could

be attcibuted to a slighr fall in sea-level, However

the clevation of the inner shingle lies within the

reach of storm swash activity.

The beach-foredune ridge plain at Morrison

beach comsists of a 2-3 km wide plain that hag

prograded northward and westward into the bay

(Fig. 1; Mocris 1976). Shells abtained from a swale

hehind one of the innermost ridges (site 19) (WK

839) dated at 4480 + 70 yr BP, indivating (hat

BEACH RIDGE PROGRADATION 155

Fig. 6. A view looking eastward along the Smoky Bay side of Eyre Island. The subtidal sand-flats are backed by

a low energy reflective beach, and multiple recurved spits/beach ridges with samphire flats and mangroves

composing the core of the island.

Fig. 7. A view southeast across Tourville Bay shows the Davenport Creek tidal channel and sandflats backed by mangrove

woodlands and the discontinuous chenier ridges (C). Active dune transgression (D) in the south is burying the

mangroves.

156 A. D. SHORT, R. C. BUCKLEY & D. G. FOTHERINGHAM

TABLE 3. Beach ridge molluscs in approximate order of abundance

a, Intertidal mollusc species commonly found in Eyre Peninsula Holocene beach ridge and swale deposits. All these

species are associated with low energy mud/sand flat situations, generally with associated Zostera beds or silted rocks

etc... Coxiella sp. is usually associated with salt pans or salty lakes. Bembicium melanostoma and Salinator fragilis

are found on mud flats at the upper intertidal zone in sheltered estuarine areas. Anadara trapezia is usually associated

with Zostera beds; it is extinct in S. Aust.

b, Rocky shoreline species (also found in Eyre Pensinula beach ridges).

c, Kangaroo Island beach ridge molluscs, primarily in Western Cove region.

Katelysia peroni (Lamarack)

Katelysia scalarina (Lamarck)

Amesodesma erycinaea (Lamarck)

Amesodema cuneata (Lamarck)

Spisula trigonella (Lamarck)

Anadara trapezia (Deshayes)

Brachidontes erosus (Lamarck)

Batillariella estuarina (Tate)

Zeacumantus diemenensis (Quoy Gaimard)

Salinator fragilis (Lamarck)

Tellina (Macomona) deltoidalis (Lamarck)

Trichomya hirsuta (Lamarck)

Cominella eburnea (Reeve)

Austrococholea constricta (Lamarck)

Phasianotrochus rutilus (A. Adams)

Sanguinolaria biradiata (Wood)

Mactra Pura (Deshayes)

Fulvia tenuicostata (Lamarck)

Ostrea angasi (Sowerby)

Bembicium cf, auratum (Quoy & Gaimard)

Katelysia scalarina (Lamarck)

Katelysia peroni (Lamarck)

Katelysia rhytophora (Lamarck)

Zeacumanties diemenensis (Q & G)

Mesodesma cuneata (Lamarck)

Fulvia tenuicostata (Lamarck)

Austrocochlea concamerata (Wood)

Nassariu pauperata (Lamarck)

Pisulatrigonella (Lamarck)

Batillaria estuarina (Tate)

Thalotia conica (Gray)

Prothalotia sp.

Bembicium meeanostoma (Gmelin)

Coxiella sp.

Phasianella australia (Gmelin)

“Bulla botanica” Hedley

Diala sp.

initiation of this substantial and continuing

shoreline progradation commenced by 4.5 ka BP.

Shore Platform Evidence

Shore platforms cut into Pleistocene beach and

dune calcarenite are widely distributed around the

western Eyre Peninsula and Kangaroo Island open

coast. Whilst shore platforms are highly

questionable indicators of sea-level (Twidale e¢ al,

1977; Hopley & Thom 1983) they do warrant brief

attention in regard to possible changes in Holocene

sea-level in the Bight and Kangaroo Island.

Three platforms in the Bight were examined for

evidence of Pleistocene sea-level (Buckley et al.

1987). Two of these platforms contained a well

defined upper, inactive Pleistocene platform,

fronted by a wide active intertidal Holocene

platform (Merdayerrah and Clare Bay; Fig. 1).

Subsequent low altitude aerial reconnaissance of the

entire Eyre Peninsula and Bight by ADS and DGF

revealed the existence of relict elevated platforms,

presumably of Pleistocene age along much of the

calcarenite coast. Active horizontal shore platforms

occupy 440 km of the Eyre Peninsula coast (Short

et al. 1986) and are usually located intertidally close

to mean sea-level. All western Eyre Peninsula

platforms examined possessed a single active

Nerita atramentose (Reeve)

Brachydontes erosus (Lamarck)

Tellina (Macomona) deltoidalis (Lamarck)

Cominella lineolata (Lamarck)

Cominella eburnea (Reeve)

Austrocochlea concamerata (Wood)

Salinator sp.

Diala sp.

Bittium sp.

platform surface. On Kangaroo Island, however,

Short & Fotheringham (1986) describe a Pleistocene

platform at Pennington Bay (site 20, Fig. 1 fronted

by two Holocene platforms at different elevations

(Fig. 8e). The upper platform (No. 1, Fig. 8e) lies

50 cm above the lower platform (No. 2). The

difference in elevation and abrupt drop from

platform 1 to 2 may be a result of a slight fall in

sea-level (~ 50 cm) producing a lowering of the level

of saturation and hence platform erosion surface.

Discussion

These data have implications both for the

evolutionary history of the low energy beach ridge

sequences and for mid-late Holocene sea-level

history in the Bight and on Kangaroo Island.

Beach Ridge Progradation

The initiation of beach ridge progradation at nine

of the 12 sites dated did not commence until 4 ka

BP and in some areas as late as 2 ka BP (Fig. 9).

Most sites indicate an hiatus between the Holocene

sea-level stillstand and inner ridge building of up

to 4 ka. This is in general agreement with Burne

(1982) who found that in northern Spencer Gulf the

BEACH RIDGE PROGRADATION I

Site 15: Brownlow

(m) '

“FF Aeolian Sand

—

3 oe ;

E ate ete NN

PEN ee

2b Shotly Sand :

100 200

is)

site 17: Cygnet River

= 1 2 Fine Sand

7 \ Salt Fat Ae

= at

2 o3kh y

- 2

a 1

200

at} aite 18: Marrison Beach

1 eh

| | i

20 40 60

Lay

bb | Site 16: Brownlow (Golf Course)

i =

Shelly Sand * "34"

aWK840 2650: GO

b.WK841 2290' 50 i

Tidal

Crock

400

600 {m)

Pleistocene

Shore Platfarm

Platfarn 1

Galcarenite

Platform

10 20 30

DISTANCE (meters)

Fig, 8. Kangaroo Island field sites. Beach ridges are numbered chronologically,

development of low energy beach ridge pluins

uwaited sufficient subtidal carbonate production

and as a result progradation did not commence until

4-3 ka BP. Belperio e¢ a/, (1988) also concluded that

carbonate secreting organisms uggrade and

prograde the subtidal sand banks in association

with the subtidal seagrass facies. Once the banks

accumulate to low water level, subsequent intertidal

und beach tidge progradation can oceur. They

found this sequence al Tourville Bay on Nyre

Peninsula as well as several sites in Spencer and St

Vincent Gulfs,

In Gulf St Vincent, Bowman & Harvey (1986)

found that the extensive Le Fevre Peninsula beach

ridges have been prograding continuously over the

past 7000 years with pluses in ridge formation

apparently linked to longshore (northward)

sediment supply, This higher energy west facing

As A, DO SHORT, ROC. BUCKLE RY AD, G, POTEIERINGH AM

system Gnodal Hb = 0.5 mi) represents a transiion

belween the low ederey beach ridges discussed in

this paper and the high eneray¥ open coast regressive

depowts which experenced rapid progrudation

following the salistand. Their mode and sequetice

of esolution of the Peninsula is therefore oe

direcily applivuble to the lower cheney systems of

all which experienced a substantial tig in ridge

development,

On the (wo sites where a sequence of beach ridges

was dated (excluding Eyre Island) the soungest

Tides dated 2010 vy BP (Glensea> and 1770 yr BP

{laura Bay), suggesting that ridges are either nat

currently prograding or older material is being

deposited during ridge building. Whilst ridge

cessaliod 4 i aercentent With sedimentation models

ivhigher energy sites do south-east Australia (Thom

etal. )984), itis surprising in ad environment where

most sediments are derived front jt sida carbonale

produetion from fronting intertidal and subtidal

sand flais, the so cullud ‘cay bonate factary' {Burne

1982). In northern Spencer Gulf, Burne found

periods of shoreline (ic. sundflat) progradation,

aliernacng with beach-ridge progradation. The

most notable shorcline progradation was between

Jand 2 ka BP, On the Eyre Peninsula, however, that

was the period of maximum beach ridge

construction. A similar contrast applied te (he

penod from 2 ka BP to presents beach ridges in the

north-east Spencer Gulf prograded, while those in

ihe Eyre appear (o have rervained static. Evidently

nore information iy required before genctal models

of S Aust. beach ridge-sand flat progradaliqn can

be clarified,

Eyre Island liss a slightly younger outer ridge

«400 yr AP) and the ishind is vontinuing to develup

new recurved spits, However this may be accounted

for by erosion and dowadrift reworking of istand

sediments father that external sund supply.

Hulucene Sea-Level

Yhe exdet timing of the Holocene sea-level

Maxi eatanot he resolved from these dites

However the date of 6330 - 90vr BP from Little

Douglas as Well as the 3450 | 90 yr date trem

Acranmin Creek lend support to the general curve

adupted by BelperiG ef af, (1984) for Spencer Gull

aid | opley & Thorn (1483) for Australia as a whole,

Phat is it dppears thae the sea-level reachyeel its

present position by 63 ka BP in the Greal

Australian Right,

Warren, dK, (982) Nate Quarernare evaporite

cambodatecdepourional covianmenisad Saul Amaral,

PHD esis, Plaideds Gil. Adeluide (uopulht

Inevaluating evidence lor relative sea-level history

during the mid to late Holocene, tectome differences

between (he three main regions, the Bight, Spencer

Gulf and Kangaroo Island must be taken inro

aecoult. The most stable resion appears to be

western Lyre Peninsult (the Bight), Teetonic stabiliry

i§ Suggested by the widespread occurrence of the

+ 2-3 Pleistocene shore pluctorms (Buckley ef a/.

1986), the continuity of the presently. active shure

platforms. the single intertidal solurion notches in

limestone clifts in Coffin Bay and the evidence from

dated beach ridge and tidal sites (Figs 4, 5), In the

Bight the s¢a-level maximum was reached at least

by S650 yrs BP AFig. 5b) and has been stable since.

In northern Spencer Gulf there ip, However, evidence

for Holucene reeronism. Belperio ef af (1984)

proposed a sea-level cucve for vorthern Spencer

Gulf which has a srillstand being reached ar ~ 6.5

ka followed by an abrupe fall ta present sea-level

at 1.2=l.6ka BP. They auribure (his relative fall te

local tectanism, The present Speucer Cull dala are

not sufficient lo.add to the Belperia ey a/. curve und

may well be tectonically or isastatically distinct, The

Gilensea bec rides aod swales slupe seaward Fig,

Ja); however (his 1s not conclusive evidence for a

slight fall iv sea-level without subsurface

morphostratigraphy. I could also represent iv

decrease in waveencrey as the T Am wide intertidal

and subtidal sand Mats accunmulated, Likewise, while

other Spencer Gull’ sites logerher with Yanwe Isiand

and Mount Dutton Ray sites nay suggest a mudesi

fall in sea-level (<1 m), errors inherent in surveying,

facies identification and elevation, dating and

establishing reliable davums, place all of (he possible

changes within the sampling error bands, Conse

quently (here iy no conelusive evidence ura small

relative (Hl) insca-tevel, Wi any has oceprred it would

be only on the order ofa few eleciinelers. Together

with the Bight data this suggests a relatively stable

sea-level site the sea-level maaima ar or prior tu

6300 yr BP

Summary aed Canchisiuans

The following sequence oF events appeiins to have

Ooeeld red ulong Me lay eferey Bight, Evre Penisulit

and Kangaroo Ihind coyists sites Hie Holoeene sed

level srillstana,

7-6 ka BP. The stillsiand was achieved by at leasi

6330 + 9041s, BP and most likely by 6,6-6,4 ka

AP based on evidence from northern Spencer Gulf

(Belperiés ef af L984),

6—1 ka BP Pollawing the stillsiand mast law energy

secliuns of caast experienecd minimal shoreline

proanmidition, Exceplons were areas where shingle

Was immediaigly available, particularly ib Spencer

159

BEACH RIDGE PROGRADATION

“puels] OOIesULY puke e[NsutuAag aAq Uo Sars aEpi4 Yea AB19U9 MO] IOJ WOTIBADTS SNSI9A Sa]Bp SUSIO[OH JO JO} 6 “A

dd SUVSA <OT x

PW ‘W }daoxa

TSW 11V WNLYd ON

yisodap jJsowuauu]

pnw acoibuey

jyoou aaosbuew

(doy) jepyszsjzu|

aarydue

jatae ys W 0

abpia yseeg

yDo4 yoeeg

‘O)

M1SW

LIWIT 39GIY HOVAE LNASAYd

NOILVOVYOOdd “XVW h

1 At A.D SIPORT. RO. BUCKLEY & DG. FOTHERINGHAM

Gull, by these areas storm shingle mdges were

developed following the stillsiamd. Whilst na dates

are available from the present data, dating of similar

ridges in northern Spenger Gulf by Burne (1982)

found proseadation conumenced around 6 ka BP

Onty three beach ridge sites (Mow Dutton Bay,

Acraman Creek and Morrison Beach) have back

barrier deposits older than 4 ka BP No beach ridges

were dated prior ta dha BP. However prior fo beach

ridve Tormation the substantial inter to sub-ndal

sand flac would have to be building during this

period,

4-2 ka BP. This was uppareucly a period of major

beach fdwe and dimited chenier development in

Spencer Guill, the Bight and on Kangaroo Istand,

The commencement of ridge construction has been

cared as early as 3930 | GO vr (WR 830) in Collin

Bay (hig. 4b) and as tate as 2190. 60 (WR RIA)

in Tourville Buy (Tig. $e) The most logical

caphinationool this sree in ridge formation ts the

development of The miler to sub-tidal carbonate

platforms and sedimenrs required to both support

wich supply sediment for ridge development possibly

coupled with a slight tall in sea-level.

24) ka BP. big. Y is nor reymescntanive of low energy

deposits as many of the samples were selected to

date the oldest ridge/swale, All that can be said is

that au rhetwo sequences ditled, the wounest ridge

dared ar 2010 and 17270 BP (Figs 3a, 3), However

there is ubundwnt evidence ol contemporary cidpe

formation, partivalirly in areas of recurved spits

(ve. Lyre Island, Acraman Creek, St Peter Island).

Possible eyplanarions forthe outer ridge ty Chat in

vreus Of shore normal sechiment supply there is a

present hiittus toridge Jormation, or pusstbly uldee

shells are bene iicerporated in outer ride deposits

lo fiveanimoemolans ave. Whalever Uhe catuse (hese

gysrems require further investigation In conclusion:

1 Low enerey bench widves and seme cheniers

occupy 6S knw 40) of che western Spencer Gull

wnd Bight coast, particularly inthe larger more

shellered biws. They alse occupy T4 kor of coast

ju the horth-easr of Kangarao Island, Most

sequenues Gomstst ol 2-1) ridees (mean 4.5 in

Spencer Gull)

2. Phe videes are composed of precdonunaimly

carbouwle shell detritus Clable 4) apparently

sourced und reworked trom the adljacent iter-

and sub-tidal sundtlats and adjacent mek

shores.

3 Ridge formation awaited the eonsuuebon vl iii

underlying inters and sub-tidal sandr fhe

sundflars were required both to predic a

platform lor ndge buildirus, as well sa beac

supply of macerial The inereasiag prayrite ae

of carbonate sedunents is reflected in ie

oulfward increase in carbonate vantemt of the

Cilensea ridges (Fig. 31).

Once the intertidal sandflat platforms were

vonstructed, ridge formation otlen was rapid) dn

places several ridges separated by waler sales

were deposited m several Hundred year. usualy

between 4—2 ha BP, However age contaniiaiion

of outer ridges by older strells may piadsk it

younger age and hence longer period ut (idee

building.

5. At present most active ridge formarion ts

vecurring Lb aeas of longshore supply, resulta

inthe construction of muluple reebrsed spits

Shore nernial suppiy may be negligible resling

IN un apparent Hidtusy in ridge Gomstrin Hon wt

conversely older material is being inci posited

to provide anomalous ages.

6. The sea-level stillstagd occurred prion 1 0.9 hat

BP and has been relatively stable siney

particularly in the Bight. There is inconclusive

evidence of a slight {<1 m) fallin sea-level in

Spearcer Giulf and a (0.5 m) fall on Kangaroo

Island in the late Holoeene.

Agkituw bedginents

AL Lield work on Kangaroo Islaad ane |be byte

Peninsula was supported by the Coastal Protec tony

Board through the Coasial Management Binh,

South Avstialiaty Department oF Prvironinicns &

Planning. The bayre Pemisula radiocerbort lining

was provided by the S.A, Dept ob Mines & bavi.

Molluscs were indentified by Phil (obeuan,

Ausrralian Muscam. Piguies were didied by

Jeanette Neeson and the manuscript typed) ty

Janetre Brennan iid Kay hoster: Mauy thanks to

all.

References

Bre iden ASTI Greeais Vi ACs, DP Hod Mi tiitay

Woaotacty $V. 40988) Awellingil-orgaitiss coma tien

dod the evotiiian of Holocene tidal sequences a

snuilieny) Austria pp itS<4972 fa, Pd de Boer et

wi (hale) Pile Latlueneed Scdimantary Lar rennenns,

und haynes CD Beadet Paik Cap.

i, Hts KR Sling WO. CEYBE A revi

WT Pilbeune nwa desels Sip (tr Ata eab ay de di

V. Hopley. (Rely Austin sea lesets i rhe fist 15,000

years. aoreview. Geeas. Pap. 3, Monugear ser Dep

Generapliy, Hanes Cook Univ, Tawrgville,

— po & Poracth He AL aed) The

aan apby of coastal carbotiale bake aod Holocene

sed levels Of nerrhern Spenver Gull, Some Averabe

Marine Geal 6), W7As

Hoary, GO. & Lager (PO83) Baeeabon danny

BEACH RIDGE PROGRADATION j6t

of macine shells in Sour}, Australia, Aust. Archaeology

17, 13-123,

«1986 fieemorphic evolution of a

Holovune beach-ridge .niolex, Le Fevre Peninsula,

South Ausirahia. 4. Cry ssud Res, 2, 345-362.

Buckies, RoC. Srogl, 4 DX ForHERINGHAM, D. G,

(1987) tan Pleistocene se level maxima.in the Great

Australian Bivht Sear 4: Yk, 86-89,

Buknr, RY, (1982) Reliti. > (ll of Holocene sea fevel

and enastal progradati vi, northeastern Spencer Gull,

South Nustcalia. BMH fr Aust, Geol. Geophys, 7,

35-45

Burterk, A... Dreers, A. ML, McKitiup, S.C, &

THowss, DP. (1977) Distribution and sediments of

mangriowy elds in Soulli Australia. Tras. R, Soe. S.

lust, WOH, 38-44,

CHtiTON, Dh, Hussey, K. J & Parke, M. E. (1981)

Global satelliile measursiicnis of water vapour, wind

speed vil wave height, Satire 294, 529-532.

Dany, B. Mirsss, ALR, Twtbare, C. RB, & BOURNE,

JA. (1979) Geology aad geamorphology pp 1-38. /7

M, J. Tyler C. R. Twidile, & dK, Ling, (eds.) “Natural

History of Kiingaroo Island" (R. Soc. S. Aust,

Adelaide)

DE DECKKIR, P., BOLLD, J. & BURNE, R. V. (1982) Pillie

Lake, Eyre Peninsula, South Australia: Modern

environment and biota, dolomite sedimentation, and

Holocene history. Trani. R. Soc. 5. Aust, 106, 169-181.

Galioway, R, W., Story. R.. Cooper, R. & YApp, G. A.

(1984) Coastal lands ot Ausiralia, CSIRO Diy. water

and Land Resources. *

‘Natural Resources Series 1.

Haris, J. R. & Gostin, VN. (Eds.) 1984 The Spencer

Gulf region, Marine Geol, 6L, 111-424,

Hesp, P, A. 1984 The fo qutton of sand ‘beach ridges’

and foredunes. Search 13. 289-291.

Hopcey, D. & THom, B. G. (1983) Australia sea levels in

the last 18,000 years: a review. Jn Hopley, D. (Ed,)

Australian sea levels in the last 15,000 years: a review

pp 3-36, Qcvas. Pap. 3, Monog. Ser. Dept Geography.

James Cook Uniy,, Townsville.

Morris, Bo 4 (1976) Investigation of heavs imineral

sands, Morrisua Beach, Kangaroo Island, Adineri/

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SHorRi, A. 2. +i98Sa) Response of a high iraugh low

energy Sandy coast to sea level transgression and

stillstand, South Australia, Geogruphical Review 78,

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— (98KS! Modes of Holocene coastal dine

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- FGI edeam, DL G, (1986) Coastal mMmorpho-

dynamics and Holocene evoludion of the Kangaroo

Island. voasi, South Australia, Coastal Studies Unit Tech,

Rept, 86/1, +s uastal Studies Unit Univ. Sydiey, Sydney).

- > ——_. » & Brentey, R. C_ (1986) Caasial

marphodynamics and Holocene evolution of the Lyre

Peninsula coust, South Australia, Coastal Studies Unit

‘Tech. Rept, $62, (Coastal Studies Unit, Univ. Syditey,

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& Hew, PA (1984) Beach and dune morpho

dynamics o! the South East coast of South Australia,

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THom, BL G.. Rowman, G. M,, GILLispre. Ra, Pesan,

R, & Baker iii, M. (1981) Radiocarbon dating of

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Monog. LH, fue Geography, R.M.C., Duntroon, ACT.

TWiDALE, C. R., Bourne. AL & Twroatn, N. (1977)

Shore platfort witd sea-level changes in the vulls region

of South Awenha Trans, R, Sac. 8. Aust. 1, 63-74.

THE OCCURRENCE OF A DIPROTODONTID MARSUPIAL REFERABLE

TO EUOWENIA GRATA DE VIS IN WESTERN NEW SOUTH WALES

BY N. §. PLEDGE*

Summary

A complete last upper molar (M?) and fragments of several other teeth found in a bore on the

Talyawalka Anabranch, east of Menindee, N.S.W., are identified as the Pliocene species Euowenia

grata De Vis. Euowenia grata is redescribed and refigured, and the new M? is compared with the

equivalent teeth of Diprotodon optatum, Zygomaturuwss trilobus, Meniscolophus mawsoni,

Nototherium mitchelli and Palorchestes azael. The Talyawalka specimen extends the range of

Euowenia grata by almost 1000 km and suggests a Pliocene age for the deposit.

KEY WORDS: diprotodontid marsupial, Euowenia, Diprotodon, Zygomaturus, Meniscolophus.

Nototherium, Palorchestes, Talyawalka, Menindee, New South Wales, Pliocene.

Transactions of the Royal Society of S. Aust., (1989), 113(3) 163-168.

THE OCCURRENCE OF A DIPROTODONTID MARSUPIAL REFERABLE TO

EUOWENIA GRATA DE VIS IN WESTERN NEW SOUTH WALES

by N. S. PLEDGE*

Summary

PLebGe, N. S. (1989) The occurrence of a diprotodontid marsupial referable to Euowenia grata De Vis in

western New South Wales. Trans. R. Soc. S.Aust. 113(3), 163-168, 30 November, 1989.

A complete last upper molar (M°) and fragments of several other teeth found in a bore on the Talyawalka

Anabranch, east of Menindee, N.SW., are identified as the Pliocene species Euowenia grata De Vis. Euowenia

grata is redescribed and refigured, and the new M’ is compared with the equivalent teeth of Diprotodon

optatum, Zygomaturuwss trilobus, Meniscolophus mawsoni, Nototherium mitchelli and Palorchestes azael.

The Talyawalka specimen extends the range of Euowenia grata by almost 1000 km and suggests a Pliocene

age for the deposit.

Key Worps: diprotodontid marsupial, Euowenia, Diprotodon, Zvgomaturus, Meniscolophus, Nototherium,

Palorchestes, Yalyawalka, Menindee, New South Wales, Pliocene.

Introduction

De Vis established the species Owenia grata (De

Vis 1887), for a skull and jaws (Fig. 3, QM F519)

found at Chinchilla, Queensland. Later he renamed

it Euowenia (De Vis 1891) after finding the earlier

name to be preoccupied by an annelid worm. At

the same time, he named a second species,

Euowenia robusta, for a set of dentaries found at

Freestone Creek, near Warwick, Qld, in Pleistocene

alluvium, Woods (1968) suggested that Euowenia

robusta is a junior synonym of Nototherium inerme

which could effectively restrict Euowenia to

specimens from the Chinchilla Sand, and to a

Pliocene age (Woods 1960; Bartholomai & Woods

1976).

Euowenia grata occurs with certainty only at

Chinchilla (Stirton ef a/, 1968). M. Archer has

informed me (pers. comm. 3.xi.1975): “In many

cases, specimens should more honestly be referred

to merely as undetermined small nototherines”, and

there is no evidence at present for any Pleistocene

occurrence. Recently, Ewowenia has been listed

(Tedford, Williams & Wells 1986) in the Plio-Pleisto-

cene Kanunka local fauna at Lake Kanunka, South

Australia, but this record needs confirmation.

The purpose of this note is to record a new

occurrence of the species well beyond its previously

known locality.

Material and Methods

In 1969, Mr D. Walker of Talyawalka Station east

of Menindee, New South Wales, presented to the

South Australian Museum some unusual tooth

fragments found in the sludge from a newly-drilled

water bore in “White Water Lake” (32°25’S,

* South Australian Museum, North Terrace, Adelaide, S.

Aust. 5000.

| Broken Hill 3

| a al 2

| <

| ne

i L.Menindee

| 2

| L-Tandou

| Peet

NEW SOUTH

SOUTH WALES

AUSTRALIA |

{

0 Km 100

Fig. 1. Locality map. White Water Lake, Talyawalka

Anabranch, New South Wales.

143°18’E) on the Talyawalka Anabranch of the

Darling River (Fig. 1),

The new specimen is very fragmentary,

comprising the complete left M’, two enamel

fragments with differing degrees of wear, believed

to be anterobuccal corners of the protolophs of M'

and M' of the same tooth row, the anterobuccal

corner of the metaloph of M', the anterior half of

M,, an unidentifiable molar fragment and a

fragment of incisor. Because of the poor quality of

De Vis’ figure (De Vis 1887), the new specimen was

for several years considered by the author to be a

possible very large new species of Palorchestes,

because it had a fairly distinct midlink between the

lophs. However, direct comparison with a cast of

the upper dentition of Evowenia grata (holotype

QM F519) by courtesy of Dr M. Archer, has now

proven its identity. The specimen also has been

THE OCCURRENCE OF A DIPROTODONTID MARSUPIAL

compared with teeth or casts ot Dipratodon

optatum, Zveormaturus trilabus, Meniscalophus

mawsoni, Nototherium mitchell and Palorchestes

azuel,

Material is from the South Australian Museum,

Adelaide (SAM), Queensland Museum, Brisbane

(OM) und British Musetum (Natural History),

London (BMNH).

The status of the genus Evoweni¢ is thought to

be in doubt and in need of mayor revision (Archer

& Wade M976), but wai this done itis felt that

Aadcdesery ton of fhe species is desirable, based

partly on this new material. Tooth notation follows

Archer (1978),

SYSTEMATICS

Euowenia grata De Vis

FIGS 2-4,

Owenia grata De Vis, 1887, p. 100, pl. I-IV

Upper premolar two-rooted, about half the length

of M* and triangular (De Vis 1887, 1888).

Protocone slightly smaller than parametacone.

Protocone and anterior part of parametacone

joined by narrow transverse loph to form short side

of a right triangle. Buccal side formed by buccal

crest from parametacone to anterior end of the

165

tooth, Third side formed by anterolingual cingulum.

Small deep central basin with rugose enamel,

Upper molars large (Tables 1, 2) and bilophodont,

with lophs convex forwards and somewhat tumid

with incipient midlink development — postproto-

crista posteromedially from protocone and pre

metacrista(?) extending anteromedially from meta-

cone. Midlink development virtually absent in M¢

but increases slowly to its greatest extent in Mé

where pre- and post-cristae are contluent fora few

millimetres vertically, M°~“ subrectangulary MM?

trapezoidal, with metaloph abour three-quarters

width of protoloph. In M! and M>, metaloph

forms distinct dog-leg bend where midlink joins it.

Anterior cingulum extends full width of tooth, but

is longer at its lingual end. Posterior cingulum

narrow, particularly on M’ where small pocket

may be formed by weak postmetacrista, Only trace

of a lingual cingulum may be seen, at bottom of

transverse valley. Buccal cingulum better developed,

across mouth of transverse valley.

Lower molars more rectangularly elongate than

uppers. Both protolophid and hypolophid are

oblique and concave anteriorly. Pratoconid end of

protolophid enlarged by development of preproto-

cristid extending anteromedially. Similarly, hypo-

conid end of hypolophid is enlarged by cristid

obliqua which forms posterior half of weak mid-

TABLE 1. Cheek Teeth Dimensions, Euowenia grata

Except for (he premolar, measurements are made in planes parallel io and normal to the lingual edge of the tooth.

SAM P14506 QM F519 (epoxy cast) SAM P14506

LM. LP* LM? LM" LM! LM’ LM"

A-p Length ~ 15.0 26.5 32.6 36.2 35.7 34.8

Ant, Diam. 19.3 — 20) 26.2 30.3 28.7 27.7

Post. Diam. — 15.7 24.2 25.9 28.4 23.2 23.1

TABLE 2. Euowenia grata Dimensions as given hy De Vis (1887)

These are not directly comparable with those to Table 1, as M”

for example is apparently measured along the outer

edge.

PL M. Pp! M’

Length 14,5 34.5 1540 38.0

Max. Breadth 85 22.0 14,5 29.5

Vip. 2. Euowenia grata De Vis. Lett ipper molar M’ of SAM P14506. Stereopair photographs: a) Occlusal; b) buccal;

¢) (itigual view, Seale = | em.

166 N. S, PLEDGE

FIG. 3. Euowenia grata De Vis. Holotype palate and dentaries, QM F519. Scale bar = 10 cm.

link. In the holotype, lowest part of midlink is intact

in last molar (De Vis 1887). Anterior cingulum

almost full width of tooth. Postcingulum divided

unequally into two parts at its highest point by fine

submedian posthypocristid.

Comparison with other species

The Talyawalka specimen may be compared with

this (Table 1) and other diprotodontoids.

The upper molar Mis the same size as that of

the holotype of Euowenia grata, and same order

of size as Meniscolophus mawsoni. It is smaller

than Zygomaturus trilobus and slightly more than

half the length of Diprotodon optatum.

The Talyawalka tooth differs from Diprotodon

(SAM P23406, Lake Callabonna) in: its much

smaller size, relatively higher crown, more convex

protoloph, slightly narrower precingulum with

greater development at the lingual end, a buccal

cingulum across the end of the transverse valley, a

narrow but strongly developed postcingulum, the

absence of cementum, and particularly the presence

of a Y-shaped hypoloph formed by a distinct

premetacrista.

It differs from Zygomaturus (SAM P29889,

Naracoorte) in: its smaller size (about three-quarters

the length), shorter precingulum, absolutely higher

lophs, thicker lophs, presence of a distinct buccal

cingulum across the transverse valley, deeper and

more acute transverse valley, distinct premetacrista,

more distinct postmetaloph-pocket, and presence

of rugose/punctate enamel.

It differs from Nototherium mitchelli (cast of

BMNH M5002) in: its smaller size, its longer and

better developed precingulum, presence of a buccal

cingulum across the transverse valley, relatively

higher lophs, presence of a premetacrista, and better

developed postcingulum.

It differs from Meniscolophus (SAM P13647, of

which M? is unknown) in slightly smaller size,

higher lophs, presence of distinct premetacrista, and

narrow postcingulum.

THE OCCURKENCE OF A DIPROTODONTID MARSUPIAL. th’

Pi. 4, Euowenla grata De Vis, Cast of left upper maxillary tooth row of holorype, OM FSi9

Ni differs trom rhe holotype of Auowenia grata

(cast of QM F519) im having a distinct buceal

cingulum with two small (one twice as big as the

other) cusps across the transverse valley, and a more

distinel posimelaloph-pocket.

I differs from Palorchestes azael (QM F773, fide

De Vis 1895) in: its larger size, relatively greater

length, lack of a forelink, presence of a strong

buccal cingulum across the transverse valley, strong

postcingulum, and absence of a complete midlink.

The new specimens (SAM P14506) agree closely

with the holotype, considering the difference in

degree of wear and preservation, In M1’, the

enamel is just breached so that a good indication

of the height and torm of the tooth ean be gained,

The lophs are high (eqnal to the width of the tooth)

aid stout so that the transverse valley is particularly

deep and narrow, a harrowness accentuated by the

bend caused by (he premetacrista, ‘Together, the

metaloph and premetacrista have a distinct Y-form,

Centra De Vis (1887), 1 found no indication of a

forelink on the upper molars of the holotype cast,

bul there is a slight protuberance i (his position

on the new specimen of M®°.

The molar cnamel is notably rugose and

punctate, similar to that of Diprotodon and

Euryzyzoma (Archer 1977). In the worn M,, frag-

Ment, the anterior cingulum is heavily abraded by

interdental appression and has a small cusp de-

veloped at the anterolingual corner; The “midlink"

is discontinuous and does not cross the transverse

valley as an integral structure:

The incisor is too fragmentary to describe, save

that it would be roughly circular in section with a

diameter of at least 22 mm, Only a portion of the

rool is preserved and, from De Vis’ deseription

(1887), it could be either upper or lower II,

Discussion

The teeth came trom a depth of about 28 m, in

a 1.5 m thick bed of coarse, yellow-brown dirly

quartz sand, below sandy clay, The age is uncertain,

but an early Pleistocene, or even Pliocene, age 1s

possible, based on the absence of the species rom

lale Pleistocene surficial deposits nearby — hinette

sands dated at 25,300 years BP at Lake Menindee

(Tedford 1967) and 32,000 years old at Lake Mungo

(Barbetti & Allen 1972), Although speculative, this

determination is supported by the Pliocene age of

the holotype (Woods 1960). The specimen extends

the known range of the taxon by almost 1000 km,

It seems that the bore drill struck a more-or-less

complete skull and jaws of Euowenia because all

tragments appear to relate to a single individual,

there being no duplication of elements.

Conclusions

These fossil teeth, recovered from a water bore

on the Talyawalka Anabranch, eas! of Menindee,

N.SW., match those of Evowenia grata (De Vis)

and, because (hey are Jess worn than the holotype,

add fo (he knowledye of the species. The occurrence

extends the known range of the species.

Acknowledgments

| thank Mr D, Walker for donating the specimen

to the South Australian Museum, Mrs Debbie

Brunker and Mrs Debbie Lowery for typing the

manuscript and Mrs Linda Blesing for drafting the

map.

References

ARCHER, M. (1977) Origing and sublanilial relationships

of Diprotodon (Diprovodontidae, Marsuplatia), Mem.

Qld. Mus, V8, 37-39.

—_—— (1978) The nature of the molar-premolar boundary

mi marsupials anda reinterpretation of the homology

of marsupial checkteeth, Jbid 18, 187-164.

& Wabe, M. (1976) Results of the Ray EB. Lemley

Expeditions, Part 1, The Allingham Formation and a

new Pliocene vertebrate fauna from northern

Queensland. /hid 17, 379-397.

Baruetii, M. & ALLEN, H, (1972) Prehistoric mar at

168 N. S. PLEDGE

Lake Mungo. Australia, by 32,000 years BP. Nature 240,

46-48.

BARTHOLOMAIL, A. & Woops, J. T. (1976) Notes on the

vertebrate fauna of the Chinchilla Sand. Appendix 2

in Bur. Min. Resour. Bull. 166, 151-152.

De Vis, C. W. (1887) On an extinct Mammal of genus

apparently new. Proc. R. Soc. Qld. 4, 99-106, pl. 1-4.

(1888) Notes on the genera Zygomaturus and

Nototherium. Ibid 5, 111-116, plate.

_ (1891) In confirmation of the genus Owenta so-

called. Proc. Linn, Soc, N.S.W. (Ser. 2) 6, 159-165, pi

13.

(1895) A review of the

macropodidae in the Queensiant Mossum

75-134, pls. 14-18.

STIRTON, R, A., TEDFORD, R. H. & WoopBuRNE, M. O.

(1968) Australian Tertiary deposits containing terrestrial

fossil vs af the

lind 1,

mammals. Univ. Calif. Publs Geol. Sci. 77, 30 pp.

TeDFoRD, R. H. (1967) The fossil Macropodidae from

Lake Menindee, New South Wales. Univ. Calif. Publs

Geol. Sci. 64, 156 pp, 4 pls

» WILLIAMS, D. & WELLS, R. (1986) Late

Cainozoic sediments and fossil vertebrates pp 42-72.

In Wells, R. T. & Callen, R. A. (Eds) “The Lake Eyre

Basin — Cainozoic sediments, fossil vertebrates and

plants, landforms, silcretes and climatic implications”.

Australian Sedimentalogists Group Field Guide Series

No. 4+. (Geol. Soc Aust... Sydney)

Woops, $ T. (1960) Posstliterous thiviatle and cave

d (Eds) The

yaits. Un Hill, O A& Denmead, ALK.

wy of Queensiand. £ Geol Soe. Aust. 7, 393-403.

— (1968) The idenuty of ihe caisnci marsupial genus

Nototherium Owen. Mem. Qld. Mus. 15, 111-116, pl.

13-14.

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 113, PART 4

PARASITOIDS OF URABA LUGENS WALKER (LEPIDOPTERA:

NOCTUIDAE) IN SOUTH AUSTRALIA, WITH DESCRIPTION OF TWO

NEW SPECIES OF BRACONIDAE

BY A. D, AUSTIN & G. R. ALLEN*

Summary

Information is presented on the large complex of hymenopteran and dipteran parasitoids associated

with Uraba lugens Walker (the gumleaf skeletonizer) in South Australia. A key to the 22 species

involved is presented, along with notes on identification and relationships with their host. Two

species of microgastrine braconids are described, Cotesia urabae sp. nov. and Dolichogenidea

eucalypti sp. nov.; both are parasitoids of the larval stages of U. lugens.

KEY WORDS: Uraba_ lugens, Noctuidae, parasitoids, hyperparasitoids, Braconidae,

Ichneumonidae, Aphelinidae, Chalcididae, Elasmidae, Eulophidae, Eurytomidae, Eupelmidae,

Trichogrammatidae, Tachinidae.

Transwelions of the Koval Sociey of & Aust, (1989), LIM4) 169% 184.

PARASITOIDS OF URABA LUGENS WALKER (LEPIDOPTERA; NOCTUIDAE)

IN SOUTH AUSTRALIA, WITH DESCRIPTION OF TWO NEW

SPECIES OF BRACONIDAE

by A. D. AUSTIN & G. R. ALLEN*

Summary

Ausrin, A. D. & ALLEN, G R. (1989) Parasitoids of Urabe (ugens Walker (Lepidoptera: Noctuidae), with

descripuian of rwo new species of Braconidac. Trons, R. Sov. S, Aust. 113(4), 169-184, 1D Noveniber, 1989.

Information is presented on the large complex of hymenopteran and dipteran parasitoids assucialed with

Uraha luzens Walker (the gumleaf skeletonizer) in South Australia. A key to the 22 species involved is

presented, along wilh nates an identificanon and relationships with their host, Tao species of micengasirine

bravonids are described, Caresio nndlive se mo, and Nolichogenidey eucelypli sp, now.; both are parasitoids

of the larval stages of Of lugens.

Key Worps Graha dueens, Noctuidas,

parasitoids, Hyperparasitoids, Bracomdar,

Ichneumonidae,

Aphelinidae, Chalcididae, Elasmidae, Eulophidae, Kuryiomidae, Eupelmnidae, Trichogrammatidac, Tachiniduc.

fntroductiva

Uraba fugeas Walker, he gumleal skeletonizer,

is 3 native noctuld moth and has been. collected

from all Scares of Australia (Turner 1944), It has

been recorded as damaging stands of cucalypt

species in eastern Australia (Brimblecombe L962;

Campbell 1962; Harris 1972!, 1974; Harris er ai.

1977°y, the Adelaide region and south-western

Western Austialia (pers, comm. F. D. Morgan;

Strelein 1988). Occasionally outbreaks of this

species can defoliate large arcas‘of native forests.

Several such ourbreaks have occurred in stands of

Eucalpytus camaldulensis Dehnh. along the Murray

Valley region of N.SW. and Victoria, where, on at

least four Occasions, More than 30,X) fia have been

affected (e.g, Campbell 1962; Harris 1974; Harris

ef al. 1977"). Apart from the widespread damage

associated with this spevies, it is also responsible

for the partial defoliation of individual eucalypt

trees planted as ornamentals in parks and gardens.

Other than the work of the above authors and

those of Morgan & Cobbinah (1977) and Cobbinah

(1983), very little has been published on the biology

and ecology of L! tugens, while even less has been

reported on its parasitoids, Brimblecambe (1962)

reported five species of primary purasitoids as

"Harris, 1A. (1972) Che effect of flooding on papulation

density of the gum leaf skeletonizer morh, Urabe lugeny

Walk. in Barmah State Forest, Fores) Commission,

Victoria, Research Branch Report, No. 25 (unpubl)

“Elarris, J. AJ Neumann, FG. & Ward, B. (1977) An

outbreak of the gum leaf skeletonizen Urabe lugens

Walker, in river red gam Torest near Barynah. Fores!

Commission, Vicioria, Research Branch Report, No.

87 sonpubl)

* Department of Entomology, Waire Agricultural

Research Tnstinite, Universiny at Adelaide, Cilen

Osmond, S. Aust, S064.

altucking LU dugens and Campbell (1962) ten species

However, in most cases these parasitoids were no!

identified Jurther than family level and, if so, their

identification was not reliable. One of us (G.R.A.)

has recently completed a major sludy on the

interaction of this insect and its parasitoids in the

Adelaide region. This work shows thal the

immature stages of U, lugens support a diverse

complex of hymenopteran and dipteran parasitoids

(22 specics — Table 1), which includes both primary

parasitoids and hyperparasitoids, In this paper we

provide 4 taxonomic framework for the information

on the behaviour and ecology of’ this: parasitoid

complex ard its wteraction with &, fugens, which

will be published elsewhere by ChR.A. Here we

present a key to identify all the parasitoid species

involved, and provide notes on their taxonomic

position and biology, including information on the

stage attacked and place of pupation, Two of the

more common species reared from U /ugens larvac.

which are members of che braconid subfamily

Microgastrinae, and are the subject of detailed

behavioural studies by G.R_A., are described here

as new.

Materials aod Methods

All lite bistery siages of U dugens were collected

froin eucalypts (mostly FE, cumaldulensis, &

feuvoxylon F. Mucll. and &. pricrocanpa (Maiden)

Maiden) between 1985 und 1988 at several sites

within 7 km of Adelaide G. P.O. They were brought

to the laboratory, held gt 20°C in rearing cages, and

larvae provided with new foliage, so that any

parasitoids present could develop and emerge

normally, Parasitoid cocoons found with ibeir dead

host in the field were also collected and reared in

the laboratory at 20°C until adults emerged,

Vi)

4.1, ALIST IN A Ci

R, ALLIN

TABLE 1 Sueterant af relationships berween UU, lugens and its parasitoids and. hyperparasitaids

Parsi

Primary (P).or Hi perparasnaid (H)

Li. levers suive Stage viteryis

pels Family Solitary (8) or Gresuniaus (C1 netacked trom

A a

Treitiourantind Sp, Trichegrammatidge Pis ene 22

Cola yrahae Braconidae BS larva larva

Doalivhogenilea eucalypt Braconidae PS larva larva

fuplecveus ap. Eulophidae BN larva larva

Casihora mer {ehneumoniclae BS lorva larva

Exorista Navivepy Tachinidae WS larva larva

Priborus sp. IchneumoniWae RS hanya pupil

Yenthopunpla rhopalaceroy lehneumociudee PS pupa? pupit

Anirocephatis ay Craleididae PS nope pupa

Brachemeria sp. | Chaleididae RS pupa pupa

Winthemia larenalis Tachinidae Iss i pupa?

Eearktorna sp Eulophidae PGS pupa pupa /purasitoid cocoon

Centrudora sy. Aphelinidae TG — parasitoid coenon

Urachymerid 4p. 2 Chaleididae Hs ~ Parasitoid covoon

Klasius aastrulienses Fla\midae HG — parusitoid cogaon

qpeviys ines, Ewlbophidae IG _— parasitoid eacoon

Podtobns sp. Fillapaidac HG = psrasitad cacoon

species indet, Eupelmidae HS parasitoid carood

Anasiatuy sp. Eupelmidac HG — parasitoid, cocoon

Mesachorus sp. Iulneumonidae HS — paHasditmd cocoon

Paraphylax sp. (chneumomdae His - parasitoid cocaon

Pleririalus sp. Precomilidae Hib - parasitoid cocoon

Parusitoids were stared in a freezer or in 70% 3,

ethunol prior to mounting on pins or card points,

Material for S.E.M. study was washed in half

strength concentrated liquid soap, rinsed in distilled

waiter, dehydrated in an alcohol seriés and critical

point dried using an Emscope CPD 750, before

being cxamined under a Cambridge Stercoscan 250

(Mk 3B) electron microscope. Terminology for

morphology follows Boutek (1988), Eady

(1968), Gauld (1984), Harris (1979), Mason (1986)

and van Achterberg (1979), The term ‘alitrunk’ is

used for the tharax plus propodeum, and ‘gaster‘

is used for the post-ptepadeal segments, The

abbreviation ‘T’ refers to the gastral tergiles.

Abbreviahons for collections are; ANIC, Austratian

National Insect Collection, CSIRO, Canberra,

WARL, Waite Agricultural Research Institute,

Adelaide. Voucher specimens of all species are

lodwed in the Waite Institute collection.

Key to the parasitoids of L'raba lugens

in South Australia

1, Two pairs of wings developed; dorsal surface: very

rarcly with slout bristles: wasp- lke i appearanee (ovi

posiror always developed in fepygle and wsuatly clearty

visible (Migs 5, 7, 29)) (Tymenoptera), , 2

Only one paar od wins developed (fore wine 28); ‘dorsal

surface with nunierods stout bristles; blow fly like in

appearance (Figs 48. 49) (scutum with several black

lensiniding] bands) (Tachinidacy --21

Pow wing with relatively complete venation (eg. Digs

1,2, 6, 19), 3

hore wing wilh pigrnemed venation Feabestt lo dmlenor

Maren (ee Paes 2, 23, 27, FB eee ee ed

More wing, with venation distal to prerosiigma wanting

(Fies J, 2), sem 2 absent (Rraconidaey..-. 4

Fore wing with) distat veins peesent and well-

pizmented; vein 2ancu a te 4, “Hy. {etc

monde), ee a i ett

rréncideum with lonwivaglisal nvedial carina, coarsely

sculptured at least anteriorly (Pig. 12); legs red to red:

yellow . . Cotesta urubue ap. nay,

Propodeum with large arinate areola and horizontal

carinae extending to lateral margins of propodeum

(Fig. 13): leas dark brown fo black with distal parts

reddish... 2... Doalichogenidea eucalypt sp. nov.

fore wing wih an dredlet (Pigs 4, 6).... 0... -- fi

Fore wihg without dn dreolet (Figs 19, 20)......7

Scatum and prapodelin coursely punctate or

ruiwulase: ovipositor very shorn (Pig, 95 oO Benimalny

Wifthoutlone rods protruding posteriorly (body dark

Hrewir to black, fews reddish), 0... er een

Custraria Uvera tooman & Giauid

Seucuim and propodenm generally unsculptired

(oxcepr for propaded) eurinae); OVIpOXiton extending

well past posteriar easter (Fig. Ty, ot pentratia wirk

pair ol lon fads (poresquama) probity poster ian Lye

(body vellow-browin with uitrher Mmarkinsh oc.

Mesochorus sp,

Body brush) yellow will block markings: EE shorn ancl

breistel hasatlly (Vig, 16)...

Nunthapinphr rhapaline eres ‘Krieger

Body nut so coluureds TH narrow Iasi (bigs 17, 18)

fe ee a)

Fore wing with radial cell} short and browd (Fig. 19):

‘TT tlat, broadening distally (Hig. 17) (small species,

lenwth 23-3.2 mm wot including 2 ovipgsilor,

body black except for T2 which is yellow-brown

Paraphylax sp.

kore wing with radial cell lane and narrow (Fig. 20).

MT tubular in basal ball, balbouws in distal half (Fig

18) (large species. lenath 6.5-8.7 mim not including

ovipesitor tor » head and alitrunk black, gasrer

reddish-brown - Frihorus ay)

eae

10.

PARASIVOIDS OF WAAR FUGENS V7

Femur of hind leg greatly expanded, juolaed

ou sepnited aloe lower margin (Fig. 24) (hal

white) ee pit eco se ey Le

Tymar of hind leg normal, smgott along lower tiangin

(Pipe 25; 35) oy ecaarpeeesse 4 ee wae ee 12

fore wing willt marginal ver much longer than

pastmarginal vein (Fla. 22); apex of hind tibia tapering

Imo strong spine (Brachpmneriay. 00.000,

Pere wing, with marginal vein about same length as

pasiparginal vein (Fig, 2 apex of hind sihia

perpendicularly (runeale (Fig. 24) (large spevics, 4.8

mm in teneth; body black, bine lee dark red-brawn

marked witth black) Antrecephalas sp,

Boy black with reel lined lenmurand tities 42-44 myn

in Jength, dag ey eet vee Brachymeria sp |

Kody black with Whikkyellow markings on rowalau yd

leps; L8-23 mim in Jength,.... Brachynerie sp. 2

- Hibd voxa developed as large [at dise; hind tibiy waph

Uistinel criss-cross patterd al setae (Fig. 25) (fore wing

with stigmal vein very short (Fig. 23). body dark.

tezula and legs excepr for hind coxa pale; body length

ot 18-27 mm, ¢ 13-19 mm) (Elasenidac)

vevtreees cs 2 Blass australensis Girault

Hind coxa, hind tibia and stiemal vein nat as above

ane, tri teet th Mee. te 13

. Pronotum (seen derselly) targe and qiradeangutiar;

dorsal surtave of abtrunk coarsely sculptured (Lig.

2h), fole Wing Venation asin Fig, 27 (hody black aod

Hon-metilhe, legs with seme pale “tnarkings)

(Buryronmdaey, oc .. eee) Burytone sp.

Pronoturny not large and quadrangular and ahrrunk

withoul such sculpraniies: body offer meruic in

OTE usttetn. ee, 345 ath

Body length greater than about 1.5 ome: waster

sepiraicd from alinnink by narrow waist (ges 28, 29)

Ai ote, & Het - oo 20

Tarsi S-segmented (ull Jigs 24, 257... , 16

Farsi 4-segmiented (Pig. 359 ito LB

Mesopleurion hot cotired and shield-lke (Ei. 28);

body rather robust with large hoad ancl alitrunk (2

aaster i laigral view Sharply angled: cr paster snvall

and (Tallened; body length of & 23-33 mm,

o [7-24 mince inetallic green ia colour with

yellow-hrawi legs and antennae) (Pieromatidae)

» Preramalus sp.

body sumewhett

- Fore wing with stigmal and postmaruinal veins very

Shor (Fiy. 34); anterior sewlellum longitudinally

strivte (Pig. 43) (hady dark with merallic ureen tinge,

11-15 mm in engi), 00. Pediohnes sy.

Vivre Wiig wilh stigmal and postinarginal veins long

(Fig 7)> scutellum sinmorh or with fine rencntate

UD UUTINE 5) ; resold

19. Dorsal head and alitrunk with scartered long bristle

like hairs (hig. 34) (body mostly dark and non-

metallic; eyes red; antenna Jegs.and broad pareh on

dorsal gaster yellow-brown), 2... Euplecius sph

Dorsal head und alitrunk with shorter liner fairs

(dorsal alitvunk with distinenve avetaltic green ane

yellow markings (Fig. 4%)), 280. & species inder

20. farsi 3-scumented (big 40); antennae S-senmented

(Figs 4), 42); fore wing very browl (Fig, 39)

(Trichograntiniatidue). .... Trichogrammd sy

farsi S-seemented (Fig, 47); antennag 6-sepmenied

(Higs 45, 46); fore wing narrower (Fiz, 43) (Aphel-

inidac). -. GUnTRodara sy

21) Hues cayering occiput (pasterion purt of head )silver.

grey, abdomet in dorsal view with anterior, medial

and posterior parts black, latlurs! areas brown (Fis.

FA) eee Winthemia lateralis (Macquart)

Hairs covering ogviput salden brown, Ist seument ot

wbdomen black, other segments blagk with parclics

of silva (Pin. 49)... Beoriste Hayiveps Muaciuart

‘Treatment af species

HYMENOPTERA

bamily Beaconigies

Cotesia urahae sp, nov,

FIGS 2, 8, 0-12, (4

Malotwpe: F, ANIC, 8. Aust., Adelaide (Mitcham),

reaied ex Uraha lugenson Kucalvpiys microdarpa,

coll, 14...J985, emerged 3,xi.)985, G. R. Alten,

Paratypes: 8 O, 20 o, same dala as holotype

except some with different suburbs of Adelaide and

dilferont dates, 179 9, orc, S. Aust, Adelaide

(Waite Institue campus), various collecting and

emergence dates during 1964, |. Hope, §¢ >,10,

5. Aust., National Park, Belair, coll. 20,x7,1964,

emerged 25.41.1964, RDM. (59 9, Jar, ANIC:

rest of material including 1% gold coated for SEM

(Wings slide-mounted) in WAR ).

Femate

Length. 2.9 mm (2.53.2 nm, r=

oVipositor.

Colour, Body black; palps yellow: lugs cxcepr far

coxa yellow-brown, hind leg infus cute distally, distal

chd of hind tibia with dark patch; anterior pleural-

sternal region of gaster dark red, wings liyaline,

stigma cvenly and darkly pivsmented, as is rest of

fore Woe venation,

Heal. Mn anicrior view vertex arched su that head

is somewhat circular; face and gena with lopewh

White hairs and associated ting collieulaie sculp-

turing; ther margins of eves siriually parallel: in

dorsal view ovelli in wide triangle, posterior uangem

of median ocellus coincident with imaginary line

across {he anterior margins af laicral ocelli: frons

and medial occiput siiooth and talrless; lemples

with white hairs avid Yssociated collivulate aculp

10) including

172 A, D, AUSTIN & G. Ro ALLEN

—

Figs \-7, 1-2, Fore and hind wings: |, Dotichogeniaea excalypul sp, Nov,, 222, Culesia urabae sp, nov., 3; 4, Lateral

view of vaster uf Dolichoxenidea eucalypt spinov., 95 4, Distal tore wing of Mesockorus sp, 9) 3-6, Cusinaria

atiera Jerman & Gaull, 9: 5, Lateral view of easier; 6, Fore wing: 7, Lateral view of gaster of Mesochors ap. 9,

Scales: Figs 1, 2, 4.and 6 = 0.5 mm: Fie. 3 = 200 um; Figs Sand 7 = 1.5 mm, Abbreviations: a = areolet; pt =

pterostigma,

turing which ig slightly coarser than on face; vertex

with few scattered short hairs otherwise smaoth;

antennae slightly longer than body, distal 4-5

segments only slightly Jonger thar wide.

Alitrunk. Seutum punctate with covering of shortish

hairs, punctation denser along courses of notauli

and along lateral margins (Fig. 8), punctation along

posterior margin becoming slightly longitudinally

elongate; courses of notauli faintly depressed, these

faint depressions broadening posteriorly; scutellum

faintly punctate; phragma of scutellum exposed

along posterolateral margins (Fig. 8); propodeum

PARASITOIDS OF URABA LUGENS 173

Figs 8-11, 8-9, Dorsal view of alitrunk; 8, Cotesia urabae sp. nov., 9; 9, Dolichogenidea eucalypti sp. nov., 9; 10-11,

Cotesia urabae sp. nov., 9; 10, Dorsal view of gaster; 11, Lateral view of gaster, Seales: = 200 am,

coarsely rugose to rugose-striate in anterior half,

generally smoath with faint ragose punctation in

posterior balf, medial longitudinal carina well

developed with associated short horizontal and

oblique carinae (Fig, 12); mesoplewron finely

punctate in anterior half with associated short hairs,

smooth and hairless in poslerior hall except for

compact group af 5-6 foveae medially; metapleuron

smooth antenorly, rugase-punctate posteriorly (Fig.

14); hind coxa faintly punctate on dorsal surface

with associated short hairs, this sculpturing

becoming coarser on ventral surface.

Wings, Fore wing with veins c and 2-SR sharply

angled, r slightly Jonger than 2-SRy cu-a almost

striate; distal part of basal cell and anterior part

of sub-basal cell devoid of hairs or almost 50; discal

cell sparsely covered with hairs; hirid wing with yein

r (spectral) present (Fig. 2),

Gasier, T1 as wide as long, broadening posteriorly,

surface in posterior two-thirds coarsely punctate to

rugose-punctate, becoming coarsely striate jn

posterior one-quarter (Fig. 10); sclerotized part of

T2 rectangular, slightly wider than T1, coarsely

Tugase-punctare With a few longitudinal striations,

sculpturing fading lo nearly smooth in pastero-

Jateral corners, longitudinal midline smooth; T3

usually rugose-punctate in anterior ane-quarter to

two-chirds, with seattered hairs, posterior part

smooth, in some specimens T3 virtually smooth

throughout but always with al least anterior margin

with band of punctation; rest of tergites smooth

wilh scattered hairs; in lateral view hypopygiurt

pointed, extending past posterior easter, with scat-

tered fine Nairs, ventroapical margin not indented

(Fig, 11); ovipositor sheaths with few apical hairs,

Male

As for female except lor length, 2.8 mm (2.6-3,1

mm, nm = 10) and sexual differences (genitalia and

development of hypopygium),

Qlther material exantined: S. Aust., suburbs of

Adelaide. various dates and vollectars, 109 9,

16c 4 -WARI (excluded from type series because

material is damaged or inadequately labelled),

Camments: The sculptunng on the propodeum and

TI-f3, the shape of these sclerites, the form of the

hypopygium and ovipositer, clearly place (his

species in Coresia Cameron. Colesia has previously

been referred to as (he g/omeratis species-eroup of

Apanteles s./. (see Mason 1981; Nixon 1965), and

is the fatgest generic level taxon in the subfamily

Microvastrinae, the latter comprising same 1300-

1500 described species world-wide (Mason 1981),

Ju Australia Colesig ts common and diverse, but

other than several species intfeduced trom Europe

and North America as bjo-control agents tor certain

lepidepteran pests (viz, G /lavipes Cameron,

(Cresson), C plure/lae (Kurdjumay), C. rubecula

(Marshall) and C. rufierus (Haliday)), che

Australian Fauna rentains unstudied, A few poorly

characterized species, which presently remain under

the name Aparteles 3,1, May turn oul to belong in

Cotesia, These species are unlikely to be conspecific

with the present species as their type localijies are

outside rhe known distribution of C urabge sp. nov.

or they are asgociated with other hosts, There are

no workable keys fo Indo-Australian species of

Cotesia. However, the key in Nixon (L974) to the

north-western European fauna can be used io

separate C, wrubae sp, nov. fram four af the seven

introduced species mentioned above. Of the other

urabae sp, noy. in that its body is strongly flatrened

dorsoventrally (see Austin 1989), while C. kuzak and

(~ rnarginiventris can only be identified reliably in

association with their hosts, Helicoverpa spp. and

Myrhitina coavecta (Walker), respectively,

Biology; C. urabae sp, nov, is a solitary, primary

eridoparasitoid and oviposits into early to inter-

mediate larval instars of its host, emerging from

intermediate to Jate instars before pupating. The

pupal cocoon is alongside the host and is sulphur

yellow-green with a surrounding silk matrix, This

species has only been reared from t/ dugens.

Dolichogenidea eacaly pti sp, noy.

FIGS 1, 3, 9, 13, U5

Holarype: 9, ANIC, S. Aust, Adelaide (Highgate),

reared ex Uraba lugens on Excalyptus camal-

dulensis, collected 1,x7.1985, emerged 17.x7.1985,

G, R, Allen, Paratypes; 18° 9, 20o°o same data

as holotype except some with different suburbs of

Adelaide and different dares; 139 9, Soo, S,

Aust., Adelaide (Wajle Institute campus), various

collecting and emergence dares during 1964, L-

Hope (59 9, Sorcr, ANIC; rest of material

including 19 gold coated for SEM (wines slide-

mounted) in WARE).

Female

Length 2.9 mm (2.53.1 mm, 1 = 10) meluding

OVIPASITOT,

Cu/our, Body black, wings hyaline woh darkly

pigmented venation, palps brown, legs black with

distal Fore legs, tarsi of mid and hind legs and ubio-

femoral joint yellow-brown.

Head, Mostly smooth except for tine colliculate

sculpluring associated with dense covering of short

PARASITOIDS OF URABA LUGENS 175

Figs 12-15. 12-13, Propodeum and first and second tergites of gaster: 12, Cotesia urabae sp. nov., 9; 13, Dolichogenidea

eucalypti sp. nov., Q; 14-15, Lateral view of alitrunk: 14, Cofesia urabae sp. nov., 9; 15, Dolichogenidea eucalypti

sp. nov., 9. Scales: = 200 pm.

176 A.D AUSTIN & GR, ALLEN

hairs; surface with Characteristic dull lustre; in

antefior view inner margins af eyes virtually

parallel; in dorsal view ovelli forming wide triangle,

posterior rangent af median ocellus coincident with

imaginary Line across the anterior margins of lateral

weelli, fine colliculate sculpturing and assacialed

pilosity slightly denser across vertex and occiput,

excenl for smooth narrow band around posterior

margins oF eyes; antennae reaching to posterior

gaster or shghtly beyond, uistal three segments

slightly longer than wide and sometirnes shghtly

compressed.

Alftrunk. Scutum coarsely punctate with punctures

mastly closer to each other than their own diameter,

except along posterior border and along. courses of

notaull, which are thus faintly indicated (Fig. 9);

scutellum smooth; scutum and sculellum densely

cavered With short hairs; metanoium rather broad,

an(erolateral margins emarginate so that phragma

of scutellum well exposed; caringe forming

propodeal arevla raised well above surface, carinae

extending laterally belaw lyorlzontal midting, these

carinac with dorsal and ventral extensions forming

cristulae, but nat enclosing spiracles (Fig. 13);

anterior part of propodeutn mastly smooth and

setose, posterior part with faint rugose-puinctate

sculpturing which becomes more obvious laterally;

mesopleuron sefose in anterior half, smooth

posteriorly; metapleuron mostly smooth, except for

ventrapostertor one-third which is rugose-puncvrate

(Fiz, 15); distal fore rarsus without spine opposed

to tarsal claw:

Wings. Fore wing venation as in Fig. t; costal and

basal cells bare posteriorly; Kind wing broad; vein

LSC+R deeply bowed; r present but faint;

cubilellan cell moderately broad: submediellan cell

minded posteriorly.

Gaster. 1) as wide as long, slightly widened In

posterior half, jateral margins slightly emarginale,

surface mastly punctate, striale-punclate along

lateral mains and stale in posterolateral corners

and along posterior margin (Fig. 13); sclerotized

part of T2 slightly wider than TL, 25 x wider than

long, mostly smooth wih faint scattered punctures:

TS slightly longer than T2 (14:11); T4-16 shorter

than T2 (R11), T7 very short, about one-quarter

length oF T2; 13-16 all smooth; T2-17 each with

single transverse row of hairs; ovipositer and

sheaths shorl, not extending far past posterior

gasier: OVipositar with strong distal attenuation

(Fig. 4}; hypopygium lacking obylous lateral creases

though weakened normally im ventral midline:

Male

As for female except as follows: Length 2.7 mm

(A528 mm, n = LW); alitrunk very slightly

flanened dorsovenirally; fore wing stigma unpie-

mented in medial area so that it is transparent: rest

of wing venation generally with less prementation

than female; TI sometimes with dense rugose-

Pulictate sculpturing merging with posterior

striations, otherwise same as female except for male

genitalia and Lacking hypopygium,

Comments: The sculptunng of the sculunmt and

propodeum, shape of Tl and T2, form of the

hypopygium, and shape and fringe of the vannal

lobe of the hind wing clearly place this species in

Dolichogenidea Viereck. Previously Delichogenidea

was considered as three related species-graups in the

genus Apanteles s./,, vic. he ultor, laevigains and

lodpipalpis species-groups (see Mason 1981; Nixon

1965, 1967). This species falls into the witor group

Which was revised by Nixon (1967) for the [ndo-

Australian region, Jn this work 2 euculypit sp. nov.

keys out to _D, cleo (Nixon) (couplel 23), a species

Known only from India and associated with a

nymphalid host Eriboae arja Felder, or with some

difficulty it keys as D. cuniae (Wilkinson) (couplet

3b), which is known only from Java and associated

with a limacodid, Cania bandura Moore (Austin

1987). Apart from having different hosts, these

species differ from 2 evcalypti sp. nov. in that 2,

voniae has an unusual striate seulpturing pattern

on T) and T2, and D, cleo has the sub-basal cell

evenly and darkly setose, the hind femur yellow in

colour, the proximal half af the ovipositar very

broad, and the mesopleuron coarsely rugose-

punctate anteriorly. This is the fourth species of

Dalichozenideu tecorded from Australia, tte others

being DO Jipsis (Nixon) comb. noy., DB niris (Nixon)

comb, noy. and Q tasmanica (Cameron) comb. navy.

Examination of the holotypes of these species shows

that they differ from & eucalypt sp, nov, in a

number of important characters, All three have the

oviposior much Jonger, being at least as long as

the hind tibia, and, if addition, D, Jipsis and DB,

tasmanice have a while spot ou the cheek and a

much reduced propodeal areola. These species can

be readily separated from 2. eucalypt sp, poy, using

the key in Nixon (1967),

Biology: D. eucalypli sp, nov, isa solitary, primary

endoparasitoid which oviposits into early to

intermediate larval instars of its host, emerging From

inlermediate to late instars before pupating, The

pupal cocoon is spun alongside the host and is white

in colour and lacks a surrounding silk matrix, This

species has only been reared from UO lugens.

Family tehneumovidae

Xanthopinpla rhopalaceras Krieger

FIG. (fi

PARASITOIDS OF URSHA LUGENS 37

This species (s easily identified by its distinctive

colour pattern and fore wing venation. In South

Australia it is Knows from two specimens (WARL),.

one collected 4s an adult in the Adelaide region,

the other reared from U. /vgeas at Keith, It is also

known from Queenstind and Tasmania where it has

been recorded from Ll /ugens, ard (he Lortricids

Epiphyas postvittane (Walker) and MeropAves

divulscina (Walker) (Brimblecombe 1962; Jumble.

ton 1940; Gauld L984) also see Townes & Chui

1970), It is a solitary, primary endoparasitoid and

emerges from dhe pupal stage of its frost. See Gauld

(1984) fur additional taxonormie information and

hst of synonyins.

Parapivlax sp.

FIGS t7, 19

Peraph ay is a large Old World genus with more

than 50 recognized species from Australia, the

majority of which are undeseribed (Guuld 1984),

Where their biology is known Porgpiviax spp. have

been recorded as primary and hyperparasitoids,

mostly of lepidopteran hests, Apart from the

charagters given in the key this species 14 notable

in comparison to other parasitoids assactated willt

UL /ugens for its relatively smooth unseulptured

body {except for propodeal carinae) and lateral teeth

on the propodeum, The species here belongs to the

cormvax species-group (see Gauld 1984) and is only

khown from the Adelaide region, where it has been

reared as an obligare, solitary hyperparasiloid

Erthorus sp.

FIGS 1%, 20

This is a distinctive species when compared to

ihe other ichneumonids associated with UL, frees:

In addition to the characters given in the key this

species has distinctive reticulatc-puncrate sculp-

(uring on the scutum, seutellum and propodeunt.

Friborus sp. is 4 solitary primary parasitoid of L

ingens, ovipositing into the larval Siages and

emerging from the pupa. dels poly known from the

Adelaide region and has been reared from its host

On various occasions since 1965 (WARL, unpub-

lished recorels),

Casinaria micra Jerman & Gauld

FIGS 5, 4

This species is a solitary, primary enduparasitoid

eisily recognized by its lore wing venation, shor

ovipasitor and colour, ft has been recarded trom

ull states in Australia aod, although it has been most

commonly associated with U fiers, C mucru has

been reared [rom species belonging ro three other

distantly related lepidopteran families — Geomet

tidac, Qecuphoridae, Notodontid&e (see Gauld

184; Serman & Gauld 1988), The pupal cocoon is

constructed near to (lerman & Gauld J98X) or

underneath (observalions in this srudy) its dead

larval hoscand is attached firmly to the jeaf surface

Tt iS grey-browi and marked with characteristic

black spo. See Jerman & Gauld (1988) for add-

ional taxonomic information and list of synonyrnos,

Mesochorus sp.

FIGS 4,7

This is a large cosmopolitan venus of printary

and hyperparasitoids of lepidopteran and colooys

teran hosts; mos} Indo-Australian species are tindes-

enibed (Gauld (984). The species recorded here Is

a solitary obligate livperparasitoid of &. fugens

through C micra, C. urahge and DB. enculyprti, and is

known only from the Adelaide region. It is a pale

coloured delicate species. with fine thread-like

antennae, The male is distinetive in having the

sxonosquama Of the genitalia extending from the

posterior gaster as 4 pair of tony rods,

Family Chaledidar

Brochvinerta spp,

PG, 22

This ts a large genus ui Australia with nearly 70

described species, the majority of whieh are

primary. pupal parasitoids of Lepidoptera (Boutek

1988), The two specics meorded here (both

unidentified) can be separated easily by the

characters in the key. Spevies | is a solitary, primary

parasitoid of GO. /daens and emerges. from the host

pupa. Species 2 16 solitary and hyperparasitie

noi known whether this relationship is facultative

or obligatory. In eastern Australia Iwo species of

Grachymeria have been reared from LU, /uvens, vie.

8. Jroggutti (Cameron) (Rrimblecambe 1962) and

A. rubripes Girwalt (Campbell 1962) (B rbripes is

considered a jubior synonym ot B fear (Watkerk:

sec Boutck 1988). However, the matertal from these

records Would have been identified ar atime when

the Australian species in the gsenus were confused

by most authors and hence might be misidentified,

Phese numes should thus be used with some care,

especially since Routek (1988) did not see any

matetial in Australian collections or elsewhere

revred trom Of fugens that he could assign. ro either

8: froggattt oc Bo teuta,

Autravephalus sp.

PGS 2b, 24

This spevies is fepecsetited here hy a site

178 A. D. AUSTIN & G, R, AIT EN

PAKASITOIDS OF URA#A LUGENS \7y

Specimen reared as a primary parasitoid trom the

pupa of UL lugers in the Adelaide region, Ti is easily

distinguished by the characters in the key, There are

more than 60 described Australian congeners which

are discussed by Bouck (1988).

Family Eurytomidae

Fueyroma sp,

FIGS 24, 27

(his is 4 large cosmopolitan genus with more

thin 60 described Australian species (Boucek 1988).

Biologically the group is very diverse including

phytophagous species, primary parasitoids (mostly

of tepidopteran hosts) and typerparasitoids, The

species recorded here develops as either a primary

Bregarious parasitoid of Ui /uvens, or as asolitary

hyperparasitoid through (2 wrehee.

Family Pleramalidac

Pferonialus sp.

FIG. 28

This specigs ts-relatively easily separated from

other Chalcidoidea associated with U, lugens by its

robust body and metallic green colour, The genus

ig Gixonomically very complex and the Australian

species are in need of revision (Bougek 1988). The

species recorded here is an obligate, solitary

hyperparasitoid reared from U, Jugens through Cc

micra, C. wrafae and D, eucalypii in the Adelaide

Tegion.

Family Flasmidae

Elasmus australfensis Girault

FIGS 23, 25

A distinctive species recarded here as an obligate,

solitary hyperparasitoid of UL fugens through C.

auera, C. urabae and D: excalypti. Previously it was

known to be hyperparasitic and occasionally

gregatious through an unknown ichneumonid

assuciated with CC /ugens in the ACT, and also

fias been collected from N.S.W., and northern and

southern Qid (type locality: Gordonyale, QId). This

ty the first record of E. australiensis from §, Aust.,

indicating that it is probably distributed throughout

south-eastern Australia. See Riek (1967) for

addifional taxonomic information and list of

synonyms.

Figs tqy-28

Aeros sp oe

Vetravephaliy ap. 25 22, Brachyieria sp. 2,

Anravephalys sp.

IG-15, Virst tergite of gaster: 16, Aanthapinpla rhapaloceroy Krieger, (97

1-20, Distal tore wings: 19, Paraphylie sp.

Family Eulophidae

Euplectrus sp.

FIGS 36, 37

Eviplectrus is a cosmopolitan genus of preparious

ectoparasiloids of lepidopteran larvae, Which 1s

represented in Australia by 13 described species

(Boutek 1988). The spevies recorded here is solitary

and only known from the Adelaide region.

altacks the early to intermediate larval stages, killing

them before they pupate while pupating itself

Undernesth the dead host.

Pediobius sp.

FIGS 33-33

This is a large, cosmopolitan genus of primary

and hyperpurasitoid species that attack a wide range

of insect groups (Boutek 1988). There are more than

30 described Australian species, mast of which do

not have associated host information (Boutek

1988), The unidentified species recorded here is

easily distinguished from other Hymenoptera

associated with Uf lugens by its distinerive venation

and sculpturing on the scutellum. Jt is represented

by four specintens (WARI) reared in the Adelaide

region from L. dusens and developed as a solitary

hyperparasitoid through Evplecteus sp. and-as a

gregarious hyperparasifoid through 1. excalypri.

Fulophidae (genus & species indet.)

FIG. 38

The two specimens reared as hyperparasitoids of

O. lugenrs through D. eucalppti in the Adelaide

région could not be identified to genus due to phe

poor condition of ihe material, They are different

from the other eulophids recorded here and can be

distinguished by the dorsoventrally flattened body

and distinctive colour pattern,

Family Eupelmidac

Anastalus sp,

FIGS 29-3)

This species is easily separated front the other

chaleidoids associated with UL /ugens by its

distinctive body shape and banded wings. The genus

is cosmopolitan ard Known mainly as parasitoids

of the eggs of Lepidoptera and Heteroptera,

17 Peraphvlis sp. 95 UR,

V7 20, Erifarus sp. G. 21-23 Fore wires: 21,

B28, Elasmus dustralieusis Giraull, GO: 24-25, Hine levs: 14,

25, Elaxemues australensis Gira, &. Scales: Tigs 16-19 = 250 pins bigs 20 aud 24 = 0.5

mm Pye 2) = ON maw Pig, 22 = 250 pans big, 23 = 200 yin; Fip, 28 = Loimm. Abbreviaiionss uy = cova t=

hide, Ob = mureinal ver: pm =

postMarginal veins R

= radial val, st = Sstigmal vem; is = aisal seMINCAS,

180 “4D. AUSTIN & GR, ALLEN

Pigs 26-35 26-27. Aurvieme yp, 4 26, Dorsal wew af atiorunk; 27, Pore wing; 28-29, Lateral view of body: 28,

Pieramulies sp, 29, -bauwarus sp., 2) 30. Forge wing of Avfesfafis sp,, 27 31-33, Dorsal view of alitvunk: 31,

Anasweiy sp. 7732. Eupelmidae, genus & species indet., oy 33, Pediobius sp., 9; 34, fore wing of Pediobivs yp.,

2, 38, Hind feo of Peeiabins sp. %, Seales, Figs 26, 27, 29, 34 and 35 = 250 wm; Fig. 28 = 0.8 mm; Pin. 30. =

200 wan; Pigs 31-33 = 150 am. Abbreviations: mp = mesopleurony pm = postmurginal vem; ke = sculums: xl =

scurellumy st = stigma) veing is = tarsal segments.

PARASITOIDS OF URABA LUGENS RI

a felt { rN

bigs 36-42. 36-37, Euplectrus sp., 97 36, Dorsal view af head and atitrunk; 37, Pore wing: 38; Dorsal view af alitrunk

of Eulophidae, genus & species indel., 2 (stippling indicating colour pattern); 39-42, Trivhogrammnia sp. 39, Pore

and hind wings, a; 40, Hind leg, 9; 41, o antenna; 42, 9 antenna. Scales: Figs 36 and 37 = 250) yim; Pigs 38 and

39 = 150 pin, Fig. 40 = 100 um; Pigs 41 and 42 = 50 #m (same seale line). Abbreviations: pm = postmarsinal

vehi, st = atigmal yein; ts = larsal seoments,

although a few are recorded as hyperparasitoids

through bravonids, Anestatus is represented in

Australia by 40 described species (Bouéek 1988),

The species recorded here is represented by three

specimens reared as a gregarious hyperparasitoid

region.

Eupelmidae (genus & species indet,)

FIG. 32

A single male specimen reared as a hyperpara-

sitoid of Uo lugens through CG urabue in the

Adelaide region could not be identified 1a genus,

but it is not Anastatus sp., from which it van be

separated by the characters in the key.

Family Trichogrammatidae

Trichogramma sp.

FIGS 39-42

The members of this cosmopolitan genus are

1k? A.D, AUSTIN & GR. ALT EN

Figs 43-49. 48-47, Centradora spc 43, Fore wing, 7-4, Dorsal view af whole body, Fs 44) 2 antenna, 46, cr

antenna: 47, Hind lee, 9:48, Abdumnest of Mlthentiatoveraly. 9) (Miaxcquartk: 49, Dorsal siew of Peoresta flaviceps

Macquart, 2 (stippima showing colour patiern in Figuees46 and 47). Seales: Fig. 43 = 200 jum: Fig. 44 = 300 yan,

Fieg 45-47 = 50 aim (same scate tine for Figs 43 and 16); Figs 48 and 44 = Lijmm, Abbreviation: 1s = tarsal segments,

obligate, primary parasitoids of insec! eggs, most

frequently those of Lepidoptera. [t is the anly egg,

parasitvid of U. lugens so far recorded, and it can

be identified by the characters in the key, as well

ay its minute size, dislinctive lore wing venation,

fore wing setal pallern, and very narrow. hind wing.

This species is a solitary parasitoid known only front

the Adelaide region, We. have not seen material of

the Jrichograrmima sp, reared [rom U. lugeris in the

Murray Valley in N.SW. (Campbell 1962), which

may be the same species to that recorded here,

Tamily Aphelinidae

Centrodort sp.

FIGS 43-47

This is a cosmopolitan genus of about 40

deseribed species (Llayar 1983), most of which are

primary parasitoids of the eggs af Orthoptera and

Homoptera, although at least one Species is

reported to be hyperparasitic (Gordh 1979; also see

Viggiani 1984), The species recorded here is an

obligate, gregarious hyperparasitoid of U fugens

PARASITOIDS OF URAHA LUGENS 1s3

through C. urahae or 2 eucalvpti. Apart from the

characters in the key and the life history stage

attacked, Cenfrodora sp. can be separated From the

other parasitoids associated with UL lugens by is

minute size, wing venation, and ovipasiter which

1% more than half the length of the guster {the

oviposiior being significantly less than half the

gastral length in Trichogramma sp.).

DIPTERA

Family Tachinidae

Winthenvia lateralis (Macquart)

FIG. 48

Previously this species has heen collected at

various localities in all states of Australia. It has

been reared from host species belonging to seven

lepidopteran: families, viz. Arctlidas, Noctuidae,

Pieridae, Notodontidae, Nymphalidag, Saturnidac

and Geometridae (Crosskey 1973; Cantrell 1986,

1989), The only record trom U fugens is from

specimens if this: study (3 specimens, Waite Institute

campus, J. Cobbinah, 1975, WARI). 4 tuteralis

oviposits onto the external surface of a hast larva.

Alter hatching the fly larva penetrates the host larva

and usually emerges from the host pupa, alchough

we were not able to confirm its’ biology in this study,

Set Crosskey (1973) and Cantrell (1986, F989) For

additional taxonomic information and list of

synonyins.

Exortsla flaviceps Macquart

FIG. 49

This species has been recorded from all states of

Australia and the N.T, (Cantrell 1985), and has been

reared from members Of nine lepiduptcran families,

Vaz. Lymeanimidas, Anithelidac, Pieridae, Aguris-

lidac, Sphingidac, Genmetridae, Notodontidae,

Lasiocampidae and Noctuidae (Crosskey 1973;

Cantrell 1986), Lt ovipeastts unto the surface of U.

fugens laryac. After hatching the fly larva burrows

into che lost ro feed and develop internally, finally

emerging from late larval instars fo pupate outside

the dead host. The colour pattern on the abdomen

and occiput of the head ts the easiest way (0 clis-

tinguish (his species trom Ff laferalis. See Caitcell

(1985) for additional taxonomic information and

list_of synonyms

Other Parasitnids

trom LU. /ugens in Queensland Brimblecombe

(1962) reared (wo ichneumonord species Which have

not been recorded in S. Aust, viz. Jrebuthe sp.

(Ichneumonidae) and Camprloneure sp. (Bracon-

tac). Also Gauld (1984) reports the following

ichneumonids as having been reared trom U dugens:

Stiromesostenus spp., Campoplex sp. and

Pristomerus sp,, but again, they were not reared

during this study and may: not be found in S, Aust.

Acknowledgments

We thank Dr lan Naumann (Australian Natiunal

Insect Collection), Mr Geatf Hollowsy (Australian

Museum) ail Dr Bryan Cantrell (Qucenstand

Deparunent of Primary Industries) for assistance

with identifications: Mr Pan! Dangerfield for the

line drawings and scanning micrographs, and Dr

Mike Keller for reading a draft of the manuscript.

This: work was supporled by a Commonwealth

Postgraduate Research Award to GRA and a grant

from the Australian Biological Resources Study

Participatory program to ADA,

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CONTRIBUTIONS TO THE TAXONOMY OF AUSTRALIAN

BUPRESTIDAE (COLEOPTERA): NEW SPECIES OF ASTRAEUS AND

STIGMODERA (CASTIARINA) AND A KEY TO ASTRAEUS (S.S.)

BY S. BARKER*

Summary

Astraeus meyricki Blackburn is resurrected from synonymy, redescribed and illustrated. Astraeus

badeni van de Poll is redescribed and illustrated and the following five new species of Astraeus

(sensu stricto) are described: A. aridus sp. nov., A. occidentalis sp. nov., A. princeps sp. nov.,

A. williamsi sp. nov., A. yarrattensis sp. nov. A revised key to Astraeus (s.s.) is provided. The egg

of A. pygmaeus van de Poll is illustrated and some biological information given. Two new species

of Stigmodera (Castiarina) are described: S. garnettensis sp. nov., S. wellsae sp. nov.; the first has

sexually dimorphic colour pattern, rare in the sub-genus.

KEY WORDS: New species, Astraeus, Stigmodera (Castiarina), Coleoptera, Buprestidae.

Transactions of (he Raval Society af S, Ause, (989), TL3(4) 185-184,

CONTRIBUTIONS TO THE TAXONOMY OF AUSTRALIAN BUPRESTIDAE

(COLEOPTERA): NEW SPECIES OF ASTRAEUS AND STIGMODERA (CASTIARINA)

AND A KEY TO ASTRAEUS (s.s,)

by &. BARKER*

Summary

Barker, 5. (1989) Contribulions to |he taxonomy of Australian Buprestidue (Coleoptera): New species

of Axrraeus and Sigmodera (Castiarina) and a key to Asrraeus (s.8.), Trans. R, Nae, 8, Aus 134), TRS-194,

30 November, 1989

Asiraéus meyvrick? Blackburn is resurrected from synonymy, redescribed and illustrated. Astraeus budent

van dé Poll is redeseribect and illustrated and the following five new species of srraeus (sens sfricto)

are described; 4. an/dus sp. nov., A. eeeldentalis sp. nov, As prineeps sp. nov. AL williams’ sp. Way, A.

Jarrattensis sp, noy, A revised key lo Asirueus (5.4) is provided, The egg of 1, pugmeeus van de Poll is

illustrated and some biological information given. Two new species of Sriginadera (Castiarinia) are described,

S. gornettensis sp, nov,, 5. wellsue sp. nov; the finest bas sexually dimorphic colour pattern, rare in the

sub-gerius,

Koy Worps: New species, Aatraeus, Sigmodera (Custiarina) Coleoptera, Buprestidae.

Introduetion

The buprestid genus sfraevs was revised by

Barker (1975) and additional species described by

Barker (1977). No further information has been

published. Fresh material has now come to hand

and five new species are described herein including

one species from the Flinders Ra., S. Aust., which

was misidentified in the revision of Barker (1975).

The separate Stalus of A. baden*yan de Poll and

A, meyricki Blackburn has been in doubt since they

were deseribed, A re-examination of the specimens

available has convinced me that they are separate

species. The key to Asyraeus (s,s,) given by Barker

(1975) requires Updating to include three new species

published by Barker (1977), five new species

described herein and one species resurrected [rom

synonymy. A new key is presented and new lovality

records of two species recorded. The egg of A.

pygmaeus van de Poll is illustrated, the first such

observalion of an Asirdeus species, together with

biological information on A. pygmaeus. Two new

species. of Srigmodera (Castiarina) are described

and illustrated; one being unusual in showing sexual

dimorphism in colour pattern, a rare occurrence in

this sub-genus,

Materials and Methods

Male genitalia were prepared and displayed by

the method deseribed by Barker (1987). Specimens

for S.E.M, were dehydrated prior to being coated

with gold-palladium, Abbreviations used in (he text

for museum and private collections following Wall

* Department of Zoology Liniversity of Adelaide

GPO. Box 498 Adeliuide, S. Aust. S001

(1979) are: BMNH_ British Museum (Natural

History), London; JHQA Mr J. Hasenpusch.

Innisfail; MNHN Museum National d'Histoire

Naturelle, Paris) NMVA Museums of Victoria,

Melbourne; SAMA South Australian Museuni,

Adelaides WAMA Western Australian Museum,

Perth GWNA Mr G. Williams, Landsdowne,

GWOA Mr G. Wood, Atherton,

Resurrection of Astracus nreyricks,

Van de Poll (1892) considered A. mepricki

Blackburn a synonym of 4, hadeni van de Poll but

Blackburn (1895) disagreed, Barker (1975) followed

van de Poll, | have re-examined the two male

syntypes of A. meyricki held respectively in the

BMNH and SAMA collections and compared them

with specimens in the SAMA identified by

comparison with the female holotype of A, badent

held by the MNHN, The pronotum of 4, badeni

is Widest medially and dilters trom A. meyrickl

which is widest basally, Male genitalia of A,

meyricki are broader than those of 4. baderi and

are rounded more abruptly al the apices (Figs 1,

JE). | conclude that 4, mevricki{Pig. 2B) isa good

species and hereby resurrect it from the synonymy

of A, badeni (Pig, 2A), Both species are redescribed

here.

Astnreus (Astraeus) williaims/ sp. n0y.

FIGS 1A, 2D

Holotype: &, Peninsular Rd, Lansdowne, N.SW,,

10.45.1987, G, Williams, SAMA 1 28219.

Allutype 9, same data as holotype, SAMA 121220.

Purarypes) NSW. S oc, Peninsular Re,

186 S. BARKER

1mm

Fig. 1, Photomicrographs of male genitalia of the following Astraeus (Astraeus) species: A, Astraeus williams sp.

nov., B. A, aridus sp. nov., C. A. meyricki Blackburn, D, A. occidentalis sp. nov, E, A. badeni van de Poll, F.

A. yarrattensis sp. nov,, and the following Stigmodera (Castiarina) species: G. Stigmodera garnettensis sp. nov.,

H. S. vallisi Deuquet, I. S. wellsae sp. nov.

Lansdowne, 17.xi.1980, 23.x.1980, 30.x.1980,

22.x.1982, 28.x.1988, G. Williams, GWNA.

Colour: Head blue-green or blue. Antennae,

segments: 1-3 coppery; 4-11 blue, Pronotum blue

laterally, purple medially. Elytra dark blue, each

elytron with following yellow markings: basal spot;

pre-medial fascia represented by mark over humeral

fold and spot near suture; post-medial fascia

touching margin but not reaching suture; small pre-

apical spot. Ventral surface dark blue. Legs: femora

dark blue; tibiae mostly dark blue, distal tip

testaceous; Ist tarsomere mostly testaceous, distal

tip blue, 2nd tarsomere mostly testaceous, distal tip

blue, 3-4 tarsomeres blue. Hairs silver.

Shape and sculpture. Head closely punctured,

without median keel, hairy. Pronotum closely

punctured; laterally parallel-sided at base, rounded

posteromedially to apex, hairy. Elytra costate,

intervals flat and smooth each with basal row of

punctures; laterally parallel-sided basally, rounded

posteromedially and tapered to sharp marginal

spine; sutural spine sharp, rounded inner margin,

humeral fold moderately developed, angled (vide

Barker 1975 Fig. 1B). Ventral surface shallowly

punctured, moderately hairy, hairs short,

Size. Males, 7.5 + O18 x 2.9 + 0.08 mm (6).

Females, 7.9 x 3.2 mm (2).

Male genitalia: (Fig. 1A). Parameres parallel-sided

from basal piece, widened medially, tapered,

rounded and tapered to apex, Apophysis of basal

piece broad, rounded apically.

TAXONOMY OF AUSTRALIAN BUPRESTIDAE 187

Fig. 2. Habitus illustrations of the following species: A. Astraeus badeni van de Poll, B. A.

meyricki Blackburn, C. A, aridus sp. nov., D. A. williamsi sp. nov., E. A. occidentalis

sp. nov., FA, yarrattensis sp. nov., G. A. princeps sp, nov., H. Stigmodera wellsae sp. nov.

{88 S BARKER

Remarks; Named after Mr Go Williams, Latis-

downe.

Asiraeus (Astraetis) yarrattensis sp, nov,

FICIS IF, 2

Holomwpe o, VYarratt State Forest, NE Wingham,

N.SAW., TEXLI98L, G. Williams, SAMA 1 2122),

Allolype: @, 10 km SSW Laurieton, N.SW,,

6.xi, 1980, G. Willian, SAMA 1 21222,

Paratvpes: NSW. | oct, same data as holotype,

GWNA3 oo, Yarratt State Forest, NE Wingham,

ZU.xPYBL, 16.01.1982, GC) Willais, GHNAS 1 cr,

Bom Bor State Foresi, SE Grattan, 14,%0,1984, G.

Wilhains, GWNAs TO. Mitwhells* Island, (Factory

Rd), & Taree, 9.xi,1988, G. Willians, GWNA.

Colour, Male, Head blue-green, dark blue basally

with purple reflections. Antennae, segments: I-3

coppery; 4-11 blue, Pronotum dark blue with

purple reflections, blue at lateral and basal margins.

Female. Head and pronotum dark blue, rest as in

male, Elytra dark blue with following yellow

markings On each elytron: large basal spot) pre-

medial fascia covering tumeral fold bul not

reaching suture; post-medial fascia covering margin

but not reaching suture. Ventral surface blue. Legs:

femora bluc; tibiae blue at proximal end, rest

(eslaceous; Ist larsofiere lestaceouss 2-4 tarsomeres

blue, Hairs silver

Shape and xculplure: Wead closely punctured,

glabrous median keel, hairy, Pronoetum closely

punctured; laterally parallel-sided basally, rounded

posteromedially, tapered to apex; median impressed

line from (iiddle to apical margin, laterally hairy.

Elyira costal, intervals flat, basally wrinkled, each

with row of punctations; laterally tapered from base,

rounded posteromedially, tapered to sharp marginal

spine; sharp surural spine, rounded inner margins

humeral fold well developed and angled. Ventral

surface closely punetured, moderately hairy, hairs

medium length,

Sizes Males, 6.9 x 2.9 om (6). Females, 74 x 31

mim (2),

Male genituliar (Fig. We) Parameres angled

outwards fron basal piece, more or less parallel-

sided, rounded to apes Apophysis of basal piece

broad, rounded apically.

Remarks: Named after che wpe localicy.

Astraeus (Astraeus) aridus sp, noy,

FIGS. 1B, 2C

Astrueus badeni: Barker 1975, p 131,

Holowpe o, Puttapa Gap, 21 km 8 Copley, 8.

Aust., Flinders Ra., 21.x.1971, §. Burker, SAMA I

21223,

Allotype: 9, same data as holotype, SAMA | 21224.

Paratypes: §, Aust; 6 co, 8 9 Q, same data as

holotype SAMA; 8 vo, 6 99, Derna Pass,

25.x.1969, N. MeFurlund, SAMA,

Colour Head, antennae, pronotam black with blue

reflections. Elytra black with blue reflections and

following yellow markings: basal spot; fascia

covering humeral fold, not reaching sulure, concave

anteriorly, often represented by two spots; short

post-medial fascia touching margin but not suture,

coneave posteriorly, Ventral surface and legs black

with purple reflections, Hairs silver.

Shape and sculpture’ Head closely punctured, na

median keel, hairy. Pronotum closely punctured:

laterally rounded and narrowed from base to apex;

hairy. Elytra costate, intervals flat, each with row

of punctures; larerally parallel-sided basally,

rounded ‘posteromedially (o marginal spine; both

spines sharp, sutural spine with rounded inner

margin, humeral fold poorly developed, slightly

angled, Ventral surface shallowly punctured,

moderately hairy, hairs medium length.

Size: Males, 8.9 + 0.10 x 3.5 + 0.07 mm (15).

Fenvales, 9.1 4 0.14% 3.6 + 0.07 mm (15).

Male genitalia: (Fig, 1B), Parameres angled

oulwards Irom basal piece, rounded anteromedially

to apex. Apophysis of basal piece medium width,

rounded apically,

Remarks: All specimens were collected on the leaves

and stems of Melaleuca glomerata F, Muell. from

the same locality, Derna Pass is a synonym ol

Pullapa Gap, The specific name is derived from

aridus L.,. dry.

Astraeus (Astracus) badeni yan de Poll

FIGS 16, 2A

Asiraeus badeni van de Poll 1889: RA, 93, 94, pl, 2, Figs

7, 7a, Blackburn 1891; 496; Kerremans 1892: 101; van de

Poll 1892; 67; Blackburn I895: 45, 46; Kervemans 1903:

a8; Carter 1929 282; Obenberger 1930: 365; Barker 1975;

13h.

Asirueus budeni dixjunctus Qbenberger 1928: 204; 1930:

465; Barker 1975: 130,

Astraeus meyricki Blackburn 1890; 1256, 1257; van de

Pall 1892; 67; Kerremans 1892; 101; Kerremans 1903: 148.

Carter 1929; 282; Obenberger 1930; 365; Barker 1975: (31,

IAXONOMY OF AUSTRALIA. BUPRESTDAL 189

Tiper Holotype 9, Gawler, MNHN,

Colour: Head, antennae, pronotum black with blue

and purple reflections. Elytra black wih purple

reflections and following yellow markings on each

elytran: basal spot; pre-tnedial fascia from marain

over humeral callus, not reaching suture, concave

anteriorly; short post-medial fascia from margin not

reaching suture, concave posteriorly: small pre-

apical spot. Ventral! surface black with bronze and

purple reflections. Legs black with blue and purple

reflections. Hairs silver.

Shape and sculpture: Head closely punctured, no

median keel, hairy. Pronotum closely punctured;

laterally parullel-sided basally, rounded, indented

to apex, hairy, Elytra costate, intervals flat each with

row Of punctures; laterally parallel-sided basally,

rounded posteromedially, abruptly tapered to sharp

nrarginal sping; sharp surural sping, rounded inner

nrargin, huniensl Sold moderately developed, angled.

Vential surface shallowly punctured, mauderately

hairy, pairs long,

Size! Males, 7.9 * 3.4 mm (6). Females, 8.4 x 3.6

mim (8).

Mate genitalia: (Fig, 1E). Parameres angled

outwards from basal piece for most of length,

rounded to apex, Apophysis of basal piece medium

Widtl, rounded apically.

Distribution: §. Aust. Gawlet, Morgan. Murray

Bridge, Nuriootpa, Vics Inglewood. W,A,; Marvel

Loch, Southern Cross,

Remarks: A, baeleni differs from congeners in the

bulbous appearance of the pronotum and the more

abruptly tapering apices of the elytra. Ip is a short

broad species. A specimen in the SAMA calleeted

by Tepper at Nuriootpa, S. Aust. has ‘ou Culfitris”

oo the label, the only specimen scen with w recorded

Hust pln,

Astroens (Astracun) mevricki Mackburn

FIGS 1c, 2B

ANeraous Meveiek: Blackburn, 1890: 1256, 1297) 1K95) 45,

Tipe Syntype o, no data, BMNIT syriype er, W,

Auswalia, SAMA,

Colour Head, aitennae, pronutim black with

purple rellections, Blyrra black with following

yellow marks on each elytron: stall basalspot; pre-

medial fascia covering humeral fold, not reachivig

suture; post-medial Mscua not reaching mann ur

suture, Ventral surface black with purple reflections.

Leys dark brown with purple reflections. Halrs

silver.

Shape gnd sculpture; Head closcly punctured,

shallow basal median sulcus, without median keel,

hairy, Pronotum closely punctured, with prominent

basal fovea, median glabrous line from base Lo

middle; laterally rounded from buse in apes, Elytra

costate, intervals flat, each with row of punctures;

Jatcrally tapered from base, rounded pusteromedi-

ally then tapered to sharp marginal spine; sharp

sulural spine, rounded mner margin; humeral fold

very poorly developed, very slightly angled. Ventral

surface shallowly punctured, moderately haicy, Hairs

medium length.

Size; Male, 10.6% 4.5 mim (2).

Male genitalia: (Vig, IC). Paramens angled

outwards from basal piece, rounded te apex.

Apophysis of basal piece medium width, rounded

apically,

Remarks: There is a temale spectinen lodged in the

SAMA possibly belonging to this species which !

identified as A, badeni (Barker 1975 p. 133). It was

collected 18 km SW Three Springs, W.A,, on

Dryandra cirisoides, 8.41.1968, NO Me Farland,

Astraeus (Asiraeus.) princveps so. nov

PIG. 2G

Rololype: 9. Prince of Wales Island, Qid, Feh

1939, SAMA | 21225.

Paratype: Qld: |, same data as holotype SAMA.

Colour; Head black with blue reflections and yellow

frontal spot. Antennae black with blue reflections.

Pronotum black with blue reflections and lnree

lateral yellow murks, Flytra black with blue

teflections and following yellow. marks on each

elytron: broad pre-medial fascia covering margin

but not reaching suture, obliquely angled anteriorly:

broad post-medial fascia covering margin bur nol

teaching suture, Veutral surtace dark blue, Legs red

brown, Hairs silver,

Shape and sculpture: Head shallowly punctured, no

median keel, few shart hairs. Pronotum shallowly

punctured, laterally rounded from base to. apex,

widest medially, few short hairs. Elytra costate,

Intervals Mat, each with row of punctures, surface

wrinkled basally; laterally tapered trom base,

rounded postcromediatly, tapered to sharp marginal

Tt & BARKER

spine; Sharp sutural spine, inner margin straight and

cured upward, with extended marginal im(erval

between spines; humeral fold absent. Ventral surface

shallawly punctured, moderately hairy, hairs

medium length.

Size: Females, 7.9 x 3.4 mm (2).

Remarks: Roth specimens were given jo Mr §.

Watkins by Robert G. Wind, insect supply dealer

and tesident-of Monterey, California. His Australian

wite had been stationed on Prince of Wales [sland

priocto World War Tl and presumably she was the

collector. They are unique to the sub-genus due to

the coloration on the head and pronotum. The

dorsal coloration may have been pink or red in fife

and now faded, The name is derived from princeps.

L., leader.

Astracus (Astracus) occidentalis. sp. wow

FIGS TD, 2b

Hotaiype: cr, 99 km S Overland Roadhouse,

Carnarvon Hway, WiA,, 25.1x,1980, on Casuarina

dielsiany, 5, Barker, SAMA, | 21226

Allotvpe:

24227

G. same dala as holotype, SAMA, |

Paratypes: W.A., 2 ors, 1 G, WAMA, SAMA.

Culow: Head. amennue, pronutum voppery-brornze.

Elytra blaek with blue and purple reflections and

allowing yellow markings on each elytron: basal

spot; pre-medial fascia covering humeral fold,

Louching basal margin bur not reaching suture,

concave anteriorly; medial spot on margin and one

near suture fi some specimens; large post-medial

Spot near margin; pre-upical spot neat suture. Hairs

silver.

Shape and seulptures Head closely punctured, thin

medial imipressec line from base, continued as.short

glabrous median keel af apex. hairy. Propotum

closely punctured; prominent basal fovea projecting

anteriorly for short distance as impressed ling,

anterior murgin projecting medially; laerully

rounded and narrowed trom base to apex, hairy.

klyira costate, intervals fat, each with row af

punctures; laterally parallel-sided basally, rounded

medially and narrowed to small marginal spine;

slurp sulural spine. rounded inner margin; humeral

old poorly developed, slightly angled. Ventrul

surface shallowly puncrured, moderately hairy, hairy

lone.

Size: Males, 11,2 x-4.4 mm (3), Females. 11,8 ¥ 4.6

mm (2).

Male genitalia: (Fig, 1D), Parameres. angled

ounvards from basal piece, shghtly rounded

medially, rounded abruptly to apex. Apophysis of

basal picee broad, rounded apically,

Remarks: The name is derived from occidentalis U..,

western.

Key to the species of Astraeus (sensu stricta)

1. Fleud with median keel... eeeeee CITT

Head without median keel, ...,,----------- 22

2) Hats silver... ee ee eee eee eee 3

Hairs tyellow. 22. 00. eee ee bet

}\. Part or all of anterior ventral surtace Ted-brown

None of anterior ‘Ventral surface red-brown... - 7

4. Gulat prosternum, mesa- und metasternum, coxae

2-3, ubdominal segment 1 red-brown... 1.0.0...

a. A. bekeri Barker

Less of vertical’ surface red browat oe iter od

5, Prosternum, coxae red-brown, ,

hoster ys (eect et On Bett tetie 6

6. Shorter than 7.5 ni, humeral told well developed,

ucutely angled . sda! «IN deur A. fraseriensis Barker

longer than 7.5 mm, humeril fold moderately

developed, aneled..,,......, 24, vbscurus Barket

7. Most or part of lew jestaccous - HEE ela

None of ley testaceous - 54 tt 14

R.. Legs 1-2 festaceous except for onirer fiarin. af femora,

lew 3 lestaceous except for femur, ....-,

erdQesd eae -- cee i eg A. dilufipes vau re Pol}

1 ess of leg than above testacequys.,---.. ----.-- vf]

9. Tibiac, tarsomeres 1-2 restaceauy.. 2. ee

ee OSE aie Al sme Barker

Less of lez than above testaceous. ptt _- Mt

10. Tibiae testaceous, distal tips blue, tarsomere |

testuveaus . AL J artittensis sp. Noy,

Less of Jeg than above lestaceous..-- == il

Il, Distal tips of tibiae tesraceous; tarsomeres l-2

(estaceous, distal tips blue... A. willie sp. nov.

Less Of Teg thin above [ebtaceous eer | J

12. Hips of tibiae and jarsomere { tesiueeouy,..... 13

Tursomere |} testa, gents

ett lp AAAS olm AL notraneeeas/]s Barked

13, Usually shortee than Fo MM, ., 0. ecs sce eens

saat AL pereeus van de Poll

Usually fohget ‘Than 7mm. ......-5,

id, Eluieral fold well developed, aculely angled...

AL enasterst. Macleay

Humeral fold ‘well developed, umyled......--

pee Pict oh samonelli Saunders

- Fumera! fold moderately developed, angled... 16

Tumeral fold poorly developed, slightly anvled-

TAXONOMY O)} AUSTRALIAN BUPRESTIDAE 191

16. Head green and coppery purple or blue-green. . 17

Head black......... da elon t t29Lanucraty 18

17. Head, pronotum green and eoppery purple... ,

peeeesy tet, _. A. intricatus Carter

Head blue- -green, pronotum with medial, cordiform

purple mark, anteriorly green, laterally blue... -.

adi nt tqmpape's -AL blackdownsensis Barker

18, Browd, rounded gpecies Aa bio A, vlobosus Barker

Elongute species............,, 4. watsoni Barker

19. Body blue; elyira with 2 “yellow fasc1d. ii. eee

ejetleg @49 A. “Fruteresihus van de Poll

Body black or coppery-bronze....., 22... --20

20, Body black; elytra with numerous yellow spots. .

axe : oA. crassus van de Poll

Body coppery- bronze: each elytron with yellow fascia

and 4-5 yellow spots. ... .4. occidentalis sp. nov.

21. Elytra with 3 yellow fuscia, red areas. ___-

CCOOP Mme eeisc em Ets Ea eae A, major Blackburn

Elytra with 2 1 yellow Taisen: no red,

i) De Aan od. Mavate his (Tv homEony

22. Body elongate and cylindrical... ... dae Rela apd

Body teardrop shaped... ...... cep eens 24

23, Pronotum conically elevated medially.

600199 mene tea JA. prothoracicus van de Poll

Pronotiltt convex medially, .... 6.04 ‘

we Medes cots et le oe Be A. elongatus van de Poll

24. Sutiral spine with rounded inner margin —~.. 25

Sulural spine with straight inner margin.......37

25. Legs red-brown colour.-.,........, 116 pnorefetete 26

Legs other than red-brown colour... ,, 222.27

26, Elytra marked with spots and faseia ..._,

Sao Rede A, macmillani Barker

Each elytron with 2 > vittae A. vitiatus van de Poll

27. Head, pronotum, legs metallic brown or bronze.

r= voce. Jlavopictus C & G

Head, pronotum, ‘legs other than brown of Brorize

ee ne ...28

28. Fiuimeral told well developed, ‘angled, tiewec dd

Humeral fold moderately or poorly developed. 30

29. Head black or coppery purple; ventral surface

coppery-purple, oo. 6.0645 .-AL adanisi: Barker

Head blue of green; ventral surface blue-green

Tee | bd, geet = ae A. simulator van de Poll

30, Humeral fold moderately developed, angled, ,..3

Humeral fold poorly developed, slightly angled...

A asTL SRA TOS Teen Comes Hd dote ole de

31. Pronoruin laterally rounded from base to apex, , ,

ee eet ey se eee A. crockerae Barker

Pronotum parallel-sided ap base, rounded, indented

10 AON LL. a A. badeni yan de Poll

32. Basal spot touching basal margin of elytron,

Did wreetewa € aD aD ve ety ed A. carnabyt Barker

Basal spot not touching basal marvin of elytron

Aecosm oP ARE T ON oho s eee O60. 33

33, Blytron with 2 spots und 2 fascia or 4 spots and a

fascia’ ....5 54 reat 34

Elytron with 6 gnats ‘and. a fascia, 7 T spots ors spors

34.

Head pmnotum ‘and “ctytra black, '

fL3900R.3.emm wo AL meprick! Blackburn

Head pronotum and elyira plac with Blue and

purple reflections... .... -A. eridus sp. nov.

35. Each elytron with 6 spots and a fascia or & spors

Vit ratte c vol fansoni yon de Poll

Each elytron with 7 ‘spots 1 Peri te DOCer 36

36, Pronotum parallel-sided ram base to ‘middle strongly

rounded and jiarrowed to apex; dorsally conver (i

lateral profile. ..22.....4. oherthuri vay de Poll

Pronulur gradually rounded laterally, narrowed from

base ta apex; dorsally flattened in lateral profite

se Ese on oc ul oa -AL carter( Barker

37, Head with frontal spor... .A. princeps sp. nov,

Head withoul frontal spot,,,....... 00222) 38

38. Head with shallow median suleus.-.,..,,,,,....

elaete sop at ‘ ect guerlingi Barker

Head with deep median SUIEVS no setew essen, oo

Head with basal median sulcus; provotum laterally

inflared, medial oval patch of hexagonal cells.

b dete oe oA, evaneus Kerremans

Head with apical median sulcus; pronotum laterally

rounded, no medial hexagonal cells. . b

caledonteus Fauvel

New locality records

A single female specimen of 4. irregularis van

de Poll was collected by Mr E, E, Adams and myself

at Blackdown Tableland, Qld, oo 17.1.1979 on the

foliage of Casuarina inephioia, This isa common

W.A. species recorded from 5. Aust. by Barker

(1977), In the revision of Astraeus (Barker 1977) t

did not record 4, jumsoni van de Poll outside of

S, Aust, except for a doubtful Queensland record.

Carter (1929) listed the species as occurrilig in the

Blue Mts. | have examined Carter specimens in the

SAMA collection from the Blue Mts and more

recent material collected by Mr S. Watkins on

Mg. 3, SDM. microwraphoof che elarion af Asrraeus

pygroens Van de Poll

192 5S. BARKER

Callitris muelleri at Wentworth Falls and Kings

Tableland in the Blue Mts and confirm them to be

A, jansoni.

Egg of A. pygmaeus van de Poll

A female specimen of Astraeus pygmaeus

captured by me 24 km east of Armidale, N.SW.,

on 19.xii.1989, laid 13 eggs onto the cardboard label

in the vial in which it was placed. These hatched

approximately 28 days later. The hard sculptured

chorion of an egg from which a larvae hatched is

illustrated (Fig. 3). Crowson (1981) indicated that

most beetles have smooth eggs with a soft surface.

A micropyle cannot be distinuished, but a row of

tubercules, each pierced by an aeropyle, is clearly

visible around the margin (Fig. 4). The eggs are oval

with a flat ventral surface. The larvae emerged

through the ventral surface and burrowed directly

into the label. The chorions adhered to the label

and covered each individual gallery. Presumably the

same happens in the field situation. These are the

first eggs of Australian buprestids that I have seen.

Fig. 4. S.E.M. micrograph of aeropyles (arrowed) on

chorion of Asfraeus pygmaeus,

Stigmodera (Castiarina) garnettensis sp. nov.

FIGS 1G,5

Holotype: o, Mt Garnett, Qld, 24.ii.1989, G. Wood,

I 21228 SAMA.

Allotype; 9, Mt Garnett, Qld, 24.11.1989, J.

Hasenpusch, | 21229 SAMA.

Paratypes: Q\d: 1 9, Mt Molloy, 9.ii.1987, G. Wood,

GWQA; 2 99, Kuranda, 6/8.i.1989, J.

Hasenpusch, JHQA; 1 9, Mt Garnett, 20.11.1989,

G. Wood, GWQA; 1 o, Mt Garnett, 21.11.1989, J.

Hasenpusch, JHQA;2 oc, 3 9 9, 22.11.1989, J.

Hasenpusch, JHQA; 5 oo", 2 9 9, same data as

allotype, JHQA; 5 oo, 6 99, Mt Garnett,

21/22.11.1989, G. Wood, GWQA; 10 oo”, same data

as holotype, GWQA; 1 9, Mt Garnett, 26.11.1989,

G. Wood, GWQA.

Colour: Male (Fig. 5A). Head, antennae and

pronotum coppery-purple. Scutellum green with

coppery-purple reflections. Elytra yellow with

following markings either black with coppery-

purple reflections, or dark blue with green

reflections: variable basal mark, continuous with

basal margin, not reaching lateral margins, reduced

to basal margin and extension along suture in one

specimen; post-medial fascia reaching margin;

apical mark. Ventral surface coppery purple. Legs

green with coppery-purple reflections. Hairs silver.

Female (Fig. 5B). Head, antennae and pronotum

green with yellow reflections, occasionally head and

pronotum coppery-bronze. Scutellum green. Elytra

yellow with following dark blue markings: broad

basal fascia; broad post-medial fascia; basal mark.

Ventral surface green with yellow reflections. Legs

blue-green. Hairs silver.

Shape and Sculpture: Head closely punctured,

median sulcus, short muzzle. Antennae, segments:

1-3 obconic; 4-11 toothed. Pronotum closely

punctured, narrow basal fovea extending forwards

to middle as glabrous line, basal notches on each

side more marginal than medial; apical margin

projecting medially, basal margin barely bisinuate;

laterally angled outwards from base, rounded one

third of distance from base, bulbous, tapered to

apex. Scutellum scutiform, punctured, flat. Elytra

punctate-striate, intervals convex, punctured;

laterally angled out from base, rounded at humeral

callus, concave, rounded posteromedially, abruptly

rounded to spineless apex; apices diverging, apical

margin subserrate. Ventral surface with shallow

punctures, edges of abdominal segments glabrous,

moderately hairy, hairs medium length. S, rounded

in both sexes.

Size: Males, 14.2 + 0.15 x 5.8 + 0.07 mm (24).

Females, 15.4 + 0.27 x 6.6 + 0.12 mm (17).

Male genitalia: (Fig. 1G). Short and_ broad.

Parameres angled outwards from basal piece,

rounded posteromedially then parallel-sided,

rounded to apex. Apophysis of basal piece medium

TAXONOMY OF AUSTRALIAN BUPRESTIDAE 193

Fig, 5, Habitus illustrations of the following species: A. Stizinodera gurnettensis sp. nov, male, B. S.

garnetlensis sp. nov. female.

width, tapered to rounded apex. Medium lobe

pointed, sides obtusely angled away.

Remarks: This is a member of the 8. cruenta

Saunders species group as well as S. flava Saunders,

S. ovala Barker, S. pallas Blackburn and S. vallisi

Deuquet. Like S. va//isi the sexes are dimorphic and

can be separated on colour and pattern differences.

In the male genitalia of S. yal/lisi (Fig. 1H) the

apophysis of the basal piece is narrower. The

specific name is derived from the type locality,

Stigmodera (Castiarina) wellsae sp. nov.

FIGS 11, 2H

Holotype: &, Portland, Vic., 21.xi.1944, FE. Smith,

NMVA.

Allotype; 9, 20 km N Moe, Vic., 8.xi.1979, A.

Wells, SAMA | 21230,

Paratypes: Vic 1 co, 1 9, Emerald, 3.xi1.1907,

Jarvis, NMVA, SAMA; 3 oo’, Grampians, Nov.

1950, B Given, NMVA; 4 oo’, Portland, 6.xii.1918,

NMVA; | ov, Ferntree Gully, 30.x.1919, NMVA; |

oO, Portland, NMVA;1 0, | 9, ‘Taruna’, Croydon,

S. HW. Fulton, NMVA; 1 9, Belgrave, 19.xi.1946, F

£. Wilson, NMVA; 1 9, Healsville, Nov. 1916,

NMVA; 19, Monbulk, 12.xi.1902, Jarvis, NMVA;

1 9, Boulora, 4.xii.1954, 4. L. Brown, NMYA; 1

®, locality indecipherable, 14.xi.1903, NMVA; 1 o,

49 9,nodata, NMVA; 4 o'o",5 9 9, Mt Rosea,

Grampians, 15.xi.1950, G, F Gross, SAMA; 1 co,

Hall’s Gap, Grampians, 14.xi.1950, LeSoue/,

SAMA.

Colour; Head and antennae blue with bronze

reflections. Pronotum blue with bronze reflections

medially, yellow laterally. Scutellum blue with

bronze reflections, Elytra yellow with following blue

markings with blue-green reflections: very broad

basal mark only touching margin at humeral callus;

broad post-medial fascia; pre-apical spade-shaped

mark, reduced in holotype and allotype to short

horizontal bar, all marks connected along suture

to apex. Hairs silver.

194 5. BARKER

Shape and Sculpture: Head closely punctured,

median sulcus, short muzzle, Antennae, segments:

1-3 oabconic; 4-11 toothed. Pronotum closely

punctured, narrow basal fovea extending anteriorly

to apical margin as impressed line, shallow fovea

at each basal angle; apical margin projecting

medially, basal margin barely bisinuate; laterally

rounded from base to apex, widest medially,

Sculellum scutiform, few punctures, flat. Elycra

punctate-striate, intervals convex, punctured;

laterally angled out from base, rounded at humeral

callus, Concave, rounded posteromedially and

narrowed to spineless apex; apices hardly diverging,

apical margin rough. Ventral surface with shallow

punctures, edges of abdominal segments glabrous,

moderately hairy, hairs short, S.; male truncate;

female rounded.

Size: Males, 10,1 + 0,18 x» 4.0 4 0.09 mm (19),

Females, 10.5 + O19 x 4.1 4+ 010 mm (16).

Male genitalia: (Fig. (1) Short, Parameres parallel-

sided basally, rounded posteromedially, parallel-

sided, rounded apically. Median lobe sharp, sides

acutely angled away, Apophysis of basal piece

medium-width, tapered, flattened apically.

Remarks; Member of the S. distinguenda Saunders

species group together with S. arida Barker, S.

blackdownensis Barker, S. garrawillae Carter, S-

grata Saunders, S$. militaris Carter, 8. subgrata

Blackburn, 5, supergrata Barker, S, watkinsi Barker,

It is closest to S. warkinsi but differs in having: blue

markings on the dorsal surface whereas those in S,

watkinsi are green, a different distribution; different

male genitalia (Barker 1988; Fig. 1B). Specimens

in the NMVA had been misidentified as 8, grate

Saunders. Named after Dr Alice Wells.

Acknowledgments

I thank the following for their assistance: Miss

Cc. M. H. von Hayek, BMNH; Dr A. Neboiss,

NMVA; Mr G. G. Burns, Mornington; Mr J.

Hasenpusch, Innisfail; Mr 8. Watkins, Caparra; Mr

G, Williams, Lansdowne; Mr G. Wood, Atherton;

Mr P. G. Kempster, Mr C. Miller, Ms H.

Vanderwoude, Dr A, Wells, Mr D, J, Williams,

Department of Zoology, University of Adelaide.

References

Barker, S. (1975) Revision of the Genus Asiraeus

LaPorte & Gory (Coleoptera: Buprestidae). Trans. R.

Soc, 8, Aust. 99, 105-141.

(1977) Asiraeus (Coleoplera: Buprestiducy; A

description of three new species and new lacality

records. (hid, Wl, 11-14,

(1988) Contributions to the taxonomy of

Stigmadera (Castiarina) (Coleoptera: Buprestidae).

Thiel. 112, 133-142.

Brackuurn, T. (1890) Notes on Australian Coleoptera,

with descriptions of new species, part v, Proe, Lina,

Soc. NSW. 4, 1247-1276,

(1891) Notes on Australian Coleoptera, with

descriptions ol new species, Joi, 5, 479-550,

(1895) Further notes on Australian Coleoptera,

with descriptions of new genera and species. Trans, R.

Soc, §, Austr. 19, 27-60.

Carrer, H. J, (1929) A check list of (he Australian

Buprestidae. Aust, Zool. 5, 265-304.

Crowson, R. A. (1981) The biology of the Coleoptera.

(Academic Press, London.)

KeRREMANS, C, (1892) Catalozue synonymigue des

pupeete decrits de 1758 & 1890. Mer. Soc. ent. Belg.

__. (1903) Coleoptera; Servicornia Fam, Buprestidae.

Genera Inseetorum 12, 1-338. (P. Wytsman, Bruxelles.)

OBENBERGER, J. (1928) Opuscula Buprestologica 1.

Beitrage zur Kenntnis der Buprestiden (Col.), Arch,

Narurgesch, 92, 113-224,

(1930) Buprestidae I], Coleoptorum Catalogus

NIT: 213-568. (W, Junk Berlin, Den Haag.)

VAS DF Pont, J. R, He N. (1889) Monographical essay on

the Australian Buprestid genus Astraeus C, et G. Tjschr.

v. Ent. 32, 79-110.

___ (1892) Note sur quelques especies d'Asrraeus, bid,

36, 67-68,

Watt, J. C, (1979) Abbreviations for entomological

collections. N. 2. Zool. 6, 519-520,

TRILOBITES AND THEIR TRAILS IN A BLACK SHALE: EARLY

CAMBRIAN OF THE FLEURIEU PENINSULA, SOUTH AUSTRALIA

BY RICHARD J. F. JENKINS* & PAM HASENOHR

Summary

An Early Cambrian trilobite from the Normanville Group in the Sellicks Hill area, Fleurieu

Peninsula, South Australia is here identified as [vshiniellus briandailyi sp. nov. The trilobites occur

in ‘black shale’ in close stratigraphic proximity to a bedding-plane surface showing trails made by

large trilobites. The environment was below storm-wave base and the traces indicate that the black

shale deposited in water sufficiently well oxygenated to support large metazoans with a presumed

oxidative metabolism. Black coloration in shales may not be taken as a necessary indication of

anoxic bottom waters.

KEY WORDS: new species, conocoryphid trilobite, trace fossils, black shale, Cambrian, South

Australia.

Transactions af the Roval Saetety af 8. Aust, (1989), 113(4) 195-203,

TRILOBITES AND THEIR TRAILS IN A BLACK SHALE:

EARLY CAMBRIAN OF THE FLEURIEU PENINSULA, SOUTH AUSTRALIA

by RICHARD J. F. JENKINS* & PAM HASENOHR

Summary

Jenkins, R. J. b. & HaAsenorr, P, (1989) Trilobites and their trails in a black shale: Early Cambrian of

the Fleuricu Peninsula, South Australia. Trans. R, Soc S. Aust. 113(4), 195-203, 30 November, 1089,

An Early Cambrian rrilobire from the Normanville Graup in the Sellicks Hill area, Fleuricu Peninsula,

South Australia is here identified as /vshiniellus briandailyi sp. nov. The trilobites occur in “black shale’

in close stratigraphic proximity to a beddiig-plane surface showing Lrails made by large rrilobires, ‘Lhe

environment was below storm-wave base and the traces indicate that the black shale depasiled in water

sufficiently well oxywenated to support large metazoans with a presumed oxidative metabolism. Black

coloration in shales may not be taken as a necessary indication of anoxic bottom waters.

Key Worns: new species, conocoryphid trilobite, trace fossils, black shale, Cambrian, South Australia.

Introduction

Jago et al, (1984) reported the first discovery of

trilobite remains in the Early Cambrian Norman-

ville Group on Fleurieu Peninsula, South Australia.

They described two specimens of a conocoryphid

trilobite from the Heatherdale Shale in the Sellicks

Hill area (Pig, 1), bul considered the material so

poorly preserved as to preclude formal description.

While exploring the same site during late 1984, one

* Department of Geology and Geophysics, University of

Adelaide, G.P.O, Box 498, S. Aust, S001,

of us (P.H.) discovered a spectacular trace fossil

made by an arthropod, Further searching revealed

two more body fossil remains of trilobites, This

paper Teports these new discoveries.

Of the new trilobite remains, one shows the

cephalon and anterior thorax and the other is a

fragmentary thorax. They were discovered on

surfaces that had split open to the weather, and the

very incomplete specimen is preserved essentially

as a ferruginised film, The more complete individ-

ual, an external mould, resembles specimen SAM

P24321 described by Jago ef al. (1984) and provides

2/0 Granilas

Hectiierenitalnel

Kangaroo (stand lreup

Kanwiantow Grown

= P| Adelman (Matern

a =

Dey alalliay Maw irriertt

CAMBRIAN

—rT Theat Fauits

——"_ Faults

TAN

MLOMb IES

io 20 40 Ww

FRE

AMI

Fig. 1. Locality thap showing find of trilobites ahd trace fossils at Sellicks Hill and the placement of this discovery

in relation to the framework of the Adelaide Pold Bele and adjacent foreland basin.

196 R, J. &. JENKINS & P. HASENOHR

Fig. 2A-C. /vshiniellus briandailyi nov. sp: A, holotype SAM P24321, with anterior glabella relatively well preserved

and composite mould of hypostome present, x3; B, C, paratype SAM P24844, B, external mould and C, silicon

tubber cast of individual made from mould, both x1.5. D, conocoryphid gen, et sp. indet, SAM P24845, fragment

photographed under a film of water, 1.5.

TRILOBITES OF THE FLEURIFU PENINSULA 17

sufficient! additional morphological information ta

justily formal deseripoion of the taxon.

The arthropod traces in near association with the

body fossil specimens of the trilobites are of especial

interes! as the host rock vepresenis a ‘hlack stale"

facies’ Thus the traces provide an unusual

palaenhiological perspective on this interesting find,

The studied speciens are registered in rhe

Palneontolugy Collection of the South Ausualian

Muscum, Adelaide (SAM).

SYSTEMATICS

Soperfainily CONOCORVYPHACEA Angelin, 1854

Family CONOCORYPHIBAE Angelin, [854

Remarks: Divisions amongst the various genera of

vonocoryphids tend to be ill-defined due to

description ol cranidia alone or because many

remamas of these thin-shelled trilobites are murkedly

distorted. Furthermore, workers have tended to

stress thal the group is palyphiyletic (Westereard

F950, Sdzuy 1961; Hutchinson 1962; Rasetti 1967).

On the basis of the evident positioning of the facial

suture on the brim, the present form is a ‘thc’

conocoryphid (Figs 2 & 4),

The family Conocoryphidae is rare in the Early

Cambrian and much more common globally in the

Middle Cambrian (Koroboy. 1973). Lochman-Balk

& Wilson (1958) considered thac the blind

Conocorypludac and Eodiscidae were characteristic

of eXtracratonic, enxinic biofacies in the Cambrian

uf eastern Nerth America and suggested that the

blind condition wis favoured by some imporcant

ecological factor,

Genus Justinrel/lus Korabov, 1986

Remarks: The better preserved of (he orivinal

specuens (SAM) P2432}) is described as having a

rearwardly directed furrow on the side of rhe

anterior part of the glabella (lage ef af, 1984):

medially this par of the glabella seems 1o have been

entires und undivided. Though the front of the

elabella of the new specimen is crushed, there

appear to be tour pairs of glabellar furrows (Fig.

3); the elabella tapers anteriorly and is somewhat

constricted at abour a (third its length. These

charactenstics arn! other aspects of the cranidium

conform to the diagnosis of the Early Cambrian

venus /vsftinieliay Korobov, 1966, Most Cambrian

conocoryphid genera show three pairs of glabella

furrows {in advance of the occipital groove). The

distance the glabella reaches forwards towatds the

frontal border, the possible presence of a raised pre-

glabella lobe, and the width of the froneal fleld, if

it is developed, are same of (he significant character-

istics important lor dislinguishing conocoryphids.

While Korobov (1966) compared his new genus

with the Middle Cambrian taxon Builiella Mathew,

1885. ir is similar ta the Carly Cambrian Pseudatops

Lake, 1940. Species of Psevdatops may show Tour

pairs of glabella furrows, but the alabella is only

weakly lapered forwards and differs from thar in

Mvshiniellus in encroaching on the frontal border,

Pig. 3. Representation of the elabella wid cence! pan ol

theeranidiany of SAM) 24844 shaw tag posiian ot ihe

four parrs.at ehabetlar Jurpaws, t=. ii udvaitee of phe

ovoipilal furrow. Gee. The riore urterion glubetlar lobes

aic strongly campressed ih an atlerforepasteriu

threanan, Consperv wid) Peni 2B;

Ivshiniotlus brianduilyi sp, sv.

FIGS 2-4

Conovoryphid sp, indet, daee er af, 1984, pp. 208-210,

‘fies, 2o-c,

Etymology: Lonouring the lite DY Brian Raily

(1931-1986) for his contribution to Australian

feology and (caching.

Diagnosis: Brim weakly inflexecd medially, crani-

dium encireted by border furrow and rather evenly

vovered by tine to coarse, pointed granules {spin-

ules) in immature specimens, glabella almese

amooth in more mature exumnples; pleura with

acicular, weakly deflexed terminal spines; eleventh

segment macroplenrous.

Material: Holotype SAM P2432], lacking reariust

parts Jago er af. 1984, liv. 2), by; two paratypes,

SAM 224322, rear thoracic segments and

Tragmentary pyeidtur (lage er wh M984 fie, Bey

SAM P24844, cranidium and parts of 14 choracie

LeAOenNts,

Ayseripiivn Tt is unecessary 16 repear te

desenpions ul (he holotype and second fragmeni

198 kK oIF

hip. 4. Reconsrruetion af eephalon cand miajol part a

Hark of fevdinelus brigndailvi sp. tus, abo 2.9

Himes dimensians OF halorype.

provided in Jago-er af (1984, pp. 209-210). The new

specimen confirms the arcuate shape of the

cephalon before, and slight median inflexion of the

border, as well as indicating that rhe facial suture

is cunlined to the brim. In the holotype, the

granulose brim is poorly preserved and is wider than

might seem obvious in the previously published

Fipures; its posterion is well defined and che wide

border furrow more fincly and sparsely eranulose

than the ceptral parts of the cranidium. The new

remains show a nearly smooth, narraw glabells

tapering forwards. with four pairs of glabells

furrows directed rearwards, The (riagmentary base

of an occipital spine is present in the holurype.

The additional individual indicates at least 14

thoracic segments and confirms that the eleventh

is Tnucropleurous. The thin pleural ridges om the

holotype show a line of 10-1 spinules or pornted

eranules on the an(erior segments and 8-9 such

iramules on the posterior ones; other Cine granules

accur between.

Meusurements: The hulotype, SAM P2434),

indicates an animal caceeding 25 mnvin length. ‘The

distorted cranidiun of SAM P24844 is 50 mm wide

and the maximum width oy ihe glabella Is 8.5 jm:

fourth iboracie segment wboul /45 ain wide,

JENKINS & T HASENOHEK

excluding the spines and with the axis about 70.2)

the width of the segment; animal in excess of 55

mm long,

Comparison: The genus includes two previously

described tormns, /vshiniellus nikolii- ard Iyshiniellus

pululus Korobov, 1966, both from the Aldanian

Stage in Tuva, southeastern Siberia, within the Ezim

Suite on the Ezim River. Both are represented by

distorted cranidia and so any comparison with the

present distorted remains must neecssarily lack

precision, 7, hbrigndai/yi resembles f. ackold in the

shape of the cephalon, save for the distinctive

medial inflexion vf the trontal border: rhe glabella

also seems narrower. 7. pa/w/as differs from both

of the former in us pointed glabella and the

somewhat quadrate shape of the fixed cheeks.

Mr J. G. Gelling has kindly provided

photographs of cranidia and loaned several nearly

complete dorsal exoskeletons of conocoryphid

trilobites collected from the Parara Limestone at

Bunyeroo Gorge in the Flinders Ranges. This

material is of a taxon different ta £ briandailyi as

the cighteenth thoracic segment is inacropleurous

Hot the eleventh, and the medial thoracic segments

have remarkably long and strong pleural spines. The

tather long glabella, which reaches a depressed

horder furrow behind the medially widened border,

has three pairs of incised, backWardly directed

furrows and a weak pair of depressions on the

exireme anterior portion} laterally, the facial sutures

cut off the major part of the brim. En its cephalic

characteristics this form from the Parara Limestone

shows similarities (oO the EBurly Cambrian genus

Arops Emmons, 1844.

Conocoryphid gen. et sp, indet.

Remarks: Vhe additional fragmentary thorax (SAM

P24845) dues not show the eleventh segment

(counted from the most anterior segment preserved)

macropleurous, and the pleural spines are more

deflexed and evidently shorter than in the preceding

form. However, the pleura show the characterise

wide-flat pleural grooves and narrow pleural ridges

of conocoryphids, ‘Plic thoracic segments are about

11.3 times as wide as long, excluding the spines, with

the axis about 0.24 the width of the segment, The

poor preservation tias largely obliterated any surface

detail,

Disarticulated thoracic segments associated with

cranidia of Clendcephius Hawle & Corda, 1947, in

material from the Middle Cambrian of France,

show the pleural terminations narrowed abruptly

and thence produced into a somewhat deflexed,

slender spine, The shape of rhe pleural terminations

in SAM P24845 is markedly similar; This specimen

coukl be a dimorphic individual of 5 briandaily

TRILOBITES OF THE FLEURIEU PENINSULA 19

or even represent a separate conocoryphid taxon,

but basically is too fragmentary for determination,

Age of trilobite finds

The occurrence of Jyshiniellus on the Fleurieu

Peninsula is of potential significance with respect

to the timing of a major stratotectonic development

within the Adelaide Fold Belt, Cambrian rocks of

the Normanville Group below the Heatherdale

Shale are of both basinal and shelf aspect (Daily

1963, 1976; Jago et af. 1986), The trilobites occur

in the upper metnber of the Heatherdale Shale high

in the section of the formation exposed (Jago er al.

1984), At Carrickalinga and more southerly parts

of (he Fleurieu Peninsula the Heatherdale Shale is

ubruptly succeeded by up to ca. 1300.m of curbidites

of the Carrickalinga Mead Formation (sertsw late

of Daily & Milnes 1971, 1972; Milnes 1986), the

basal interval of the thick psammific and/or pelitic

sediments of the Kanmantoo Complex, Sandstones

of subtidal or lidal origin (Backslairs Passage

Formation) succeeding the Carrickalinga Head

Formation record a phase of basin Silhing. Jenkins

(1989) identifies this overall basinal deepening and

shallowing as the ‘Ramindjeri/Jadliaura strarotect-

onic cycle’ and (he organic-rich sediments (Foster

et al 1988) of the Heatherdale Shale may be

considered as The eusiule’ phase Marking the scary

of rapid subsidence associated with renewed litha-

spheri¢ extension,

The likely age of /vefuiel/as lo Siberia is fate

Aldanian (Korobov 1973; Jago ef af 1984) or medial

Early Cambrian, and the comparable genus

Pseudafags also oecurs in the Imid Early Cambrian

of eastern North America syd the British (sles. The

present Finding that 4 brivadaity; and ia

conocoryphid from Bunyeroo Gorge are differed

taxa Weakens che claim of Japo er vf (1986) that

conoeeryphids suggest a votretation between che

Heatherdale Shale and higher parts of the Parant

Litiestone in te Flinders Ranges, though on a

tecloric basis the Onset of deposition of the dark,

nodular offshore facies of the latter formation mity

well have corresponded wilh (he deepening signalled

by the Mearliendale Shale (see Jenks 1989),

Trace Fossils

Ab upper-surface beddina plane at che trilebite

silt shows a relatively well preserved arthnopod (rail

extending over a distance of BO vin, parts of two

other traile probubly made by large trilobites and

& poorly preserved section al! the back-flled burrow

produced by a worn dike creatine (Figs #6), This

surface has heen preserved due lo an unusual

CUCULTES EAN

The arthropods evidently moved over a relatively

fine-grained and soft, muddy substrate with the

ends of their limbs making deep impressions. Had

this surface been subsequently buried by further silr,

the cleavage now present in the rock due to

deformation (Cambro-Ordovician Delamerian

Orogeny) would probably preclude splitting along

the particular bedding surface, However, the traces

were buried by a 3.5 cm thick bed now comprising

intergrown phyllosilicates and uinor tine (— 0.1 mim

diameter), angular quartz grains, which are

commonly amalgamate and show undulose

extinetion, The quartz is probably detrital. XRD

analyses of the rock confirm visual observation that

the bed is inhomogeneous with a (less weathered?)

fraction comprising predominantly illite 2M),

minor muscovite M , and kaolinite-smectite, and

a second (weathered?) fraction including illite arid

Ca montmoriilonite. Iron rich opaques are

concentrated in parts, Mite 2M is consistent wih

a paragenesis involving deep burial and alteration

of possible older mineral phases, The bed is

continuous along strike over an exposure distance

of 60m and thickens slightly lowards the south (up

to 4.5 cm), tn places, obvious eranular components

that are apparently ‘reliets’ of the original

sedimentary parucles grade in sive from ~ 0.5 -

12 mm near the base of the bed to ~ 0.1 mm al

the top,

The granular aggregates of phyllosilicates

posubly represent the tnghly altered components

ol an airtall cuff that wixed with a small amount

of detrital material and formed a slurry which

buried and moulded the trilobite traces. During:

modern Weathering 4 ling of parting, developed

subjacent to the Underlying silistone such that avery

thin veneer of granular material still adheres jo and

completely covers the traces,

The trail traced over 80 em consists of sub

parallel rows of rather closely spaced iniprints of

ambulatory limbs and averages 75 fo 80 aim in

width. The Individwal imprints wend ro be ser

obliquely or en-echelon and the best preserved are

about 1 mm tony. Pils urace is af the form

itlusteated by Seilacher (1955, fig, la, b) and was

evidently made by a perambulating trilobite with

Irs body axis alned in che direction of travel. In

one area il is overlapped by a ericely segmented

trace nade hy & Wolfielike anual, Nearby

exposures iy the Hesiherdale Shale show much

hetier preserved examples of whit is probably this

Same race OCCUTTINg Lh assomation wilh nuiterous

burtows of a relatively large furm of Planolites

Nicholson 1872. The segmented trace is probably

Of endogenic ongin aid well preserved examples

show @ reeqlar back-filling. of large ovate faecal

pellets arranged Irninsversely

200 RK. J.B. JENKINS & P. HASENOHR

Pig. 5. Bedding plane below tull horizon, showing trace lossils made by trilobites, A-C, and structure, Dy artibuted

fo ain unknown worn-like animal. A-A’, trace made by perambulating trilobite with body aligned in direction

of travel B-B', macks made by large trilobile moving, obliquely; C-C', indication of a trilobire travelling relatively

quickly, Seale bar equivalent to 20 em.

pil f

/ f

LIL SE ff

Lip fe v

PLILEEEL SPELL E /) f a — ia ao

aed - Cc 9

p 20 cm % i Fi ; ae ao® d

es LLL SPITS

Fiv, 6. Representation of bedding surface portrayed in Fig. 5 indivaling (he several sets of markings made by trilobites

(A-A’, B-B’, C-C") and another unknown animal (D).

TRELOBITES OF THE FLEUKIFL: PESENSULA alt]

The two other traces show relatively widely

spaced series of imprints interpreted ay the

inipressions Of che tips of the telopodites of

trilobites In the larger trace. the matvidual imprints

ae up to 20 mm Jong and on one side of the

complete trail dre arranged in ratbec Wregular rows

positioned in an eeechelun manner Phis sore of

marking is well expluined by Seilacher (1945S, fig.

le V) and indicates a trilobite moving obliquely

erab-wise, perhaps partly in response lo a eentle side

current. Che tins of the limbs which formed the

racks on elher side al the itace spanned

approximately 23 cm, indicating a quite large

animal, The imprints of the remaining tre are

widely spaced and suggest a smaller animal

travelling relatively quickly with the body aligned

in dhe diteetion of travel. These marks. are tol

especially disanenve (aml wete only seen in cera

Hating) but dre comparible with sont ol the lane

HHobile (rails Occurring in the late Eariy te Middle

Cambri Kangaroo ishind Group of Daily (1956),

waethe berch couse of Kangarog Intima,

Che body fossil rennains of the tilobites desenhed

Jictern occurred tespectively. 1.6m siratigraphically

below (he surface with trilobite trails (SAM P24844)

and #8 cm above (his suclaee (SAM P2484), The

taces suvges! (hat (rilobites were live jnhabitaats

of the horton environment

Patacobjological signifivance

The thinly laniinated argillites of the upper

member ( ~240 nm thick) of the Hewtherdale Strate

are almost cutirely far bedded and certainly

jocomulated at depts below storm wave base (cf,

daeo er df, (986). A single rippled surfidoe observed

riray reflect localised bottom currents. Considering

the thickness of the succeeding Larbidites, che depth

of water in the deposibonal envirgiment was

prabably of the arder of hundreds of retres.

Jago et al. (1984) cite and illustrate apparent

tiuderacks in the Heutherdale Shale Oheir fig. 4)

ws possibly indivating a shallow environment of

depusition bul (hey concede that the eracks may

have formed due to syiaeresis, DAiasiiski &

Walton (1965) illustraie their tigs 12 & 113)

compurdble eracks in turbidites.

Trilotile trails oveur abundantly in parts of the

late Barly (& Midulle Citrbrinn sequence on the

north coasr of Kangaroo Island (Sprisuz 1985; Daily

eral \979). Lxarmples of such rails up bo LU-32 em

in Width gre especially dumerqus in iitervals of

(ese forks deposited in tidal and subtidal facies.

This assoeiahion of t(rdobite wails ii shallow-water

sediments is common world wile duriuig the older

Vulacozpie and constinites the Cruzer facies of

Seilueher (46d). Mis u is apparenc that crilobires

had anordinary oxidative kind of metabolism such

as is characteristic of modern, free living arthronads

and metazoans in general,

The wtthropod tras in the Beylherdl: Stale

suggest relatively larce animals moving over the

bottom in seme numbers. Lhe implication is thar

the bocly fossil remidins of urilabites were part ul

un indigenous papulation and not isolated

individuals somehow transported inte the enviror-

ment by chance. Presuming sn oxidative meta-

bolism for (he (rilobites, and [he worm-like crealures

responsible for various burrowed horizons, the

holtom waters of the Heatherdale Shale

enviroment were clearly not uniformly anoxic and

must have been at lease moderately well oxygenated

al the fines Gf broturbation, Chis comtradiers the

commonly treld opinion that black shales

characterize oxygen starsed basins; fluctuating

oxvgen tensions may be more likely (Marris 1980);

Kuufiiman 1982; Kuispert 1982; Wetzel 1982;

Conway Morris 1985).

By measunney the displacement of a eonveniently

sived plaster model of a trilobite, the soft tissue

mass of the anjmal which made the 23 cm wide trail

tay be estimated at ~ 500 9, With yeferehee to fie.

#6 of Alexander (1979), the oxygen consumption

of wnanimal of this size may have been about 0.045

ml ein hie’ Jor for the whale organism, 23 al

hr . Runnevar (1982) demonstrated that the

surface area or rhe respiratory Organs (eg. gills) of

aqueous animals iy approximately proportonal to

the body mass; this sugeesis a cespiratory exchange

surface of ~S500cm) for the trilobite. Apptivation

of equation 13.5 of Alexander (1979, p. 279)

indicates that the partial pressure between the sea

water and blacd aver the respiratory surface needed

to supply the required OX Veen consupipsiarn is -0.06

atm. Assuming that the respiratory pigment in the

animal's blood was saturated at 0.02 aim. the

(OUI Oxy Ben COnventtation: i the seu Waree

Hecessury far (he tnlobile was - (LOR aim or ~ 38%

PAL (~60 mm He of 0). This vahte lies within

the medial part of the speerrum of oxygen conech-

Hations in the oxygen midtimuto layer of modern

oceans, effectively 2.8% PAT. 10.60% PAL (Barnes

& Hughes (982, fie, 1,9),

There is corvently a renewed wave of speculicion

concerning the possible icrense ol atmospheric

oxyeen during the late Prevambliriad atd ity

relationstip to Me cinergerice Of Ne metacoa ane

higher plant life (Runnegar 1982a, 6 aind spers

comm; Glaessner 1983, 1984), The present exanple

Sugeest that Siniple observation of dark coloration

in parts of Precambrian sequenees (eu, arey anc

blick phyllites in the “Tarrensian’ Series Ot the

Adelaidean) is unlikely 1 give relluble inlocmation

OW ambient wsyeen tenons, The coloration of

402 R. J.P JENKINS & P. HASENOHR

shales Will also reflect diagenetic processes and the

éffects of inapient metamorphisre, for exaniple

fécrystallizatian of Naematite to specularite may

cause beds to become almos! black (Clemimey &

Badham 1982).

! briendaily! is remarkable for the tong genal

spines, the attenuated spines on the pleurites and

iS Macropleurous eleventh segment. Similar long

Spines in adontopleurid trilobites have been

considered to have functioned in helping to support

the animals on ihe sea bottom, either during

‘resting’ ur while they were feeding with the aid of

turrenls passing, het ween che substrare and the lower

side of the thorax (Wittington 19S6a, b: Clarkson

1969, for a different opinion see Bergstrém 4973).

Such an adaptation is consistent with L brianedilyi

inhabiting a soft muddy bottom, Because of the

lack of streamlining brought about by the spines

the trilobite was probably not ati ace swimmer and

this avcards. with the finding of irackways in vlase

proximity to its temains, However, the crackways

indicate larger animals than the body fissil

Specimens and there is no way of telling: whether

the iruees were made by more mature indivaduals

of conaderyphids or orher trilobites kitewn bo reach

large sive locally, such as Kedlichia Cossmann. 1902

(sve Conway Morris & Jenkins 1985), A host of

modern day créatures have adapted to inliaht caves

and other dark places and beeume secondarily

blind, and loss of sight in conoguryphids would

have posed no disadvantage if these primal denizens

of the deep were bottom dwellers who lived below

the effective perelration of light, sustained

primarily on settled detrjpus,

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REDISCOVERY OF PRIMOVULA (P.) HELENEAE CATE AND

DESCRIPTION OF A NEW SPECIES OF CRENAVOLVA FROM SOUTH

AUSTRALIA (MOLLUSCA: GASTROPODA: OVULIDAE)

BY K. L. GOWLETT-HOLMES* & N. J. C. HOLMES*

Summary

The first live collected specimens of the South Australian allied cowrie Primovula (P.) heleneae

Cate, 1973 are recorded. This species, previously known from a single shell collected prior to 1908,

is redescribed, and details of the animal and its habitat given. Crenavolva cruenta sp. nov., 1s

described from Spencer Gulf, $.A.; it most closely resembles C. striatula and C verconis but is

distinguished by its more elongate form, more produced ends and colour pattern.

KEY WORDS: Gastropoda, Ovulidae, South Australia, Primovula heleneae, Crenavolva, new

species, habitat.

Transactions af the Reyal Saciery af S. Aust, (1989), L304) 205-214.

REDISCOVERY OF PRIMOVULA (P) HELENEAE CATE AND DESCRIPTION

OF A NEW SPECIES OF CRENAVOL¥A FROM SOUTH AUSTRALIA

{MOLLUSCA; GASTROPODA: OVULIDAE)

by K. L, GoWLETT-HOLMES*® & N. J.C, HOLMEST

Summary

COW LETT-HOL bs, K, b. & Horns, N. J.C. (1989) Rediscovery of Prinvrovele (Pd helenewe Cate and

description ala new species af Crenuvalva (rom South Australia (Mollisca: Gastropoda: Ovulidae). Trans.

R. Soc. 8, Aust. 1134), 205-214, 30 November, 1989,

The first five collected Specimens of the South Australian ullicd cowrie Prinrcvala (27 feleneae Cave,

1973 wre recorded, This species, previously known from a single shell collected prior to 1908, is redeseribed,

and details of the wnimal and its habitat given, Cren@volve crvenra sp, Nov, is deseribed from Spencer

form, more produced ends and colour pattern,

Key Wonns; Gastropoda, Ovyulidac, South Australia, Primovuly heleneae, Crenavolva, new species, hubiiar,

Introduction

Three species of the allied cowrie family Ovulidac

are recorded from South Australia: Creravolva

verconis (Colton & Goudlrey, 1932), Primovula (P)

heleneae Caie, 1973, and an Ac/yvelva sp. (Coleman

W9R1). All are regarded as very rare, The first two

species are known from single specimens dredged

dead prior to 1908 (Vereo 1908, as Oviilea formosa).

Ol Douglas Point in northern Spencer Gulf,

S. Aust, one of us (NICH) found a number of

allied cowries living on fans of the gorgonian corals

Muplexaura sp. and Mehinugergia sp. in 10-15 m

depth, These specimens were identified as

Primovula (P) helerieae Cate, 1973, (living on

Euplexaura sp), and an undescribed species of

Crenavolya, (living on Echinegergia sp,), Here we

describe the new species of Crenavolva, redescribe

F (PF) heleneae, and include details of the animal

and its habitat.

Materials and Methods

The material reported here is deposited in the

South Australian Museum, Adelaide (SAM),

Australian Museum, Sydney (AM), Museum of

Victoria, Melbourne (NMYV), Northert) Territory

Muscum of Arts & Sciences, Darwin (NTM),

Queensland Museum, Brisbane (QM), Western

Australian Museum, Perth (WAM), Tasmanian

Museum and Art Gallery, Hobart (TM) and

National Museum of New Zealand, Wellington

(NMNZ). Living material was collected by

“SCUBA” and “Hookah” diving and using a Smith-

Melntyre grab dredge. Underwater photographs of

the living animals were taken using Nikonnos IL

and Nikonnos V cameras with macro extension

* South Australian Museum, North Terrace, Adelaide,

S, Ausr. 5000

) Kinhill Mating Scenes, 200 East Terrace, Adelaide,

S. Aust. S000,

tubes and clectronie flash units, Wet material was

preserved in 75% methylated ethanol saludo oe

2% formalin/propylene glycol solution, Radulae

Were prepared for examination under scanning

electron microscope (SEM) afler (he method of

Bandel (1984), Colour descriptions follow Kornerup

& Wansehe: (1978).

Primoyvula (Primovula) helerete Cate, 1973

FIGS I, 3, 4,

Ovule farmoasa; Verws, 1908, p. 342 (non Actans & Reere,

RAB),

Prosuiunta verconis: Couion & Godley, 1932; po a6, pl.

1, fig, TA (in part).

Pellusimriia verconis: Tredale, 1935, f, 105; Coren, TYS9,

p. 368, (in part),

Primovula (Primovula) vercanis: Schilder, 1941, p. 107,

(in part).

Neaspinia (Pellasininia) verconis: Allan, 1956, p. 130,

pl. 14, Ne. 40 (in part),

Primovula (Primovula) helencue Cale, |873a, po 4a—4,

li, B9,

Type: Holotype (SAM 1915943), part of the type

series examined by Cotton & Godfrey (1932); a

paratype of Prosiminia vercenis.

Type locality: Holotype dredged in Gill St Vincent

or Spencer Gull, S. Aust., by J. C. Verco; the

holotype of PB yerconis was dredged of f St Francis

Island, S. Aust, by J C. Vereo.

Other material: SAM D17973 (4 + eggs, i spiril,

SAM DI7975 (2, dry) Douglas Poilit, northeri

Spencer Gulf, S. Aust., N.C, Holmes, 171.1987;

SAM D18434 (13, dry), SAM D18435 (8, in spirit)

Douglas Point, northern Spencer Gull, §. Aust,

N, J.C, Holmes, 5.v, 1988,

Species description: Small allied cowrie, adall length

9.15-13.80 mm, mean 10.77 mm + J.004 Elongare;

both ends slightly produced, posterior end mare so;

widest part of dorsum in posterior 4 (Fig. 3B),

- GOWLETT-HOLMES & N. J. C. HOLMES

Fig. 1. Primovula (P) heleneae, two adults, one with partially extended mantle, on the gorgonian coral Euplexaura sp.

Fig. 2. Crenavolva cruenta sp. n

, two adults with partially extended mantles on the gorgonian coral Echinogorgia sp.

REDISCOVERY OF PRIMORULA (PR) HELENEAE CATE 2U7

Dersim (Fig. 3A) sculptured with numerous

cransverse Wavy striae; funiculurn prominent, with

three weak transverse ridges, Base (Fig. 3B) convave,

outer lip and patietal-columeflar lip Nat and inward

sloping; aperture narrow with a strong fossula, both

sides strongly calloused. Parietal-columellar base

smooth but wih body whorl striae visible beneath

the callus. Outer lip with weak transverse ridges

posteriorly, inner margin slightly dentate

Posteriorly, Outer margin with three to ten slight

fo prominent nodules at posterior end, mare

prominent in smaller shells; usually with one to

three slight nodules at anterior end,

Posterior canal somewhat produced, with a

basally projecting terminal knob dividing canal inte

distinet Jeft and right channels (Pigs 3C, 3D).

Auiterior canal shallow “U-shaped, sliglitly Narng

and jurned to the right, bordered on both sides by

distinct and slightly projecting terminal ridves.

Shell pastel red to pale orange re ercyish yellow,

anterior and posterior tips usually orange; paler

band across widest part of dorsun\s colameliar

vallus, outer Jip and labial callus all paler than

dorsum,

Animal mantle lobes (Fig. Lb with oo papillae;

translucent pastel red to dark orange, with irregular,

reddish brown patches varving in size, area

immediately around each pateti not patterned,

remainder of mantle patterned wih slightly raised,

while dots, Poot pinkish white; anterior dorsal foot

with a series GF prontinent, vertical lamellae;

posterior dorsal foot with a longitudinal, violel

brown (6 brown stripe, Tentacles with while up for

1/6 length, violet brawn for almost half length,

remainder pinkish white with basal 1/5 from eve

swallen, with violet brawn stripe from eve to base.

Siphon and preboscis violet brown to brown,

Radula (Fig. 3F) with central teeity narrow at

base, will large, broad, flal heads, outer edge with

many perpendicular, sharp cusps, central cusp very

large, mairow at base, Widening for about half

length, where there is usually a lateral denticle on

ar least one side, then natrowlnie apically fo sharp

point) with eight or qine small cusps on either side

of central one Firs} lalyral teeth elongate. strongly

curved, heads tricuspidate, cusps Lineven, central

cusp largest. Secand Jateral teeth elongate, with

broad, Nat heads, heads with aboul five long,

curved, sharp-tipped cusps.

Eee mass (Fig, 4A) with 57 colourless,

transparent, jellylike capsules laid on a single

branely of the host sequentially in an elongated

spiral and fused together, Fach capsule contains

minty White-shelled embryos, shells wider than long,

shell lengli 150-230 um, shell round, punctate with

thin operculum (Figs 4B, 4).

Range: The new material examined here is fram

nothern Spencer Gulf, S. Aust. Cate (1973a) pave

the type locality of # (PB) heleneae as the “S. end

ol St Vincent Gull, South Australia, (34°5S'S

138°10'B)", but the holatype is labelled in Verco’s

handwriting "“G: St Vincent or Spencer Gulf,

dredved”, so Cate's (1973a) restriction of the type

locality is suspect, The range of this species is

assumed to be central S. Aust. It is noteworthy thar

Cotton (1935) refers to the occurrence of

Crenavolva verconis in southern W.A., but we can

lind no specimens to substantiate it, Care (1973)

refers Cotton's (1935) reference (0 a Queensland

species, Mrimovula (P) platvsia Cate, 1973 tas Ovula

Jormosa Vereo), but does not mention if le

cxamined the specimens,

Habitat: On fans of (he gorgonian coral &uplewaure

sp, (Caldaria; Octocorallia: Plexauridae), in areas

OF strong tidal movernerit.

Remurks: The specific identity of the host coral

(SAM H355) could pot be determined (PL Alder-

slade pers. comm,), The white spicules (Pig. §C) are

typical of Euplexeuna, ln northern Spencer Gulf,

colonies (Fig. SA) are multibranehed fans In one

plane perpendicular to the strong tidal currents, (ey

ire attached by & thickened base to a large shell

fragment or other piece or pieces of solid debris

in the substrate; a poorly sorted, silty, shelly sand.

Colonies are relatively large (ca 30 cm |ileh}, and

Widely and irregwlarly dispersed at 1-20 m depeh_

Distribution appears to be associated with tidal

current velocity and substrate siltiness, Buplevaneg

SP. appears Lo be less toleraw of silt than Behine

gorgia sp., and is less common. Each Explexmne

sp. colony usually hosts two to five individuals of

P (PF) heleneae. Polyps are usually withdrawn

during daylight on all of the colony regardless wr

the presence of PF (FP) helenewe. Orientation of (he

shell is always with the longit\dinal axis parallel ro

the branches of the host, with po observed

preference fora head up of livad down attitude. Na

particular position on (he host is favoured, and

adults and juveniles are present together, Fags have

been observed in Jate autumn and early winter, laid

on a single branch of the host and pot altered by

the parent, although adults were present on the Lost.

The colour pattern of ihe extended mantle lohes

Of F(R) heleneae is almost identical to that of

Euplexaura sp, The background colour of the coral

is pastel ted, and rhe withdrawn polyps are red to

reddish brown, The pattern of reddish brown

patches an the mantle lobes of PP) heleneae

mimics the position of the polyps on the earal, ane!

an animal with ihe mantle lobes fully extended is

208 K. L. GOWLETT-HOLMES & N. J. C, HOLMES

very Well camouflaged. The very close mimicry by

P. (P) heleneae of its host suggests that it is an

obligate predator and that its distribution will be

limited by that of Euplexaura sp. The distances

between colonies of Buplexaura sp. (10’s of metres)

suggest that adulis of PR (P) heleneae would not

move between colonies, so dispersal is probably by

a free-swimming veliger stage of short duration.

Crenavolva cruenta sp. nov.

FIGS 2, 7.

Holotype; SAM D18431, dry adult specimen

10.45 x 3.60 x 3.00 mm, collected on the gorgonian

coral Echinogorgia sp. at 15 m depth, 1.6 km E. of

Douglas Point, northern Spencer Gulf, S. Aust.,

32°5)'4R"S, 197°49'12"E, by N. J. C. Holmes,

5.v.1988.

v. 3. Primovula (P) heleneae (SAM D17975), A. dorsal view, x 7.5; B. ventral view, x 7.5; C & D. lateral views,

x 7; E. size range, x 2.5 (SAM D18434); F. radula, scale bar = 10 pm (SAM DI7973).

Paratypes: Same locality data as holotype. Spirit

specimens: SAM D18433, 14 juv., 4.65-9.60 mm,

24 ad., 7.35-11.65 mm, Dry specimens: SAM

D18432, six juv., 6.30-9.15 mm, 27 ad., 7.15-12.35

mm. SAM D18437, 11 ad., 6.70-12.60 mm. AM

156094, two ad., 9.55 and 10.90 mm respectively.

NTM P1384, two ad., 945 and 10.20 mm

respectively. NMV F52836, two ad., 9.35 and 9.95

mm respectively. QM MO.19385, two ad., 9.40 and

9.70 mm respectively. TM E17744, two ad., 9.15 and

9.35 mm respectively. WAM 152/88, two ad., 8.30

and 10.30 mm respectively. NMNZ MF. 49494, two

ad., 8.65 and 10.60 mm respectively.

Other material examined: Two broken specimens

in spirit from Douglas Bank northwest, northern

Spencer Gulf, S. Aust., 32°47°18"S, 137°49°12"E,

E. Oks, Sept. 1987 (SAM D1i7976); Backy Point

REDISCOVERY OF PRIMOVULA (P) HELENEAE CATE 209

Fig. 4. Primovula (PR) heleneae (SAM 117973), A. egg mass on branch of Fyplexaura sp., x 4; B. embryonic shell,

dorsal view; C, embryoni¢ shell, showing aperture and operculum, scale bar = 10 ym,

cast, northern Spencer Gulf, S. Aust., 32°55'00"S,

137°51'24"E, E. Oks, Sept, 1987 (SAM D17977).

Diagnosis; Small allied cowrie, adult length

6,70-12.60 mm, mean 9,51 mm + 1.2. Elongate;

both ends produced. Dorsum sculptured with

numerous transverse wavy striae; funiculum not

prominent; aperture narrow with strong fossula;

Outer lip outer margin with slight to prominent

terminal nodules; both canals bordered by slightly

projecting terminal ridges. Shell greyish orange to

brownish red; tips red to orange; dorsal bands and

patches usually present, white, pale yellow to pale

orange; columellar callus, outer lip and labial callus

yellow to light orange; fossula pinkish white.

Animal mantle lobes translucent pale greyish pink

with dark red spots and small and large white

papillae; foot ruby, underside with median

longitudinal very pale pinkish white Stripe; tentacles

with white tip for 4 length, remainder of tentacle,

siphon and proboscis ruby.

Description of Holotype: (Figs 7A-D) Shell

clongate; both ends produced; widest part of

dorsum in posterior 2/5, Dorsum sculptured with

numerous transverse wavy striae; funiculum not

prominent, smooth. Base somewhat concave, outer

lip Mat, inward sloping; parietal-columellar lip

convex; aperture narrow with a strong fossula, both

sides strongly calloused. Parietal-columellar base

smooth with faint transverse striae present on

central inner part, Outer lip with rounded transverse

ridges, becoming weaker anteriorly; inner margin

smooth, outer margin with four slight nodules at

posterior end, three slight and one prominent

nodule at anterior end.

Anterior canal shallow “U"-shaped, slightly

flaring, bordered on both sides by distinct and

slightly projecting terminal ridges. Posterior canal

somewhat produced, shallow “U-shaped, strongly

flaring, bordered on both sides by distinct and

slightly projecting terminal ridges, outer lip ridge

terminating in a nodule, parietal-columellar lip

ridge with two nodules, one terminal,

Shell orange red, anterior and posterior tips

bright red; light yellow median band across widest

part of dorsum, continuing onto base; light yellow

diffuse band midway between median band and

posterior end, continuing onto base; light yellow

patch almost midway between median band and

anterior end. Outer lip and labial callus yellow:

columellar callus translucent yellow with dorsal

colour pattern visible. Interior of posterior and

anterior canals red, becoming orange red at very

tips. Fossula pinkish white.

210 K. L. GOWLETT-HOLMES & N. J. C, HOLMES

Fig. §. A. Euplexaure sp. colony in situ, x 0.8; B. Echinogorgia sp. spicules, scale bar = 100 pm (SAM H356); C.

Euplexaura sp. spicules, scale bar = 100 pm (SAM H355).

Etymology; “Cruentus” (L.) meaning bloody, with random, dark red spots, rows and groups of

pertaining to the main colour of the shell and the — small, simple white papillae and occasional much

animal. larger, white, elevated, rounded papillae with wart-

like knobs, mantle more opaque white at base of

Additional Characters from Paratypes: Animal papillae. Foot uniform ruby, underside of foot with

mantle (Fig. 2) lobes translucent pale greyish pink median, longitudinal, very pale pinkish white stripe;

REDISCOVERY OF PRIMOVULA (PR) HELENEAE CATE 211

Fig, 6. Echinogorela sp. colony in situ, x 0.75.

anterior dorsal foot with series of prominent,

vertical lamellae, becoming less distinct posteriorly,

disappearing about '4 length of foot from anterior

end, Tentacles with white tip for 4 length, remain-

der ruby with basal Ys from eye swollen. Siphon

and proboscis ruby,

Radula (Fig. 7F) with central teeth narrow at base,

with large, broad, flat heads, outer edge with many

perpendicular, sharp cusps, central cusp very large,

tapering apically to sharp point, with sharp, lateral

denticle on either side at about half length; with

five or six small cusps on either side of central one.

First lateral teeth elongate, strongly curved, heads

tricuspidate, cusps uneven, central cusp largest.

Second lateral teeth elongate with broad, flat heads,

heads with eight long, curved, sharp-tipped cusps

Fariation: Paratypes differ from the holotype in

size, shell colour and sculpture (Fig. 7E), Shell

colour is from greyish orange to brownish red:

anterior and posterior lips red to orange; dorsal

hands and patches white, pale yellow 10 pale orange,

and yary from very prominent (often forming three

distinct bands) to almost absent, median band being

most conspicuous; columellar callus, outer lip and

labial callus yellow to light orange.

Funiculum varies from smooth to with up to five

weak nodules on inner edge. Outer lip outer margin

nodules vary from very slight with one prominent

nodule at anterior end, to three prominent nodules

at posterior and two prominent nodules at anterior

end. Posterior end usually much more produced in

small specimens, almost rostrate. Posterior canal

parietal-columellar lip ridge occasionally with three

or four nodules.

Range; Northern Spencer Gulf, S. Aust.

FHlabitat; On fans of the gorgonian coral

Echinegorgiu sp, (Cnidaria: Octocorallia:

Paramuriceidae), in areas of strong tidal movement.

Comparison with other species: The new species

belongs in Crenavalva, which is distinguished from

other genera in the Ovulidac by the small, elongate

shell with median angulation, narrow, produced but

truncate ends, dentale, curved aperture, striate

dorsum and ridged outer lip (Cate 1974), C eruenta

most closely resembles C. striatula (Sowerby, 1828)

and C. verconis, the congeners recorded from

Australia, Cate (1973a) records two subspecies from

Australia, C. s. traillii (A, Adams, 1855) and Cs,

tinctiira (Garrard, 1963), both from Queensland. C

cruenta can be distinguished from C. striatula and

C. verconis by its narrower, more elongate form,

more produced ends, the shape of its more elongate

aperture, and shell colour pattern.

Remarks: The specilic identity of the host

gorgonian coral (Eehinogorgia: P, Alderslade pers.

comm. SAM H356) could not be determinded: the

red thorn scale spicules (Fig. 5B) are typical for the

genus. In northern Spencer Gulf, colonies of this

species (Fig. 6) are sparsely branched fans in one

plane, attached by a slightly thickened base to shells,

fragments or pebbles in the substrate: a poorly

sorted, silly, shelly sand. Colonies are relatively

small (ca 15 em high), and appear much more

tolerant of silt (han Euplexaura sp., being widely

212 K. L. GOWLETT-HOLMES & N. J. C. HOLMES

Fig. 7. Crenavolva cruenta sp. nov. holotype (SAM D18431), A. dorsal view, x 6.6; B. ventral view, x 6.6; C & D.

lateral views, x 6.6; E. size and colour pattern range, paratypes, x 2.25 (SAM D18437); F, radula, paratype, scale

bar = 10 pm (SAM D18433).

but patchily distributed in 10-20 m of water, often

in quite high densities (up to 20 colonies/m’).

When C. cruenta is present, each Echinogorgia sp.

colony usually hosts one to three individuals. From

observations made in daylight, the host’s polyps are

usually withdrawn on all or most of the colony

when C. cruenta is present. Orientation of the shell

is always with the longitudinal axis parallel to the

branches of the host, with no observed preference

for a head up or head down attitude. No particular

position on the host is favoured, and adults and

juveniles are present together.

Colour pattern and form of the extended mantle

lobes of C cruenta are almost identical to

Echinogorgia sp. The background colour of the

gorgonian coral is red: the polyps are reddish orange

when withdrawn, yellow when extended. The thorn

scale spicules in Echinogorgia sp. protrude through

the surface of the colony, giving it a rough texture.

These protruding spicules help retain a ragged layer

REDISCOVERY OF PRIMOVULA (P) HELENE AL CATE 213

of mucus and detritus upon the surface: this

nucous layer is white, and patchily masks the red

background colour The pattern of reddivh brown

patches and white papillae an the mantle lobes ol

C. erventa mimics the ragged mucus on the

vorgonian coral, and an animal with the mantle

lobes fully extended is very well camouflaged on

ity hose. The very close mimiery of C. ervente to

Its host suggests that it is an obligate predator and

that its distribution is limited by thar of

Eehinogergia sp. The distances between colonies of

Echinogorgia sp, (often <0.5 m) and the suitability

of C. cruente§ camoullage suggest that the latter

would be able to nove across the substrate between

colonies, so dispersal is probably only partly by a

lree-swimming velier.

Discussion

The generic names used in this paper follow the

sysiem and modifications proposed by Cate (19734,

1973b, 1974). The distinctions between some of the

genera used there are inadequate,

The two species of Ovulidve previously named

from, Aust: Creravolva verconis and Primovuta

(PR) feleneue, are both based an damaged, dead

shells. Vereo (1908) had three specimens und

recorded them as Ovula formosa A. Adams. &

Reeve, 1848, a species described from Borneo, The

deseripvion and photographs of the holotype of O.

Jormosa [= Delonavolya_forniosa (A, Adams &

Reeve, 1848)] in Cate (19734) were compared with

the holotype of CG verconis, with the holotype and

other specimens of P (P) kelencae, and with the

differ sivnificanily from them, D. Jormoasa is bright

purple with vellow tips: the fossula is not prom:

ihent; the whole shell is-much narcower overall, with

a narrower outer lip than either C. crwenta, C

verconiy or F(R) helenewe, and a more pointed

and which lacks the basally projecting terminal

knoh of PR (P) heleriewe

Coton & Godfrey (1932) described Verco’s

specimens as a new species, Prosimnia vercanis,

Their desctiptian obviously was based on all the

specimens, although the holotype is clearly defined

as the shell trom St Francis Island (SAM DI3476,

= DJ0I74). Cate (1973a) re-examined the Lypes of

& verconis, which riow consisted of two specimens,

(one specimen of the original series is missing,

presumed lost). and discovered that they were no!

conspecific. Cate avknowledged the designation by

Cotton & Godfrey (1932) of (he specimen registered

SAM DI3476 (= SAM. DI0I74) (very worn and

faded), as the holotype of Crenavelya verconis

(Cotton & Godfrey, 1932), and used Cottan &

Godfrey’ (1932) paratype (SAM D15943), a worn,

faded and broken specimen, us the holotype of

Primovule (P) helenege. The live collected material

from upper Spencer Gulf was compared with the

types of both species and, although both types are

damaged, the material callected from Euplexaura

sp. was clearly conspecific with P (P) heleneae. and

all of the new material was different from the

designated holotype of GC. verconiy.

According to Coleman (1981), the specimen

illustrated by: Coleman (1975, fig, 262) as Neosimuia

verconis (Cotton & Godfrey, 1932) is actually a

specimen of an unnamed species of Ac/yvolve.

Acknowledgments

We are grateful ta Dr B. R. Wilson for his

considerable assistance and critical comments, and

to Mr PB. Alderslade for identification of the

gorgonian corals, Mt W, Zeidler, Dr -C. Watts, Dr

TE. Wells and un anonymous referee are thanked

for critical comments, We thank Mr R. Thomas,

Mr D. Evans, Mr M. Moore and Ms J. Young, all

of Kinhill Marine Sciences, for assistance in the

held. The photograplis (other than Figs 1, 2, 7, SA,

6) were taken by Mrs J, Forrest and Ms A. Rentrey

(SEM). The colour plates were fiinded by the

Research Fund of die South Australian Museum.

2l4 K. L. GOWLETT-HOLMES & N. J. €, HOLMES

References

ALLAN, J. (1956) “Cowry shells of world seas.” (Georgian

House, Melbourne).

BANDE, K. (1984) The radulae of Caribbean and other

Mesogastropoda and Neogastropoda. Zool. verhandl.

Leiden 214, 1-188, pls 1-22.

Cate, C. N. (1973a) A systematic revision of the recent

cypraeid family Ovulidae (Mollusca: Gastropoda).

Veliger Suppl. 15, 1-116.

(1973b) The Ovulidae: Replacement names for

four ovulid homonyms. Veliger 16(2), 238-239.

____—s«(1974) The Ovulidae: A key to the genera, and

other pertinent notes (Mallusca: Gastropoda). Ibid.

16(3), 307-313.

COLEMAN, N. (1975) “What shell is that?” (Hamlyn,

Sydney)

_ (1981) “What shell

(Lansdowne, Sydney).

is that? 2nd Edition.”

Corton, B. C. (1935) “Combing the southern seas.”

(Rigby, Adelaide).

(1959) “South Australian Mollusca: Archaeo-

gastropoda.” ( S. Aust. Govt Printer, Adelaide).

& Goporrey, F. K. (1932) South Australian shells

(including descriptions of new genera and species). Part

LI. S. Aust. Nat. 13, 35-86.

GARRARD, T. A. (1963) New species of Mollusca from

eastern Australia. J. Malac. Soc. Aust. 1(7), 42-46.

IRLDALE, T. (1935) Australian cowries. Aust. Zoal. 8(2),

96-135.

Kornerup, A. & Wanscier, J. H. (1978) “Methuen

Handbook of Colour. 3rd Edition.” (Eyre Methuen,

London.)

ScHILDER, F. A. (1941) Verwandtschaft und Verbreitung

der Cypraeacea. Archiv Mollusk. 73(2/3), 57-120.

Verco, J. C. (1908) Notes on South Australian marine

Mollusca, with descriptions of new species. Part LX.

Trans. R. Soc. S. Aust, 32, 338-361.

DEVELOPMENTAL BIOLOGY OF THE AUSTRALOPAPUAN HYLID

FROG LITORIA EUCNEMIS (ANURA: HYLIDAE)

BY MARGARET DAVIES*

Summary

The egg mass and tadpole stages of Litoria eucnemis are adapted for a stream-dwelling habit. The

egg mass is compact and adherent whilst the tadpoles have suctorial mouthparts and shallow fins.

The eggs have a large yolk reserve used by the tadpole for nutrition between stages 19 and 26 after

hatching. At 16-24°C, L. eucnemis metamorphoses 74 days after egg laying and 69 days after

hatching.

Trammactions ul the Royal Society af S Aust, (989), TIB4) 2S-20

DEVELOPMENTAL BIOLOGY OF THE AUSTRALOPAPLAN HYLID FROG

LITORTA EUCNEMIS (ANURA: HYLIDAF)

by MARGARET Davies*

Summary

Davirs. M. (1989) Develupmental biptogy ul the Anstralopapuat tiylid thoy Litera eecneniy (AnuTa

Hylidae), Trans. Rk, Soc, S. Ase, 113(4), 215-220, 30 November, (989,

The eve mass and radpole stages. of J oma egenentis are adapted for a stream-dwelling habit. The cog

Mass J compact and adherent whilst the tadpoles have suctorial mouthparts ani shallow fins, The caps

have a large yolk reserve used by the midpole tor nutrition between stages 19 and 26 wfler hatching, At

16-24°C. 1, ehore ms Metamorphoses 74 days alter eye laying and 69 days afer hareliine.

Kiy Wrebs: Liter eucaesns, tadpoles, developmental biology, Joie adayrarons

introduttion

fuloria encnemis (Lonnberg) is disrribured in

rainforest streams act widely-separated localities in

New Guines (Tylec 1968; Zweifel 1980) and on the

Cape York Peninsyla in northern Queensland

(Zweifel 1980, Couger 1986). The species has had

un unstable nomenelatural history in Australia

(Tyler & Watson 1985) and the names /iteria

variously been applied 10 Australian representatives

of the species.

Life history data are confined to 4 comment that

the eggs are dark brown and that nothing appears

distinctive about the tadpole (Menzies 1974).

A pair of £. eucnemis bred in the laboratories

of the Zudlogy Department, University of Adelaide.

The tadpoles were reared to metamorphosis and the

development of the spevics is reperted here

Matevialy and Methods

Specimens of 1. ewcnenris were collecred by M.

Mahony, J. Read, M. Schmidt and T, Haff at

Charmiltin Ck, Qld (17°43 145931‘) in late August

1988 and reached Adelaide on |ix.1988. A pair of

frogs, amplectant upon arrival, Was (ransferred to

a glass aquarium with a pond of water atone end.

The aquarium was muimiained at oem temperature.

Spawn was removed from the pond, and placed

in dechlotinated, acrated jap water in a glass

aquurium 25%25x8 em for 24hr at 30°C. The

aquuzium was then maintained at room temp-

erature, aerated and the walter chiinged daily.

Tadpoles were fed on lightly-boiled lertuve leaves.

The culture was divided in two on 26.48.1988. To give

the ladpoles roweh surfaves on which to attach,

alga-covered stones were introduced to the aquaria

when the spawn hatched,

* Deparoment of 7Goloyy, Universiry af Adelaide, GPO.

Bax 498, Adclaide S. Aust. S001

Samples of embryos and larvae were vollected as

shown in Tuble PF and preserved in Tylor’s fixative

(Tyler 1962),

Measurements of developmental stages were

taken usity vernier cullipers reading, fo 0.05 min.

Hiustrations were made with the gid-of a Wild Ms

stereodissecting microscope and camera lucida.

Developmental stages were classified acyardins:

to Gosner (196()),

Results

Aniplevuss Amplexus was axillary (Fig. 1), The

pair was amplectant upon arrival in the tbordtory

possibly ue response to the reduced aie pressure wi

the cargo hold of the aeraplane (M, J ‘Tyler & M.

Davies unpubl. obs.),

&ggs: Oviposition took place overnight, 2 and

3.ix, 1988 Temperature of the water in which the

egps were laid was 18,5°C, The egy tatss of 843 Guys

was 67.5. mim at its greatest width und compact

Individual jelly capsules were tightly adherent to

each other (Fig. 2). The animal pole was dark brown

and approximately half of each egg on the vegetal

hemisphere was white. Only one membrane was

detected in each capstle:

Mean diameter of six eggs was 2.25 mm (range

2,10-2.56 mm) and of the capsules, 2,98 nim (range

2.40-3,60 mm). On the day following oviposition,

the eges were in lare cleavage (Stave 9), Within 24

hours they hadl progressed to (he neural fold stage

(Stage I4) and within a further 24 hours. the

embryos were at Stuge 17. The cinbryos were

perched on a large chovolate-coloured yolk sae (Fig.

3); larpe crescentic adhesive organs were located

anterodorsally, A prominent stomoducal pic was

loculed posteradorsally io rhe adhesive organ (lig

3),

On Tax. d9RK, rhe embryos remiined in the jelly

capsules. The adhesive Organs were beginning: to

divide ag head flexion conuneneed, The storbodaent

netch was prominent and jhe pronephiros wus

apparent laterally; the will plate was vleurty

216

delineated. The tail was differentiated and the

embryos were at Stage 18.

Larvae; The embryos hatched on 81x.1988 al

Stage 19. The head was flexed ventrally and the

adhesive organs situated ventrolaterally as two

raised pigmented suckers. A stomodaeal depression

was dorsomedial to the suckers, The primordia of

two pairs of gill filaments were present and eye

bulges were apparent (Fig. 4a). The pronephros was

M, DAVIFS

still apparent. A large yolk sac provided nutrition

for the embryo.

By 9.1x.1989 the embryos were at Stage 20. The

two pairs of gills were now filamentous, the anterior

pair with four to five branches, the posterior pair

with one to three, Ventral suckers were ventrolateral

to an open mouth. The optic bulges were well

developed hut opaque. Nutrition was supplied by

a large yolk sac (Fig. 4b),

TABLE |. Dimensions of developmenral stuges of Litoria eucnemis.

Age Slage Body length Total length il

(days, date) A, range in A, range in

parentheses (mim) parentheses (mim)

6 19 312 5.07 6

(8.1x.1989) §2,.96-3.18) (4.95-5.25)

7 20 3.43 6.6 3

(21x. 1989) (3.3-3.3) (6.4-6.8)

8 20 3.47 8.03 3

(10.1x.1989 (3.25-3.9) (7.8-8.2)

y 21/22 3.65 8.68 \2

(LLix A989) (3.4-4.05) (8,0-9,4)

12 a4 10,96 Ms

(15.1x.1989) (9.9-12.0)

14 28 12.3 13

(17.18.1989) (11.0-13,2)

23 26 17.07 10

(27.1x. 1989) (16,1-18.1)

i at 16.8 3

(5.x.1989) (16.7-17,0)

42 30 42 1

(26.*.1989)

42 4 7,92 24.5 5

(26.x.1989) (7.0-8.4) (24,1-25,2)

42 a OAS ate: 2

(26.x. 1989) (8.8-9,35) (26.0-28,5)

42 33 Ya 27,9 !

(26.x.1989)

42 44 9.1 28.9 3

26,.,.1989) (8.7-9_6) (27,0-30,8)

50 a4 B98 27,5 4

CXL TMD) (8.7-9.2) (26.5-24,9)

50 35 985 2k A 2

(A.NL1989) (97-100) (27,5-28,7)

50 36 10,4 33.7 2

(3.81. 1989) (10.3-10,5) (33.5-33.9)

54 6 9.65 30,3 z

(7x1. 1989) (9,3-10.0)) (29,3-31.3)

54 a4 7 29.6 2

(7.x), 1980) (24-101) (2K.()-31.2)

$4 4) Wd 34.6 |

(Txt A9RY)

57 41/42 9.8 20.4 |

(10.81.1989)

38 42 10.6 31.2 |

(xi 989)

6] 3 Hk 22.1 |

(4.x. 1989)

6l 4S 10,5 14.0) |

(14.n1.1989)

46 1.2 - 2

(16.x1.1989)

DEVELOPMENT IN LITORIA EUCNEMIS 7

Fig. 1. Amplectant pair of Litorta eucnemis.

By 10,ix.1988, the larvae remained at Stage 20 bul

olfactory pits were now clearly visible. Larvae had

reached Stage 21 within the next 24 hours. The

cornea was (transparent, pigmentation of the

adhesive organs was fading and the gills were more

filamentous (Fig. 4c). Ridges could be distinguished

on the lateral and posterior margins of the mouth,

foreshadowing the labial teeth and papillae. The tail

had elongated; nutrition was still supplied by yolk.

By 15.ix.1988, the external gills were completely

covered on both sides by the operculum. The

adhesive organs were degerating, although some

pigmentation remained. The horny beak was

keratinized, but the tooth rows had not yet formed.

Nutrition was still provided by yolk and the embryo

was at Stage 24. Some tadpoles were now free

swimming.

hig. 2. Dorsal view of a spawn clump of Litoria eucnemis.

Greatest diameter ix 67.5 mm.

The spiracle had begun formation by 17.ix.1988.

It was situated ventrolaterally and the opening

directed posteriorly. Two upper and the two lower

entire tooth rows were keratinized. The gut was

loosely coiled but yolk-filled, and embryos were at

Stage 25.

Larvae were nol sampled again until 27.ix,1988.

The spiracle had elongated and its orifice was now

directed posteradorsally. Mouthparts were com-

plete, the larval gut had formed and the tadpoles

were feeding on vegetable matter. The development

of the limb buds indicate that the tadpole was at

Stage 26.

eS |

Fig. 3. A) Dorsal and B) anterior views of embryo of Liraria eucnemis at Stage 17, Scale bar represents 1 mm.

218 M. DAVIES

Fig. 4. Litoria eucnemis larvae: A) lateral view at hatching, Stage 19; B) lateral view Stage 20; C) lateral view Stage

21. Scale bar represents 1 mm.

DEVELOPMENT IN. LATORTA EUCNEMIS 214

dt (nea Yl } Duy Ae

Fig. 5, A) Dorsal and 8) lateral views and €) mouth disc of a lacva of Litarid edcnemis ar Stage F1. Seale bur represents

5S mm (A,B) and 1 mm (C).

The next sample of larvae, taken on 5.x.) 988, was

at Stage 27. The body of the larva was covered with

brown pigment dorsally, and was pale ventrally. A

disrupted fine of pigment granules bordered the

dorsal myotomes and further pigment granules were

scattered laterally and on the dorsal fin.

Dimensions of developmental stages of Ly

eucnemis are shown in Table |, A larva at Stage 31

preserved on 26.%.1988 is illustrated in Fig. Sa.b, The

anus was dextral and the spiracle elongate and

sinistral. The dorsal fin way shallow and tapered

to a genily-rounded tip, General body colour was

brown with faint pigment granules on the tail

myotomes and the dorsal fin.

The mouth disc of a larva at Stage 31 is shown

mn Fig. Sc. There are two upper and three lower rows

of labial teeth, Both the second uppec and the first

lower row of teeth have a median gap. There are

two rows of labial papillae forming a wide band

around the lateral and pasterior murgins of the

mouth.

Metamorphosis: By 10.xi 1988, the lett forearm

had protruded in one larva and the first froglet ut

Slage 46 was recorded on 16.x7. 1988, a toral of 74

days after the egws were laid and 69 days afler

hatching. Water temperatures varied throughout the

pertad of development from 16.4 -23.8°C,

Larval behaviour: Larvae frequently avereuated,

mostly beneath the stones within the aquaria or on

the surface of the lettuce. ‘They attached theriselyes

by their suctorial mouthparts to the floor of the

aquaria during water changing,

70 NM

Discrission

Nhe develupment of Litoria enonenus is wrusual

amongst Australian hylds tutilising a lange yotk

reserve and showing relatively lite development ol

the larval ut (the speaes bilehes at stage 19 hurt

the gut docs not form until stage 26, 9-19 dityn utter

hatching), Such development is more representative

of dircet-developing species or those (hit spend a

considerable period of Iheu development within the

cee capsule [as Avartans sphaunecotus (Moure

1961: Anstis }981)]. Amougst Ansiralian species,

this mode ot development is restricted fo

leprodactytid frogs. £. ewcnens is a New Guinea

species that extends its range wito Australia, and irs

relationships lie with the new Guinean hylid fauna,

in particular the momtune species of Liter and

Nictinysies. Such species lay huge unpigmented

eves: tadpoles, where known. exhibit love

adaptations (Menzies 1974; Ther & Davies 1979;

Zweifel 1983). However, complete life bistiory data

gre unknown for these species, The exes of L.

eucnemis ure Jame in Comparison with other

Australian hylids (see Lyler 1989 for review), direetly

related to the made af early development. [Montine

New Guinea Litata and Nyediniystes species alsa

are described as huving hinge eggs (Menzies (974),

The single adherent e2 mass is an adapuon in

Lirarta eucnenis lo 4 orig lifestyle: (he tooth rows

and orher sttwerires are typical of other Australian

hylids (Marin & Watson 1974), but dig suctorial

moulli disg is an adaptation to fast-flowing waters,

ASIES

Amongst Australopapuan amuriins, there ire

several kinds of lofic adaprationts of (He MOUTH cise.

Nhe vstreme claborations exhibited by Aitartd

ndjnols, bo rheocala (Liem W874), L. urhek heed

(Menzies 1974), and Nvetourstes teed (see Lyler

1989 for Plate) consist of a large Suetormil Arai

oveupying the entire ventral preoeular reyian of the

tadpole. Juuducev/us spp have umbrella-shaped hips

hanging vertramedially (Licm & Hosoter 1973)

whilst a2. subvlendulosa (he moutliparts form a

funnel in whieh all labial tooth rows and

piznentation af the homty beak are absent, Oral

dise tubercles have keratinised tips whilst unusual

black filaments sre suspended anteriorly i the

pharyiny (lyler & Anstis 1975). Lhe less elaborate

acaprations occupying & lesser Proportion of the

anleroventral surface of the Gtdpote ane exhibjted

by f encnemnts are shared by L, copland? and L.

theirlana (Tyler ef al, 1983), L. deseenes (Martin &

Watson 197), Afeeistofaris fipmariay Clyler eral

1979) and Aiivopives balbus (Watson & Martin

J97A9).

Orher torrent adapiions include the narrow (ail

tins and the ventrolateral spiricular opening,

Ack rowledgiients

thank Michael Maliony for the provision of the

adult frogs, Michael. Tyler assisted with ulpote

rearing and critically read the manosedpr “This

work was supported by Liniversity of Adelaide

Deparment of Zoology funds,

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Anatralia” 3rd cdi (Reed, ‘Syelney),

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Lira, Po oS. 9TH) A review Of the Liferte tn ony

spemes BOP and a desenpned Of a few spouts it

Citurie Tom serthers Queenshaids Australia (Agora,

Hylidae, Mew Od Muy. (71), 181-168.

—_ & Hosen, We (M74) Frags of the genus

Tundaoss hes vith descriptions of Awe fiw species

jranura! beplodactylidae) Aten, Qa Miia. 1603),

444457,

Martin, ALA. & Wavptoem, Gy be (ISL) Life history as an

aid (oy senerie delineation in Wie lnnly Myhidae, Cuprerer

(UY), FR T9,

Misges bol. (1974) Handbook Gf eonmen New brute

frogs, WAL Leology Tastinite, Handbook not,

Nionar DA. UY6L) Fons of eistern New Soh Wiles.

Bult Am Mus. dat, Host, W234, 14-3R6

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—___()908) Papuan hytid frogs of (Ne genus Lyle feed

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8, Choke Ch AL & Dawes, M, (1989) Repracuertye

Wiolowy ar the fross of the Magehe Creek system,

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— J& Taviks, My, (979%) Kedefiniion und by

evolutionary ongio of the Austalopapnian frog genus

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Foul, IT, V3AS-350.

& Wasson, Gok. (O84) On ihe nomenchiare of

oy hytid tree frog trom Queenston Phan Ae Sec 8.

dase WO84), 19s t94,

Woavvsos, Ch FA MARIN AL AL (1973) Lite besten, larval

morphology and pelavodships of Australian Jepto-

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(983) Two ny hylid tows trany Papow New

Hine anu) a discussion af The Nverariisreny: Aaya

spewies wimpy, di dite, Novildtes 2754, 12h

brops? PViking O'NaTh,

HYBRIDISATION OF LITORIA CHLORIS AND L. XANTHOMERA.

(ANURA: HYLIDAE)

BY GARY R. BEARDSELL

Summary

Hransactions af the Rewil Society aj S$. Aust, (1989), Disa) 221-224.

BRIEF COMMUNICATION

HYBRIDISATION OF LITORIA CHLORIS AND L. NANTHOMERA. (ANURA:HYLIDAB)

A study of intraspecific variation among the three

disjunct populations af Litoria chloris (Boulenger) of

Eastern Australia showed that the northern population

differed in morphology from the olher two populations!.

Iscenzyte electrophoresis demonstrated extensive

Benetic divergence between the northern and southern

populations resulting if the recognition of the Horthern

population as @ tlistinet species, L. xanihomera’.

Farly development and larval nerphology of /. ehloris

have been described! and (he larvae of 4, xanthomera are

identical to those of L. vhloris’.

Here I (est genetic compatibility through a fertile cross

between a captive male L. chloris and a female L.

yonthomera.

Two adult male L. ch/oris were acquired from Finch

Hatton Gorge, Mackay, Queensland in January 1984, and

one adult female of L. xenrhomiera Irom Crystal Falls near

Cairns, Old on 10.xi1.1987, The frogs were housed in an

indoor terrarium. Amplexus occurred on 28.i1.1988.

Embryos were reared in an aquarium with rainwater to

a depth of 24cm. Water temperature was 18-29°C, Larvae

were ed pond algae, commercial fish food and hoiled

lettuce. When they approached metamorphosis they were

transferred to a terrarium, The frogléts were fed on house

flies and flour moths.

Measurements were made using yernier calipers reading

(0.0.1 nim and an ocular micrometer to 0.1 mm. The larval

mouth dise was drawn under a stereoscopic microscope

while the larval and froglet drawings were inked directly

Onto enlarged photographs, that were later treated with

polassium jodide and a sodium thiosulphate fixing bath

to semove the photographic image and iodine stain

respectively’. Developmental stages follow Gosner',

Calling, commenced on. 17.xi.1987 and continued on

Most nights through to mid-January when the female Z.

xanthamera became gravid. Amplexus pceurred on

Pip. |. Developmental stages of a fertile cross between

Litoria chlorts and L, xanihomera, A, Stage 29; B, Stage

34; C, Stage 42. Seale bar = | mim.

28.11.1988 at O800 hr, and the hale continued to call for

one minute while the female produced a soft call, The

amplectant pair was then placed into a smaller breeding

aquarium.

Oviposition occurred from 0630 ro 0700 hr of 29,1), 1988.

Approximately 600 eggs were laid as a fat, jelly-like mass

on the surface of the water; mean ovidiameter = 1.5 mm,

capsule diameter = 4.4 mim, (n = 6). The animal pole

was dark brown, the yegetal white. The fertile eggs began

to sink and adhere to the botiom and sides of the

aquarium in the first hour,

Dimensions of the embryos and larvae are shown in

Table {. Hatching began 1,iii.1988 al stages 20 & 21, The

embryos were dark to mid brown; length! 6.5 mm, Each

embryo had three pairs of external gills with numerous

branches,

On 3 iii. L988 the larvae were at stage 23, The operculum

developed and melanophores appeared along the mid-

dorsal ling, over the tail musculature and as circular

patches over the dorsal surface of the abdomen. The larvae

were pale brown and the coils of the intestines visible

through (he abdominal wall. The larvae reached stage 25

at 1700 hy (4.ii1.1988).

AL stage 29 (Fig, 1A) the larvae had a total length of

15.5 mm and a body length of 5 mm, Most larvae

developed @ lateral curvature of the (ail of varying degrees

of severity between stages 28-30 in weeks 4 & 5. Artis

point there was marked variation in growth: some larvac

artained a total length 2x that of others, As the larvae

approached stages 34-38 between weeks § and 7 the

severity of the fail bend j/ic¢reased in(o an S-shape in many

individuals (Fig. 2), At stage 34 body colour was light

brown wilh a transparent area over the abdominal wall,

Pigmentation was apparent on the dorsal tail (in and tail

musculature (Fig. 18). On day 49 (he largest larva

mMeasurect 52 jam in stage 40 (Vig. tC),

The mouth dise (Fig. 3) of a larva in stage 34 has the

typical hylid. formuta of

hig. 2, Abnocmal tail bend in a hybrid larva al Stage 40

Total length 52 nom.

222

TABLE |, Dimensions of developmental stages of a cross between Litoria chloris and L. xanthomera.

Means (mm) with ranges in parentheses

AGE STAGE SAMPLE EMBRYO DIAMETER

(Hours)

CAPSULE DIAMETER

BODY LENGTH TOTAL LENGTH

0 2 6 15

(1.5-1.6)

15 3 — —

2.75 4 = =

3.75 8 — —

6 10 —~ —

10.75 Wel — =

11.75 ul = =

12.5 12 = =

15 13 — —

(Days)

I 18 5 44

(4.2-4.8)

2 21 6 6.5

(5.9-6.9)

3 23 5 1

(7.0-7.3)

4 25 5 19

(7.7-8.0)

8 26 5 —

37 33 4 =

38 38 4 —

45 40 3 =

51 46 4 —

In many larvae extra papillae with labial teeth appeared

between the upper rows of labial teeth (Fig. 3).

Four froglets completed metamorphosis 20.iv.1988 and

had snout-vent (S-V) lengths of 14.4, 14.6, 14.8 and 15.1

mm. The remaining 52 surviving larvae completed

metamorphosis by 26.iv.1988. The majority of juveniles

metamorphosed with pelvic girdle abnormalities (Fig. 4A)

to varying degrees of severity. Approximately on third of

the juveniles displayed no external sign of skeletal

abnormalities (Fig. 4B). The colour of juveniles was mid-

- 43 10.3

(4.1-4.4) (9.8-10.6)

- 12.1 38.2

(10.2-12.3) (36.7-39.1)

= 134 39.3

(13.1-13.7) (38.9-39.9)

= 22.4 52.1

(21.6-22.7) (50.1-52.8)

= 147 52.1

(14.4-15.1) a

brown with a very fine, diffuse speckling of white

disappearing after one week.

The embryo and larval lifespan extended over a period

of 51-56 days at 18-29°C. The froglets were measured at

monthly intervals after completion of metamorphosis (Fig.

5). Within a few days they developed a white supralabial

stripe disrupted in some individuals and complete in

others, The mid-brown colour changed to a light mustard-

brown within two weeks of metamorphosis. By 5.v.1988

rare

a yn

11 cal

yay. ‘diayodinyul mire

marvacun tormenta

Fig. 3. Hybrid larval mouth dise at Stage 34. Scale bar

= | mm.

Fig. 4. A, Pelvic girdle abnormalities in a three month old

hybrid froglet (S-V 23 mm); B, Normal three month old

hybrid froglet (S-V 27.2 mm).

223

the supralabial stripe developed a green tinge while green

pigmentation developed on the supraobital skin.

The thighs developed the orange of L. xanthomera by

May-mid June, the supralabial stripe became indistinct

concomitantly, with a green tinge over the brown on the

body parts that are green in the adult life. After eight

months all 25 surviving froglets continue to grow

regardless of the presence or absence of pelvic girdle

abnormalities. On 6,xii.1988, the largest fraglet had a

snout-vent length of 37 mm and had developed nuptial

pads.

The life history pattern of the hybrid conforms to the

pattern of Australian hylids",

The larval life span of 51-56 days is in agreement with

the 41 days recorded for L. chloris' and 58 days for L.

infrafrenuta’ considered usual among tropical species

adapted to life in ephemeral waters”.

More importantly, development of a lateral curvature

of the tail in larvae at four weeks manifesting as pelvic

girdle abnormalities in newly metamorphosed froglets

supports the premise of genetic differentiation attained

by L. xunthomera and ils recognition as a separate taxon

from L. chloris*. The development of some apparently

normal froglets suggests that these species are closely

related.

t Bt]

= |

‘ n-s

? 35 n=5

= ns5

z n=6

2 n-5

184t n=10

0 1 2 a 4 5 6

Time from Metamorphosis (months)

Fig. 5. Growth rate of hybrid offspring from a cross

between Litoria chloris and L. xanthomera. (Mean and

Tange; n = sample size).

I thank Mr M. J. Tyler and Dr M. Davies who critically

read the manuscript. Special thanks are due to Richard

Perkins for his expert skills in photography and

introducing me to the technique of bleach-outs and the

science department of Ingle Farm High School for the

use of various equipment and their general interest in the

frogs, Thanks are due to my brother Kym for typing the

manuscript.