VOL. 109,

28 JUNE,

PARTS | & 2

1985

Transactions of the

Royal Society of South

Australia

Contents

Incorporated

Shiel, R. J. & Koste, W. New species and new records of Rotifera (Aschelminthes) from

Australian waters - - - - - - - - -

Greenslade, P. J. M. Some effects of season and geographical aspect on ants

(Hymenoptera: Formicidae) in the Mt Lofty Ranges, South Australia

Spratt, D. M. Spirura aurangabadensis (Ali & Lovekar) (Nematoda: Spiruridae)

from small Dasyuridae (Marsupialia) - : 3 : a :

Barnett, B..& Schwaner, T. D. Growth in captive born tiger snakes (Nofechis ater

serventyi) from Chappell Island: implications for field and laboratory

studies - - - - - - - - - - -

Davies, M. & McDonald, K. R. A redefinition of Uperoleia rugosa (Andersson) (Anura:

Leptodactylidae) - - - - - - - - -

Edmonds, S. J. A new species of Phascolosoma (Sipunculay from Australia -

Tyler, M. J. & Miller, C. A. Surface architecture of the dorsal epidermis in Australian frogs

Long, J. A, A new Cretaceous chimaerid (Pisces: Holocephali) from South

Australia = - - - - - - - - - -

Brief Communications:

Schwaner, T. D., Edwards, A. & Miller, B. Distribution of Lerista terdigitata (Sauria:

Scincidae) in South Australia - - > = : re b

Smith, J. A. Tides of the Onkaparinga estuary, South Australia - - -

Glover, C. J. M. Additions to the fish fauna of South Australia - - - -

Shepherd, S. A., Clarkson, P. S. & Turner, J. A. Studies on southern Australian abalone

(Genus Haliotis) V. Spawning, settlement and early growth of H.

scalaris - - - - - = = 3 : p “

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

TRANSACTIONS OF THE

ROYAL SOCIETY OF SOUTH AUSTRALIA INC.

CONTENTS, VOL. 109, 1985

PARTS | & 2, 28 June

Shiel, R. J. & Koste, W. New species and new records of Rotifera (Aschelminthes) from

Australian waters - - - - a “ : es . :

Greenslade, P. J. M. Some effects of season and geographical aspect on ants

(Hymenoptera: Formicidae) in the Mt Lofty Ranges, South Australia

Spratt, D. M. Spirura aurangabadensis (Ali & Lovekar) (Nematoda: Spiruridae)

from small Dasyuridae (Marsupialia) - - - - - -

Barnett, B. & Schwaner, T. D. Growth in captive born tiger snakes (Nofechis ater

serventyi) from Chappell Island: implications for field and laboratory

studies - - - - - - - - - - -

Davies, M. & McDonald, K. R. A redefinition of Uperaieie mgosa (andersson) (Anura:

Leptodactylidae) - - - . x si 4

Edmonds, S. J. A new species of Phascolosoma (Sipuncula) from Australia - -

Tyler, M. J. & Miller, C. A. Surface architecture of the dorsal epidermis in Australian

frogs - 2 - - = - - 7 = = = 2

Long, J. A. A new Cretaceous chimaerid (Pisces: Holocephali) from South

Australia - - - - 7 si =! a 2 * :

Brief Communications:

Schwaner, T. D., Edwards, A. & Miller, B. Distribution of Lerista terdigitata Sag:

Scincidae) in South Australia — - - - - - -

Smith, J. A. Tides of the Onkaparinga estuary, South Australia - - -

Glover, C J. M. Additions to the fish fauna of South Australia - é - :

Shepherd, S. A., Clarkson, P. S. & Turner, J. A. Studies on southern Australian abalone

(Genus Haliotis) V. Spawning, settlement and early growth of

HI. scataris - - - - - - - - - - -

PARTS 3 & 4, 29 November

Zeidler, W. A new species of crustacean (Syncarida: Anaspidacea: Koonungidae),

from sinkholes and caves in the south-east of South Australia - -

Ludbrook, N. H. Trigonioididae (Mollusca: Bivalvia) from the Cretaceous of Lake Eyre

North, South Australia - - - - - - - +

Jackson, J. E. Larvae and pupae of Lectrides varians Mosely and Leptorussa

darlingtoni (Banks), (Trichoptera: Leptoceridae) - - - -

Wells, A. Four new species of Hydroptilidae (Trichoptera) from the Alligator

Rivers region, Northern Territory - - - - - - -

Davies, M., Mahony, M. & Roberts, J. D. A new species of Uperofeia (Anura:

Leptodactylidae) from the Pilbara Region, Western Australia - -

Lansbury, I. The Australian Naucoridae (Insecta, Hemiptera-Heteroptera) with

description of a new species - - - - - - -

Cann, J. H. & Gostin, V. A. Coastal sedimentary facies and foraminiferal biofacies

of the St Kilda Formation at Port Gawler, South Australia - -

Spratt, D. M. Redescription of two trichurid nematode parasites of vertebrates in

Australia and Papua New Guinea - - - - -

Skinner, S. Australian and New Zealand species of Elachista and Halothrix

(Elachistaceae, Phaeophyta) - - - = “ e A #

McNamara, K. J, The spatangoid echinoid Linthia from the Late Eocene of southern

Australia - - - f 2 2 7 5 e 2. .

Lange, R. T. Spatial distributions of stocking intensity produced by sheepflocks

grazing Australian chenopod shrublands = - - - = -

Brief Communications:

Sadler, T. & Pledge, N. S. The fossil sea urchin Fellaster incisa—an extension of range

Shepherd, S. A. & Hobbs, L. J. Mae a and growth of the blue-throated wrasse Pseudolabrus

tetricus - - - - - - -

Shiel, R. J. & Koste, W. Records of rotifers Seatais on cladocerans from South Australia

Insert to Transactions of the Royal Saciely of South Australia, Vol, 109, Parts 3 & 4, 29 November, 1985

103

109

12]

143

151

161

167

175

177

179

NEW SPECIES AND NEW RECORDS OF ROTIFERA (ASCHELMINTHES)

FROM AUSTRALIAN WATERS

BY RUSSELL J. SHIEL & WALTER KOSTE

Summary

One hundred and sixty-seven taxa are added to the Rotifera recorded from Australian waters,

bringing the total to over 600. New taxa described and figures are: Lepadella patella var. nov.,

Lecane tasmaniensis sp. nov., L. unguitata var. nov., Asplancha brightwelli asymmetrica ssp. nov.,

and Testudinella husseyi sp. nov. In addition, Hexartha oxyuris (Sernov) is redescribed and

refigured and its synonymy discussed.

NEW SPECIES AND NEW RECORDS OF ROTIFERA (ASCHELMINTHES) FROM

AUSTRALIAN WATERS

by RUSSELL J. SHiBL* & WALTER Kosret

Summary

Sebi Ro TM Rost, W (E983) New species md mew records of Rollers (Asehelitiies) Irvin Ausrruiit

wanes. Ces. ke Sue

Se ltrs, WY), LAS, 28 tue, 198s,

One Wiindred ane sikty-suver lasa are added to the Romer recorded trom Australian wuers, (ima

The fot) Lo over HOO, New taxa deserthed and figured are: Lepadetla patella var. nov., Lecane losploniensix

Sp no, 2 ineuiran var nov, Asplanchne brivhtwelli asvnmeiricu ssp. wor, and Testruddineld hussev)

Sp nov. Thaddition, Hevarihra axvuris (Sernoy) is redesecribed yud func amd irs svronvniy Uneussed

Kis WokbS: Retilera, few spuctes. new fecords, Atstralia

Introduction

Althongh knowledpe of the systernaries and

ceology oF Australian Rotifera tas improved

considerably in recent years, i) remains fragmentary,

Preliminary taxonomic works (Shich& Koste 1979,

Kosice & Shiel 1980a, b) stemmed trom a srudy of

the zooplankton of the Murray-Darling basin, from

samples collected alone the Mapela Creek,

Northern territory (Roste (981, Koste & Shiel 1983,

fait ev a, W984) Und in southwest Western Australia

(Brock & Shiel 1983, Koste ev a/, 1983). The rotifer

fuuna oF most of Australia renains wnstudied.

While this ean be attributed in part to a shortage

of srotiferalogists’ rather than of rotifers (ef.

Dumont 1953), there are undoubtedly mare

collectors Of zooplankton in Australia than the

rotiler records indicate, Systematic difficulties

appear to be a major obstacle.

We tioted earher (Koste & Shiel 1980a) that major

systematic works deal with the northern hemisphere

fauna; there is limited information on the Rotifer:

OF the southern continents. Indeed, a global revision

of the Class is desirable, particularly sinee the

recorniion of ecorypic variation. in response to

habitat variability. In the brachionids, for example,

extensive intraspecific variation in size, lorica

morphology or spine lenwth has been demonstrated

im response to habilal temperature. pH, salinity or

predators, (ae. Gilbert 1967, Green 1977, Gilbert

& Stemberger 1984). This raises the diffieulty: of

specHic determination when the extent ol intra

speciic variability is pot Known far most rotiter

genera, and accounts in part for the proliferaion

OF “species” based on very limited material

fo help overcome lasonomic problems we

commenced a comprehensive itlustrated key fo the

* Department ot Botiny, Linversity of Adefidey, Box 498,

PO, Adekinde, S. Aust SOD,

| Ludwig Brill Staisse 5, Qyitkenbrick, D-4570, bederul

Kepubtic of Cherpnerey,

plunktonic monogonont Raritera recorded from

Austialian waters. In view of comples jolter

wsseniblages, particuladly i tropical Ausiralia, we

have found it necessary lo exiend the key to inelude

Racullitively or occasionally plankianie tsa,

including Bdelloides. The first diaenostic keys have

been prepared (Koste.& Shicl in press a,b). bur an

entire revision may fake several years. Here we

upslale the species list and describe new axa thom

our (and other) collections.

In the firsr cheeklist, Shieh & Respe (979)

recorded 331 taxa, iy 73 eenera. A Surther 106 taxa

were listed by Koste & Shiel (9800), One of these,

ciled a5 Filinia holfinann’ (se) (<0. hefinanyy

Koste, 1980). Was arbenrary ir ice Buropeun species.

To the 436 known tid We add here a furrher 167,

colkued trom the literature since 1980, fray

material made available to 1s by others, and trom

our collections. An important contribulion was

made by Berzins (1982), wlio provided a list of

rotifers identifed from volleetions wiade in

Australian waters over a petiod of 30 years. Many

of these are Hirst recards (or the conrinent, and a

Humber al species are new, However several species

described as new appear to be synonyms, arwluers

of preservation of misidentifications. These ale

discussed in the text. Infrasubspecitic tua hitherto

deseribed as for Var, ure deseribed in nation tu

the nomimite ssp. simply to document variabslity

and we notnamed, Moye deluded ecolovical dara

and/or material will facilitate a larer revision,

The format follows that of Kosic & Shiel (ysab)

All records are listed systeamariely followin

Donner (1965) for the Bdelluiden apd Koste (978h)

for the Manugononta, Locality (or nearest! named

feature) and date of collection are viven. Ranpes

or waler quality are eiven where available Published

revords are ciled by author, unpublished reverds by

collector, Only new taxa are figured. TWpes ane

paraiypes are lodged wale the South Suseraljan

Nluseum (SAM).

2 R. J. SHIEL & W. KOSTE

SYSTEMATICS

Order BDELLOIDEA

Family Habrotrochidae

Habrotrocha bidens (Gosse), 1851

Loc: Bombala, N.SW., in tree stump mould;

28.xii1.1949, 8.i1.1950.

Lit: Berzins (1982).

H. cucullata Murray, 1911

Loc: Bombala, N.SW., creek moss; 8.1.1950.

Lit: Berzins (1982).

H. fusca (Bryce), 1894

Loc: Bombala, N.SW., tree stump mould;

28.xi1.1949.

Lit: Berzins (1982).

H. ligula Bryce, 1913

Loc: Bombala, N.S.W., tree stump mould;

28.xi1.1949.

Lit: Berzins (1982).

H. microcephala (Murray), 1906

Loc: Melbourne, Vic.; moss and lichens; 1.i1.1951.

Lit: Berzins (1982).=H. constricta according to

Donner (1965). The latter is known from N.SW.

(Murray 1911).

H. tranquilla Milne, 1916

Loc: Bombala, N.SW., tree stump

28.xii.1949,

Lit: Berzins (1982).

H. tripus (Murray), 1907

Loc: Yarra R., Ivanhoe, Vic., in terrestrial lichens;

11.1951.

Lit: Berzins (1982).

Family Philodinidae Bryce, 1910

Embata hamata (Murray), 1906

Loc: Yarra, R., McMahon’s Creek, Vic.; 19.iii.1976.

Lit: Berzins (1982).

Dissotrocha hertzogi Hauer 1939

Loc: Hunter, R., near Gundy, N.SW.; 11.xi.1953.

Lit: Berzins (1982).

Pleuretra brycei (Weber), 1898

Loc: Geraldton, W.A., moss/humus; undated,

(?vi.1953).

Lit: Berzins (1982).

Macrotrachela aculeata (Milne), 1886

Loc: Bombala, N.SW., tree stump mould;

28.xi1.1949,

Lit: Berzins (1982).

M. angusta (Bryce), 1894

Loc: Bombala, N.SW, tree stump mould;

28.xii.1949.

Lit: Berzins (1982).

mould;

M. concinna (Bryce), 1912

Loc: Geraldton, W.A., moss/humus;

(?vi.1953).

Lit: Berzins (1982).

undated

M. gunningi (Murray), 1911

Loc: Melbourne, Vic., moss/lichens; 1.111951.

Lit: Berzins (1982).

M. nana (Bryce), 1912

Loe: Yarra R., [vanhoe, Vic., moss/lichens; 1.i.1951,

141.1951.

Lit: Berzins (1982).

M. smithi Milne, 1916

Loc: Bombala, N.S.W., moss; 8.1.1950.

Lit: Berzins (1982).

M. speciosa (Murray), 1907

Loc: Melbourne, Vic., moss/lichens; 1.11.1951.

Lit: Berzins (1982).

Mniobia circinata (Murray), 1908

Loc: Melbourne, Vic., moss/lichens; 1.11.1951.

Lit: Berzins (1982).

M. conarus Berzins, 1982

Loc: Heyington, Melbourne, Vic., moss/lichens:

14.13.1951.

Lit: Berzins (1982).

M. lineata Rahm, 1932

Loc: Heyington, and Yarra R., [vanhoe, Vic., moss;

14.14.1951, 14.11.1976.

Lit: Berzins (1982).

M. magna (Plate), 1889

Loc: Bombala, N.SW., moss/lichens; 8.1.1950.

Lit: Berzins (1982).

M. ocypetes Berzins, 1982

Loc: Melbourne, Vic., moss/lichens; 1.111951.

Lit: Berzins (1982).

M. scarlatina (Ehrenberg), 1853

Loc: Melbourne, Vic., moss/lichens; 1.ii1.1951.

Lit: Berzins (1982).

Family Adenitidae Bryce, 1910

Adineta cuneata Milne, 1916

Loc: Bombala, N.SW., tree stump mould;

26.xii.1949,

Lit: Berzins (1982).

Order PLOIMIDA Hudson & Gosse, 1886

Family Epiphanidae Bartos, 1959

Epiphanes macrourus (Barrois & Daday), 1894

Loc: Lake Grace-Kalgoorlie-Norseman area of

southwest W.A.; viii.1978; x.1981.

Lit: Geddes et a/. (1981), Koste et al. (1983).

AUSTRALIANS ROTIPLIRA

Family Brachionidae Wesenbers-Lund, 1889

Brachionus cauddtus personaius Abistrom, 1940

loc: Winthuria Billabong, Magela Ck, ear

Jabiluka, NTs 15.iv.1980,

PEL 5.44, 285°C, 6.2 mel! O,, 23 wSsocm |.

Lil: Koste (L981,

B. fulcatus {. reducrus Kosie & Shiel, 1983

Loe: Winmurra Billabong, near Jabiluka, Magela

Ck, NTs Asti 1980,

Lite Koste & Shich (1983),

B. kastel Shiel, 1983

Loc: Sheepwash Billabong, Yea, Vier 19.1.1982,

PH 7,0-7,7, 24°C, sat. Os, < 50-200 88 em |,

soll magnesium bicarbonate waters, high trans-

pareney (= 1 NTU).

Lit: Shiel (1983),

B pinneenauy Kosie & Shicl, 1983

Loe: Cue, southwest WIA Viih.1978,

hits Geddes et af (1981),

B quadridentatus ancvlognathus (Schimarda), |859

Loe L. Bullenmerri, View 22.iy,.1983,

Colk lJ. Powling.

B yucdridentatus rhenanus (Laulerborn), 1893

Loe: L. Bullenmerri, Vies 22x, 1983

Coll bd. Powling.

B quadridentatus ininor Koste & Shiel, 193

Loe: Billabongs, Magela Ck, NoT 13.vi.1979; Lake

Grace area, southwest WA. sii LORD,

Lit: Koste er al (1983),

Keratella quadrara dispersa (Carlin), 1943

Loc: Yarra R. near Ivanhoe, Vies 14.71.1976,

Lit: Berziis (1982).

Family Euchlanidae Bartos, 1959

E. pvrifarmis Gosse, 185]

Loe. Moorabool R., Ballan, Vies 9vil.1954,

Lit: Berzins (1982),

Diplerchlanis clegaus (Wierzejski), (893

Log: L, Catani, Me Buffalo. Vie. (2. paludosu

Hauer, 1936); 25.11.1955,

Lily Berzins (1982),

D. propatula ingcradacivla (Maer), 1965

Lover Magela Ck, N.Ry 1S,iy L980: Yarra R. east of

Warburton, Vics 19iif.1976.

Lit: Koste (1981), Berzins (1982),

Manfredium endactlonun (Gosse), 1886

Loc: Magela Ck. NI, 1S.iv.1980; Lake Grace-

Norseman area, southwest WAL: viii L978, 4.1981,

Litt Koste (981), Koste ef af. (1983),

hamily Mytilinidae Bartos, 1959

Mytitina acanthophora Hauer, 193%

Loe: Solomon Dam, Lake Moondurrat,

Undated.

Col: P. Hawkins, T. Orr,

M. bisuleata (Lucks), 1912

Loe: Yarra R., Dight’s Falls, Vie 12.i1,1976,

Lit: Berzins (1982).

Old.

Mt ventralis \. longidacivla Wallert, 1965

Loe: Magela Ck near Jabiluka, NoTr (S.)y. 1980.

Lil: Koste (1981),

Lophocharis curvata Berzins, (982

Loc; King Parrot Ck, Kinglake, Vie; 18.4,1953,

hits Berzins (1982). Inadequately figured and

deseribed.. May be a formalin artelact.

7. oxysrernin (Gosse), 185]

Loe: Jagkson’s Creek, Moorabool R.. View 6.111.196),

Ovil 195d,

Lit: Berzins (1982).

Family Trichotrlidue Bartos, 1959

Trichorria tetractis var. similis (Stenroos), 1898

loc: Mavela Ck near Jabiluka, NJ; 15.iv.1980.

Lit: Koste (1981).

Macrochgetts altamirai (Arevalo), L918

Loc: Riddel’s Ck, Sunbury, Vie 06.11.1961.

Lit Berzins (1982) (syn: M. australiensis Berzins),

M. danneeli Koste & Shiel, 1983

Low Bullitlo Billabong, Magela Ck, N-T; (O.X/,1980,

PH 3.38, 30.9°C, O, 4.15 ppm, 2948 em |.

Lit! Koste & Shiel (1983).

Family Colurellidae Bartos, 1959

Colurella colurus (Ehrenberg), 1830

Loe, Moorabool R., Yarra R., Mt Donna Buang,

Views WWI L9S4, 25.iN,1962, 12,711.1976,

Lit: Berzins (1982),

€. tesselata (Cilaseott), 1983

foc: Coghill’s Ck, Vies (yi.1953,

Lit. Berzins (1982),

©. hicuspidata uncinuta (Ehrenbery), 1832

Loe: widespread, Victoria, perennial.

Lits Berzins (1982),

Lepadella acuntinata sexcostata (Bartos), 1955

Loe: Coghill's Cky L. Catani, Mt Buflalo, Vie:

[bvi.)953, 25.41.1955,

Lit: Berzins (1982).

L. ucuiminada seplemvostara Berzins, 1982

Loe: L, Catani, ML Buffalo, View 25.11.1955.

Lit: Berzins (1982),

4 R. J. SHIEL & W. KOSTE

L. amphitropis Harring, 1916

Loc: Barker’s Ck, Castlemaine; Yarra R., Vic.;

21.iv.1953, 19.11.1976.

Lit: Berzins (1982).

L. amphitropis victoriensis Berzins, 1982

Loc: Middle Tarwin R., Vic.; Ol.v.1953.

Lit: Berzins (1982).

L. apsida Harring, 1916

Loc: L. Euramoo, Qld; viii.1978. Magela Ck, N-T.;

15.iv.1980.

Lit: Green (1981), Koste (1981).

L. buangensis Berzins, 1982

Loc: Mt Donna Buang, Vic.; 25.ix.1962.

Lit: Berzins (1982). The species is poorly figured.

The resemblance to L. patella (Muller) was noted

by Berzins (1982 p. I, Fig. 5) and we consider

L. buangensis a synonym.

L. chorea Berzins, 1982

Loc: Hunter R. near Gundy, N.SW. (?11.xi.1953).

Lit: Berzins (1982). Also very poorly figured, and

possibly a formalin artefact.

L. decora Berzins, 1982

Loc: Loddon R., Vic.; 10.vi.1953.

Lit: Berzins (1982).

L. latusinus f. mucronata Koste, 1981

Loc: Magela Ck near Jabiluka, N-T.; 15.iv.1980.

Lit: Koste (1981).

L. lindaui Koste, 1981

Loc: Winmurra Billabong, Magela Ck, NT;

15.iv.1980.

pH 5.44, 28.5°C, 6.2 mg! O,, 23 pS cm I,

Lit: Koste (1981).

L. minorui Koste, 1981

Loc: Leichhardt Billabong, Magela Ck, NT;

15.iv.1980.

pH 5.55, 30.1°C, 5.53 mg 1 O,, 58 BS cm-!.

Lit: Koste (1981).

L. oblonga (Ehrenberg), 1834

Loc: Sheepwash billabong, Yea, Vic.; 17.iv.1976.

Coll: R. J, Shiel.

L. ovalis n.f. Koste, 1981

Loc: Nankeen Billabong, Magela Ck, N.T.;

15.iv.1980.

pH 5.47, 29.9°C, 5.45 mg * 'O,, 44S cm-!.

Lit: Koste (1981).

Lepadella patella patella (Muller), 1786 n. var.

FIG. la, b

Loc: Buffalo billabong, Magela Ck, N-T; 8.xii.1980.

Coll: R. D. Tait.

SAM V3941

In this sample were four Lepadella with a circular

lorica which resembled L. patella (Miller) described

Fig. 1. Lepadella patella (O. F. Miiller) var. nov. a,

ventral, b, transverse section.

by Wulfert (1960) from a sphagnum pool in middle

Germany (see also Koste (1978b): PI. 59, Fig. 2K).

Whereas the latter form has a circular foot-opening,

the N.T. form has a semicircular opening with a

quadrangular termination at the end of the ventral

plate. The tropical form is markedly larger: Wulfert

(1960) states that L. patella has a lorica length of

70 pm, lorica width 62pm and toes 21 pm, whereas

the Buffalo Billabong form is 98m long, 92 ~m

wide, with toes of 24 pm. Both taxa live in acid

biotopes, In the absence of additional information

we document and figure the variant form as an

ecotype, and await more intensive treatment at a

later date.

AUSTRALINS ROTIPERA 5

1. patella biloba (Hauer), 1958

Loe: Yarnup Swamp, near 1, Unicup, southwest

W.A. 8 XL98E.

170°C, 1600 KS em |,

Litt Koste ef al (1983).

1. quadricdrinata (Stentaos), 1898

Loc, Coghil’s Ch, View WW.vi.1953,

Lit: Berzins (1982),

L. quadricarinata sexcarinaty Klemen, 1959

Loe: Covhills Ck, Viey 1.vi.1983.

Lit: Berzins (1982).

1, rotenburgi (Lacks), 1912

Loe: Riddel’s Ck near Sunbury, Vics A.t 6.411.196),

Lit: Berzins (1982),

L. iriha Myers, 1934

Loc: Strathpine pool, Brisbane, Qld: 21.iv.1950,

Litt Berzins (1982).

Helerolepadella hererodacivla Fadeew, 1925

Loe: Magela Ck neat Jabiluka, NoTs 15.iv. 1980

Lilt Koste (l9S8t).

Family Leeanidae Bartos, 1959

L. fHentimonostvlad inopinata t. svarpoda (Hauer),

1929

Low Cianis, N.Old

Colh Ry Hantond (Det. 7.

Fernando).

svaipeda by C. HH.

L. hifured entome Berzins, 982

hoe: Creswick Ck, Clunes, Vies Lhiv.1953.

Litt Berzins (1982),

1, bulla goniata (Marring & Myers), 1926

Loe: Jackson's Ck, Sunbury, 06.11.61; Moorabool

R., Ballan, Vics O.viil9S4,

Litz Berzins (1982),

L, copes (Marring & Myers),

Loc: Yarra R., near Yarraglen, Wes 18.11.1976,

Lite Berzins (1982).

L. decipiens (Murray), 1913

Loe: Cairns, Old.

Coll POA. Tyler (Det. ©. HL Pernandy).

1. clachis Harring & Myers, 1926

how: Mine Valley Billabong near Jabiluka, Not;

13.1979,

Coll Re Shiel.

1. dumellata thalera (Warring & Myers), 1926

Loc: boresidale Lavoon near Perth, W.A.

Coll: J. van Alphen-

1, hinteris australis Bereiis, 1982

Loc: L, Catani, Mt Buffalo, Vier 25.11.1955.

Lily Berzins (1982). Doubtful, poorly figured.

Formalin artelac,

L. lunaris consiricta (Murray), 1913

Loe: R. Murray, Blanchetown, S.A. 7.v,1982.

Coll: R. J. Shiel.

1, fnaris perplexa Alistram, 1938

Loe: Yarra Re, Yarra Glen, Warburton, Vie. 14, 18,

19.111,.1976,

Lit: Berzins (1982).

L. monostvla (Daday), 1897

Loc: L. Furamoo, Qld; viti.d978.

Lit: Green (1981),

L. obtiosa (Murray), 1913

Loe: L. Euramoo, Qld, viii.1978; Coghill’s Ck, Vic:

ILvi1953.

Lit: Green (1981), Berzins (1982),

L. opias (Harring & Myers), 1926

Loe: Yarra R., MeMuhon’s Ck, Vie 19401976,

Lit: Berzins (1982),

L, perpusilla (Hauer), 1929

Loe: Yarra R., near Ivanhoe, View 14.11.1976.

Litt Berzins (1982).

L. pyrtformis (Daday), 1905

loc: Magela Ck near Jabiluka, Nc, 1S.v.L980; Lb.

Evramoo, Bromfield Swamp, Atherton

Tableland, Qld, viii,l978; southern Vie.,

Hhivl9S55, Tivi.l953, 6.n7.1961,

Lit: Koste (1981), Green (1981), Berzins (1982).

7, sculate (Harring & Myers), 1926

Loe: Magela Ck floodplain, N2T, 13.6 1980,

Jat: Kaste (1981),

L. tethis (Warring & Myers), 1926

Loc: L, Euramoo, Atherton Tableland, Qld;

Vili. 1978.

Lit: Green (L981).

L, (MJ) unguitata Fadeew, 1925 var ney.

FIG, 2a, b

Loe: Billabong of Coleman R.. Cape Yorke, Qld.

264.1983,

Coll: B. VY. Timms,

SAM V 3944

A single individual of this species was identified

only from a figure given by Hauer (1938, Fig. 7a, b)

(see also Koste 1978, Fig. 80, Gu-e). The species is

rare, bul seems to be cosmopolitan, Wulfert (1966)

deseribed variable eeotypes fram the Sokoto R.. W,

Africa, and fron Indian waters, The characteristic

anlenor projections of the lorica were not visible

or were absent on the Cape Yorke variant.

Measurements: Ventral lorica length 1042m, ventral

lorica Width 83 am, dorsal lorica length 94 xm,

dorsal lorica width 72 am, toes inel, claws 40 pin,

claws 12 pain,

6 R. J. SHIEL & W. KOSTE

‘

Fig. 2. Lecane (Monostyla) unguitata Fadeew, var. nov. a, ventral, b, dorsal.

Lecane (s. str.) aculeata var. arcula (Harring), 1914

Loc: Eildon Res., Bonnie Doon, Vic.; 10.xii.1962.

Lit: Berzins (1982). (=L. arcula (Harring)).

Lecane aspasia Myers,

Loc: L. Catani, Mt Buffalo, Vic.; 25.11.1955.

Lit: Berzins (1982).

L. boorali Koste & Shiel, 1983

Loc: Meekatharra, W.A.; 20.viii.1978.

Lit: Koste et al. (1983).

L. clara (Bryce), 1892

Loc: Moorabool R., Ballan, Vic.; 9.vii.1954.

Lit: Berzins (1982).

L. curvicornis nitida (Murray), 1913

Loc: Magela Creek floodplain near Jabiluka, N-T.;

15.iv.1980.

Lit: Koste (1981).

L. formosa Harring & Myers, 1926

Loc: Hunter R. near Gundy, N.SW.; 11.xi.1953.

Lit: Berzins (1982).

L. hastata (Murray), 1913

Loc: Lake Grace-Meekatharra area,

W.A.; xi.1981.

Lit: Koste ef al. (1983).

southwest

L. inermis (Bryce), 1892

Loc: Creswick Ck, Moorabool R., Serpentine Ck,

Vic. I.iv.1953, 9.vii.1954, 22.iv.1953.

Lit: Berzins (1982).

L. levistyla (Olofsson), 1917

Loc: Magela Ck floodplain near Jabiluka, N.T.;

15.iv.1980.

Lit: Koste (1981).

L. mira (Murray), 1913

Loc: Strathpine pool, Brisbane; 21.iv.1950.

Lit: Berzins (1982).

L. mylacris Harring & Myers, 1926

Loc: Creswick Ck, Clunes, Vic.; I1.1v.1953.

Lit: Berzins (1982).

L. ploenensis (Voigt), 1902

Loc: Dunwich, Qld; 03.x.1959.

Lit: Russell (1961).(=L. signifera var. ploenensis

(Voigt) after Wiszniewski (1954)).

L. pusilla Harring, 1914

Loc: Bromfield Swamp, Qld; viii.1978., Sunbury,

Vic.; 6.111.196].

Lit: Green (1981), Berzins (1982).

L. pyrrha Harring & Myers, 1926

Loc: Dam, Beaufort, Vic.; 04.xi1.1953.

Lit: Berzins (1982).

AUSTRALIAN ROTIFERA 7

L. rhvtida Harring & Myers, 1926

Loc: Yarra R., MeMahon’s Ck, Vic; 19.ii1.1976.

Lit: Berzins (1982).

L. stichea t. intrasinuata (Olofsson), 1917

Loe: Winmurra billabong, Magela Ck, N.T;

15.04.1980.

Coll: R, D. Tait.

L. subtilis Harring & Myers, 1926

Loc: Winmurra billabong, Magela Ck, NW;

1S.iv,1980,

Colk R. D. Tait.

Lecane tasmaniensis sp. nov.

FIG. 3 a,b

Marerial: 1) contracted females, sample No. 1120,

in formalin.

Holotype: loricate female, sample No. 1120. Coll:

22.1v.1984, R, J. Shiel. SAM V3942.

Paratypes: SAM V3943.

Type locality: Roadside pool, button grass plain

near turnoff to McAuliffe’s Weir on Lake St Clair

road, Tasmania (42°11 'S/142°56'E).

Description: Lorica outline (Fig. 3a) elongate, oval;

anterior dorsal margin straight; triangular cusps or

projections at the external angles extend past ventral

margin; dorsal plate unmarked, wider than ventral

plate; ventral plate with distinct transverse ridge

across first toe segment from which parallel lines

run towards the anterior margin (Fig. 3a); posterior

segment rounded caudally; coxal plates medium,

rounded; first foot joint rectangular, distinct;

second foot joint square; toes less than half body

Vig. 3. Lecane tasmaniensis sp, noy, a, ventral, b, dorsal.

8 R. J. SHIEL & W. KOSTE

length, parallel-sided ending in short claws with

basal spicules.

Measurements (pm): Total length 155-168; dorsal

plate length -115, width -86; ventral plate length

-126, width -79; anterior points -58; length of toes

incl. claw -61; claw 10-12.

Discussion: In habit, the new species resembles L.

acronycha Harring & Myers, 1926 and L.

curvicornis (Murray), 1913; however the

measurements of these taxa, in the same sequence

as given above are: L. acronycha -290; 111-162, -136;

125-182, 95-113; -80; -102; -12 and L. curvicornis

-280; 110-120, 95-112; 130-145, 105-116; -63; -79; 8-13.

L. tasmaniensis is markedly smaller in most

respects, and is further characterized by the

distinctly visible first toe segment under a transvers

ridge, with nearly quadrangular lines above it

(Fig.3 a).

L. tudicola Harring & Myers, 1926

Loc: Dam, Beaufort, Vic.; 4.xi.1953.

Lit: Berzins (1982).

L. venusta Harring & Myers, 1926

Loc: Coghill’s Ck., Vic.; I1.vi.1953.

Lit: Berzins (1982).

Family Proalidae Bartos, 1959

Bryceella voigti Rodewald, 1934

Loc: Bombala, N.SW., moss and tree stump mould;

8.1.1950.

Lit: Berzins (1982).

Proales doliaris (Rousselet), 1895

Loc: Riddel’s Ck, near Sunbury, Vic.; 6.ii1.1961.

Lit: Berzins (1982).

Family Lindiidae Remane, 1933

Lindia annecta Harring & Myers, 1922

Loc: Mt Buffalo, Vic.; 10.xii.1962.

Lit: Berzins (1982).

L. truncata (Jennings), 1894

Loc: Yarnup Swamp, near L. Unicup, southwest

W.A.; 8.x.1981.

Lit: Koste et a/. (1983).

Family Notommatidae Remane, 1933

Monommata maculata Harring & Myers, 1924

Loc: Magela Ck, NP, 14.iv.1980; L. Grace-

Norseman area, southwest W.A.; ix.1981; dam

near Chillagoe, N. Qld, 12.vii.1983.

Lit: Koste (1981), Koste ef a/. (1983), Qld material

coll. B. V. Timms.

Dorystoma caudata (Bilfinger), 1894

Loc: Yarnup Swamp, southwest W.A.; 8.x.1981.

Lit: Koste ef a/. (1983).

Ttura viridis (Stenroos), 1898

Loc: Reservoir, Trentham, Vic.; 5.xii.1984.

Coll: [. J. Powling.

Eosphora anthadis Harring & Myers, 1922

Loc: near L. Logue, Eneabba Rd, southwest W.A.;

30.1x.1981.

Lit: Koste et al. (1983).

FE. thoa Harring & Myers, 1924

Loc: Forestdale Lagoon, southwest W.A.

Coll: J. van Alphen,

E. cf. thoides Wulfert, 1935

Loc: Buffalo Billabong, Magela Creek near

Jabiluka, N.T.; 8.xii.1980.

Lit: Koste & Shiel (1983).

Resticula gelida Harring & Myers, 1922

Loc: Yarnup Swamp, southwest W.A.; 8.x.1981.

Lit: Koste et al. (1983).

Notommata cf. triangulata (Kirkman), 1906

Loc: Magela Creek floodplain near Jabiluka, N.-T.;

15.iv.1980.

Lit: Koste (1981).

Cephalodella apocolea Myers, 1924

Loc: Magela Ck floodplain near Jabiluka, N-T.;

15.iv.1980.

Lit: Koste (1981).

C. auriculata (Muller), 1773

Loc: Creswick Ck, Tarwin R., Vic; iv, v.1953.

Lit: Berzins (1982).

C. euderbyi Wulfert, 1940

Loc: Serpentine Ck, Bears Lagoon, Vic.; 22.iv.1953.

Lit: Berzins (1982).

C. exigua (Gosse), 1886

Loe: Clunes, Gippsland, Mt Buffalo, Vic.; iv, v.1953,

xii.1962.

Lit: Berzins (1982).

C. gracilis (Ehrenberg), 1832

Loc: widespread, Vic.; prob. perennial.

Lit: Berzins (1982).

C. hoodi (Gosse), 1896

Loc: Tarwin R., Vie.; 1, 7.v.1953.

Lit: Berzins (1982). Berzins also recorded C. remanei

Wiszniewski from Castlemaine, Vic. (=C. hoodi.

For synonymy see Koste 1978b p, 351).

C. intuta Myers, 1924

Loc: Magela Ck floodplain near Jabiluka, N.T.;

15.iv.1980.

Lit: Koste (1981).

C. misgurnus Wulfert, 1937

Loc: Magela Ck floodplain near Jabiluka, N.T.;

15.iv.1980.

Lit: Koste (1981).

AUSTRALIAN ROTIPLRA 4

C. megalocephala (Glascotr), 1893

Loe: Wentworth Falls, N.SW. 22.xi,1953.

Lit: Berzins (982),

C. mversi Wiszniewski, 1934

Loc: Scrubhy Ck, Whittlesea, Vics 30.v.195S,

Lit: Berzins (1982).

C. nana Myers, 1924

Loe: Creswick Ck, Clunes, Viex 1liy1953,

Lit: Berzins (1982),

C. parasitica (Jennings), 1894

Loe: Diggers Ck, MI Kosciusko, N.SW.; [Liv L96L.

Lit! Berzins (1982),

C. sterea (Gosse), 1887

Loe: Magela Ck lMoodplain neer Jabiluka, NoT;

[SAV 1980,

Litz Koste (L981),

C. runtiloides Hauer, 1935

Loe: Bromfield Swamp, Qld; viii 978.

Lit: Green (1981),

Family Trichocercidae Remane, 1933

Trichoacerca hidens (Lacks), 1912

Loc: Creswick Ck, Clunes, H.iv,1953; L. Catani, Mt

Bullalo, 10.sii.1962; Yarra R. Warburton, Vic.;

19.11.1976.

Lits Berzins (1982).

Ll exdonta (Hituer), 1938

Loe: Moorabool R., Ballan, Vig; 9vii.1954,

Litt Berzins (1982). (=7. iyers? Hauer, 1939),

L fusiformis Levander, 1894

Loc: Magela Ck Hoodplain near Jabiluka, NT;

WSuiv.1 980.

Lit: Koste (1981),

T. gracilis (Tessin), 1890

Loe: Solomon Dam, Qld. Undated.

Coll Hawkins.

Lf rernis (Gosse), 1887

Loc: Yarra R., Warburton, Vie; 14, 19,i11.1976.

Lit: Berzins (1982),

T inermis (hinder), 104

Loc: Sheepwash Billabonys, Yea, Viej 8.1982,

Colk R. I. Shiel,

LT. insulana Wiauer, 1937

Joe: Magela Ck floodplain near Jabiluka, NJ1.;

HSuiv. L980,

Lit: Koste (1981).

T. intermedia (stenroas), 1898

Loe: Bromticld Swamp, Qld; viti1978.

Lit: Green (1981).

T. macera (Gesse), IS8&6

Loe: L. Catani, Mt Bulfalo, Vies 25,1/,1955;

Gaumboola Stn Dam, Cape Yorke, 12.yit. 1983.

Lit: Berzins (1982), Qld material coll. B. V. Timms.

IT) montana Hauer, 1956

Loc: Winmurra Billabong, Magela Creek, N.T;

15.iV.1980.

Coll: R. D. Tait.

ZT. musculus (Hauer), 1936

Lou; Diggers Ck, Mt Kosciusko, N.S.W., Liv. L961:

Lake Grace-Meekatharra area, southwest WA;

ix, X.1981,

Lis Berzins (1982), Koste e/ al, (1983).

T. ratius cristata Harting, 1913

Loc: Sunbury, Whittlesea, Vie; 6.v.1955, 30.j11,1961.

Lit: Berzins (1982).

7, rulineri (Donner), 1953

Loe: Dam near Chillagoe, Old; 12.vn.1983.

Coll: B. V. Tinims.

T. sulcute (Jennings), 1894

Loe: Avoca R., Gowar East, Vie; 23.iv.1953,

Lit: Berzins (1982),

L uneinata (Voigt), 1902

Loc: Tabrabucea Ck, Barrington, N.SW,; 12.x1,1953.

Lits Berzins (1982).

T. vernalis (Hauer), 1936

Loe: Serpentine Ck, Bears Lagoon, Vic. 22.iv.1953.

Lit: Berzins (1982).

Family Gastropodidae Remune, 1933

Ascomorpha saltans indica Wulfert, 1966

Loc: L, Barracoota, Vic., 20.13.1975,

Lit: Berzins (1982).

Family Synehactidae Remiane, 1933

Syachacta lackowilziana Lucks, 1912

Loc: Moorabool R,, Vie Yi. 954; R. Murray,

Tailem Bend, S.A. 10.%.1983.

Lit: Berzins (1982), S.A. material coll. R. J. Shiel.

Polvarthra major Burekhardt, 1900

Loc: Coliban Res., Kyneton, 06.711.1961; L. Catini,

Mr Buffalo, 25.47.1955.

Lait: Berzins (1982),

P ininor Voigt, 1904

Loc: Magela Ck floodplain near Jabiluka, NI:

IS.iv. L980,

Lit: Koste (1981).

Family Asplanehnidae Harring & Myers, 1926

Asplanchna brightwelli asymmetrica spp. Wov.

FIG. 4a, b

Material: 12 females, formalin preserved, sample

#870; 9 females, sample #100S,

Teonotpe: Wophi, adult female, Fig. 4, sample 4876,

coll. 47.1981 P. Hawkins,

Pararypes: SAM. V3945.

Tipe localiny Solomon Dani, Palm Island, near

Townsville, Qld (1844'S, 14635 (BE).

10 R. J,

SHIEL & W, KOSTE

eT ¢ cerers.

—-

—

_—

]

Fig. 4. Asplanchna brightwelli asymmetrica ssp. nov. A, trophi apical, length 130 m, B, trophi compressed under

coverslip.

Description: Typical saccate illoricate form;

horseshoe-shaped vitellaria belonging to the A.

brightwelli-intermedia-sieboldi group (Koste 1978,

Salt et al. 1978, Gilbert ef al, 1979). Trophi (Fig.

4a, b) asymmetrical, rami (r) with short medial

inner tooth only on left side, rudimentary swelling

on right; right ramus smaller, more tapered and

arched than that of typical form (cf. Fig. 5). In

addition, the right ramus lacks a second uncus (U,

and a lamella (la) before the ramipoint (rp).

Measurements: Comparative measurements/

features of f. typ. are given in parentheses after

those of the n. ssp.: Trophi length: 130 wm (104-156);

apophysis (Fig. 4: ap, sap): equal (equal); fulcrum

(f): equal (equal); inner marginal teeth: 1 small on

left only (2 small symmetrical); lamella before

ramipoints (la): left only (2 symmetrical); rami (r):

conspicuously asymmetrical (symmetrical); uncus

(U,): equal (equal); uncus (U,): left only (2

symmetrical); subterminal rami tooth: equal (left

only).

Discussion: It was not possible in the preserved

material to count the nuclear number of gastric-

and yolk glands, which are used additionally to

verify Asplanchna taxa, however the features of the

Fig. 5. Trophi of Asplanchna brightwelli (Gosse)

(ap=apophysis on the bulla of the ramus; f= fulcrum;

it=inner teeth protruding from rami inner margin;

la=lamella behind ramal points; m=manubrium;

r=ramus; rp=ramipoints; sap=subapophysis;

strt=second rami tooth; u, =first uncus; U,=second

uncus).

AUSTRALIAN ROTIPE RA '

characleristic asymimetrieal trophi in our opinion

distinctly separate this taxon from 4. brivhewelli

(Gosse), which it most resembles.

Asplanchia herricki De Guerne, WR88

Loc: Albert Park, Vic,

Litt Evans (1951),

hamily Dicranophoridae Remane, 1933

Alberta naiclis Roustield, 1886

Loe: Magela Ck floodplain near Jabiluka, Ni.

1S.iv. 1980,

Lik Koste (981).

Aspelta aper (Harring), 1913

Loe! Yarra R., View [41ii.1976,

Lit; Berzins (1982),

AL crrcinaior (Ciosse), 1886

Loe: Plenty R. Morang 8,, Vie 30~.1954,

Lil: Berzins (1982),

A. neboisi Berzins, (982

Loc: Yarra R. near Ivanhac, View 14iih1976.

Litt Berzins (1982),

Balatro calvus Claparede, 1867

Loc: Winmurra Billabong, Magela Ck, NT;

TS.iv. 980.

its Koste (YR1),

Dicranophorus epicharis Warring & Myers, 1928

Loo: Magela Ck foodplaii near Jabiluka, Nef:

1S.iv, 1980,

Litt Koste (1981),

D. halbachi Koste, 198)

Loe: Winmurra Billabong, Magela Ch, NTs

IS.iv 1980,

Lit: Koste (98h).

D. hercules. Wisniewski, 1932

Loc: Mayela Ck floodplain near Jabiluka, N-T.

[S,iv, 1980.

Litk Koste (98h,

2. robustus Harring & Myers, 1928

Loc: Norseman-Kaleoorlie area, southwest WA.

XDORL,

Lit: Koste et af (1983),

Encentriam inium Berzins, 1982

Loe: Bombala, N,SW, Ip inould on a eucalyptus

stump. 28,xii, 1949,

Lit: Berzins (1982). Doubiful species on the basis

of poor figures.

kL. prosdendrus Berzins, 1982

Loe: Bombala, NSW, In mould on a euealyptus

Stump, 28.11.1949.

Lit; Berzins (1982). As above, figures are inadequate

for a new species’ description. Doubtful record,

Wierseisktella velox (Wisniewski), 1932

Loe: Yarnup Swanip, W.A4 8 x, 1981.

Lil: KRoste er al. (1983).

Fumily Testudinellidae Baitos, 1959

Poinpholyx tubulosa Berzins, |Y82

Loc: L, Purumbeie; Vie.; 29.vii.1953.

Lit!’ Berzins (1982). Poorly figured, ?fornialin

artefact. =P contplenata Gosse.

Testudinella ereeni Koste, 198]

hoc; Leichhardt Billabong, Mavel Ck, NT:

IS.AV.1980,

Lit: Koste (1981),

Testudinella husseyi sp. nov.

PIG. €:la, b

Material: \3 tormalin preserved females, sample

AYORN.

Holowpe: 1 adult female, sample #998, coll.

AWLIISS BV. Tins. SAM V3946,

Paraivpes; SAM V 3947,

Tipe locality: Dune lake near Cooktown, Cape

Yorke, North Qld (15°28'S, 145°15'B).

Deseription: Shape of lorica oval; anterior margin

with five tooth-shaped relatively long, keeled

projections, two on either side of a median shorter

tooth, In contrast, 7) patina patina (Hermann),

1783 (see Fig. 6:2) has only a soft rounded

projection. The new sp. has at the anterior ventral

lori margin a median, relatively broad notch

between two. short keels, which is unknown in

related species.

Measurements: Lorica length 140-150 pains lorica

width 112 pm; width of semicircular foot-opening

IS-TR pm: median noth in ventral plate Ham deep,

Diseyssjon’ The morphological group T patina-

ohlet-thucronata (after Koste 1978) have variable

short tongues at the anterior dorsal margin of (he

lorica. The new species most resembles 7 allel, but

differs in the form of the dorsal and ventral anterior

lorica margin. 7) oh/ei Koste has only three more-

rounded (Ltongue-shaped) elongations of the dorsal

anterior margin [see Koste 1978a, b, ef. also 7

patina trilobata Anderson & Shephard and 7. kostei

De Ridder (De Ridder 1983)].

Etymology: The new species is named alter Charles

G, Hussey, of the British Museum (Natural

History), in recognition of his work on the Rotifera

of the Uk.

T. munda Berzins, 1982

Loc: Creswick Ck, Clunes, Vies U.iv, 1953,

lat: Berzins (1982). Not distinguishable trom 7

aimphota Hauer on the basis of the figures and

regarded here as synonymous.

R. J. SHIEL & W. KOSTE

100 pm

Fig. 6:1 Testudinella husseyi sp. nov. a, dorsal, b, ventral.

6:2 Testudinella patina (Hermann) from Tasmania, length of lorica 245 pm, width 216 pm, ventral view.

\USTRALIAN ROLTIFERA

i “+

P Mid dT ile) gas

f C

Hexarthig osvuris (Sernoy) dy lateral, b, ventral, ¢, ventral armlike appendage, d, lrophi apiertl, e, rao ciel

Hilerwin, fy rammiand fulerim lateral, @, manubrium and uneus teeth, ti, cue projections of A. axveris trom: Tasmania,

1, cuudal projections of 77. barracvaticn Berzins, 1982, a-d from Tasmanian population, ef hom WA. population,

Vig?

4 R. J. SHIEL & W. KOSTE

T. neboissi Berzins, 1982

Loc: Creswick Ck, Clunes, Vic.; I1.iv.1953.

Lit: Berzins (1982). Inadequately figured. Resembles

T. emarginula (Stenroos) and must be considered

a doubtful record.

T. patina intermedia (Anderson), 1889

Loe: L. Barracoota, Vic.; 20.1.1975.

Lit: Berzins (1982).

T. semiparva Hauer, 1938

Loc: Magela Ck floodplain near Jabiluka, N.T;

15.iv.1980.

Lit: Koste (1981),

T. vidzemensis Berzins, 1943

Loe: Creswick Ck, Clunes, Vie.; [Hiv.1953.

Lit: Berzins (1982).

Family Flosculariidae Harring, 1913

Octotrocha speciosa Thorpe, 1893

Loc: Buffalo Billabong, Magela Ck, N-T.; 8.xii.1980.

Lit: Tait ev al. (1984),

Sinantherina ariprepes Edmondson, 1939

Loc: Island billabong, Magela Ck., N‘1; 15.iv.1980.

Coll: R. D, Tait.

Family Hexarthridae Bartos, 1959

Populations of a species of Hexarthra found in

W.A. (4788) and Tasmania (#1146) were determined

to be:

H. oxyuris Sernov, 1903

FIG. 7a-i.

Syn: Hexarthra fennica var. oxyuris (Sernov), 1903

Loe: L. Barracoota, Vic.; 20.i,1975; Norseman-

Albany area, southwest W.A., x.1981; Farm dam

8 km east of St Helens, Tas.; 24.iv.1984; peaty,

acid water, 14°C

Lit: Berzins (1982), W.A, material coll. M. A.

Brock, Tas. material coll. R. J. Shiel.

Regarded as a variety of H. fennica (Levander), 1892

development and unci teeth number. The only

difference between H. fennica and H. oxyuris is in

the caudal armature. The caudal spine may be

absent, as in Hf. fennica, or varying between

individuals (both Tas. and W.A. populations) as in

Fig. 7h (cf. Fig. 7i for a population described as

H, barracootica Berzins, 1982). Most of our

specimens had 7/7 unei teeth (cf. Fig. 6d, g),

however some had 6/7,'7/8. Koste (1977) describes

the variability: 7/7, 8/7. 7/8, 6/6, 6/7. Therefore,

H. barracootica, which was described by Berzins

(1982) as “very like H. oxyuris ... but trophi with

only 6/6 teeth” apparently is a synonym of H.

oxyuris. Further, the ventral armlike appendages of

both taxa have the same number of bristles (8) (Fig.

7c, ef. Berzins 1982 Fig. 60a)

Family Collothecidae Bartos, 1959

Collotheca tenuilobata (Anderson), 1889

Loc: L. Catani, Mt Buffalo, Vic.; 25.11.1955.

Lit: Berzins (1982).

Acknowledgments

We gratefully acknowledge the support of the

Australian Biological Resources Survey to collect

in Tasmania. The word-processing and photo-

copying facilities of the Department of Botany,

University of Adelaide were used in MS production.

Interested collectors are thanked for material, and

access to unpublished and published work: J. van

Alphen (Murdoch University), P. Arumugam

(University of Adelaide), Dr B. Berzins (Lund),

Prof, C. H. Fernando (University of Waterloo),

Prof. J. Green (University of London), Dr R.

Hamond (University of Melbourne), P. Hawkins

and T. Orr (James Cook University), Ms lf. J.

Powling (Melbourne), R. D. Tait (Sydney), Dr B. V.

Timms (Avondale College), Dr P. A. Tyler

(University of Tasmania). M. J. Tyler and an

by Hauer (1941), this rotifer is characterized by anonymous referee are thanked for critical

intra-population variability in caudal spine comments.

References

Brock, M. A. & SHieL, R. J. (1983) The composition of

aquatic communities in saline wetlands in Western

Australia. Hydrobiologia 15, 77-84.

BLRZINS, B. (1982) Contribution to the knowledge of the

Rotatoria of Australia, (University of Lund: Lund.)

Donner, J. (1965) Ordnung Bdelloida. Bestimmungsb.

Bodenfauna Europas.

Dumont, H. J. (1983) Biogeography of

Hydrobiologia 104, 19-30.

Evans, J. (951) The rotifer record of Victoria. Proc.

Microsc. Soc. Vict. (W951), 4-7.

Fapeew, N. (1925) Materialen zur Kenninis der

Rotatorienfauna Russlands. Trav. Soc, Ni. Charkow 50,

3-12.

rotifers.

Gipert, J. J. (1967) Asplanchna and posterolateral spine

production in Brachionus calyciflorus. A rch,

Hydrobiol. 64, \-62.

& STEMBERGER, R. (1984). Asplanchna-induced

polymorphism in the rotifer Kerarella slack. Limnol.

Oceanogr, 29, 1309-1316.

, Birky, C. W. & Wurbak, E. S. (1979)

Taxonomic relationships of Asplanchna brightwelli, A,

intermedia and A. sieboldi, Arch. Hydrobiol. 87,

224-242,

Green, J. (1977) Dwarfing of rotifers in tropical crater

lakes. Ergeb. Limnol. 8, 232-236,

(1981) Associations of rotifers in Australian crater

lakes. J. Zool, Lond, 193, 469-486,

AUSTRALIAN ROTIFERA 15

Hauer, J, (1938) Die Rotatorien von Sumatra, Java und

Bali nach Brac ouissen der Deutschen Limnologischen

Sunda- -Expedition. Archiv. Hydrobial. Suppl. 15,

507-602.

—_— bean Rotatorien. aus dem “Zwischengebiet

Wallacea’”’. 1. Teil Das Genus Pedalia Barrois, Int. Rev.

LES. Hydrobiol 41, 177-203.

Kosit, W. (1977) Uber drei neue Formen des Genus

Hexarthra Sehmarda 1854: H. jenkinae ( nakurun.t.,

H. brandorffin. sp. und H. polydenta soaplakeiensis

n. ssp. Gewdss. Ahwass. 62/63, 7-16.

— (19784) Uber Testudinella ohlei Koste, 1972, cin

Radertier der U-Ordnung Flosculariacea aus der

Ciuinea-Brasilianischen Region der Neotropis. rcs.

Hydrabiol. 82, 359-363.

(Y7Kb) Die Rédertiere Mitteleuropas. 2. vols.

Borntrucker, Stuttgart

(1981) Zur Morphologie, Systematik und Okologie

yon neuen monogononten Radertieren (Rotatoria) aus

dem Uberschwemmuneseebiet des Magela Creek in der

Albgator-River-Region Australiens, NoT. Teil. 1.

Osnabrucker aren Mitt. 8, 97-126.

— & SHIEI, J, (1980a) Preliminary remarks on

the chi aa of the rotifer fauna of Australia

(Notovaeca). //vdrobiologia 73, 221-227.

& - (1980b) New Rotifera from Australia.

Trans, R. Soe. 8. Aust. 105, 133-144.

& ~ (1983) Rotifer communities of billabongs

in northern and south-eastern Australia. Hvdrobiologia

104, 41-47,

—K& - (in press a) Planktonic and

semiplanktonic Rotitera from) Australian inland waters

1. Introduction to rotifer morphology and ccology, Aus.

J. Mar breshwaier Res.

Tau, R, D,, Sears,

(in press b) Planktonic and

Semiplanktonic Rotilera from Australian inland waters.

Hl. Systematics and aa Ibid.

& Brock, M, (1983) Rotifera from

Western Australian wetlands ik descriptions of two

new species. Hydrohioloxia 14, 9-17.

Morray, J. (1911) Australian Rotifera collected by the

ee rit Antarctic Expedition, 1909, 4. Roy. Micrase.

Soc. 31, 164-174,

hh ME De (1983) Recherches ecologiques et

biogeographiques sur les Rotiferes de la Basse

Casamance (Senegal). Rev. Mydrobiol. trap, 1, 41-55.

Russetl, C. R. (1961) The Rotatoria of Queensland,

ha ee alia. Trans, R. Soe. VE 1, 235-239,

SALI, W., SARBADINI, Fo and ComMins, M.

1578) Trophi ToNsbaKoe: relative to food habits in six

species of rotifers (Asplanchnidae). Trans. Amer,

Mierose, Soc. 97, 469-485.

Srire, RK. J. (1983) The genus Brachionus (Rotifera:

Brachionidae) in Australia, with a description of a new

species. Proe. R. Soc, Viet. 95, 33-37.

& Kosph., W. (1879) Rotifera recorded from

Australia, Trans, R, Soe. 5. Aust 103, 57-68.

R. J. & Rosre, W. (1984) Structure

and dynamics of zooplankton communities, Alligator

Rivers Region, N-T., Australia. Mydrobiologia 113, 1-13.

WULEERT, kK. (1960) Die Raidertiere saurer Gewiisser der

Dibener Heide. 1. Die Rotatorien des Zadlitymoors und

des Wildenhainer Bruchs. Arch, Mydrobiol, 56, 261-298,

(1966) Rotatorian aus dem Stausee Ajwa und der

Trinkwasser-Aubereitung der Stadt Baroda (Indien),

Limnologica (Berlin) 4, 53-93.

SOME EFFECTS OF SEASON AND GEOGRAPHICAL ASPECT ON ANTS

(HYMENOPTERA: FORMICIDAE) IN THE MT LOFTY RANGES, SOUTH

AUSTRALIA

BY P. J. M. GREENSLADE

Summary

Ants were sampled for a year in Eucalyptus baxteri-E. obliqua forest at Bridgewater near Adelaide

with two transects of pitfall traps, one on a well isolated northwest-facing slope, the other on a more

shaded southwest-facing one. The fauna was typical of humid parts of South Australia in that the

total of 32 species included both cryptic ants (active mainly within soil and litter) and epigacic

species (active on and above the soil surface) nesting either in soil or arboreally. Catches of

epigaeic species had a pronounced maximum in late spring and early summer. Catches of cryptic

species did not show such marked seasonal variation. The two transects differed substantially. On

the northwest slope catches reached a maximum in February; species with distributions extending to

areas drier than Bridgewater predominated, Eyrean elements were present in the fauna, and

Iridomyrmex species and associated taxa were frequent. On the southwest slope catches declined in

late summer and species extending into areas wetter than Bridgewater, with highest catches earlier

(in November), were predominant, Eyrean elements were less, and Bassian elements were more

frequent than on the northwest slope; a higher incidence of cryptic and other ground-living ants

compensated for a reduction in frequency of Iridomyrmex and associates. Community structure on

the southwest slope, but not on the northwest, resembled that of an ant community that has been

studied in Eucalyptus obliqua woodland in southern Victoria. As a whole this Bridgewater ant fauna

is a transitional one on the elevation gradient of rainfall in the Mt Lofty Ranges. The presence or

absence of Myrmecia forficata on one hand, and of M. pyriformis and Iridomyrmex purpureus on

the other may serve as indicators of this transition. On this site associations of invertebrates (ant

communities) differ in composition according to aspect within a single floristic and structural

vegetation unit. This introduces a brief discussion of the conservation of invertebrates and their use

in environmental assessment.

SOME EFFECTS OF SEASON AND GEOGRAPHICAL ASPECT ON ANTS

(HYMENOPTERA: FORMICIDAE) IN THE MT LOFTY RANGES, SOUTH AUSTRALIA

by PL J. M. GREENSL ADI *

Summary

Gut i sstatn, POL ML (985) Sume etfeers af season and geourtphiedl aspect on ants Cyinenapleri:

boride) in tle ML Lotty Ranues, South Austatin Chere. RSee 8. lust TOU 1), 17-23, 28 fine, YAS

Aubs were siumpled tora year in Lvewvpris haviere€, obliqua forest at Bridgewater near Adelaide

With 1Wo trabseers of pitfall (raps, ane ana well isolated northwesi-taci2 slope, (He orher on a mere

shaded southwest-taeing ane The fauna was typical of Municl party of Soult Australia iy Hil the Golat

OF 32 species included both eeyptic ants (detive maimbhy within soil and liner) and epivaeie species [aerive

on and above (he soil surface) esting either in soil or arboreally. Caivhes of epigavie species hac a pronounced

Manton dy hte spring and summer, Carches of cryplie species did not Show’ stich marked seasonal variation,

The mwo transects differed substantially, On the northwest slope caches reached aamasinnim in Feby iui

species with distrnbuoons extending jo areas drier (han Bridgewater predominated, Eyrean elements were

Present wi the fauna, and idnavines species andl assiviated taxa were frequen, On (he sourhwesr slope

catehes devlined in fate suritiner und species extending into areds Wwetrer dui Bridgewater, wath highest

guiches cuier Gin November), were predominayt, Eyrean elements were less, and Bassian elements were

More frequent than on the northwest slope; a higher incidence of eryplic and other eround-living wals

compensated tor a reduction in frequency of /edemrnex and associates. Community steucture on the

souTfWest slope, but not On the Horthwese, resembled thar of an ant commuaity that Nas been studieal ay

Lucalypius ohliqua woadhaad in southern Victoria, Asa whole this Bridgewater au fauna isa Leansitional

one on the elevational gradient of caimfallin We Mt Lofts: Ranges. The presence or absence of Ww rnerie

fariodra on ole hand, and of AW. periforeis and fridenwrriex, purputeds on Ue other may serve as indesiers

OF This eansition, On this site dssociations of inver(ebrates (ant communities) differ in coMposibon acca dine

fo uspect within a single (loristic and siruciucal Veeention unit, This introduees a bref discusaen al the

Conservation ot iaveriebrales and their use in environmental assessment.

Introduction

The ant communities of South Australia’s lorests

and woodlands have been deseribed recently in

general terms (Greenslade, in press). Sorne of the

ants of one open forest lovality in the Me Lotty

Runges are discussed here in preater detail.

Although Gentilli (1972) stressed the complexity of

the climatic influences that alfect sourhern South

Australia, the climate of the Mt Lofty Ranges near

Adelaide is unequivocally mediterrancan. Summers

ure hoi and dry, winters cool and wet, imposiny a

double stress on biota. Climate can be bots

moderated and accentuated by wspecl. The

distributions of the Avcelypius species that make

up the forests and woodlands of the Mr Lalty

Ranges are determined primarily by soil type and

the eradient of increasing precipuation that

accompanies increasing elevation, but Specht &

Perry (1948) showed that aspeet is involved as well.

Compared with south-facing slopes, north-facing

ones are likely to be waiter in winter bul excessively

hot dnd dry in the summer. Aspect also affects

invertebrates in this cea. tr influences the

distribution of mounds of the termite Nasweiterimes

BOSIRO Division of Sails, Private Bay No. 2

Osmond, So Aust, 306d,

Glen

exitiosus (Hil (Wood & Lee 1971) and the structure

af ant populations and vommnqunities (Greenslade

1975, in press; P. J. ML Greenslade & C. 1, Johnston,

wutpubl.). Effects may be comples, Mie taleraction

of aspect, vegetation and insolation determines the

soil climate which, in turn, affects leading members

of an} conmmunities with secondary effect® on

subordinate species.

In 1983-84, Dr G, H. Baker used pirfull traps al

Bodgewater near Adelaide for a year to sumplea

population of the introduced nhillipede

Ommnataiulus moreleni (Lucas) and to monitor its

potential ground-living predators. This paper

reports on the ants taken in these traps. They were

divided between a well insolated nurthwesr-facitne

slope and a more shaded southwest-lacing one so

their catches presen! another opportutiity to

investigate the interaction between the effects al

season and aspect.

Previous work involving invertebrares on tris site

has dealt with the biology af OQ. werelesii (Baker

1978a-c) and the inputs of dead plant material re

the forest Moor and its disposal by termites (Lee &

Butler 1977; Lee & Correll 1978; Lee & Wood 1968)

and other soil biota (Hartson & Veitch 1983), Data

anid iaterial provided by Baker and Hurtsan have

been used to show how different methods of

sampling soil fauna can yield differing results

18 P. J. M. GREENSLADE

(Greenslade & Greenslade 1983). For example, the

number of ant species captured in pitfall traps

reaches a maximum in summer when temperatures

are high and ants are most active on the soil surface.

On the other hand numbers extracted from soil and

litter samples, for the most part different species,

are greatest in spring and autumn, indicating direct

or indirect limitation by low temperatures in winter

and by low soil moisture and high temperatures and

saturation deficits in summer.

Site and Methods

The Engelbrook Reserve at Bridgewater consists

of an open forest of Eucalyptus baxteri and E.

obliqua with a sclerophyll shrub layer, growing on

an acid yellow duplex soil. Mean annual rainfall is

¢.1000 mm (cf. summit of Mt Lofty, 1690 mm). Ants

and other surface fauna were sampled with pitfall

traps, consisting of plastic jars, 9 cm diameter by

9 cm deep. They were used without killmg agent

and preservative and a lid was supported above

them to prevent the entry of rainwater. Two transects

each of 20 traps 10 m apart were laid out, c. 350 m

above sea level, one on a northwest-facing slope,

the other adjacent to the first, on a slope that faced

southwest. The vegetation on the two slopes was

very similar apart from a slightly higher density of

eucalypts on the more shaded southwest slope

(G. H. Baker, pers. comm.). The site is described

in more detail by Hutson & Veitch (1983).

Traps were cleared at weekly intervals from March

1983-March 1984 and each week ants were pooled

from all traps. These traps were not primarily

intended for ants and are not very effective for

them, due to predation within and escape from the

traps. Consequently the data used here are monthly

mean numbers of species per transect or total

records of species in traps.

Results

The total of 32 species given in Table 1 is not a

complete list for the site. Many other species are

known to be present but a large proportion are

cryptic, the workers foraging mainly within soil and

the litter layer. Therefore they are, in general, not

susceptible to pitfall trapping and only three cryptic

species were recorded in these traps: Amblyopone

australis* and the Heteroponera and

Sphinctomyrmex species. The Amblvopone was

*For the few species for which specific names are available,

authorities are given in Table 1, Numbers within genera

refer only to this site, Voucher speciments are retained

in my collection pending further studies and are to be

deposited in the Australian National Insect, CSIRO,

Division of Entomology, Canberra.

trapped frequently and in substantial numbers,

suggesting an unsuspected amount of foraging

activity on the soil surface. All other species are

epigaeic, that is active on the soil surface and, very

often, on vegetation. All but one or two, of these

nest in soil. One exception is /ridamyrmex sp.1

which nests above the ground in hollow eucalypt

branches and /ridomyrmex sp.4 may nest arboreally

as well since I know it only, as in this case, from

alate queens. This combination of cryptic and soil-

and arboreal-nesting epigaeic ants is typical of

communities of forests and woodlands in the wetter

parts of South Australia (Greenslade, in press).

Seasonal trends in captures of the cryptic species

(Fig. la) are broadly consistent with the pattern

already noted for ants extracted from soil and litter

samples. Together, the epigaeic species showed a

simple summer maximum (Fig. Ib), although there

are differences between the two transects. On the

well insolated northwest-facing slope activity, as

indicated by pitfall captures, extended throughout

the summer (Fig. Ic); on the shaded southwestern

slope catches fell from December onwards (Fig. !d).

(a)Cryptic spp.

(b)Epigaeic spp.

(d) Southwest aspect

Mean species / week

‘ )

AMJJASONDUFM AMJJASONDJE™M

Months

Fig, 1. Mean weekly catches: (a) cryptic species; (b) other,

epigaeic species; (c) epigaeic species, northwest aspect;

(d) epigaeic species, southwest aspect.

This is contrary to expectation. From other studies

(Greenslade 1975, in press) it might be predicted that

catches on the northwest slope would increase

earlier in the spring than on the southwest slope,

only to decline sooner in the late summer as the

habitat became excessively hot and dry, inhibiting

foraging. It remains possible that this applies to

MI LOFTY RANGES’ ANTS

TABLE L. Ants trapped at Bridgewater. 1983-84. Total records on slopes with northwest and southwest aspects. South

Australian distribution: W, occurring only in the wertest, most humid parts of the State; D, occurring mainly in areas

drier (han this Bridwewater locality; M, at about the midpoint of their range on the rainfall yradient at Bridgewater.

Records

South -

Australian Northwest Southwest Total

Ant Species distribution aspecl aspecl

Myrmeciinae

Mivrinecia nigriscapa Roger W ~ z 2

M. forficate (Pabricius) W — 4 4

M. pyriformis F. Smith D 4 3 6

M. pilosnla F. Smith M 6 6 12

Ponerinae

Amblyopone ausiralis Brichson M 12 7 29

Heteroponera intbellis ( I:mery) Dp _ 1 |

Rhytidoponera ‘metallica’ (Vv. Smith) D 3 = 3

Sphinetomyrmes sp. M — | ]

Myrmicinae

Epopostruma sp. M _ 1 |

Crematogaster sp.) Ww I 7 5

Crematogaster sp.2 Db 2 | 3

Meratioplys sp. Db | — 1

Pheidole sp.) W 2 7 19

Pheidole sp.2 Db 6 | 7

Chelaner sp.\ W - § 5

Chelaner sp.2 M — 2 2

Chalaner sp.3 ? I - I

Gen et sp. indet.* ? l — |

Dolichoderinae

Iridomyrien purpureus (F. Smith) dD 2 a 2

Iridomyrmex sp. Ww 10 14 25

Iridomerme’ sp.2 D 3 I 4

Iridomyrmex sp.3 D 2 | 3

Iridomyrmex sp,4 Db l — i]

Dolichaderus sp, D 5 1 6

Formicinae

Melophoris sp. D | — I

Melophorus sp.2 D | _ l

Nafoncus sp. D 2 — 2

Camponotus suffusus FP. Smith M - I |

C. ‘ceansobrinus’ Erichsou Db 18 y} 20

C, innexus Fore! b 3 _ 3

Cumpanolys sp.2 ? I _ 1

Camponotus sp.4 ? -- | I

Toral

Species 23 21 32

Reeords 88 8Y 177

*A damaged alate queer

some species but can be tested only for those that

were relatively frequent in both transects, ie.

Myrmecia pilosula, Amblyopone australis and

fridonirmex sp... In neither transect however do

the summed frequencies of these species differ

significantly from the overall trend for the whole

fauna with both transeets combined: northwest

xy) = 4.60, p>0.05; southwest, My = 3.95,

p>0.03, The differences between Figures le and Id

therefore indicate that the two sets of traps sampled

different combinations of species, ie. different

communities.

All but four of the species in Table 1 can be

placed in one or other of three groups according

to their known distributions in relation to rainfall

in South Australia (from material in my collection

and that of the South Australian Museum). These

groups ure defined in the caption to Table | which

compares transects. Treated as a contingency table

this shows significant heterogeneity. Group W

predominates in catches on the southwest slope,

group D on the northwest slope; the community

here seems the best adapted to exploiting high

summer lemperatures and withstanding dry

cwonditions (Table 2),

The phenology of the epigaeic members of

groups D, M and W is shown in Fig. 2 where catches

from both transects are combined. Compared with

froup D, group W is active earlier in the spring and

has an earlier maximum; catches decline through

P. J. M. GREENSLADE

TARLe 2. Catches in the two transects with species grouped according to their distributions in relation to rainfall

in South Australia: W, D, M,—see legend to Table 1 for explanation.

Transect Distribution type Totals

W M D

Northwest 13 18 54 85

Southwest 50 27 10 87

Totals 63 45 64 172

2 Fs 554

the summer but extend further into the autumn,

Group M is essentially a single species, Myrmecia

pilosula (see Table 1), whose seasonal distribution

in traps is closer to group D than group W.

% Records

AMJJASONODJFM

Months

Fig. 2. Seasonal occurrence of epigaeic ant species

according to their South Australian distributions. See

caption to Table | for explanation of distribution types

D, M, W.

Table 3 describes the ants in each transect in terms

of their biogeographical distributions and their role

anid status in communities. Here, distribution refers

P<<0,001

to the Australia-wide range of genera and species

groups and not, as in Table 2, to the South

Australian distribution of individual species. ‘Role

and Status’, is explained in detail by Greenslade (in

press).

Taking the biogeographical affinities of these

communities first, there is only a small element of

species belonging to Eyrean groups, centred on arid

Australia, and it is most evident on the northwest

slope. Conversely, Bassian taxa, characteristic of

cool southern and highland climates, are best

developed on the southwestern slope.

Turning to community structure, there are

biologically significant differences between transects

in most rows in Table 3. The majority of Australian

ant communities share a basic framework of

epigaeic soil-nesting species consisting of dominant

Iridomyrmex species (row la in Table 3),

subordinate formicines, typically members of the

genus Camponotus (row 2), and species of

Melophorus that forage when most other ants are

inactive (in row 3a) (Greenslade 1979). Commonly

accompanying these genera are broadly adapted,

opportunist species of the Rhytidoponera

‘metallica’ complex. As the frequency of cryptic and

arboreal-nesting species increases in more humid

climates, this /ridomyrmex-dominated core

becomes less important, as can be seen here.

Iridomyrmex, Camponotus, Melophorus and R.

‘metallica’ contributed about half the total catches

in the northwestern transect with 45 records of 11

species (Tables 1, 3). On the southwest slope the

incidence of these genera was halved: 21 records of

only six species. To compensate, on the southwest

slope there was an increase in catches of cryptic ants

and of ground-living Pheidole and Crematogaster

species (in rows 4 and 5 in Table 3): 45 records of

seven species compared with 23 records of five

species on the northwest slope.

This northwest-southwest difference is illustrated

further by comparing the Bridgewater communities

with the results of pitfall-trapping on Wilson’s

Promontory in southern Victoria. Here A. N.

Andersen (unpubl. ms) sampled ants in Casuarina-

Leptospermum heath and adjacent Eucalyptus

ME LOPTY RANGES’ ANTS 4 |

PAW) 3. Siicrure of ent communities at Bridgewater, Taxa (eenera ani species groups) are classed uecording to

their biopeoxraphical distrimiaions and role ane staius in communities. Role and status is described in detail by

Greenslade (in press). Dashes (--) ndicate cells that are wnoecupiuble ar apparently lave ne-occupants in the Australian

Gat futind, ex. cell C3 ty wnoecupiable: it is not possible to have a ‘climate specialist’ that is widely distributed in

respect lo climate. A fourth disiributtonal carezory “Torresian’ (ie. northern tropical) is recognised but tt is not

represented by any species in these communities so the fourth column has been omitted, Fractions tnéicate Auinber

of species/number af records,

SE en A ARS Rs A i ed 1s

Role and Bioveographical distribution Totals

Status ; ’ ane pan

A. Eyrean B Bassian C Widely distributed

Northwest Southwest Northwest Sourhwest Northwest Southwest Northwest Southwest

aspect aspect aspect aspect aspecl aspect aspect aspect

} Dominant

Dolichoderinae 2/5 ial ale tis 2/3 ie 3/18 3417

ay [riclomyrmex Tridamyrntex fridomyrinen Tridamprines

b, Other venera “- v5 val nil 1/5 Va

Dotichoideru

2 Subordinate

cium ponotine nil nil 3/22 3/4 3/22 3/4

hormicinge Cumponotys

3 Climate 4/3 Va) 2/3 BYR — 5/6 Ws

specialists Meranoplis Epopostrunia

Melophorus Chelaner

Notoncus

4 Cryptic and/or i 1/12 1/17 0/0 2/2 1/12 3/19

predacious on Amblyapane Heteroponera

micro-arthropods Sphinciomvrmex

5 Generalised — = 4 4/26 4/11 4/26

myrmicines Crematozaster

Pheidale

6 Opportunists — nil 3 iraa) 13 O/0

Rhwridoponera

7 Large solitary nil 2-10 4/l4 nil 2/10 4/14

foragers Myraiectr

Totals 5/8 Fal 7/40 (he S5 10,39 10/33 22/K7 21/89

SSS

baxter? Woodland. Raintall was similar to that at

Bridgewater, ¢ 1000 mm per amium, but evenly

distributed through the year. Although the soil, a

siliceous sind, also differs there are similarities in

ant faunas, Among Myrnrecia for example, MZ.

prrvornis, ML nigriseapa and M. pilosula were all

(rapped at Wilson’s Promontory (compare Table [).

Andersen partitioned his communities according

to the structure in Table 3, Using his raw data,

Wilson's Promontory—Bridgewater ants can be

compared by means of a correlation coefficient

where the oecupiable cells in Table 3 provide 21

paired observations (Table 4). At) Wilson's

Promontory, as at Bridgewater, there were no

Isbit 4. Comparison of the structure of ant communities at Bridgewater, South Australie, und Wilson's

Promontory, Victoria (see text). Correlation caefficients (r) far frequencies in pus of conmmunities partitioned as

in Tihle 3. Sienilicanee: *,

BERS SOLOS, << OO, OOO,

TT SSSSSSSSSSSSSSSSSFSMmmsmFmFmFfsfhshFhsFeFeF

Bridgewater Wilson's

Promontory

Localities =

and Northwest Southwest

habits aspeet aspecl Woodland

Wilsons’ Heath O40 0.40

Promontory

Woodland OAT (51!

Bridgewater Southwest (70%

aspect

eee

22 P, J. M. GREENSLADE

Torresian taxa so the fourth column is omitted. The

closest similarities were between adjacent habitats

at both localities (Table 4), in each case due to a

large proportion (42%) of shared species:

Bridgewater 13/31 (Table 1), Wilson’s Promontory

14/33. The heath community differed from both

Bridgewater communities and this can be related

to differences in vegetation structure and

composition. However there is a_ statistically

significant correlation between ant communities in

woodland at Wilson’s Promontory and in the cool,

shaded, southwest facing transect at Bridgewater.

In contrast, emphasising the difference in structure

of the two Bridgewater ant communities, and

despite the number of species they share there is

little similarity in structure between the northwest-

facing transect and Wilson’s Promontory woodland.

Discussion

These results from pitfall traps conform with

other phenological observations on ants in the Mt

Lofty Ranges (Greenslade 1975, Greenslade &

Greenslade 1983), showing a pronounced summer

maximum in catches of epigaeic species and greatly

reduced activity in the winter. The same seasonal

trends have been described for ants taken in pitfall

traps in similar mediterranean climates in Western

Australia (Majer 1978, 1980).

Different ant species have different ranges on

rainfall gradients in South Australia, giving complex

overlapping patterns of distribution and

associations of species. Consequently the change

from communities of predominantly epigaeic, soil-

nesting ants, dominated by /ridomyrmex in dry

areas, to weakened /ridomyrmex-dominance and

stratification of cryptic, soil-nesting and arboreal

epigaeic species in wetter climates is essentially a

continuum. But differences between these two

transects suggest that there may be critical points

on this continuum,

Iridomyrmex purpureus is not found in the

highest parts of the Mt Lofty Ranges which reach

700 m above sea level. Myrmecia forficata does

occur here but not M. pyriformis. At a lower

elevation in the ranges, at 300 m at Belair, a

population of 1 purpureus was studied in

Eucalyptus leucoxylon-E, odorata woodland

(Greenslade 1975). Here it occurred with Myrmecia

pyriformis, as on the northwestern slope at

Bridgewater. A further Belair ant community has

been investigated recently (O’Dowd, in press) at

445m with FE. obliqua (one of the species at

Bridgewater): M. jforficata is present but M.

pyriformis and I. purpureus are not. At 350 m the

Bridgewater ant fauna seems to be transitional; it

contains both Myrmecia species as well as J,

purpureus which must be close to its cool, wet limit

since nests are restricted to open, sunny situations

on and near a track,

The same switch from /. purpureus and M.

pyriformis, to M. forficata without I. purpureus is

to be seen on Kangaroo Island. The first pair of

species occur at the drier eastern end of the island

while M. forficata alone is found in the moister

maritime climate to the west (Greenslade 1976, in

press). The two Myrmecia species show similar

relative distribution patterns in relation to

environmental moisture in southern Victoria (A. N.

Andersen, pers. comm,

The Bridgewater ant communities have further

significance in the context of conservation and

environmental assessment. Greenslade &

Greenslade (1984) suggested that terrestrial

invertebrates should not be ignored in this field since

they may integrate environmental variables in ways

quite different from plant associations.

Consequently it cannot be assumed that a

vegetation type supports one characteristic

invertebrate community, and invertebrate

distributions may lead to insight into critical factors

in an environment. The differences between ant

communities of the two Bridgewater transects are

a good example since they exist within a single

floristic and structural vegetation type and they

demonstrate the importance of aspect.

Acknowledgments

Thanks are due to Dr G. H. Baker for the

material from his traps at Bridgewater and to Alan

Andersen, John Buckerfield and Penny Greenslade

for their comments on the manuscript of this paper.

References

Baker, G, H, (1978a) The distribution and dispersal of

the introduced millipede, Ommmatoiulus moreletii

(Diplopoda; Iulidae), in Australia. J. Zool, Lond. 185,

1-11.

(1978b) The post-embryonic development and life

history of the millipede, Ommatoiulus moreletii

(Diplopoda: I[ulidae), introduced in south-eastern

Australia. bid. 186, 209-228.

——— (1978c) The population dynamics of the millipede

Ommatoiulus moreletii (Diplopoda: lulidae). [bid. 186,

229-242.

Gentiiti, J. (1972) Australian Climate Patterns. (Thomas

Nelson (Australia); Melbourne).

GREENSLADE, P. J, M. (1975) Short-term change in a

population of the meat ant /ridomyrmex purpureus

(Hymenoptera: Formicidae). Aus?. J. Zool. 23, 511-522.

MT LOFTY RANGES’ ANTS 23

— (1976) Distribution of two forms of the meat ant

Iridomyrmex purpureus (Hymenoptera: Formicidae) on

Kangaroo Island and Yorke Peninsula, South Australia.

Ihid. 24, 557-564.

— (1979) A Guide to Ants of South Australia, (Sourh

Australian Museum: Adelaide),

(In press) Ants. /n Wallace, H. R. (Ed.) “The

Natural History of South Australia’s Forests and

Woodlands”. (Government Printer: Adelaide).

& GREENSLADE, P. (1983) Ecology of soil

invertebrates. da “Soils: an Australian viewpoint”

Division of Soils, CSIRO, pp. 645-669 (CSIRO,

Melbourne/Academic Press: London).

—— & —-- (1984) Invertebrates and environmental

assessment. Environment & Planning 3, 13-15 (Dept.

of Environment & Planning: South Aust.),

Hurson, BR. & Verve, L. G. (1983) Mean annual

population density of Collembola and Acari in the soil

and litter of three indigenous South Australian forests.

Ausi, J. Ecol. 8, 113-126,

Lit, kK. BE. & Buiter, J. H. A, (1977) Termites, soil

orgame matter decomposition and nutrient ¢ycling.

Evol. Bull. (Stockholm) 25, 544-548.

— & CORRELL, KR. Lb. (1978) Litterfall and. its

relationship to nutrient cycling in a South Australian

dry sclerophyll forest. “lust, J. Zool 3, 243-252.

& Woop, T. G. (1968) Preliminary studies of the

role of Nasutitermes exitiosus (Hill) in the cycling of

organic matter in a yellow podzolic soil under dry

sclerophyll forest in South Australia. Travts, int. Congr,

Soil Sci. 9(2), 11-18.

Maser, J. D. (1978) Studies on invertebrates in relation

to bauxite mining activities in the Darling Range. A

review of the first eighteen months of research. Env,

Res. Bull. No. 3 (Alcoa: Australia),

—(1980) Report on a study of invertebrates in relation

io the Kojonup fire management plan. Dept. Biol. Bull.

No. 2 (W.A. Institute of Technology: Perth).

O’Dowp, D. J. (In press) Seasonal patterns in the activity

of ants in Belair Recreation Park, South Australia, /#

Greenslade, P. & Majer, J. D. (Eds) “Supplementary

papers to the Mediterranean and Related Ecosystem

Source Book”. Depi, Biol Bull. (W.A. Institute of

Technology: Perth).

SpecuT, R. L. & Perry, R. A. (1948) The plant ecolory

of part of the Mount Lofty Ranges. 7rans. R. Soc. S.

Aust. 72, 91-132.

Woob, T. G. & LEE, K. EB. (1971) Abundance of mounds

and competition among colonies of some Australian

termile species. Pedobiologia 11, 341-366.

SPIRURA AURANGABADENSIS (ALI & LOVEKAR) (NEMATODA:

SPIRURIDAE) FROM SMALL DASYURIDAE (MARSUPIALIA)

BY DAVID M. SPRATT

Summary

Spirura aurangabadensis (Ali & Lovekar, 1967) Quentin & Krishnasamy, 1975 is described from

the oesophageal mucosa of the dasyurid marsupials Antechinus stuartti Macleay, Planigale ingrami

(Thomas) and Sminthopsis leucopus (Gray). This is the first member of the genus reported from

Australian native mammals, although the species has been described from a microchiropteran

(Taphozous kachhensis Dobson) in India and from tupaiid (Tupaia glis (Diard)) and lorisid

(Nycticebus coucang (Boddaert)) primates in Malaysia.

SPIRURA AURANGABADENSIS (ALI & LOVEKAR) (NEMATODA: SPIRURIDAE)

FROM SMALL DASYURIDAF (MARSUPIALIA)

by Davip M, Spratt

Summary

PAT,

1, SL (OSS) Spine veruneaduedenses (Al & Lovehun) (Nemiiodi: Spirutidie) fran sural

Dosti (Miarsupialio), Jeans, A. Seo, 8. tee MYL), 28-29, 28 ine, [VRS

Sprury aurangubadensts (Al & Lovekat, 1967) Quentin & Krishnasamy, 1975 is deseribed fran the

ogsophageal njpucosa of the diasyurid marsupials Aarechines stuart Macleay, Plenigale ineramni (Thomas)

and Sminthapsrs deveopus (Gray) ‘This is he first member of the genus reported from Australian native

imaimitmals, alihouyh the species hay been described from a microchiropreran (laphozous kuchhensts Dobson)

in Ioutia and from (upeiid (Tupaia elis (UNard)) and lonsid (Nvercebes concone (Boddaert)) primates in

Malaysia

S. avrangabadensis is similse in moephology and in host disreibiinon to S, eeianensis (Orulepp, (924)

Chityood, 1938 from platyrrhine primates and cicelphid mrsupials in South America. Cephalic structures

ih (hird-stage larvae OFS. aurunvaebadensis (rom 4. styartii are, in conteast to larvae of S. euianensis,

identical to those i adult forms, offering turther evidence that 8. velanensiy is the more highly evolved

member of the genus, Although the penus Spirura iy probably of Gondwanaland origin, 118 undikely that

S. auransubadensts entered Australia via this route, but rather from the north with Chiroptera or with

Windborne Inseata, Its occurrenee in Australian marsupitls is interpreted as secondary rranster

KEY Words Seirvne Nematoda, distribution, hosts, morphology, Dasvuridae.

Introduction

The spiruroid nematodes of Australasian

marsupials are in need of a thorough taxonomic

revision, A preliminary assessment of the group,

based on examination of type specimens held in

Australian institations and on material collected

since 1967, revealed that one species exhibited

morphological relationships more akin to those

froma microchiropteran, primates and didelphid

marsupials than to those from other Australasian

marsupials, Details of this parasite, from the small

dasyurid marsupials Ariveehinus stuartit Macleay,

Planigale ingrami (Thomas) and Sminthapsis

lencopus (Gray) form the basis of this report.

Matertals and Methods

Native mammals were examined diving the

period March 1967-December 1983 from the

following regions: Arnhem Land, Northern

Territory; Queensland; southeastern New South

Wales; northeastern Victoria, the Australian Capital

Territory; northern ‘Tasmania.

Nematodes recovered by the author were fixed

in hot, 10% neutral buffered formalin and cleared

in lactophenol,

The following abbreviations lor morphological

features of adult worms and third-stage larvae are

used in the text:

L. Length of body, MW Maximum width of body,

Po Length » width of pharyox, NR Length from

* Division of Wildlife & Rangelands Researeh, CSIRO,

HO. Hox 84, Lyneham, A-CCT. 2602.

cephalic extremity to nerve ring, EP Length from

cephalic extremity ta exeretory pore, DO Length trom

cephalic extremity to deirids, MO Length of

muscular oesophagus, GO Length of glandular

oesophagus, ‘TO Total length of oesophagus,

VCB Length from cephalic extremity to ventral

cuticular boss (large (hurnb-like swelling on ventral

surface of body), V Length from cephalic extremity

to vulva, RS Length of right spicule, LS Length

of left spicule, T Length from caudal extremity to

anus or cloaea, GP Length from cephalic extremity

LO genital primordium of third-stage larva

Measurements were made with the aid of an

Ocular micrometer, drawing tube, and measuring

wheel, and are presented in micrometers unless

otherwise stated. The range of measurements is

followed by the mean, in parentheses. Illustrations

were made with the aid ot # Lenz drawing device.

Specimens have been deposited in the Australian

Helminthological Collection (AHC), Queensland

Museum (QM), South Australian Museum (SAM),

United States National Museum Helminihological

Collection (USNMHC) and Division of Wildlife &

Rangelands Research, CSIRO (WL HC),

Results

SN characteristic spiruroid nematode was

encountered rarely, atlached to the oesophageal

mucosa of small dasyurid marsupials, Number of

hosts infected/number of hosts examined fram each

locality are shown in parentheses, [he parasite was

found in Antechinus suartii at Mt Nebo (1/5), Qld,

and yt the same host at Nadgee Nature Reserve

26 D. M, SPRATT

(section formerly part of Nadgee State Forest)

(6/54), N.SW., in the same host at Jervis Bay (2/28),

A.CT., in Sminthopsis leucopus at Timbillica State

Forest (1/9), N.SW., and in Planigale ingrami at the

Smithburne River (1/2), near the Gulf of

Carpentaria, Qld.

In view of the taxonomic complexity suspected

in the nominal species A. stuartii, collection details

of specimens from this host from different

geographical localities are presented separately.

Spirura aurangabadensis (Ali & Lovekar, 1967)

FIGS I-11

Material examined: from Antechinus stuartii, Nadgee

Nature Reserve, near Eden, N.SW., P. Haycock and E.

Walter, Id, 19, 3.xii.1981 in WL HC N1438; 19, 5.iii.1982

in WL HC NI519; I¢ posterior, 7.vi.1982 in WL HC N1630;

Id, 19 anterior, 29 posterior, 8.111.1982 in AHC No, 13651;

Id, 12, 7vi.l982 and 19 10.iv.1984 in SAM Nos, V3557,

V3558, V3559.

From A. stuartii, Jervis Bay, coastal A.C-T., J. Kenagy

and P. Haycock, Id, 19, 12.ix.1983 in USNMHC No.

78252; Ig, 2 L3, 7.xii.1983 in AHC No, 13652.

From A. stuartii, Mt Nebo, Qld, D. M. Spratt, 1¢, 19

anterior, 19.vii.1973 in WL HC N167.

From Sminthopsis leucopus, Timbillica State Forest,

near Eden, N.SW., E. Walter and P. Haycock, Id, If,

1.xii.1983 in AHC No. 13653.

From Planigale ingrami, Smithburne River, near

Normanton, Qld, L. Owens, 19, 4.1.1979 in QM No. GL

4613.

Description

Short, robust nematodes, attenuated anteriorly

and generally reflexed in anterior %4 or 4 of body.

Both sexes bearing large, thumb-like swelling on

ventral surface of body (ventral cuticular boss) at

point of reflection; threads of host connective tissue

and infiltrating cells passing under ventral cuticular

boss and surrounding body of nematode at this

point, boss apparently serving as holdfast for

nematode in lumen of oesophagus. Cuticle thick,

deeply wrinkled or folded anteriorly, with

prominent transverse striations throughout.

Cephalic region with prominent elevated cuticular

shield or plate formed as extension of cuticle of

pharynx and hiding inner circle of cephalic papillae

and amphids in en face view (Fig. 1). Pharynx large,

laterally compressed, indistinctly divided into two

parts, armed with six robust teeth originating at

base and protruding well beyond oral opening.

Teeth simple and non-bifid at distal extremities,

terminating in smooth, sharp points. Teeth indented

on non-oral margin when viewed in transverse

section at level of oral opening; oral opening dorso-

ventrally elongated, with large lateral pseudolabia

(Fig. 2). Four pairs submedian cephalic papillae

arranged in inner circle of four large and outer circle

of four smaller papillae. Amphids large, opening

at base of elevated lateral pseudolabia. Oesophagus

divided into short muscular anterior and long

glandular posterior regions, terminating in short but

distinct valvular region (Fig. 7). Nerve ring near

junction of muscular and glandular oesophagus.

Large conspicuous deirids anterior to nerve ring.

Excretory pore observed in only two specimens,

near level of deirids in male, well posterior to nerve

ring in female.

Measurements

Male (7 complete specimens, 1 anterior end): L

5.4-8.5 (6.5) mm; MW_ 180-390 (294); P

50 x 30-80 « 50 (60 x 40); NR 160-275 (232); EP 110

(observed in one specimen only); D 120-170 (146);

VCB 1180-1770 (1572); MO 170-300 (242); GO

2380-4620 (2986); TO 2600-4920 (3218). Spicules

unequal, dissimilar, RS 170-230 (208) and LS

488-612 (545) in length (Fig. 11). Gubernaculum

boat-shaped, narrowed distally, broad proximally

with in-rolled lateral edges, 80 in length (Fig. 6).

T 124-240 (193), with narrow caudal alae but

without cuticular bosses on ventral surface.

Generally four pairs large pre-cloacal papillae

(Fig. 5) (one specimen with three papillae on right

side, four papillae on left side); two pairs large post-

cloacal papillae, one pair immediately posterior to

cloaca, one pair near tail tip (Fig. 9); single, small,

median papilla near tail tip and pair of minute

subterminal phasmids.

Female (8 complete specimens, | anterior end,

2 posterior ends (Fig. 8)): L 7.9-15.4 (10.9) mm; MW

312-645 (445); P 60x30-90x60 (75x43); NR

170-370 (243); EP 412 (observed in one specimen

only); D 100-200 (157); VCB 1690-3150 (2174); MO

230-350 (268); GO 3052-6010 (3822); TO 3332-6360

(4090), Proximal vagina with thick cuticular lining,

surrounded by thick muscular wall and convoluted

distally, VY 3518-5460 (4055), vulval opening

surrounded by small, irregular, densely-packed

cuticular bosses (Fig. 10). T 120-240 (174),

terminating in three minute cuticular digits. Eggs

in distal vagina 34 x 25-40 x 30 (38 x 27), containing

larviform embryos.

Third-stage larva (2 complete specimens): Small

ventral cuticular boss present in anterior one-third

of body (Fig. 4). Cuticle thick, with prominent

transverse striations throughout length. Cephalic

region as in adults (Fig. 3). Pharynx similar to that

in adults, however teeth less sclerotised, walls thin-

ner and clearly divided into anterior and posterior

parts. Oesophagus divided into short anterior

muscular and very long posterior glandular regions.

Nerve ring in anterior region of muscular oeso-

phagus. Excretory pore conspicuous, opening into

elongate, terminal excretory duct. Subventral glands

conspicuous, each with prominent nucleus and

LURK ANG ABTDENSTS FROM DASYURIDAL

SPIRUR A

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Figo LIL. Spinura aurangabadensis Irom small dasyurid marsupials. 1, Cephalic end, adull, en face view. 2. Cephalic

end, adult, lateral view. 3. Cephalic end, third-stage larva, lateral view, 4, Third-stage larva, lateral view (arrow

ventral ouriedlar boss). 5, Caudal end male, lateral view, 6, Gubernaculum, latero-ventral view. 7, Ocsophiago-

testinal junction, Jateral view. 8, Caudal end female, lateral view, 9, Caudal end male, ventral view. 10, Cuticular

bosses around vival opening, 1. Right and left spicules, ventral and latero-ventral views respectively, Scale lines:

Figs 1.4, 20 wm: Pigs 2.6, 9-11, 50 pm; Figs 4, 5, 7, 100 nin,

28 D. M. SPRATT

nucleolus. Deirids slightly anterior to nerve ring.

Genital primordium small, just anterior to

oesophago-intestinal junction in one specimen,

extending anterior and posterior from junction in

other specimen. Tail terminating in single

papilliform structure. Lateral alae absent.

L 2.55, 2.47 (2.51) mm; MW 97, 113 (105); P

46x 24, 45x 20 (46x 22);NR 120, 96 (L08); EP 310,

270 (290); D 105, 92 (98); VCB 813, 738 (776); MO

154, 157 (156); GO 1482, 1198 (1340); TO 1616, 1354

(1485); GP 1510, 1150 (1330); T 85, 96 (91).

Discussion

The laterally compressed pharynx, dorso-

ventrally elongated oral opening with large lateral

pseudolabia and the ventral cuticular boss in both

sexes are characteristic of the nematode genus

Spirura (Spiruridae). Cephalic, cuticular and genital

structures in specimens from small dasyurids in

Australia are unique among members of the genus

Spirura and are identical to those occurring in S.

aurangabadensis originally described from a

microchiropteran (7aphozous kachhensis Dobson)

in India (Ali & Lovekar, 1967) and redescribed from

tupaiid and lorisid primates (Tiipaia glis (Diard) and

Nycticebus coucang (Boddaert)) in Malaysia

(Quentin & Krishnasamy, 1975).

Specimens, particularly males, from Australian

hosts are similar in size to those from 7. kachhensis

but generally smaller in all measurements than those

from primates. However, my measurements and

those of Quentin & Krishnasamy (1975) indicate

that this is due, at least in part, to the age and/or

state of maturity of the nematodes and does not

justify erection of a new species for the Australian

material. In all other respects except one (males with

two pairs post-cloacal papillae and single median

subterminal papilla contra males with three pairs

post-cloacal papillae) specimens from Australian

hosts are identical with those from Indian and

Malaysian hosts and are here recognised as

conspecific and determined as S. aurangabadensis

(Ali & Lovekar, 1967) Quentin & Krishnasamy,

1975,

Spirura aurangabadensis is distinguished from all

species of Spirura except S. guianensis (Ortlepp,

1924) Chitwood, 1938 (syn. S. tamarini Cosgrove,

Nelson & Jones, 1963) by the presence in both sexes

of six robust teeth originating at the base of the

pharynx and protruding well beyond the oral

opening, and the presence of a cuticular cephalic

plate or shield formed from an oral extension and

outfolding of the cuticular wall of the pharynx, and

hiding the inner circle of cephalic papillae and

amphids in en face view.

These two features serve also to distinguish S.

guianensis from the other 19 species of the genus

recognised by Quentin (1979). The similarities

between S. aurangabadensis from microchiropteran,

primate and dasyurid marsupial hosts in India,

Malaysia and Australia and S. guianensis from the

oesophagus of platyrrhine primates of the genus

Saguinus in South America (Ortlepp, 1924;

Cosgrove ef al. 1963; Thatcher & Porter, 1968) and

from the same site in the didelphid marsupials

Caluromys philander (L.), Marmosa_ cinerea

amarare Thomas and Philander (Metachirops)

opossum (L.) in Guiana (Quentin, 1973), are

striking.

Nevertheless, S. aurangabadensis is readily

distinguished from S. guianensis by the following

suite of characters: (i) pharyngeal teeth terminating

distally in uniformly tapered, sharp points rather

than a bifid extremity, (ii) absence of five ventral

cuticular denticles on cephalic plate, (iii) cephalic

papillae of inner circle Jarger than those of outer

circle, (iv) presence of fine cuticular bosses

surrounding vulva, (v) absence of longitudinal

cuticular bosses on ventral surface of male tail, and

(vi) much longer (2) and morphologically distinct

left spicule.

On the basis of host distribution and comparative

study of larval and adult cephalic structures it was

postulated that the genus Spirura originated in

lemurids in Madagascar and Africa and that it

diversified principally along two evolutionary

pathways in Old World host groups which first

appear in the palaeontological record during the

Tertiary (Quentin & Krishnasamy, 1975; Quentin,

1979). In the most primitive line the border of the

cephalic cuticular collarette is rounded in dorsal and

ventral regions in both larval and adult forms.

Cephalic structures remain stable and five groups

containing 17 species are recognised (Quentin,

1979). Evolution among these groups of species is

based principally on differentiation of genital

features in male worms.

In the more advanced line the border of the

cephalic cuticular collarette is drawn out into dorsal

and ventral points at least in the infective larval

stage. In the four species recognised in this group

cephalic structures are not stable and in the most

specialised form, S. guianensis, structures in

infective larvae are distinet from those in fourth-

stage larvae and adult forms (Quentin, 1973) and

similar to structures seen in adults of less specialised

members of the line (Quentin & Krishnasamy, 1975;

Quentin, 1979). Third-stage larvae of S.

aurangabadensis in A. stuartii exhibit cephalic

features similar to those in adults and to those in

third-stage larvae of S. guianensis, supporting the

SPIRUKRA AURANGABADENSTS FROM DASYURIDAE 24

argument that S. guianensis is the more highly

evolved member of the second line (Quentin &

Krishnasamy, 1975; Quentin, 1979).

These authors argued that the occurrence of the

most highly differentiated form, S. guianensis, in

South America was due to its geographical isolation

in ancient platyrrhine primates. Its presence in

neotropical marsupials was attributed to secondary

invasion of ecological causation. The occurrence of

S. aurangabadensis in Australian marsupials is

interpreted also as a consequence of secondary

transfer. The less specialised form of cephalic

Structures occurring in third-stage larvae and adult

forms suggests that the species has not undergone

lengthy geographic isolation in marsupials in

Australia. On the contrary, although the genus

Spirura is probably of Gondwanaland origin it is

unlikely that S. aurangabadensis entered Australia

via this route, but rather from the north with

Chiroptera, or with windborne Insecta (e.g. Locusta

sp.) which probably serve as natural intermediate

host of the parasite.

Acknowledgments

Thanks are due to the following for collecting

specimens or supplying small dasyurids for

dissection: P. Haycock, J. Kenagy, K. Newgrain,

L. Owen and E. Walter. Drs |. Beveridge and J.

Calaby offered valuable criticism of earlier drafts

of the manuscript and Mrs. J. Rudd competently

typed all drafts of the work.

References

Au, S. M. & Lovekar, C. D. (1967) On a new Spirurid

Taphozoia aurangabadensis n.g., n. sp. from a microbat

Taphozous kacchensis. Ind. J. Helminthol, Year 1966,

18, Suppl. pp, 68-73,

Coscrove, G. E., Nerson, B, M. & Jones, A. W, (1963)

Spirura tamarini sp. n, (Nematoda: Spiruridae) from

an Amazonian primate, Tamarinus nigricollis (Spix,

1823). J. Parasitol, 49, 1010-13.

Orxtiopp, R. J. (1924) On a collection of helminths from

Dutch Guiana, J. Helminthol. 2, 15-40.

Quentin, J. C. (1973) Presence de Spirura guianensis

(Orulepp, 1924) chez des Marsupiaux neotropicaux.

Cycle evolutif, Ann, Parasitol, 48, 117-33.

(1979) Place zoologique de Spirura rothschildi

Seurat, 1915, Nématode parasite du Macroscélide, dans

Vevolution du genre Syurura. Bull Mus. rain. Hist. nat.

Paris 4° ser, 1, section A n® |, 1051-56,

& KRISHNASAMY, M, (1975) Nématodes Spiruru

parasites des Viypaia et du Nycticébe en Malaisie. Ann.

Parasitol, 50, 795-812,

THATCHER, V. E. & Porter, J, A, (1968) Some helminth

parasites of Panamanian Primates. Jrans. Amer.

Mivrase. Sac., 87, 186-96.

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 109, PART 2

GROWTH IN CAPTIVE BORN TIGER SNAKES (NOTECHIS ATER

SERVENTYI) FROM CHAPPELL ISLAND: IMPLICATIONS FOR FIELD

AND LABORATORY STUDIES

BY B. BARNETT & T. D. SCHWANER

Summary

Captive born tiger snakes (Notechis ater serventyi) from Chappell Island, Bass Strait were fed

house mice ad libitum. Growth rates (total length and weight) were twice that of mainland tiger

snakes and 5-10 times that of natural populations on other islands. The pattern of increase in weight

was exponential during the first six months of the study. Growth in length for the first six months

was best explained by a linear model; both linear and logarithimic models explained growth in

length during the second six months. Regression analysis detected a significantly greater increase in

length relative to weight in the first six months of growth, but increase in weight relative to length

during the second six months of growth. An estimate of gross growth efficiency indicated that

metabolic rate was relatively constant for these snakes under prescribed conditions. Patterns of

growth measured under controlled conditions may provide an insight into the evolution of life

history traits in natural populations of tiger snakes.

GROWTH IN CAPTIVE BORN TIGER SNAKES (NOTECH/S ATER SERVENTY 1) FROM

CHAPPELL ISLAND: IMPLICATIONS FOR FIELD AND LABORATORY STUDIES

by B. BARNETT*® & TD. SCHWANERT

Summary

Bakst)), BOS Ser ani TD. C98) Crowilh i eaptive hor byer snakes ¢Nereehis ier ee very LAA

Chappell Ishind: (iploaions toe lel sud liboratory studies, Imei A Nn So dase WM 21 4é

PR bine, RS

Capuius bop tiger snukes (Norwehis ater seevent yy) trom Chappell hud, Bass Strait were sed Nase

mice de bia, Growth rates (otal length and werehl) were eee (hal of mainjand tiger snakes and #1)

limes tharof natural popilations on othe islands. The paren of increase Wr wel Ar was exponential duping

the (rst six months and linear during the second six meanths of the study, Growth in lengthy for lhe tirsr

Six months was best explauried by a linear model; boitt linear and logarithiinic madels explained grawth

in lengih during the second six months, Regression analysis detected # significantly greater increase i leneth

relative lo weight in the first six months of growth, but inerease in weight relanve to length durim: the

second six Months of growth, Aq estimate of gross growth efficiency indicated that metabolic rate wus

telatively constant for these snakes under prescribed condjhous, Pallernms of growth measured under eonolled

conditions may provide ai msight inte the evolution of bite history reuis in natural populations of liver snakes.

Key Woris Snakes, growth, captive husbandry, Nodecsis ater serventye

Introduction

Chappell Island, Bass Strait supports a popula-

tion of tiger snakes (Norechis aler serventyi) in

which individuals reach adult body lengths

approaching 2 m (Worrell 1958). The great size

attained is almost twice the average length of

individuals on (he mainland (Shine 1978), and is

rivalled only by a few other island populations of

tiger snakes in Bass Strait and the Nuyts

Archipelago, South Australia, Although large adult

size in these snakes has been attributed to their habit

ot eating muttonbirds (Worrell 1958), no data exist

to quantify their feeding habits or to delermine

Lrowch patterns trom birth to adulthood, The only

previous studies Of growth and maturation in tiger

snitkes involved dissection of Museu specimens or

estirnates of these parameters from field sampling

of the mainland species, NV. seadatuy (Shine 1978).

Here we report a one year study of feeding and

growth ja brood of new-born, captive tiger saakes

from Chappell Island. The results are compared

with previous studies of growth mates in tiger snakes,

and with recent (unpublished) evidence for growrh

in a populanon of marked river snakes on the

Franklin tslands im S. Aust

Materials and Methods

A subset ol six snakes Was taken from 4 brood

of 31 neonates born to a captive Chappell Island

fiver snake of 1420 min (otal length, on 3.7.83,

" (6 Suspension Street, Ardeen, Viv 3022.

PSouth Austrafian Museum. North

Adeluide, S. Aust SCXW).

Vorrace,

Selection of the subsel was biased by (he need to

pair differen! colaur morphs far mdividual

identification in cages, and to achieve equal

representation of sexes, However, comparisons ol

individual weights and total body lengths wiih those

of the brood means for these parameters satisfied

the null hypothesis that each individual was

sampled from the same statistical population

(t[-05,d?=30] 2.042; ts weieht=0.46—1,231; ts

length =0,49—0.68; Sokal & Rohlf 1981).

The-six neonates were maintained in pairs in three

particle board vages measuring 440% 480% 450 yim.

The lid of each unit contained a glass window

measuring 420% 250 fin. Ligh! was provided only

from *“True-life’ fluorescent tubes suspended above

the caves, Bach cage was fiued with a blue, 40 watt,

incandescent light bulb contralled by a thermostat

fo maintain a constant temperature range of

28-30°C, Relative humidity was 70%. chrouphour

the study.

Snakes were fed entirely on house mice (Mfuis

inusculus), Neonates accepted day-old mice directly,

or mice rubbed with faces of seincid lizards (their

uulural prey). Snakes were fed older, larger mice

commensurate with inefeasine body sizes and their

ability to consume Jager prey, During the 12 month

period, the stakes were fed we Hbifiprto the extent

Ihal rejection of food jndicated repleuan. Daily

reconis were kept of rhe quantity of foud accepted.

Snakes were weighed and measured monthly,

Prior to weighing snakes were denied food so that

monthly Weights would be comparable among

individuals, This action reduced (he feeding tine

ol gach snake by 39-40 days over the 12 mons.

Complete cata sets for monthly groweh in length,

weight and amount of food consumed are deposited

in the Library, South Australian Museum.

Length and weight measurements were examined

by least-squares regression to determine a line of

best fit to the data, and an equation describing

growth was generated. Exponential (y=ae”), linear

(y=a+bt), and logarithimic (y=a+blnt) equations

were used to generate the best fit to the growth data,

where y=length (in mm) or weight (in g), t=age

(in months), a=the y intercept, b=the slope and

e=the natural logarithim of 2. For each linear

regression the largest value of the squared

correlation coefficient (r) was used as the criterion

to determine which equation best fit a given set of

data.

Growth rates were determined using the

instantaneous coefficient of growth, (G), calculated

from the equation:

G=InY,—InY,/t,—t,

where Y,=initial length or weight, Y,=final

B, BARNETT & T. D. SCHWANER

length-weight relationships were tested by regression

analysis (Sokal & Rohlf 1981).

Gross growth efficiency (GGE) was calculated by

dividing the average increase in weight of snakes

by the average weight of ingested prey, for a given

month, and multiplying by 100.

In these analyses data sets are treated without

separate calculations for males and females. This

was due to the very low degrees of freedom rendered

by such a division of the data, and by the noticeable

homogeneity in variance of mean length and weight

for all but one individual.

The latter was female No 5, which ate only

minimally between months 4-6 but thereafter fed

normally and achieved a size similar to other snakes

by the end of the study.

Results

Growth in weight: Neonates increased from a

mean weight of 8.7 g at birth to 1266.0 g at 12

TABLE 1. Monthly rate of growth in weight and total length of captive Notechis ater serventyi.

Weight Length

Group Growth Rates Group Growth Rates

No. Age Mean Mean

snakes (mos) weight (g) SD (g) Range (g) (%/mo) (g/mo) length (mm) SD (mm) Range (mm) (%/mo) (mm/mo)

6 0 8.67 0.25 8.34- 8.98 = — 289.7 6.4] 280- 298 _— =

6 ] 18.85 1.63 17.29- 21.91 77.7 14.6 372.8 12.53 355- 388 25.2 73,1

a 2 52.44 11.97 34,84- 67.48 102.3 $3.7 508.0 39.29 446- 549 30,9 157.2

6 3 83.61 22.98 43.11- 106.83 46.6 39.0 611.0 63.95 491- 677 18.5 112.8

6 4 123.68 34.04 61.36- 154.18 39.1 48.4 715.0 79.18 568- 799 15.7 112.4

6 5 196.17 44.04 117,09- 233.03 46,1 90.5 829.8 80.48 681- 914 14.9 123.6

6 6 344,37 72.00 227.76- 430.93 56.3 193.8 949.8 77.29 814-1041 13.5 128.3

6 7 479.74 74.75 336,35- 550.48 33.2 159.0 1040.2 73.88 903-1117 91 94.6

6 8 670.69 84.04 503.19- 734,49 33.5 224.7 1137.8 65,17 1017-1205 8,9 102.0

6 ) 875.37 104.80 679,42- 968,74 26.6 233.1 1213.6 61.21 1112-1268 6.5 78.3

6 10 1004.70 123.47 774.90-1130.50 13.8 138.4 1270.5 47.21 1186-1308 4.6 58.2

6 11-12) 1266.00 153,62 1021,10-1480.10 23.1 292.7 1366.8 45.66 1290-1414 7,3 99.9

nee TEU UttIdd Ua EISdEIS UES SSNSSSSSS SSS

length or weight, t;=age in months at Y, and

t,=age in months at Y, (Forsythe 1984). Relative

growth rate was expressed as percentage increase in

total length or weight per month, and was calcu-

lated by multiplying Gx100. The velocity of

growth, or actual increase in any length (L) or

weight (W) was calculated as GL, for rates in mm

per month, or GxW, for grams per month (see

Forsyth 1984). Differences in slope values for

months (Table 1).

Growth rates were highest during the first six

months, with an overall mean rate of increase of

90% for the first two months and 61.4% for the

next four months. Growth was exponential during

this period and best described by the equation:

W(g)=47.2e°"'

However, correlation coefficients (r) were similar

for all models (Table 2; p’s>0.5, Z-test, Zar 1974).

TABLE 2. Comparison of regression equations for growth in weight and total length of Notechis ater serventyi.

values are in parentheses.

Months

0-6 6-12 1-12

y =47.20-78" (.94)y = 719.3600 (.67)y = 320.7500 (.48)

Weight y= —1.44+2.2Int (.75)y = 564.7 + 210.81nt (.79)y = 230.6 + 156t (.10)

y= —35.34+5It (.86)y = — 578.6 + 156,3t (.98)y = — 190.9 + 110.6t (.92)

y = 435.6" (.60)y = 1076") (.69)y = 758.100" (.38)

Length y =353.1+2.41nt (.76)y = 313.6+2.81nt (.99)y = 224.64 6.11nt (.94)

y = 320.54 101.6 (.99)y = 498.5 + 78.61 (.99)y = 308.7 + 99.9t (.99)

CAPTIVE

Aller six months growlh rates cdeelined (fable {)

with a mean growth rate of 26.0%month from the

6th to the 2th month, Growth for all snakes during

this latter period was linear and best deseribed by

Ne equation:

Wig) = — $78.61 1561

Linear and logarithimice models (lable 2) were

similar (p=,09),

Although the pereenr growth rites (Smo)

generally declined alter the first six months, actual

mean weight per mont (2/mo) increased markedly

at this time (Table 1), Monthly increases in actual

weight averaged 37.6 2 for the first tive manths: for

the last seven months, this increase was 207,0 2, This

observation is inqportant for the discussion ta follow

because the increase in actual weight per muuth is

correlated with a marked increase in feeding rate

but not witha similar inerease in monthly growth

in length (see below),

Growth in lengthy Neonates increase from a fotal

length of 290 mm at birth to 1367 mm at 12 months

(Table 1). Proportional increases in lenge (o/mo)

were highest during the first six months, with an

overall mean ot 28% for Lhe first nwa months and

15.6%) lor Lhe next fourmonths. Therealier, length

increased by an average of only 7.4% per mou

(Table 1). Unlike proportional weight gains,

monthly increases it length were poorly fitted by

an exponential curve (Table 2). Varialion in monthly

length for the first and seeond six manehs,

respectively, and lor the entire study period was

explained well however, by logarithmic and,

particularly, linewr growth models (Table 2).

Correlation coefficients tor all linear madels

differed significantly (p's <.05) from exponential

models (Table 2), but only thelinear equation for

the first six months differed trom the loparithimie

model (p04),

It is well known that Jeneth inereases

approximately as the cube roor of weight (Brocds

1945), and thal length them must be jinereasing

consisiently in proportion to weight. However, the

average rate of iierease iy length for these snakes

appeared ta be ata much lower rie, proportionally,

To the iment increase in weight particularly during

the second six months of the study.

Lenerh-welght

relationships: The previous

analysis suggested bwo different growth plrases for

weightand length, one during the lirst six months

and another from six to 12 months. In particular,

actual increases int weight (2/mo) were highest

during the last sever months, whilst proportional

(“a/mo) Weight wains declined, A similar pattern

of decline in proportional growth in length during

(he last six months also was observed, but aetual

Increases in dengrh (mmeémo) did nor dilfer

Lah R SNAKE GROW LH 33

markedly Thtoughout the study,

From these observations it would appear thal

Weight Increases Master relative lo length during the

latter sin months, This was tesred using rewression

analysis of length versus Weight lor the two growth

phases, months J-6 and 7-12, respectively, Fig. 1

shows the plotted curves atid their equations. The

curve for Lhe second six months of growth ts steeper

than the eurve for the Jirst six months, and the

reyression slopes (3.63 and 2.95, respectively) are

highly significantly different (PIL,8]=16,8 p< 005)

This difference occurs at a total length of 800-

900 mm.

10 6 months re

Wore AzeLengHt®® ~~ of

6 Pp

WEIGHT »® 100 |g)

2 vy To 5S months

a Moss 631 Lenyth29

: e

— ~——_____.

3 . 5 f %F 8&0 20

LENGTH = 100 imm)

Fig |, Lope log plot of length ‘weighs relanonship far both

Browth phases Wil commons for tints given. Grawih

nhiases based on weyhl ineraise (2/ma) herween cponltis

3-6 (Fable 1).

faod Intuke: Average weight of food consumed

per month varied fran 26 2 during the first month

af the study to 390 2 ducting the last hwo months.

Overall averave weight ol ingested prey for the first

five pronrhs was 94 2 However, lor the latter six

MONS wverdve consuMmplion of prey rese

dramatically to 476 pg, five times thal tor the

previous months. This sudden iheréase jn average

weight Of food consumption coincides precisely

with the inerease in weight observed at the same

time (Table [; Fig, 1), and occurs at an average oral

length of about 800-900 tom.

Gross growth efficiency. Whe overall mean GOE

for Chappell [sland tiger shakes fed ad fibitin on

Mice Was 48.6 (SD=9.4). With the exceptiun of the

iwitial teeding interval, GIGUE was nearly constant

over the entire siudy period, This value cornpures

favourably with similae calculations for other

Organisms (Forsythe 1984), and suggested that

metubolic rate was relatively constant for these

snakes under the preseribed conditions.

Discussion

Shine (1978) plotted monthly body lengths and

estimated age classes and growth rates of mainland

tiger snakes (Notechis scutatus) from N.SW. Both

sexes matured in about 24 months at body (snout

to vent) lengths of 650 mm (Shine 1978); total length

(estimated from a proportion of 16.5% of tail to

total length) would be about 760 mm. Growth in

length after 14 months was approximately 550 mm

SVL (640 mm total length) and the relative rate of

increase (adult length/length at birth) was 2.9.

Similar values for the captive brood from Chappell

Island were 1140 mm SVL (1366 mm total length)

and 4.7, respectively. Thus, the Chappell Island

brood, when fed ad libitum, grew at almost twice

the rate of mainland snakes under natural

conditions. Furthermore, one pair of the Chappell

Island tiger snakes (a male and a female) exhibited

courtship and mating behaviour during the IIth

month of the study at body sizes of 1116 mm SVL

B. BARNETT & T. D. SCHWANER

(1337 mm total length) and 1077 mm SVL (1290 mm

total length), respectively. Apparently (under

laboratory conditions) maturity is reached in

Chappell Island tiger snakes at total lengths almost

twice that of mainland tiger snakes. '

An extensive mark/release/recapture programme

for black tiger snakes (Notechis ater niger) on West

Franklin Island was begun in 1982. Fig. 2 shows

SVL classes for snakes when initially marked, and

ranges and means for monthly growth in length

prior to recapture. The overall average monthly

increases in body lengths (SVL) are only 5.5 mm

for males and 3.3 mm for females. These estimates

lack any pattern of declining growth in adults (SVL

vs average growth per month, r= —0.026, n=5]).

The smallest body size for a snake observed in

courtship and mating on the Franklin Islands was

a female 1002 mm SVL. Given that this size

represents adulthood, the mark recapture data

suggested that sexual maturity in females could be

s 5

500 700 900 1100 1300

SVL CLASS(IN MM)

Fig. 2. Mean monthly growth in length (SVL in mm) for marked-released-recaptured tiger snakes from West Franklin

I. Dots represent males: open circles, females. Solid and dashed lines connect means for males and females, respectively,

in each size class.

CARVING TIGER SNAKE CROWT EY 4

aelneved i about 20 years (about 12 years in males)

on the Franklin Islands.

Several extrinsic factors obviously affeet growlh

in the natural environment (Andrews 1982), Primary

among these are (1) the availability of foad, (2) the

effects of temperature, and (3) interspecific and

INtraNpecii¢ competinon and/or predation,

Prey types vary markedly amone tiger snake

populations, particularly on islands, Diets are

composed almost entirely of frogs in most mainland

aredts (Shine 1978), whilst island populations (excep!

for Rangaroe Land Tasmania) are devoid of frogs,

Tiper snakes on many islands, cat small mammals,

birds, and/or lizards (Schwaner, in press). Little is

known of the effects of differing quantity and

quality of prey umonyg natural populations of tiger

snakes (Shine 1978). However, a high correlation

between manunal body size and mean weight of

available prey exists for several offshore island

populations of Uxer snakes in South Australia

(Schwimmer, im press).

Seasonal vanalions in temperature apparently

affvet the feeding habits of tiger snakes on the

branklin istands (Sehwaner, am prep.) Adults

(> 1000 mm SVL) do not feed during winter

tnonths apparently because they ate unable ro digese

large prev and apparently do nor eat small prev,

Under laboratory conditions these snakes

revurilated larice prey al body leriperatures lower

than these measured when the prey was ingested.

Survival of island Geer snakes during wititer may

depend heavily on their ability to store body tit,

On the Fraoklin [stands (and on Chappell Island)

adult tiger snakes feed primarily on matronbirds

(Puffiins teruirasuris), a highly seasonal prey

(Woryyel] 1958; Schwaner, in press), Adult female

snakes are gravid during wits season, bur Shine

(1978) ancl Schwaner (in press) indicated that gravid

tiger snakes do pot feed. Alternatively, (he size al

which [he laboratory reared snakes were observed

to inerease markedly in weight relative to length

(Lic. Tas approximitely the sive at transition from

juveniles to adults in both the Chappell island

populition and the tiger snakes on the Franklin

Ishines. Increased weight co lenpth ratios suggesting

fat depasirion may therefore be related to the onset

Ol reproductive maturity in these snukes

Most islatids supporting populations of tiger

snakes tack natural predators, Kangaroo [sland bias

goannas (karanus rosenberal) and the undescribed

pywy coppertead snake Clistrefaps spo, borh

known precditiors of snakes. On Reewsby Psland

near Port Lincoln, goannas have been introduced

duclmay have reduged direcly numbers of tazer

stakes (Mirtschine 1982), One St Peters Island of f rhe

Western coust of Eyre Pemnsulu near Ceduna,

nouns may have reduced the mumbers of riper

snakes indirecrly by feeding on mutonbud eggs C1.

Mertens, in litt, 3.i11,.84), Mainland populations at

tiger snakes appeas to reach smaller maxiniun body

sizes than most island populations but no direct link

with predators of competitors has yet been

established as a correlated factusr,

I there was an optimum natural environment for

feeding and wrowth in black tiger snakes, growth

rates in those environments would be expected to

exceed those of the ariifietal laboratory

environment, Recognition that actual growth rates

in natural populations vary due to particular

extrinsic factors (e@. variable food availability) can

be accamplished by comparative studies among

different island populations both in the field

(Sechwaner, in press) and in the liboritlery,

Whereas variable growth rates appear to be an

adaptive strategy for eclothermic vertebrates (Pough

1980), add one Which is highly affected by extrinsic

factors, @ Most Import question concerns

intrinsic interpopulational (intraspecific) differences

in prowth rates, Andrews (1982) has suggested that

one method of defining the relationshijy between

growth aod filness can be explored using

experimental manipulation of food availability in

luboratory setrings. Certain differenees in growl

rates may Occur between sexes, as adaptations to

particular Jife s(yles (or ditferctitial resource

availabililies) and by wide variation among

individuals. reared under similar conditions

(Andrews 1982). Among ishinc populations tiger

snakes difter in maxinia) body size by as much as

an order of magnitude (Schwaner in press), Also,

within and among populations, sexual dimorphism

in maximal body size is extreme for some

populations, with males greater than fernales, whilst

in other populanens femules equal males ut body

sive, Or are Slightly largen This stocky has

Uemornstrited that Heer snakes fed va bin craw

at prediviable nes. Similar comparative studies of

growth it captive snakes from other astanie

populations with different muximal body sizes may

reveal whether these rates ane adaptive or are simply

environmentally determined,

Acknowledgments

Paris of Chis study were supported by grants trom

the South Austridian Wildlite Conservation Fund,

and by the Austrian Research Grants Scheme. We

thank D, Bradford and R. Seymour for the use of

Theil tegression analysis progiamme and computer

fala Schwaner typed the manuseripe,

36 B. BARNETT & T. D. SCHWANER

References

ANDREWS, R. (1982) Patterns of growth in reptiles. Jn

Gans, C. & Pough, F. H. (eds.), “Biology of the Reptilia,

Vol. 13?” (Academic Press: New York.)

Bropy, S. (1945) Bioenergetics and growth with special

reference to the efficiency complex in domestic animals.

(Hafner Press: New York.)

ForSYTHE, J. W. (1984) Octopus joubini (Mollusca:

Cephalopoda): a detailed study of growth through the

full life cycle in a closed seawater system. J. Zool. 202,

393-417.

MIRTSCHIN, P. (1982) The Goulds goanna, an Australian

native, alien to Reevesby Island. S. Aust. Nat. 57, 18-20.

& Davis, R. (1983) Dangerous snakes of

Australia: an illustrated guide to Australia’s most

venomous snakes. (Rigby: Adelaide).

PouGu, F. H. (1980) The advantages of ectothermy for

tetrapods. Am. Nat. 115, 92-112.

SCHWANER, T. D. (in press) Population structure of black

tiger snakes, Notechis ater niger, on offshore islands

of South Australia. In Grigg, G., Shine; R. & Ehmann,

H. (eds.): “Biology of Australasian Frogs and Reptiles?’

(Surrey Beatty Co.: Sydney.)

SHINE, R. (1978) Growth rates and sexual maturation in

six species of Australian elapid snakes. Herpetologica

34, 73-9.

SOKAL, R. R. & ROHLF, F. J. (1981) Biometry. (W. H.

Freeman & Company: San Francisco).

WorRELL, E. (1958) Song of the snake. (Angus &

Robertson: Sydney.)

ZAR, J. H. (1974) Biostatistical Analysis. (Prentice-Hall,

Inc.: Englewood Cliffs.)

A REDEFINITION OF UPEROLEIA RUGOSA (ANDERSSON) (ANURA:

LEPTODACTYLIDAE)

BY MARGARET DAVIES & KEITH R. MCDONALD

Summary

Topotypic Uperoleia rugosa were examined and their morphology, osteology and advertisement

calls described.

A REDEEINITION OF UPEROLETA RUGOSA (ANDERSSON)

(ANURA: LEPTODACTYLIDAE)

hy MARGARET Davins® & Kerr R. MCDONAL Lt

Summary

MX Me Dome ati ROR

Hin BO Soe Ss

LW ih

Hiyinclaers tuhted

(IYSS) A hedehimtion ot Cperolde vigose (Nidersarn) ON

Vist WW 2), 37-42 OK Tie HONS,

lopolypie Cyperalend tigesda Were amined and cher morphology, astcology and advertisement calls

tleseribedt

RIY Worn Morptolany, ostealopy, adverltisenion) calls, Gperotend send.

Introduction

Proys al the leptodactvlid genus Uperaleru Gray

are small, short dimibed, lossorntl wainials exhibiting

limited thorphologieal diversity A. recent revision

and subsequent papers by TWler, Davies & Martin

(l9slahc) resulled in the recognition of 18 species.

Che revision was concerned principally with Those

species Oecurring in that part of the continent

investigured by che authors (lie Kimberley Division

of Western Austrahia, and the Northern Territory);

airention fo 1axa gecurring in the eastern staiws was

eomlined fo un esamination of type material of

species described from that area,

Because Of [he Limited imterspeestie

morphological diverpence, in Uperoleig, species

voncepls rely heavily upon bivlugical data,

particularly cell information,

Uperalvia rugosa (Andersson) is one of the

‘astern species recognised by Tyler et af (1Y8ta).

dinve dis description in 1916 it has been reported

widely fram Queensland, New South Wales and

Victoria (Barker & Gripe, 1977, Cogeer e/ al, 1983),

The rype locality of OL rugosa is reported to be

Colosseum Qld by Andersson (1916). However, a

label accompanying the type specimen stares it was

volleeted at Mp Colosseum, Colosseum is situated

onthe plains at the foot of ML Colosseum and in

19]2 was served by a now abandoued railway sic,

Duriig u visi to Lhe area in December 1984, one

ofus (K.R.MeD.) obtained a series of specimens and

recorded the advertisement call. The availability of

his (opotypic material permits a study of skeletal

structure. Here we describe the material,

Materials and Methods

Specnuens examined here ure deposited in the

Natuirtistoriska Riksmuscel, Stockholm (NRAM)

* Dept ol Zoology, University oF Adelaide, ChPOL Bo

$98) Advluide, S. Aust. SOOT.

i Queernslionl National Parks Ae Wildlife

Pallarendi, Townsville, Qld +800

Serving,

and the Sourh Australian Museum Adelaide

(SAM).

Methods of measurement follow Tyler er al,

(9S8la), Measurements taken were: eve diameter (E)

eye fo naris distance (F-N), internarial span (IN),

snout to vent denech (S-V) and vibia lengrh (PL).

Osteolouical data were Obtained from cleared arid

Alizarin) RedeS stained specimens alter the method

of Davis & Gore (1947). Ostealogical deseriptions

follow Trueh (1979).

Culls were recorded with a Uher 4000 Report

Monitor tape revorder and an AKC DI9O ES

microphone Wel bulb air temperatures were

ineasuredl wilh a Sehultheiss rapid-reading

thermomerer within 3 em of the valling site.

Calls were analysed by means of & sound

spectrograph (Kay Model 6061-B Sona-Graph) with

the overall response curve maintained in the PL-!

position. Temporal and spectral characteristics of

calls were determined from wide-band (300 Hz

bandpass) spectrograms. Because of consistency

between calls both within and between individuals,

only single representative.calls of each male were

analysed and mean values valeulated,

Uperaleia rugosa (Andersson)

FIGS 1-6

Peendophirvne riwosa Andersson, t9t6,

Verenskapsakac. Handl. 32, 9. yy, 13,

kK. Sven.

Uperaleia rugesa: Parker, 1940, Navitates Zool, 42) p, 70

(pape Wher Davies & Marin T98l, Aust. 1 Zool.

Suppl Sen 79 po 17; Cagger 183, Reptiles and

Auwplibians of Austratia, 3rd edin, p. 86; Cogper

Cameron & Copger 1983. Zoolagial Catulogue of

Auseralia Vol, | Amphibia and Reptilia, po 34 (part),

Mareril examined

NRAM 1630, Tolotype, creer 9) ME Colosseum

(24°24. 15) 379 SE Old, xn 1912, BE. Miéhers: SAM

RI7OS2-3, R27O54-5 (cleared and stained), culling et ,

We site Of former Colosseum railway suecion (Ee2b.

ISL 37°), (2ANiL19S4, KR. R. MebDonald.

38 M. DAVIES & K. R. MCDONALD

35 pes 1985

Fig. 1. A lateral and B dorsal views of the head of male Uperoleia rugosa (SAM R27052),

Definition

A small species ( ¢d& 19-22 mm, ¢ 21 mm)

characterised by moderate hind limbs (TL/S-V

(0.35-0.39); fringed toes with a trace of basal

webbing; maxillary teeth absent; prominent

subarticular and palmar tubercles; unexposed

frontoparietal fontanelle; mating call a short note

of four pulses.

Redescription of species (based on SAM R27052)

Maxillary teeth absent. Vomerine teeth absent.

Snout short, slightly truncated when viewed from

above (Fig. 1A) and in profile. Eye to naris distance

greater than internarial span (E-N/IN 1.43),

Canthus rostralis inconspicuous and_ straight.

Tympanum not visible externally (Fig. 1B).

Fingers moderately long, slender, unwebbed but

slightly fringed with prominent palmar and

subarticular tubercles (Fig. 2A). Prominent

supernumerary subarticular tubercles present.

Fingers in order of length 3>4 >2 >1. Hind limbs

moderately long (TL/S-V 0.36). Toes moderately

long, fringed with a trace of basal webbing (Fig.

2B). In order of length 4 >3'>5 >2 >1. Metatarsal

tubercles moderately large but prominent.

Subarticular tubercles rounded, Dorsal surface

faintly tubercular. Parotoid and inguinal glands well

developed; coccygeal glands poorly developed.

Submandibular gland prominent, discrete (Fig. 1A).

Cloacal flap with tiny fimbriations. Ventrally, throat

and thorax granular, belly smooth. Male with

unilobular, submandibular vocal sac and glandular

nuptial pad.

In preservative, dorsum brown with darker brown

markings between eyes, in scapular region and on

posterior portion of dorsum. Dorsal tubercles

tipped with cream. Parotoid and inguinal glands

with few cream patches. Inguinal and post femoral

patches pink after one month in preservative.

Nuptial pad white.

Ventral surface cream with few small, isolated

areas of brown pigment. Submandibular region and

thorax, except for small medial patch, dark grey.

Colour in life

Dorsum grey/brown with darker brown

markings. Dorsal tubercles tipped with yellow;

parotoid and inguinal glands yellowish;

submandibular gland yellow. Bright reddish orange

inguinal and femoral patches.

Ventral surface white with brown areas of

pigment, Submandibular region dark grey to black

sometimes extending to pectoral region.

Dimensions (in mm)

Snout-vent length 20.9; tibia length 7.6; eye

diameter 2.6; eye-naris distance 2.0; internarial span

1.4.

Variation

The addition material ranges 19.75-21.7 mm S-V.

Relative lengths of tibia are consistent (TL/S-V

0.36-0.39). E-N/IN ranges 1.24-1.67. The specimens

vary little morphologically from the described

specimen.

The snout is more rounded in two of the

specimens, and the belly is faintly granular in

others. Cream patches on the dorsal glands are

shghtly more prominent in a few specimens and the

pigment patches on the ventral surface are less

discrete in one specimen.

The material varies little from the holotype,

although dorsal colouration and patterning can be

observed. This feature conforms with Andersson’s

description (see Tyler ef al., 198la). The

pigmentation of the ventral surface was not reported

REDEPINELION OF UPEROLEIA RUGOSA 3y

Pig, 2. A palmar and B plantar views of the hand and foot of male Uperoleia rugosa (SAM R27052),

by Andersson (1916), and is not apparent in the

holotype.

Osteology (based on SAM R27055)

Skull well ossified, sloping anteroventrally.

Sphenethmoid not ossified dorsomedially or

ventromedially; small portion ossified posteriorly

to medial extremities of palatines, with ossified

portion extending about 1/3 length of orbit in

ventral view. Prootic not fused with exoccipital.

Exoccipital confluent posteromedially in dorsal

view but not fused ventromedially. Crista parotica

short, stocky, not articulating with otic ramus of

squamosal lying alongside lateral extremities.

Grooves of carotid canal present on [rontoparietals,

medial to epiotic eminenves.

Frontoparietal elements extensive, crenate

medially, overlying sphenethmoid anteriorly, more

widely separated anteromedially. Frontoparietal

fontanelle barely exposed medially; anterior margin

formed by sphenethmoid elements at level about

1/5 posteriorly on length of orbit. Posterior margin

40 M. DAVIES & Kk.

R. MCDONALD

Fig. 3. A dorsal and B ventral views of the skull of male Uperoleia rugosa (SAM R27055).

undefined because of lack of anteromedial

ossification of exoccipital.

Nasals large, closely applied medially for 4/5

length, slightly separated posteromedially (Fig. 3A).

Anteromedial extremities crescentic. Maxillary

processes moderately acute, directed ventrally, not

in bony contact with pars facialis of maxillary.

Posteromedially nasals in contact with anterior

extremities of sphenethmoid. Palatines moderately

slender, angled at about 45° to sphenethmoid,

reduced laterally, not reaching extremities of

maxillary processes of nasals.

Parasphenoid robust. Cultriform process broad,

irregularly truncate (Fig. 3B), expanded slightly

posterolaterally. Alary processes moderately long,

moderately broad, angled slightly postero-laterally,

reaching medial extremities of medial ramus of

pterygoid. Medial ramus of pterygoid broad,

anterior ramus in long contact with well developed

pterygoid process of palatal shelf of maxillary.

Cartilaginous quadrate present between base of

squamosal and quadratojugal. Quadratojugal

robust, in firm contact with maxillary. Squamosal

stocky with tiny zygomatic ramus and long,

unexpanded otic ramus.

Maxillary and premaxillary edentate; palatal shelf

moderately deep with well developed palatine

processes, not abutting medially. Pterygoid process

well developed. Alary processes of premaxillary

perpendicular to pars dentalis, slender, inclined

medially. Pars facialis of maxillary moderately deep;

preorbital process absent. Vomers absent. Bony

columella present.

Pectoral girdle arciferal and robust. Omosternum

absent, xXiphisternum present. Sternum

Fig. 4. A lateral view of pelvic girdle and B dorsal view

of vertebral column of Uperoleia rugosa SAM R27055,

cartilaginous, Clavicles slender, curved, closely

applied medially. Coracoids well developed, robust,

widely separated medially. Scapula_ bicapitate,

shghtly shorter than clavicles, Suprascapular about

2/3 ossified.

Eight non-imbricate presacral vertebrae (Fig. 4B).

Sacral diapophyses poorly expanded. Relative widths

of transverse processes III >1V > sacrum > II >V=

VI>VII=VIIIl. Bicondylar sacrococcygeal

articulation. Crest on urostyle extending approxi-

mately 1/2 length. Hium with tiny dorsal

prominence and moderately large lateral protuber-

ance (Fig. 4A).

REDERINTHON OF UPEROLEIA RUGOSA WI

Humerus with strongly developed anteroproximal

erest. Phalangeal formula of hand 2,2,3,3; small

bony prepolles, palmar sesamoid. Phalangeal

formula of foot 2,2,3,4,3. Well developed bony

prehatlus,

Pariahton

A second lopotypic specimen was examined

(SAM R27056). The specimen varied in the nature

of the extremities of the trontoparietal elements;

unterior extremities were (runcate and medial edges

were straight, not crenate. The nasals were in

lenuous contacl with the frontoparietals. The

exoceipitals were not confluent posteromedially. In

other features, the skeleton approximated the above

description closely.

ldvertisement cull

The advertisement calls of three individuals were

recorded at Colosseunt on 12.41.1984. The call is

KHz

ipRy

0°5

sec

hig. 5. Sonigram of mile adverrisement call ot Uperuleia

rugosd SAM (R27083). Temperature 228°C,

Take L dédvertisentent call characterishes of Uperalen tugosa af Colasseun, Queensland,

Specimen No. of pulses Duration Pulse repetition Domiunas| Wet-bulb

(ms) ale (pulses see |) frequency lemp. "C

(Hy)

SAM R27052 4 120) 33.33 2500 22.6

SAM K27053 4 120 33.33 2500 22.8

SAM K27055 4 110 36,55 2750 22;2

Meni + 116.7 34.40 2583 22.5

eo SSS

uw short nore of four pulses, pulsed at about 34

pulses see | and with a duration of about

H7 msec, The dominant frequency is about

2,583 Hz, A sonogram of the advertisement call is

shown in Vig. 5, and characteristics of the calls are

listed in Table |,

Habilgl

Frogs were located on the lower slopes and

adjacent plains of Mt Colosseum. The lower slopes

ure disturbed forest of /. tessedlaris F. Mucll., £

terevticornis Smith, £. erehra, and Melaleuca

quinquenervia (Cav.) S. T. Blake. Most of the plain

is heavily disturbed and is mainly grassland with

scattered large tress of &, fereticornis (Fig. 6).

The frogs were found around temporary sunimer-

lilled pools and were calling in short grass on banks

1-6 m trom water, Culling was heard only following

a thunderstorm of approx. 25 mm rain,

Other frogs heard and seen in the area were:

Litoria caerulea, L. fallax, Lo rubella, 1. rathii, Lb.

inerinis, Lo nesta Lo albogutiata, Ranidella et,

desertioala. Adelotus brevis, Limnodvnastes peroni,

L. terraeregiiue, 1. ornatus, L. tasmeniensis and

Bufo marinus.

Andersson (1916) described the

Colosseum as open forest.country,

habitat at

biz. 6 Habitat which calling mutes of Cperoleia pygose

were found near former railway station, Colosseum,

Old

old.

42 M. DAVIES & K. R. MCDONALD

Discussion

In revising Uperoleia, Tyler et al. (198la) redefined

the three named eastern Australian species, U.

rugosa (Andersson), U. fimbrianus (Parker) and U,

laevigata Keferstein, all from the type specimens.

They resurrected U. fimbrianus from the synonymy

of U rugosa in view of their experience with

restricted distributions amongst taxa elsewhere in

Australia, and because of its larger size.

They also indicated the existence of a number of

undescribed species in eastern Australia that await

resolution. Cogger (1983) considered the eastern

Australian species to be composite.

Since the revision of Tyler ef al, (198la), U.

lithomoda, Tyler, Davies & Martin, has been shown

by Tyler & Davies (1984) to have a wide-ranging

distribution across the continent. Evidently not all

species of Uperoleia have restricted distributions,

and hence the status of U, fimbrianus rests solely

on its large size.

The data presented here clarify the status of U.

rugosa: a step vital to resolution of the status of

U. fimbrianus and of the identity of other eastern

Australian taxa. Cogger ef al. (1983) gave the

distribution of U. rugosa as N.E. coastal, S.E.

coastal, Murray-Darling basin, Bulloo River basin,

Qld and N.SW. (A.C.1.). The ACT. records

probably are based upon the work of Robertson

(1982', 1984a,b). However, the species studied by

him is “U, rugosa [type B]” sensu Littlejohn (1967), a

toothed species not conspecific with the untoothed

U. rugosa and currently under investigation by

Davies & Littlejohn (in prep.).

Here we have demonstrated only the identity of

U. rugosa from S.E. coastal Queensland. The extent

of the geographic range of the species awaits

clarification of the status of U. fimbrianus (Davies

& Littlejohn, in prep.).

Acknowledgments

We are grateful to Dr G. Astrom (Naturhistoriska

Riksmuseet, Stockholm) for access to the type

specimen of U, rugosa, and to Dr M. J. Littlejohn

and P, Harrison of Zoology Department, University

of Melbourne for preparing the sonograms for us.

Michael J. Tyler critically read the manuscript and

is thanked for support and encouragement.

This work was supported by University of

Adelaide, Department of Zoology Research Funds.

References

ANDERSSON, L. G. (1916) Results of Dr E. Mjébergs

Swedish Scientific Expeditions to Australia 1910-1913.

IX. Batrachians from Queensland, A. Svenska

Vetenskapsakad. Handl. 52(9), 2-20.

Barker, J. & Grica, G. (1977) A field guide to

Australian frogs (Rigby: Adelaide).

CocceR, H. G. (1983) “Reptiles and Amphibians of

Australia’, 3rd edtn (Reid: Sydney).

, CAMERON, E. E. & CocGcer, H. M. (1983)

“Zoological Catalogue of Australia Vol. 1, Amphibia

and Reptilia” (Aust. Govt Publishing Service;

Canberra),

Davis, D. D. & Gore, V. R. (1947) Clearing and staining

small vertebrates. Fieldiana: Techniques 4, 1-16.

LITTLEJOHN, M. J. (1967) Patterns of zoogeography and

speciation in southeastern Australian amphibia.

pp 150-174. In A. H, Weatherly (Ed.) ‘Inland Waters and

their Fauna’ (A.N.U. Press: Canberra).

'Ropertson, J. G. M. (1982) Territoriality and sexual

selection in Uperoleia rugosa (Anura: Leptodactylidae).

Ph.D. thesis. Dept of Zoology, A.N.U. (unpubl.).

Ropertson, J. G. M. (1984a) Acoustic spacing in

breeding males of Uperoleia rugosa (Anura:

Leptodactylidae). Z. Tierpsychol 64, 283-297,

— (1984b) a technique for individually marking frogs

in behavioural studies. Herp. Rev. 15(2), 56-57.

Trues, L. (1979) Frogs of the genus Telmatobius in

Ecuador with description of a new species. Copeta

1979(4), 714-733,

Tyter, M. J. & Davies, M. (1984) Uperoleia Gray

(Anura: Leptodactylidae) in New Guinea. Trans. R. Soc.

S. Aust. 108, 123-125.

— & Martin, A. A. (198la) Australian

frogs of the leptodactylid genus Uperoleia Gray. Aust.

J. Zool. Suppl. Ser. 79 1-64,

&— (1981b) New and rediscovered

species sof frogs from the Derby-Broome area of Western

Australia. Rec. West, Aust, Mus. 9(2), 147-172.

& - (I98lc) Frog fauna of the

Northern Territory: new distributional records and

description of a new species, Trans. R. Sac. S. Aust.

105(3), 149-154,

A NEW SPECIES OF PHASCOLOSOMA (SIPUNCULA) FROM

AUSTRALIA

BY S. J. EDMONDS

Summary

A new species of sipunculan, Phascolosoma kapulam sp. nov. dredged from off the coast of New

South Wales, is described. It is distinguished from other species of Phascolosoma by the shape of its

introvert hook.

A NEW SPECIES OF PHASCOLOSOMA (SIPUNCULA) FROM AUSTRALIA

by S. J. EDMONDS*

Summary

EDMONDS, S. J. (1985) A new species of Phascolosoina (Sipuncula) from South Australia, /rans. BR. See.

S. Aust. 109(2), 43-44, 28 June, 1985

A new species of sipunculan, Phascolosoma kapalum sp, nov, dredged from off the coast of New

South Wales, is described. It is distinguished from other species of Phascolosoma by the shape of its introvert

hook.

Key Worps: Sipuncula, Phascolosoma kapalum sp. noy., Shape of introvert hooks.

Phascolosoma (phascolosoma) kapalum sp. nov.

FIGS 1-2

Phascolosoma Leuckart, 1828; Fisher, 1952: 422; Stephen

& Edmonds, 1972; 270.

Type locality: Stn K77-23-06, 33°40'S, 151°56'E,

off coast of New South Wales; dredged at 710 m,

during cruise of “Kapala”, coll. P. Colman & D.

Brown, 6.xii.1977.

Type specimen; Australian Museum Sydney: W

17004

Paratype: South Australian Museum: E 1694

Description:

Trunk: Length 13-32 mm, maximum width near

mid-region 4-7 mm. Subeylindrical to spindle-like

and sometimes covered or partly so with coating

of very fine particles of adherent mud. Thin walled.

Longitudinal musculature grouped into 24-32

anastomosing bundles, usually visible externally.

Introvert: Slender and almost completely retracted

in all specimens. Length 2-% trunk. Finger-like

tentacles present which do not appear to surround

mouth of dissected specimen. Numerous rows of

single-pointed introvert hooks, 0.075-0.095 mm

wide basally and with vertical height 0.070-

0.090 mm. Hook with a lateral extension of its basal

portion on side away from tip of hook. Hook

uniformly brown except for clear streak (running

from tip to mid-basal region), very narrow

anteriorly but wider basally. Twelve or more basal

bodies at base of hook on same side as tip.

Papillae: Most prominent at anterior and posterior

regions of trunk and at base of introvert. Mostly

hemispherical but may be bulbous, subclavate or

conical, Central pore usually surrounded by 5-7

pigmented plates and sometimes elevated so that

papillae appear mamillate. Largest hemispherical

forms about 0.3 mm in diameter, subclavate forms

0.3-0.4 mm in height.

* South Australian Museum, North Terrace, Adelaide,

S.Aust. 5000.

Fig. |. Phascolosoma kapalum sp. nov., entire specimen

(scale bar = 5S mm),

Refractor muscles: Four, consisting of a strong

ventral pair arising from muscles 3-6, 4-7 or 5-8

in posterior third of trunk and a weaker dorsal pait

more anteriorly from muscles 6-7, 6-9 or 7-9.

Dorsal and ventral muscle on fused

anteriorly,

each side

44 S. J. EDMONDS

0°05

Fig. 2. Phascolosoma kapalum sp. nov., introvert hook

(scale bar in mm).

Nephridia: Two; long, tubular and thin walled,

extending to mid-region or posterior half of trunk

fixed to body wall for about %4 length and opening

to exterior at about same level as anus or just

anterior to it.

Alimentary canal: Short oesophagus and long

coiled intestine. One fastening muscle to anterior

intestine. Spindle muscle, arising anteriorly from

below anus, is fixed posteriorly. Intestinal caecum

and wing muscle. Contractile vessel without villi.

Brain with 2 eye spots.

Systematic position

This species of Phascolosoma falls in the

subgenus Phascolosoma s.s. (Stephen & Edmonds

1972). It is distinguished from all other members

of the sub-genus by the structure of its introvert

hook, especially its basilateral extension. The

species is not included in Edmonds (1980).

References

Epmonps, S. J. (1980) A revision of the systematics of

Australian sipunculans. Rec. S. Aust. Mus. 18(1), 1-78.

FISHER, W. K. (1952) The sipunculid worms of California

and Baja California. Proc. U.S. natn. Mus. 102, 371-450.

STEPHEN, A. C. & EDMONDs, S. J. (1972) The Phyla

Sipuncula and Echiura. (Trustees Brit. Mus. (Nat. Hist.):

London) 529 pp.

SURFACE ARCHITECTURE OF THE DORSAL EPIDERMIS IN

AUSTRALIAN FROGS

BY MICHAEL J. TYLER & CHRISTOPHER A. MILLER

Summary

Two distinct forms of surface architecture of the epidermal squamos epithelium occur in Australian

frogs of the families Hylidae and Leptodactylidae: one consisting of an elaborate pattern of

microplicae; the other a reticulum. Each species exhibits only one form but species that are closely

related phylogenetically and ecologically may differ in the nature of the epithelial architecture.

SURFACE ARCHPPECTURE OF THE DORSAL EPIDERMIS IN AUSTRALIAN FROGS

by MICHAEL I Tyter & CHRISTOPHER AL MITLER®

Summary

fyiiin, M

bow Mb rR, CoN. C1985) Suite arebifcetire Of The clorsal epidermis i AUseratignt [ips

Tha. Bo Sees. dust (O9O, beets, 28 dune, JYss,

Two distine) forms of surtice arehiieeture of rhe epidermal squamous epithelium over in Australien

Hrovs OF the hantilies Pivlidad and Leptodactylidae: one consisting of an elaborate paren of micropticne;

the other arericulim, Maeh speeies exhibits only one form bul species Hat are closely related pliylosenetigally

and ecologically may differ in the nature of the epithetial surface aretirectiite

Ray Woks: Props, SEAM, epidermis, uliriseraeuine.

Introduction

The dorsal skin of Hogs is an extremely complex

Tissue, containing oeous and granular glands

whose contents are diseharged to the external

surface Vin duets (Elkan, 1968), The mucous glands

ure under the control of syinpathetic nerves; the

mucus liberated assists temperature control via

evaporative cooling, und, by matitaining & moist

surface, also enables the dorsal skin to Punetion as

urespiralory Surface Challywhite, 1971). Lp contrast

the secretions of the eranular vlands of many

species dre toxic, ted afford these animals

protcerian against some predators (Daly & Myers,

1967),

Lhe ventral skin is ealancdtular in the vast majority

of species and either smooth or distinetly granular.

Functionally it is involved in water uptake, and

species fable fo estreme water loss adopt a stance

in Whieh grandlar shin is adpressed tothe substrate

during periods of aetive restorarion of body wither

(Stille, 1958: Johnson, 1969),

The histology af frog skin has been reviewed by

Tilias & Shapiro (1957) and Elkai (968), These

studies have documented gross structure and

deserthed (he whinds dispersed throughout the skin,

In addition, the secretions of agerceations of glands

Jo form paroloid lands (as in Bufo species) or calf

Hands (io che Australiin leptodactylid species

Limdvrastes deaerili) have been the subject of

speeie studies (Low, 1972; Cronk & Tyler, 1981),

As is typleal far vertebrate animals, the outer

epidermis is composed of squamous epithelial cells

irarieed as at ostrardn corneum, Eleetron

microscopiy Stivlies of the ouler sarlace of

squamous epithelia in various vertebrates hits

revewled an vhuborare patter of elevated or

depressed whorls and ridges (blowkes, 1974;

Falvenbuch & Knutson, T9735; Andrews, L976;

Kalleibach, Hardite & Susan, 1980),

Departmental Zoclowy University of Adelaide Buy 498

GPO, Adglardy, S Ast SDT.

Hlere we have examined the ullrasiructure of

dorsal epidermal cells in Australian frogs, to

establish the nature of the diversity in surhitee

urchileclure and {ts possible relationship to the

systematic arrangement ol [hose species. It iiclides

more venera and species than any previows survey

Of skin architecture of the Anura, and is the fest

investigation of Australian species.

Materials and Methods

Aduli representatives of the following 17 wenera

and 30 species were examined: lde/ones brevis, laa

darlingioni, Crinia georwiand, Cyclorani atistralts,

C. euttripes, C. longipes, C. ingeulosus, ©) melee,

C. platveephalus, Heleioporis evret, Leclirwnlis

fleteheri, Limnodyvnastes convexiusculus, Ly

duinerili, b. ornatus, Lo spenceri, L. Tsiminensts,

Litoria adelaidensis, L. albaeautiata, 1} bicalar L.

caerulea, L. ehloris, L, coplandi, L. dahl, 1,

dentata, to ewtngt, 1. Jattax, b, Jreveineli, b.

eractlenta, Lo mernis. Lo pallida, LE perani, bl,

raniformis, Lo rathi, bo rithelta, Lo turatert

Mevistolats lienarins, Mixaphives fiseiolanns,

Neobulrachus pictus, N. sutor, Notaden bennett,

N. inelanascaphus, Pseudophryne pilroni 2

guentheri, Ranidella riparia, RK. suntfera,

Rheobatrachus silus, Taudacivlis aeunrosiriy, 1

diurnus, Uperalera inuadata aud GL trachvider ila,

Vhe selection of species reflects an allempl lo

sample diverse venera, bul was subjech to the

availability of material.

Portions (35 nim) of dorsal skin were remioved

from frogs thac had been killed by contael wilh

3% solution of chloral hydrate, fixed ia 3"

formalin whilst still ina relaxed state and stared

in G5 alcohol, The tissues were eriticulepoint dried

using (he liquiel CO. “anv! procedure it a Polaran

E 3000 CLP, drier, or else simply air-dried i a

desiccalor Specimens were coated with LS ni

carbon und 20 nm pold/patladium (R0:20), and

Viewed om BTR C scanning eleeron imicrascape

al 20 Ky,

M. J. TYLER & CG. A. MILLER

Vig. |. SEM view of squamous epithelial cells of skin of Cvelorana maini showing hexagonal shape.

Fig. 2. SEM view of dorsal skin of: A. Litoria freycineti showing microridges on the cell

surtace. B. Crinia georgiana showing a reticular cell surface. C and D, Sloughing of

skin of Uperoleia trachyderma. In D the central portion is the aperture to a dermal

gland duct surrounded by intercellular connections.

SURLACE ARCHITECTURE OF FROG SKIN AT

Results

In the species examined, the surface cells of the

stratified squamous epithelium are polygonal

(usually hexagonal) (Fig. 1). Upon these cells two

forms of surface architecture occur. One is a

complex pattern of curved elevated ridges

corresponding to the “microridges” of Hawkes

(1974) and the ‘‘microplicae” of Andrews (1976)

(Fig. 2A). The second form is in the arrangement

of an extremely dense reticulum (Pig. 2B). It

appears similar to the type of surtace that

Kaltenbach ef a/. (1980, p. 323) described as having

“a spongy appearance”.

The incidence of these two forms of surface

architecture of the squamous epithelium in the

species and genera examined is summarised in

‘Table 1.

To establish the existence of intraspecific

variation, dorsal skin from six Literia ewing? and

six Ranidella riparia was examined. Within cach

species there was no detectable variation in surface

architecture. We did not observe any difference in

the quality of preparations associated with the

drying techniques. We examined a vyariely of

preparations in which the outer epidermal cells

could be seen to be lifting away from the underlying

cells and which we attributed to represent normal

ecdysis. Examples are shown in Figs 2C and 2D,

The projections from the surface of the cells in Fig.

2D are considered to represent tmtercellular

Tani t 1. dneadence of two forms of dorsal, dermal urchiteciure amongst 17 senera and 50 species of Australian frogs.

Genus Family No. of spp. Surface architecture

examined Microridges Reticulum

Adelotus [ 1 0 brevis

Assa JM ] 0 dartingtoni

Crinia | l 0 georgiana

Crelarcna H 6 ausiralis langipes

cultripes platycephalus

maculosus

maint

Heleioporus | | 0 evel

Leciiriodus | ] 0 Heicheri

Limnodynastes i 5 0 convexiusenlus

dumeriti

spenceri

LSTA NSES

OOPNALUES

Litoria ia 19 adelaidensis alboeutiata

bicolor chloris

cuerulea dahlii

coplandi pallida

dentata foruieri

ewingl

fallax

Jfreycineti

gracilenta

Mmernus

Peron

raniforms

rohit

rubella

Megistolous l | 0 henerius

Mixophyes l I 0 fascialatius

schevill

Neobhatrachus L: 2 pictus a

sutor

Noladen i} 2 0) bennett

melanoscapans

Psendophrene i} 2 0 hibront

guenthert

Ranidellu 1 2 0 ripatia

signifera

Rheobutrach ts I ] situs 0

Taudacivluys I Ps 0 UCUEIFOSTFIS

cdinernus

Uperolera ft. 2 () inyadata

trachyderina

H = Hylidae; b= Leprodactylidae

48 M. J. TYLER & C. A. MILLER

connections. None of the preparations examined

demonstrated any evident intermediate stages,

documenting the change between the presence of

the intercellular connections and the formation of

the final surface architecture.

Discussion

The results demonstrate that two forms of surface

architecture of the squamous epithelium of the

dorsal epidermis occur in the species of Australian

hylid and leptodactylid frogs examined.

The results also indicate that there is no

significant variation within species, and that the

changes associated with ecdysis reveal at the surface

intercellular connections comparable to those

reported by Kaltenbach er al. (1980). We have not

observed amongst the preparations examined any

surface architecture not resembling the two types

reported above. We have no evidence to suggest that

either form of cell surface is an intermediate stage

in the cytogenesis or cytomorphosis of the other,

but we cannot exclude the possibility of an age-

related change taking place so rapidly that it was

not represented in our large sample.

Of the nine polytypic genera, two (Cyclorana and

Litoria) include some species with cellular

microridges and others with cell surfaces in the form

of a reticulum. Nevertheless in the majority of

species in each genus the surface bears microridges.

This division separates species that are considered

phylogenetically closely related: the reticular

surfaced L. alboguttata and L. dahlii were placed

by Tyler & Davies (1978) in a species-group with the

microridge surfaced L. raniformis. Similarly the

reticular surfaced L. pallida and L. tornieri have

been shown by Davies, Martin & Watson (1983) to

be very closely related to the microridged L. inermis.

Of the two reticular surfaced Cyclorana species, C.

platycephalus is certainly phylogenetically remote

from all congeners, but C. /ongipes is very closely

related to the microridge surfaced C. maculosus.

Because of the difference in these last two species

which also occupy identical habitats and have

similar life histories and habits, it is apparent that

the difference is not associated with any major

ecological shift, nor is of any systematic

significance. The fundamental similarity amongst

diverse vertebrate classes precludes any obvious

common function beyond that of increasing the

surface area. Particularly elaborate architecture has

been observed on the surface of the oesophagus of

the rhesus monkey and rat cornea (Andrews, 1976),

That author noted that the microplical and

interplical grooves would serve to hold mucus,

Whereas this structure may have a_ protective

lubricating function in areas such as the oesophagus

where the cells are alive, on the outer surface of frog

skin it is more likely to improve the efficiency of

gas exchange, simply by increasing the available

surface area.

Acknowledgments

This study was undertaken with the support of

a grant to M.J.T. from the Australian Research

Grants Scheme. Technical advice received from the

staff of the University of Adelaide Electron Micro-

scope Centre is greatly appreciated. We are also

grateful to P. G. Kempster for assistance in the

preparation of the figures, and to the referees of

the manuscript for constructive comments.

References

ANDREWS, P. M. (1976) Microplicae: characteristic ridge-

like folds of the plasmalemma. J. Cell Biol. 68, 420-429,

Crook, G. A. & Tyier, M. J. (1981) Structure and

function of the tibial gland of the Australian frog

Limnodynastes dumerili Peters. Trans. R. Soc. S. Aust.

105, 49-52.

Daty, J. W. & Myers, C. W. (1967) Toxicity of

Panamanian poison frogs (Dendrobates): some

biological and chemical aspects. Science 156(3777),

970-973,

Davies, M., MArTIN, A. A. & Watson, G. F. (1983)

Redefinition of the Litoria latopalmata species group

(Anura: Hylidae). Trans. R. Soc. S. Aust, 107, 87-108.

Evias, H. & SHapiro, J. (1957) Histology of the skin of

some toads and frogs. Amer. Mus. Novit. (1819), 1-27.

ELKAN, E. (1968) Mucopolysaccharides in the anuran

defence against desiccation. J. Zool, 155, 19-53.

FAHRENBACH, W. H. & KNUTSON, D. D. (1975) Surface

adaptations of the vertebrate epidermis to friction, J.

Investigative Dermatol. 65, 39-44.

Hawkes, J. W. (1974) The structure of fish skin. L.

General organization, Ce// Tiss, Res. 149, 147-158.

JOHNSON, C. R. (1969) Water absorption response of

some Australian anurans. Herpetologica 25, 171-172.

KALTENBACH, J. C., HARDING, C, V. & SUSAN, S. (1980)

Surface ultrastructure of the cornea and adjacent

epidermis during metamorphosis of Rana pipiens: a

scanning electron microscopic study. J. Morph. 166,

323-335,

LintywHite, H. B. (1971) Thermal modulation of

cutaneous mucus discharge of evaporative water loss

in the frog, Rana catesbeiana. Z. vergl. Physiologie 73,

84-104.

Low, B. S. (1972) Evidence from parotoid-gland

secretions. Jn: W. EF. Blair (Ed.) ‘Evolution in the genus

Bufo’. pp. 244-264. (University of Texas Press: Austin).

Stitte, W. T. (1958) The water absorption response of

an anuran. Copeia (1958), 217-218.

Tyter, M. J. & Davies, M. (1978) Species groups in the

Australopapuan hylid frog genus Litoria Tschudi. Aus?

J. Zool. Suppl. Ser. (63), 1-47.

A NEW CRETACEOUS CHIMAERID (PISCES: HOLOCEPHALD FROM

SOUTH AUSTRALIA

BY J. A. LONG

Summary

Edaphon eyrensis sp. nov. is described from a single left mandibular toothplate from the Aptian

Bulldog Shale, west of Bopeechee Siding, northern South Australia. E. eyrensis is characterised by

an angular oral margin and by the shape and arrangement of the fourth tritors. Comparisons are

made with other Cretaceous and Tertiary chimaerid dentitions.

A NEW CRETACEOUS CHIMAERID (PISCES: HOLOCEPHALI FROM SOUTH

AUSTRALIA

by J. A. LONG*

Summary

1 Ht,

Wee OS Taste HOOT) AY SA, 8 Sule, (SS

oN (URSA tow Crelieeous ebipierid (Pisces: Floloceplald tron Soul Austialin. Trea. AL

Edephodon eyrensis sp. noy, is described from a single lel) mandibular toothplate from the Aptian

Bulldog Shale, west of Bopecchee Sidiny, northern South Australia, £, evremsis is characterised by an anpular

oral margin und by the shape and arrangement of the four wWitors. Coniparisons are made wwillt other

Cretaccous and Tertiary: vhimaerid demitions.

Kry Warps Pisees, Holocephali, Cretaceous, Sauth Australia, new species, Edaplon evrensiy,

Hei Pata,

Introduction

rhe chimaerids reached a peak of diversity

during (he Mesozoic, and though tumerous genera

are Known from the Jurassic and Cretaceous of

North America, USSR, and Europe (Newlon,

1876, Woodwurd, 1892, 1912; Thussukoll, 1912;

Obruchey, 1964) the only ones described from

Australia are Edaphodon sweetl, & inirabilis and

Ischvodus mortoni from the Tertiary (Chapman &

Pritchard, 1907; Chapman & Cudmore, 1924), The

renus Adaphodon ranges from Early Cretaceous 10

Pliovene, with most species being Late Cretaceous

in age. The specimen described here was found on

the floor of Lake Phibbs, just south of Lake Lyre,

and undoubtedly came from outcrops of the nearby

Aptian Bulldog Shale (Ludbrook, 1966). Li is

therefore not only the first record of a fossil

chimaerid from the Mesozoic of Australia (and the

only chimeerid fossil from South Australia) bot also

one Of (he earliest species of Adaphodon.

Cretaceous chimaerid toothplates are also known

from New Zealund but these belong to

Cullorhynchus hector’ and lsehyodus brevirostris

(Newton, 1876).

Systematic Palucontology

CLASS HOLOCEPHALI

ORDER CHIMAERIDA

SUBORDER CHIMAEROIDE!

FAMILY CHIMAERIDAE

GENUS EDAPHODON Buckland, 1838

Type species. Chinaera manielli Buckland (835,

Cretaceous ol England (Ward, 1973).

Remarks; Whe specimen is reterred to Edaphodon

because of its broad syimphysial facel which

" Depurtnient af Geology, The Australian National

Universis, POL Bos 4, Canberra City, ACT, Austeatlay

2601

expands in breadth posteriorly, the number and

adbrangement of its tritors and the absence of a

thickening on ils Outer face (Hussakoff, 1912.

p. 202),

kdaphodon eyrensis sp. nov,

FIGs 1, 2, 3f,

1982 A Jarue laothphie of Lekplodoan Dong, py. 7)

Wsd Eduplodon sp. bong & lirner p24,

Etymology, After Lake Eyre, near where the

specimen was found.

Didgnosis. An Edaphodon having. a mandibulat

Loothplate twice as Jong as broad with four tritors:

of which the two posterior and the single

syimphysial ones are large, and the lateral median

one is small, Oral margin angular with anterior ends

of outer and lateral median tritors forming right

angles.

Material. Only one Specitien, (he holotype SAM

P24770, maximum length =10 mm,

Occurrence. From the floor ol the Lake Phibbs

approximately 21 km west of southwest from

Bopecechee Siding, west of Marree, South Australia.

Lower Cretaceous Bulldog Shale (Aptian),

Deseription, The bone is weathered, with the

semidentine of the tritors being chalky. Despite this

the overall shape of (he toothplate and outline of

the tritors are well preserved.

The oral margin of the mandibular toothplate

(Bigs. 1, 2, 3) is quite angular as each of the two

lateral (ritors (outer titer, OT, lateral median tritar,

LMT) form fight angles with the bone in between

them. Overall the toorhplate is rhomboidal with its

breadth exactly half the length. The two large tritors

in the posterion hall (outer tritar, OT; mesial tritor,

M/S) are each abouts quarter as broad us long and

i contact for close to hall their lengths. The median

50 J. A. LONG

Fig. 1. Edaphodon eyrensis sp. nov. Holotype, SAM P24770. Aptian Bulldog Shale, South Australia. A, oral view;

B, oral margin; C, aboral view. Natural size.

NEW CRETACEOUS CHIMALRILD Si

lateral tritor (LMT) is situated ventral to the mesial

tritor and in cross-section is disposed with its long

axis at right angles to the mesial tritor. The exposed

anterior end of the lateral median tritor narrows

to a point. The symphysial tritor (ST) is broader

anteriorly than the other tritors, but is relatively thin

in cross-section, Although the aboral surface is

poorly preserved it is strongly convex anteriorly

becoming flatter in the posterior half. The

symphysial facet (sf) is broadest posteriorly, with

the ventral margin being gently curved.

Discussion

Toothplates of chimaerids show a wide range of

shapes and tritorial arrangements which led early

workers to confuse various genera and species

(Hussakoff, 1912). A series of mandibular

toothplates of Edaphodon mirificus Leidy, 1856

from the Cretaceous of North America show

changes attributable to growth and intraspecific

variation. With increasing age the mandibular beak

becomes broader and the median lateral tritor

becomes more posteriorly situated (Hussakoff,

LMT

hig. 2. A, Eduphodor eyrensis sp. nov., Cretaceous, South Australia. Holotype in oral view. B, E. ssveeri Chapman

& Pritchard, Mioeene, Victoria. Composite restoration of left miundibular toothplate based mainly on NMV P160769.

LMT, lateral median tritor; MT, MT1, M2, mesial tritors; OM, oral margin; OT, outer tritor: sf, symphysial facet;

ST, symphysial tritor

52 J

1912, Fig. 6). This indicates that the basis for

comparing the mandibular toothplates of different

species relies on overall shape and the relative size

and position of all the tritors. Tritors are often

damaged or worn on fossil toothplates and

therefore some variation in the shape of exposed

tritorial surfaces is expected (Fig. 3).

Fig. 3. Comparison of mandibular toothplates for various

species of Edaphodon. A, E. mirificus; B, E.

stenobryus, C, E. agassizi; D, E. latigerus; E, E.

bucklandi; F, E. eyrensis sp. nov; G, E. sedgwicki.

A-D, G from Hussakoff, 1912. E from Casier, 1966. Not

to scale.

The other Australian species of Edaphodon are

E. sweeti Chapman & Pritchard, 1907 and £&.

mirabilis Chapman & Cudmore, 1924 both from the

Miocene and Pliocene of Victoria. Fig. 2 shows a

comparison between E. eyrensis and E. sweeti. The

mandibular toothplates of E. sweeti described by

Chapman & Pritchard (1907) were not complete, but

good material has since been found from the

Pliocene Grange Burn Coquina, Victoria, and the

. A. LONG

shape of the toothplate and arrangement of tritors

can be restored (NMV P160769). E. sweeri differs

from £. eyrensis in having a composite outer tritor

with 3-4 bony ridges dividing it; two large mesial

tritors, more slender form, and proportionately

longer symphysial facet. The oral margin is quite

angular, as in E. eyrensis, but anteriorly the

symphysial beak is more elongated and is concave

on the aboral surface. E. mirabilis is known only

from palatine toothplates, which have a long,

slender form but are quite robust in overall structure

(Chapman & Cudmore, 1924). It is unlikely that the

short mandibular toothplate of E. eyrensis belongs

with this type of palatine plate.

Edaphodon eyrensis has a broader, shorter

mandibular toothplate than most species (Fig. 3),

except for E. stenobryus Cope 1875, from the

Cretaceous of North America, which has a breadth/

length ratio around 0.7 (Fig. 3b). Some species from

the Cretaceous of Europe and North America also

have rostrally produced beaks (E. sedgwicki Agassiz

1843, E. latigerus Cope 1869; Fig. 3D, G; E.

mantelli, Woodward, 1912). In the development of

large, almost equidimensional outer and mesial

tritors E, eyrensis resembles E. sedgwicki (Fig. 3G),

which is the only other species older than E. eyrensis

as it occurs in the Neocomian of England

(Hussakoff, 1912). This would suggest that the

simple tritorial arrangement of these two species

is a primitive pattern for the genus.

Younger species have mandibular toothplates in

a variety of forms with many having additional

tritors present (e.g. &. bucklandi, E. agassizi, E.

sweeti). All of the Edaphodon dentition types stem

from the primary strengthening of the mandibular

symphysis by the widening of the symphysial facet,

which characterises the genus. Extreme thickening

of the mandibular symphysis and robustness of the

toothplates is seen in the Upper Jurassic

Pachymylus (Woodward, 1892), a possible precursor

to the line of Edaphodon.

Acknowledgments

Thanks to Prof, J. Warren (Monash University)

for drawing my attention to the specimen and

kindly allowing me to work on it, and to Prof, Ken

Campbell (Australian National University) and Dr

Ken McNamara (West Australian Museum) for

critically reading the manuscript and offering

helpful comments. Dr Tim Flannery (Australian

Museum) is thanked for the loan of Edaphodon

sweeti material which he diligently collected from

Grange Burn. Mr Neville Pledge (South Australian

Museum) gave helpful information about the

geology of the fossil locality. Thanks also to Mrs

NEW CRETACEFOUS CHIMAPRID 53

Mary MacDougall for typing the manuscript. This

work was carried out under the tenure of a

Rothmans Postdoctoral Fellowship in the Geology

Department of the Australian National University.

References

AcAssiz, L. J. R. (1843) Recherches sur les poissons

fossiles. Neuchatel and Soleive, 5 Vols., 1-188.

BuckLAND, W. (1835) A notice on the fossil beaks of four

extinct species of fishes, referable to the genus

Chimaera, which occur in the Oolitic and Cretaceous

formations of England. Proc. Geol. Soc. Lond., 2,

205-206.

— (1838) On the discovery of fossil fishes in the

Bagshot sands at Goldworth Hill, 4 miles north of

Guildford. /bid., 2, 687-688.

Casipr, E. (1964) Fauna ichthyologique du London Clay.

Brit. Mus, (Nat. Hist.), London, 496 pp.

CHAPMAN, F., & Cupmore, F. A. (1924) Some Cainozoic

fish remains with a revision of the group. Proc. R. Soc.

Viei., 36, 107-162.

& PrevcHArp, G. B. (1907) Fossil fish remains

from the Tertiaries of Australia. /hid., 20, 59-75.

Cope, E. D. (1869) Descriptions of some extinct fishes

previously unknown. Proc. Boston Soc. Nat. Hist., 12,

310-317.

Hussakorr, L. (1912) The Cretaceous chimaeroids of

North America, Bull, Aim. Mus, Nat. Hist., 31, 195-228,

Leipy, J. (1856) Notice of remains of extinct vertebrated

animals of New Jersey, collected by Prof. Cook of the

State Geological Survey under the direction of Dr. W.

Kitchell. Proc. Acad. Nat. Sci, Philadelphia, 8, 220-221,

Lona, J. A. (1982) The history of fishes on the Australian

continent, Jn P. V. Rich & E. Thompson (Eds), “The

Fossil Vertebrate Record of Australasia’, pp. SI-85.

(Monash University Off-set Printing Unit; Melbourne).

& TURNER, S. (1984) A checklist and bibliography

of Australian fossil fishes, /a M, Archer & G, Clayton

(Eds), ‘Vertebrate Zoogeography and Evolution in

Australasia’, pp. 235-254. (Hesperian Press: Perth).

Lubprook, N. (1966) Cretaceous biostratigraphy of the

Great Artesian Basin, South Australia. Geol. Szerv. S.

Aust, Bull. 40, 223 pp.

NEWTON, E. T. (1876) On two chimaerid jaws from the

Lower Greensand of New Zealand. Quart. J, Geol, Soc.,

Lond., 32, 326-331,

(878) Chimaeroid fishes of the British Cretaceous

rocks. Mens. Geol. Surv. ULK., Monogr. 4, 1-62,

OprucHEy, D. V. (1964) Subclass Holocephali

(Chimaeras). 7a Y. A. Orlov (Ed.), ‘Fundamentals of

Palaentology’. Vol, 11; Agnatha, Pisces, pp. 353-399,

Warp, D. J, (1973) The English Palaeogene Chimacroid

fishes, Proc. Geol. Assoc., 84, 315-330.

Woopwarp, A. 8. (1892) On some teeth of new

Chimaeroid fishes from the Oxford and Kimmeridge

clays of England. Ann, Mag, Nat. Hist, 10, 13-16.

(1912) The fossil fishes of the English Chalk.

Palaeontogr. Soc, Mongr, 264 pp,

DISTRIBUTION OF LERISTA TERDIGITATA (SAURIA: SCINCIDAE) IN

SOUTH AUSTRALIA

BY T. D. SCHWANER, A. EDWARDS & B. MILLER

Summary

Many recent new distributional records for reptiles in South Australia reflect inadequate field

surveys and poor documentation of the state’s herpetofauna. Here we report a significant range

extension of the fossorial, scincid lizard, Lerista terdigitata (Parker, 1926) in South Australia.

BRIEF COMMUNICATION

DISTRIBUTION OF LERISTA TERDIGITATA (SAURIA: SCINCIDAE) IN SOUTH

AUSTRALIA

Many recent new distributional records for reptiles in

South Australia!’ reflect inadequate field surveys and

poor documentation of the state’s herpetofauna. Here we

130° 134° 138"

eu, Morris

Io Kolinahstd

. ina

~ e

os R °

Qu. 4, ._- 400

Vig. 1. Distribution of Lerista terdigitata. Area under the

dashed line in the inset denotes the previously recognized

range of the species in Australia, Solid circles denote

S.A. Museum specimens.

‘Miller, B. & Schwaner, T. D. (1982). Trans. R. Soe. 8.

Aust. 106, 79-80,

*Schwaner, T. D. & Miller, B. (1984), (bid. 108, 215-216.

ISchwaner, T. D. & Miller, B. (1984). /bid. 108, 217-218.

report a significant range extension of the fossorial, scincid

lizard, Lerista terdigitata (Parker, 1926), in South Austraha.

The range of L. terdigitata was “islands, coast and

hinterland of the Great Australian Bight’ (Fig. 1).

However, specimens recently collected from near Billa

Kalina Hstd, 29°55'S, 136°11'E (SAM R20991-92, R21022),

and from the Mt Morris foothills, near Amata aerodrome,

26°08 'S, 131°05'E (R25883), in the Musgrave Ranges in

the far north-west of S, Aust., extend the distribution of

L. terdigitata about 600 km N of its previously recognized

range.

Although the Mt Morris specimen (a male, SVL=

51.5 mm, TL=64.0 mm) has the normal sealation of L.

terdigitata from more coastal localities*’ the pattern 1s

markedly different (Fig. 2). Bold stripes replace the

longitudinal series of spots on the dorsum, and much

larger spots (not flecks) are found on the venter, Specimens

from Billa Kalina Hstd are patterned like the coastal forms.

Thus, it is not known whether there is a grade of patterns

between the two populations.

The occurrence of L. ferdigitata in the Musgrave Ranges

suggests that the species also may be found in adjacent

areas of the Northern Territory and Western Australia.

The presence of yet another distributional record in the

extreme northwest of S. Aust.” illustrates the need for

herpetofaunal surveys in this region.

We thank Peter Bird for specimens from Billa Kalina

Hstd, and Wolfgang Zeidler for the specimen from Mt

Morris. Roman Ruchle provided Fig. 2, and Lila Schwaner

and Diana Massacci typed the manuscript.

‘Copper, H. G. (1983). “The‘Reptiles and Amphibians of

Austraha!” A. H. & A, W. Reed, Sydney.

‘Storr, G. M., Smith, L. A. & Johnstone, R. E. (1981).

“Lizards of Western Australia, [. Skinks. University of

Western Australia Press & Western Australian Museum:

Perth.

Fig. 2. Lerista terdigitata, (R20992, upper) from Billa Kalina Hstd, and (R25883, lower) from Mt Morris.

T. D. SCHWANER & A. EDWARDS, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000, and B.

MILLER, 10 Yarrow Crescent, Hope Valley, S. Aust. 5090.

TIDES OF THE ONKAPARINGA ESTUARY, SOUTH AUSTRALIA

BY J. A. SMITH

Summary

The Onkaparinga estuary is situated approximately 30 km south of Adelaide on Fleurieu Peninsula,

and extends some 10.5 km from its mouth at Pt Noarlunga on Gulf St Vincent to the township of

Old Noarlunga. In recent times the estuary has come under renewed interest as a recreation site

which has led to the dredging of sections in Pt Noarlunga. There is, however, some concern for the

environmental effects of extending these works to other regions.

BRILE COMMUNICATION

TIDES OF THE ONKAPARINGA ESTUARY, SOUTH AUSTRALIA

The Onkaparinga estuary is situated approximately

30 km south of Adelaide on Vleuricu Peninsula, and

extends some 10.5 km from tts mouth at Pt Noarlunga

on Gull St Vincent to the township of Old Noarlunga.

In revent times the estuary has come under renewed interest

as a recreauion site which has led to the dredging of

sections in Pt Noarlunga. There is, however, some concern

for the environmental effects of extending these works to

other regions,

During the period May to July 1983 pwo pressure sensing

lide gauzes were deployed in the estuary (Pig. 1), one

S00 m upstream from the mouth (A) and the other | km

downstream trom the head (B). This enabled a continuous

record of levels to be obtained at cach end of the estuary

over (his period (hig. 2). Water heights were taken from

each record alt hourly intervals and used to derive tidal

constams by a least squares fit technique. The estuary

constants were then used ina comparison with constants

derived for Pt Adelaide (Quter Harbor) using data from

(he same observation period.

The constants for the four major primary constituents

(Table 1) show a veneral decrease in amplitude and increase

in phase lay as the tidal wave progresses from the gull

“165

-15

W 15 JUNE 41983

1g8'E

| r i

i [ \

ag / GUL Ef

) r¢ ne 1c Oo PST VINCENT

PorRT { ( f v2 r J al BS

noarmLunba\\)) \ = 4

mh ‘e

Vif) _)

Vi / J

( 13 Af i

\ { fr

D SN

rd ORKAPARINGS \

| RIVER ]

| ey i

r nM i \ |

‘6

| oLe i)

} NOARLUNGA _

| (Wu yy)

N ; NS

\ —

lig. 1. Onkaparinga Estuary, South Australia; A, Site of

tide gauge near mouth; B, Site of tide gauge near head;

C, Shallow region. Mean river depth given in om.

(Lh4-2.5).

23 27

Fie. 2. Waiter levels for period Tth-271h June 1983; (a) Old Noarlunga (Bin Fig, 1); (b) Port Noarlunga (A in Fig, 1):

nan

and up the estuary. On the other hand the two major

secondary (shallow water) constituents show an increase

in amplitude and no obvious trend in the phase lag under

the same conditions.

TABLE 1. Tidal constituents for the Onkaparinga Estuary

and Port Adelaide.

CONSTITUENT GULF! MOUTH? ~~ -HEAD*

A g g Ag

Ol 17.3 19.1 11.8 69.0 9.8 94.3

KI 25.7 47.3 17.8 82.5 12.7 121.1

M2 51.7 106.0 19.8 124.1 11.1 176.1

82 50.0 173.2 19.0 190.4 12.8 248.0

Mk3 0.7 158.6 3.9 97.4 4.2 243.4

MS4 0.1 24.4 4.1 258.1 2.5 360.0

'Pt Adelaide (Outer Harbor); “Pt Noarlunga (A on

Fig. 1); ‘Old Noarlunga (B on Fig. 1) A Amplitude of

tidal constituent (em) g Phase lag of tidal constituent (°).

The times for these constituents to travel the length of

the estuary were also calculated from the respective phase

differences, (Table 2), and found to be similar for the

primary constituents. The values were, however, much

larger than expected for progression with a shallow water

wave speed ( W@H, where g is the acceleration of gravity

and H the mean water depth),' and demonstrated the

importance of friction in the estuary system.

TABLE 2. Wave travel times

CONSTITUENT ZB, TT.

Ol 25.3 1.81

Kl 38.6 2.57

M2 52.0 1.79

82 57.6 1.92

g,-g, The phase difference between stations A and B

("), ET. The wave travel time = g.-g), (hrs), @ The wave

-2

speed (°/msh). o

This observation prompted a theoretical study using a

finite difference numerical model to simulate the estuary

flow.?

The results from the simulation suggest that the region

extending from 1.5 to 2.5 km upstream from the mouth

forms a critical section (C in Fig. 1) with respect to the

tidal propagation. This is a particularly shallow region

of the estuary with only a narrow channel connecting the

deeper waters up and downstream at low tide, and with

parts exposed even at high tide. Flow within this section

appears to be largely friction dominated and, although

it forms only 10% of the estuary length, it accounts for

> 60% of the wave travel time and > 30% of the wave

attenuation. The model results also indicate that the head

of the estuary has mean water level approx. 7 cm higher

than the mouth—also due mainly to the frictional effects

in this very shallow section.

On the basis of these results several predictions can be

made about the possible effects of further works in the

estuary. In particular, dredging in regions other than the

critical section discussed above would probably have little

effect on the water flow or levels in the estuary, but even

minor changes to this critical section would result in

significant changes to the water movements. For example,

in one particular model run this section was ‘numerically’

deepened by extending the already dredged channel

downstream through this region. The results indicated this

would increase the tidal range by > 20%, decrease the

travel time by > 30% and decrease the mean water level

at the head by 3-4 cm. Although these changes do not

appear great, the increased tidal range with its associated

stronger tidal currents and lower, low tide (decreased by

15 cm at the spring tide) could have damaging effects on

the seagrass and other vegetation that presently exists in

the estuary.

' Dyer, K. R. (1973). Estuaries: A Physical Introduction.

(Wiley: London).

> Smith, J. A. (1983). B.Sc.(Hons) thesis, Flinders Univ.

S. Aust. (unpublished).

J. A. SMITH, The Flinders Institute for Atmospheric and Marine Sciences, The Flinders University of South Australia,

Bedford Park, S. Aust. 5042.

ADDITIONS TO THE FISH FAUNA OF SOUTH AUSTRALIA

BY C. J. M. GLOVER

Summary

Four marine species (Pristis sp., Ophisurus serpens, Hyporhamphus regularis regularis, Scorpis

lineolatus) are recorded from South Australia for the first time. A freshwater species (Amniataba

percoides) is recorded from the State for the first time on the basis of a known collection.

BRIEF COMMUNICATION

ADDITIONS TO THE FISH FAUNA OF SOUTH AUSTRALIA

Four marine species (Prislis sp., Ophisurus serpens,

Hyporhamphus regularis regularis, Scorpis lineolatus) are

recorded from South Australia for the first lime. A

freshwater species (A ninialaba percoides) is recorded [rom

the State for the first time on the basis of a known

collection.

Three of the marine species are essentially temperate

water forms Whose presence in South Australian waters

is probably permanent. The occurrence of the other warm

waler Species (Pristis sp.) is unexpected and it evidently

constitutes a vagrant (see earlier reports!).

Amniataba perevides Was been cecorded widely in

inland waters of northern Australia. It was listed for South

Australia in the 1920’s*?, but (hat record was neither

confirmed by any known collection nor sustained by

subsequent authors.

All specimens are deposited in the South Australian

Museum (SAM).

Vig. 1. Snout of Pristis sp. (P ztisron !), SAM F206d,

TL Qotal length) 399 mm

Family Pristidae. Prixtis sp. (probably PB ziysron Bleeker,

ISI).

The rostral process (Fig. |) ola specimen caught 16 km

offshore from Glenelg, (approx. 34° 59'S, 138° 20°B), in

}936, by E.G. Kelly, is registered in the South Australian

Museum. The intact specimen would have been about

1 300 rim total leneth,

The rostral process has 29 pairs of teeth. This is

consisten| with thal of PB zi/sron, one of the three pnistid

species recorded in Australian waters’.

Pristids are found mainly in the world's tropical and

sub-tropical waters’. In Australian P gi/sron has been

recorded in coastal, estuarine and fresh inland waters of

Queensland, New South Wales and western and northern

Western Australia", its range reportedly extends to the

East Indies, India and Sri Lanka’.

The present record is the first for the family Pristidae

from Australia’s southern waters,

Fig. 2. Ophisurus serpens, SAM T4566, TL 885 mm.

Family Ophichthidae. Ophisurus serpens (Linnaeus, 1758),

A specimen (Pig. 2) was collected from a rock pool at

Coobowie (approx. 35° 03'S, 137° 44'B), on 7.1.1970, by

M. J. Stanton

©. serpens has been recorded previously in Australia

from off N.SW., Victoria, Tasmania and W.A.*. It is

recorded elsewhere in the Indo-Pacific and eastern Atlantic

Oceans, and in the Mediterranean’,

Family Hemiramphidae. Ayporhamphus regularis

regularis (Giinther, 1866)

Fifteen specimens (SAM 4803) were netled in Angas

Inlet, off the Port River (approx. 34° 48'S, 138° 32'B),

on 301.1984, by S. Aust. Dept of Fisheries ol'Gvers. Two

additional specimens (Fig. 3) were hooked at the same

locality on 23.ix.1984 by T, BE. Lloyd,

Hi, regularis occurs only around the southern half of

the Australian continent, where it has been recorded from

southern Qld, N.SW., Vic. and south-western W.A,!".

Two distincL sub-species are recognised, one on the

continent’s south-east coast (H. regularis ardelia), the

other on its south-west coast (#.. regularis revularis)'”.

M, F. Goman advised me (pers. comm., 1984) thar the

S. Aust, specimens are #7. regulariy reeularis. This record

therefore represents the most easterly documented

oecurrence of the western sub-spectes.

— LESS

KF Hyporhamphugs regularis regularis. SAM '4831

ig. 3.

TL 285 mm {smallest specimen)

Fig. 4. Scorpis lineolatus. SAM F4761. TL 316 mm.

Family Scorpidae. Scorpis lineolatus Kner, 1865.

Two specimens were captured by speargun between

Victor Harbor and Seal Island (Seal Rock), (approx. 35°

34'S, 138° 38'E), on 31.xii.1983, by competitors in a

national spearfishing competition. One specimen is

illustrated (Fig. 4): the other is SAM F4762.

S. lineolatus has been recorded only in Australia, from

Qld, N.SW., Vic., Tas. and at Lord Howe Island.°.

The present S. Aust. record represents the species’ most

westerly documented occurrence.

Family Teraponidae Amniataba percoides (Gtinther, 1864).

Two specimens (Fig. 5) were netted at the Neales River

crossing on the main road south-east of Oodnadatta

1Glover, C. J. M. (1984). Trans. R. Soc. S. Aust. 108(2),

133-135,

2waite, E. R. (1921). Rec. S. Aust. Mus. 2(1), 1-208.

3Waite, E. R. (1923). “The Fishes of South Australia?’

(Govt Printer: Adelaide).

4Munro, I. S. R. (1956). Fisheries Newsl. 15(9), 15-18.

‘Lindberg, G. U. (1974), Fishes of the World. (J. Wiley

& Sons: New York).

‘Whitley, G. P. (1948). Fisheries Bulletin No. 2, 1-35.

(Western Australian Fisheries Department: Perth).

7Marshall, T. C. (1964). Fishes of the Great Barrier Reef

and coastal waters of Queensland. (Angus & Robertson:

Sydney).

Fig. 5.

(largest specimen).

Amniataba percoides. SAM F4792. TL 54 mm

(approx. 27° 54'S, 135° 46'E), on 3.1984, by a South

Australian Museum party.

A. percoides is endemic to Australia where it is recorded

widely dispersed in inland waters of Qld, Northern

Territory and W.A.''!2, An early record from S, Aust.27

was not substantiated by any hitherto known collection.

The present record is the species’ most southerly

documented occurrence!?"!,

Those persons mentioned are thanked for collecting and

donating the specimens to the South Australian Museum.

Thanks are also extended to Dr G. K. Jones (S. Aust. Dept.

of Fisheries) for forwarding portion of the H. regularis

regularis and S. lineolatus collections, and Dr M. F.

Gomon (Museum of Victoria) for identifying H. regularis

regularis.

‘ Last, P. R., Scott, E. O. G., & Talbot, FH. (1983).

Fishes of Tasmania. (Tasmanian Fisheries Development

Authority: Hobart).

Wheeler, A. (1979). Fishes of the World. (Ferndale:

London),

Collette, B. B. (1974). Rec. Aust. Mus, 29(2), 11-105.

"Lake, J. S. (1978). Australian Freshwater Fishes.

(Nelson: Melbourne).

Merrick, J. R. & Schmida, G. E. (1984), Australian

Freshwater Fishes, (J. R. Merrick: North Ryde, N.SW.).

'3Allen, G. R. (1982). Inland Fishes of Western Australia.

(Western Australian Museum: Perth).

Cc. J. M. GLOVER, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000

STUDIES ON SOUTHERN AUSTRALIAN ABALONE (GENUS HALIOTIS)

V. SPAWNING, SETTLEMENT AND EARLY GROWTH OF H. SCALARIS

BY S. A. SHEPHERD, P. S. CLARKSON & J. A. TURNER

Summary

This note describes the spawning cycle of Haliotis scalaris Leach and the settlement density and

growth of juveniles of the species at West Island (35°37°00°S, 138°35°00°E), South Australia.

This spawning cycle differs markedly from that previously described for the species at Tiparra

Reef, South Australia and is therefore of considerable interest.

BRIEF COMMUNICATION

STUDIE

SPAWNING, SETTLEM

This noe deseribes the spawning eycle of Huliotis

sedlarty Leach and the setthement density and growth of

juveniles of the species at West Island (35°37 00'S,

38°35 00"L), South Australia. This spawning cycle differs

markedly from that previously deseribed for the species

at lipara Reet', Sour Australia and is therefore of

considerable interest.

Samples of Y-12 sexually mature female A, scalars

65-80 tim long were collected at Abalone Cove, West 1,

apabout monthly defervals from bebruary 1983-July 1984,

Vine cotire visceral mass was preserved in 10% formalin

and sea-warer and later seetioned in the laboratory. Cross-

sections of the gonad and digestive gland were traced on

transparent plastic, the cut-out sections weighed, and the

respective areas of gonad and digestive gland calculated.

A pvonad index was then calculated with the formula

Gonud ides 100 (Area of gonad), (Total area of

section). The annual reproductive cycle af this species at

West 1. proved to be synehronous within the population

so that measurement of oocyte-ova diameters to

distinguish the stapes of the reproductive evele was

unnecessary (see Shepherd & Laws! for details of the

method).

Sea surface lomperature data were obtained with a

mercury thermometer ar about monthly intervals at Wes!

{

A hitnd lens, designed tor ise underwater with about

J© magnification (the optics are described by Shepherd

& Turner in prep.) was used to search for MH. sealariy on

eristose coralling alwal substrate, (he preferred substrate

for setdement of this species (Shepherd & Turner in prep),

Searches were done for 60-100 minutes at about monthly

mtcevals inthe boulder habitat ar West 1, at 4-5 m depth

and the lengths of all individuals to about 25 mm long

recorded to 0.) mim.

bi 1 Gipped— inn Wionthly sen Surfice temperatures

(smoothed curves; (mid) monthly distribution of

tonid indices Winh standart errors; (lawer) density of

ainall AY. sewluris SS mim al study site,

ON SOUTHERN AUSTRALIAN ABALONE (GENUS HALIOT/S) Y.

INT AND EARLY GROWTH OF H. SCALARIS

Changes in the mean monthly gonad index of //,

sealaris trom February 1983-May 1984, together with sea

surface temperature data are given in Fig. tl. The index

increases from winter to mid-summer and deelines

ubruptly from late summer through the autumn. The

inercase indicates maturation and increase in size of

oocyles in the gonad, and the declitie of the index indicates

the onset of synchronous spawning in the population.

Spawning appears to have commenced later in 1983 Than

in 198%4.and to have been more complete In 1984 the gonad

never became completely spent but commenced to inercase

in size again in July.

Thus 4. sealaris bas a late summer to aulunin spawning

season (February-May) which is synchronous throughout

the population, and appears to begin at about the time

of asimum summer sea temperuture. This spawning evele

contiasts strikingly with the asynchronous cycle of At

scalaris at Tiparra Reet where spawning potentially occurs

Throughout the year,

15 tap AF

1 Mal BS

29 Mar 83

[| May Oa

a 2) be

wi OS =

i ize dd Gully Wo

ot ff -

ly Be

NUNE

HO Le

ot — oe

aI han BS

y —fw ae

ot WO bel Wa

| - = —_—o—_—

b 15

VENGTH fr

Vig. 2. Length frequency distributions for A. sealaris al

West Island from February 1983 to May 1984,

OF the various environmental factors that are known

to influence the spawning eyeles of abalone ep. wri

perature, photoperiod? and food abundance’ only the

liter shows a marked difference between the two sites.

Drill food algae are seasonally in shoct supply.al Tiparca

Reef but abundant throughout the year at West L, © but

this is unlikely to account for the differences in spawning

between the sites. Purther studies of AV, sealaris are

neeussiy, especially in Other parts of its geographic Hane,

to elucidate the problem. //, rubra Leach also shows

marked, but unexplained, differences in spawning cycle

herween these two sites.! The only other abalone species

thal we know of with a similarly variable spawning eycle

between localities is //aliatis rufescens Swainsan.””

The density of small H. scalaris (measured in mean

numbers of individuals <5 mm recorded per 15 min

searching time is given in Fig. 1. Maximum settlement

apparently occurred in March 1983. The relatively high

densities recorded from July to September 1983 are of

larger individuals (3-5 mm) and do not indicate recent

settlement.

20 }

7 Y=2.99 + 0.0043 X +0.000087 x? Ks

22 R= 099 a

: AY

io} 7

Z 10 4

5 + 4+

5 ; pe

| part

MAM J J A S ON DJ FMAM

1983 1984

Fig. 3. Plot of mean length (with standard errors) of 1983

cohort of H. scalaris from March 1983 to May 1984.

A polynomial regression of best fit to the means is

shown.

Length frequency data from February 1983-May 1984

(Fig. 2) show that very small H. scalaris (1-3 mm long)

were present from February-June 1983 indicating

'Shepherd, S. A. & Laws, H. M. (1974). Aust. J. Mar.

Freshw. Res. 25, 49-62.

Kikuchi, 8. & Uki, N. (1974). Bull. Tohok. Reg. Fish.

Res. Lab. 33, 69-78.

‘Kim, Y. & Cho, C. (1976). Bull. Korean Fish. Soc. 9,

61-68.

4Giese, A. C. (1959). Ann. Rev. Physiol. 21, 547-576.

settlement in that period. This is in good agreement with

the spawning season described above. No small H. scalaris

were found in the 1984 spawning season, suggesting

settlement was very poor and not detectable.

Shepherd (in prep.) associated the similar poor

recruitment of H. /aevigata Donovan at West I. in 1984

with the lower maximum summer sea temperature in 1984

compared with 1983. The settlement failure of H. scalaris

in 1984 may also be associated with lower summer sea

temperatures in that year or with the incomplete spawning

or a combination of them.

The change in mean size of the 1983 cohort from March

1983-May 1984 (Fig. 2) enables an estimate to be made

of the growth of the cohort in the first year. The equation

of best fit empirically fitted to the data is given in Fig. 3.

On the basis that settlement occurred between | February

and 30 June 1983, a mean birth date for the cohort can

be fixed at 15 April 1983. From the regression (Fig. 3) the

mean length of the one year old animal is therefore about

18.5 mm. Comparison of this growth rate with that of

other haliotids® suggests that it is relatively rapid for a

species whose maximum size (at West I.) is only about

100 mm.

We are grateful to C. H. Deane for assistance in the

field and Dr W. G. Inglis for comments on the manuscript.

Funds supporting the study were received from Fishing

Industry Research Committee (FIRTA).

‘Shepherd, S. A. (1973). Aust. J. Mar. Freshw. Res. 24,

217-257.

°Giorgi, A. E. & De Martini, J. D. (1977). Calif. Fish and

Game 63, 80-94.

‘Lee, T. Y. (1974). Publ. Mar. Lab. Busan Fish. Coll. 7,

21-50.

‘Shepherd, S. A. & Hearn, W. S. (1983). Aust. J. Mar.

Freshw. Res. 34, 461-75.

S. A. SHEPHERD, P. S. CLARKSON and J. A. TURNER, Department of Fisheries, 135 Pirie Street, Adelaide,

S. Aust. 5000.

VOL. 109, PARTS 3 & 4

29 NOVEMBER, 1985

Transactions of the

Royal Society of South

Australia

Incorporated

Contents

Zeidler, W. A new species of crustacean (Syncarida: Anaspidacea: Koonungidae),

from sinkholes and caves in the south-east of South Australia - 63

Ludbrook, N. H. Trigonioididae (Mollusca: Bivalvia) from the Cretaceous of Lake Eyre

North, South Australia - - - - - - - - 77

Jackson, J. E. Larvae and pupae of Lectrides varians Mosely and Leptorussa

darlingtoni (Banks), (Trichoptera: Leptoceridae) - - - 83

Wells, A. Four new species of Hydroptilidae (Trichoptera) from the Alligator

Rivers region, Northern Territory = - - - - = 97

Davies, M., Mahony, M. & Roberts, J. D. A new species of Uperoleia (Anura:

Leptodactylidae) from the Pilbara Region, Western Australia - 103

Lansbury, I. The Australian Naucoridae (Insecta, Hemiptera-Heteroptera) with

description of a new species - - - - - - - 109

Cann, J. H. & Gostin, V. A. Coastal sedimentary facies and foraminiferal biofacies

of the St Kilda Formation at Port Gawler, South Australia - - 121

Spratt, D. M. Redescription of two trichurid nematode parasites of vertebrates in

Australia and. Papua New Guinea - - - - - - 143

Skinner S. Australian and New Zealand species of Elachista and Halothrix

(Elachistaceae, Phaeophyta) - - - - - - - 151

MeNamara, K. J. The spatangoid echinoid Linthia from the Late Eocene of southern

Australia - - - - - - - - - - 161

Lange, R. T. Spatial distributions of stocking intensity produced by sheepflocks

grazing Australian chenopod shrublands - - - - - 167

Brief Communications:

Sadler, T. & Pledge, N. S. The fossil sea urchin Fellaster incisa—an extension of range 175

Shepherd, S. A. & Hobbs, L. J. Age and growth of the blue-throated wrasse Pseudolabrus

tetricus - - - - = = 5 Z s 4 5 177

Shiel, R. J. & Koste, W. Records of rotifers epizoic on cladocerans from South Australia 179

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. 109, PART 3

A NEW SPECIES OF CRUSTACEAN (SYNCARIDA: ANASPIDACEA:

KOONUNGIDAE), FROM SINKHOLES AND CAVES IN THE SOUTH-EAST

OF SOUTH AUSTRALIA

BY WOLFGANG ZEIDLER

Summary

A new species of syncarid crustacean, Koonunga crenarum sp. nov. is described from sinkholes and

caves near Mt Gambier, South Australia. The species occurs in surface vegetation and to a depth of

40 m. Superficially the new species resembles K. cursor but females are at least twice as large,

reaching lengths exceeding 20 mm; the sternal process of the male petasma lacks a posterior

projection and the eyes are absent. A brief description of the development of the male petasma and

sensory organ on the antennule is given.

A NEW SPECIES OF CRUSTACEAN (SYNCARIDA: ANASPIDACEA: KOONUNGIDAE),

FROM SINKHOLES AND CAVES IN THE SOUTH-EAST OF SOUTH AUSTRALIA

by WOLFGANG ZEIDLER*

Summary

Zvipnnr, W, (1985) A new species of crustacean, (Syncarida: Anaspidacea; Koonungidae), from sinkholes

and eaves in the south-east of South Australia. Trans. R. Soc. S. Aust. 109(3), 63-75, 29 November, 1985.

A new species of synearid crustacean, Koonunga erenarum sp. nov. is described from sinkholes and

caves near Mt Gambier, South Australia. The species occurs i surface veyetation anc to a depth of 40 m,

Superfisially (he new species resembles A. curser but females are al least bwice as large, reaching lengths

exceeding: 20 nmi} the sternal process of the male petasma Jacks a posterior projection and the eves are

absent. A brief description of the development of rhe male petasma and sensory organ on the antennule

is given.

Kry Worns; Synearida, Anaspidacea, Koonunga erenarum sp. nov., South Australia, tiwanomty-

Introduction

In Australia, anaspidid synearid crustaceans are

known to occur throughout Tasmania and Victoria

and an interstitial species, Psammaspides wilfiamsi,

has been found in the New England area of New

South Wales (Schminke 1974), The Tasmanian

fauna is relatively diverse and to date eight species

have beer: described, Most of mainland Australia,

however, is too arid to support anaspidid syncarids

and besides P williams/ the only other described

species are Koantinge cursor Sayce, 1908 and

Stygocaris giselae Schminke, 1980, Several

undescribed species of Koonunga are known lo

accur in Victoria (Drummond 1959) and also on

King Island and in north-western ‘Tasmania

(Williams, W. D, 1974), Ivis likely that more species

remain to be discovered as aquatic habitats are

surveyed in more detail. A. cursor occurs

sporadically throughout southern Victoria, usually

in small permanent or semi-permanent swamps and

in pools in streams which flow only after heavy rains

(Drummond 1959) and it has recently been recorded

from similar habitats in north-western Tasmania (De

Deckker 1980). S. giselae is an interstitial species

and is only known trom the type locality, Battle

Point, Tambo River, Victoria (Schminke 1980),

The species described here was found in sinkholes

and caves in the Mt Gambier area, South Australia

(Fig, 1). It was first recorded by Zeidler (1983) being

the first record of a syncarid from South Australia.

It belongs to the family Koonungidae nd

superficially resembles A. cursor

Materials and Methods

Specimens Were collected from a number of

sinkholes and caves in the south-east of S, Aust

* South Australian Museum, North Terrace. Adelaide, 5,

Aust S000,

(Fig. 1) with a hand net from amongst algae and

aquatic plants near the surface or by entrapment

ina glass jar while scuba diving, The “1” nuniber

given with each Joeality refers to the identification

number used by the Cave Exploration Group of

South Australia to distinguish each cave and

sinkhole in the lower south-east. A total of 236

specimens (60 ¢ and 176 2) were vollected and

examined,

Specimen length is measured from the tip of the

rostrum to the tip of the telson. Specimen width

is measured as the width of the third pleonite

segment (usually the wides( segment), Pleon length

excludes the telson as the limit of the telson is

obscured by spines.

Material reported here is deposited in the Sourk

Australian Museum, Adelaide (SAM), Australian

Museum, Sydney (AM), Museum of Victoria, Mel-

bourne (NMY) and the Tasmanian Museurn are

Art Gallery, Hobart (TM). All of the specimens are

preserved in 75% alcohol or 2% lormuldelwoes

propylene glycol solution. Of the types only the

holotype, allotype and paratype uv (SAM C3992)

have been dissected (partially) amd dissected

appendages have been preserved with the carcass,

The following abbreviations are used in the text.

Mxp. = mianxilliped (thoracopod I, P 7 «

pereopods 1-7 (thoracopods 2-8), PL I-S |

pleopods |-5, LHS = left hand side and RDS -

right hand side when viewed dorsally,

Avenpned erengrum sp. nav.

FIGS 27

Holotype: SAM C3989 (2), collected among:

surface aleac and aquatic plants in unnamecl

sinkhole known as “Possil Cave” (L812), appros,

3.3 kim SE of famanoola Caves on Princes Hway,

S Ause. by W. Zeidler, §i1.19#2-

64 W. ZDIDLER

Aflotype: SAM ©3990 (er), collectedl from sume

locality by P. Hore, 841,798),

Pararypes: SAM C3992 (4) collected P. Horne,

Rb TOR], sume Lovaliry; SAM ©3999) (9), AM

PIS067 Le), PASVES lo}, NMV J1O809 (2), J10810

it), TM G2848 fo), G2849 (¢) collected by W-

Yeidler, Liii,1982, from same lovalicy

Orher material examined: SAM C3993 (11 9"s), Collected

from type lovality, al 10m depih by P, Horne, 261.1981;

SAM C35% 415 2's), from rype localiry PB Horne,

S981; SAM C3995 (20 9's), frum type lovatiry, W-

Zeidler, |. 1982: SAM C3996 (82 2%, d5 "s), from type

localiy, W. Zendler and &, L. Gowlett, 16..1),1984; SAM

C3997 419), Tank Cave (£230), just B of type locality,

collected near surface by P. Horne, 4.7).1983; SAM C3998

Oo) Allendale Sinkhate (111), ceore of tata roads

Allendale Est approx. 20 kim S of Mt Gambier, colleetect

in (ota) darkness at 27.5 m depth by & Horne, 1.11.1982;

SAM (3999 (4 92'5), Benara Slakhale (L32-33), near

“Bena”, 6 km WSW of Mt Gambier, collected at 19 m

depth, by B Horne, 127.1982. 34M ©4030 (1 2), trom

sume lovaliry, at 5 eo depih, P Horne, 27-xi.1984; SAM

4000 (2:2°s), Rilsbys Hole sinkhole (46), approx. 05: km

W oor “Burleiz"—12 ken SW Of MU Gainbier, collected

a6 30m depth by P Hore, 671.1983, SAM C400l (12 9%,

ads) from same tovality, at 3 im depth, P. Horite,

13, 1983, SAM C4002 (1), Devils Punchbowl sinkhole

(147). appro. 2 kn SW of *Barnoolut"—approw 15 km

SW at Mt Ganibier, collected from weed af 13.7 m depth

by PY) Borne 707,198), SAM C4003 (4 2S), from same

Jocahity, al 40 m depth, P, Home, 1i,1981; SAM C4004

() 9) Walnut Cave (1.54) approx, 1B km SW of Mi

Gambier and 4 kim due W af MLSchank, collected at 4m

deprti hy P Hare, 2ie 1981) SAM ©4005 (2 2's), Gums

Koad Cave (L63), approx, 4,5 kim NE of Kangorone=

20 kim SW of Mt Gambier, collected tear surface by P

Home, 26.68.1942; SAM C4 (2 ¢'s), Bomilebrush

Sinkhole (L064), in Caroline Forest, approx. [5 kin SSE

at Mr Gambier, collected at 1m depth hy P Horne,

12.4) (R2) SAM C4007 (19), Hereford-Streamt Cave (L7U,

appios, & kan E of Ewes Ponds—23 km SSE of Mi

Gambier, collected near surface by P. Horne, 22.41,1982;

SAM C4008 {le fet) Mudhote sinkhole (6.97), Tantae-

noola Forest, approx. 6 km SE of Tantanoola Caves,

oollected at 6m depth by B Berne, 3741 1982; SAM Can09

(293%. 140), Alleviis Cave sinkhole (L84/85), just E of

Mudhols collected peur surface ty P Eon, 40,x1),1983;

SAM 4010 (12), Mckay Shate sinkhate (L125), 3 kim

W of Valley bake, Mc Cramer, collected at 2 m deprh

by B Morne, toi 1982; SAM C4011 (8 2), Mushroom

Cave (1,152), upproa. 2 kin E ot Ewens Ponds, collected

at dn by B Wore, 22.15.1982; SAM C4012 (1 2), untanied

sinkhole (L144), in. Me Gambier Forest, appre. 2 km NW

of Mi Gambier Airport, ooticcted ear supfage hy PF

Flor, '4.°)7982; 54M Caos (i a, 1 ¥), The Shall

suikliole CLT58), approx, 3 kin Wool ANendale East,

cullocted at Sm depth by B Horne, (4,jii, 1984, SAM C4014

(tL 2h from same tovcalily, al 3 om depih, P) Horne

41,0984; SAM C4005 11 9), The Bullock Hobe sinkbale

(0.163). on “Bar oebue™ property—apprmm. 4 kin SW ar

Mi Cqunhier, collected at 27,5 m depel by P Hone

G.ULI9R2s SOM C4027 (4 9's 1 fd), Glencue West Cave

IL 77} approx, ThA NE of Tantanoola Caves, collected

allo depth by A, Cox att Po Horne, 29.517,1984; SAM

14029 (1-9), Eouletaests Cave (L19/20), ML Gambier

liege) a9 in dep by AL Coy, 23 i, 198s.

Speenmiens have alka been sighted in Morgans

Cave (L394) and The Pines Cave (Lal), moth near

Alleyns Cave; Hells Hole sinkhole (L40) just north

of Botilebrush Sinkhole and Simpsons Hole

sinkhole (L42) between Devils Punchbowl! sinkhole

and Gums Road Cave (Fig. 1)-

Description of holotype: Female measuring

18.3 mm total length, 3.4 mm width. General

appearance illustrated in Pig. 2. First thoracic

somite Fused with head. Pereon with seven free

somites and pleon with six free somites and a telson,

All body samifes with one pair of appendages.

Anus opens at posterior limit of pleontie 6. Pleon

(excluding telson) longer (7.7 mm) than pereon

{6.2 mim) and broader than pereon and head. Head

length equivalent to first 344 pereoniles, broader

than pereonites 1-4, equal to pereonite 5 and

narrower than pereonites 6 and 7, Pleonite 6 longest

body segment, pleonites 1-3 broadest. Pleonite 6

with row ol six well Spaced spines near dorsal-

pastenior border interspersed with 1-2 fine setae

Body segments progressively more robust from

anterior to posterlor to about pleonite 4,

Heads rectangular (22 « 28 mm) with short,

pointed postrum.and distinct antero-lateral Incislon

above atlachmenr of antenna, Pronounced short

mid-lateral transverse sulcus. Byes absent bur

pigimen) more concentrated near antero-lateral

incision,

Antennule (Fig. 44): peduncle of three segments,

outer Nagellurn of 56 (LHS) and 48 (RHS) segments

and inner flagellum of 16 (LHS) and LS (RHS)

segments. Qurer Mlagellum about 3.5% length of

inner, Pedunele and outer flagellum as long as 3/4

body length. Basal segment of peduncle broader,

almosi as jong as following two vombined, Basal

segment of ouret flagelluin with oblique dorsally

serrated inner margin, Presence or absence ol

atatocyst in basal sexment of peduncle could not

he determined with certainty.

Antenna (Fig. 4b): slightly more than 2/3 length

antennule; peduncle of 4 segments and single

flagellum Of 35 sezments (LHS & RHS). Basal

segment short, sezements 2-4 elongate, reclangular,

seement 3 longest,

Upper lip (Fig. 3a): rows ef short bristles on both

sides af extremity and small central depression on

aboral surface near extremity.

Mandibles (Figs 3c-{}: I-segmented palp; middle

se2ment greatly enlarged, aboul twice length basal

Seemenl; fenminal segment small, reunded;:

featherest selae along inner margin of segment 2,

apically on segment 3. Base of mandibles stout,

ending co well developed molar arid incisor provess,

Molar process with small grinding surface

surroutided by numerous spiniform setae, Incisor

process of left mandible with seven denticles

arranged in “S “shaped row, denricte nearest molar

NEW SYNCARID CRUSTACEAN 65

#Tantanoola

® 5

"Kongorong

141°30'

Valley Lake

Mt Gambier

lue Lake

SOUTH AUSTRALIA

VICTORIA

oMt Schank

Allendale East

e es

“Ewens Ponds

Fig. 1. Localities where Koonunga crenarum sp. nov. has been collected (@) or sighted (0). Scale bar — 5 km.

reduced. Incisor process of right mandible with five

denticles arranged in “C” shaped row overlapping

left mandible when viewed aborally. No evidence

of secondary cutting plate or spine row.

Lower lip (Fig. 3b): bifid with deep central groove;

inner lobes well developed; outer lobes upright

almost perpendicular to aboral surface. Distal

margin of lobes and part of lip covered by long

setae, particularly on aboral surface.

First maxilla (Fig. 3h): two lobed. Outer lobe with

small, one-segmented palp with three long, terminal

spinules, partially feathered along inner margin.

Extremity of outer lobe obliquely truncated with

twelve strong, chitinous (?) spines, some stouter

than others and two smaller feathered spines near

the aboral surface. Inner lobe about 1/2 width of

outer lobe with one long, stout feathered spine near

oral surface surrounded by eight smaller feathered

spines.

Second maxilla (Fig. 3g): smaller than first con-

sisting of four lobes; inner one smallest, others

increase successively in length and width. Inner two

lobes covered with short bristles or setae. Inner lobe

with six teathered spinules of varying lengths

terminally, Other lobes with more numerous (>10)

but similar terminal spinules which fan out laterally

to oral and aboral surface.

Maxilliped (thoracopod 1) (Fig. 4e): stout limb of

seven segments flexed posteriorly between merus

and carpus. Coxa shorter and wider than following

segments with two adjacent branchial lamellae

(epipodites) near outer, aboral corner. Basis with

exopodite of two joints resembling branchial

lamellae along outer, aboral margin and with several

fine setae near distal, oral margin. Ischium with

slight expansion distally, slightly wider and longer

than basis, with several long, fine setae on oral

surface near inner margin and clustered on outer

aboral corner. Merus inflated proximally, longer

than any other segment, only slightly narrower than

coxa; with long fine setae scattered over oral surface

but more numerous near inner margin. Carpus,

smallest joint with row of long setae along distal,

oral margin for inner half and on oral and aboral

hf W, ZEIDLER

. a :

Fis

Fig, 2, Koonunga crenarwn sp. nov, halolype ?, Scale bar

~ 2 min,

outer distal margin, Propodus robust, slightly

shorter than merus, with Iwo oblique rows of strong,

spines on oral Surface and tuft of long setae on

aboral, distal, outer corner. Dacrylus small,

rounded, armed terminally with one large and three

smaller strong claw-like spines and few setae near

inner margin,

Pereopods (thoracopods 2-8) (Figs 4f-l)> similar in

structure fo Mxp but with basis expanded proxi-

mally, almost as wide as coxa and becoming

progressively oiore slender with all segments more

clongale. PT slightly shorter than Mxp but slightly

longer than P2 & P3 of equal length, P4 as tong

aus Mxp and jongest pereopod. PS shortest, Pé

slightly longer than PS bul slightly shorter than P2

or B3. P? similar in Tenth to PL. All pereopods

Mexed beoween merus.and carpus and in backward

position except foe P6 which flexes inwards and P7

witich flexes forwards. Coxa with setose fobe near

inner, distal commer barely recogrisable on PL but

progressively developed to maximum on P6 but

absent on PT. PIG, cone with Iwo unequal

enipodites as in Map. P'I-3, pasis with multi

segmented exopodite consisting Of large basal

segment reaching well past ischium and flagellum

of 14-16 segments (Table 1) each segment bearing

wo long, feathered setae. P6, basis without expodite,

P7 without epipodites or exopodite, Dactyl claws

similar to Mxp bul PS-7 have one additional claw.

Spermatheca: two small, avoidal plates, slightly

raised from the sternum just forward of base of PF

on last pereonite (Pig. 5a),

Pleopods (Pigs Sb-f): all of similar structure,

lacking endopodites but with long multi-segmented

cxopodites consisting of short-stout basal joint

followed by longer more slender one and flagellum

of 15 segments in Pl 1-4 and 13 segments io PI §,

cach bearing two long feathered setae. Pl i-3 equal

in length and longest pleopods. Pl 4 slightly shorter

and PIS shortest, about 3/4 length Pl 1-3. Pl 1-3

also flattened laterally, Pl 4 & 5 round in

cross-section.

Uropods (Pig. Sh): peduncle stout, rectangular, as

long as telson and almost 3/4 length pleonite 6

(measured ventrally), projecting to slightly more

than 1/3 telson length; few scattered short spines

on dorsal surface and near outer margin. Outer

ramus almost 1 1/4 peduncle length, slightly longer

than inner ramus, with long feathered setae along

inner and outer margins. and row of short strong

upturned spines ucar outer dorsal margin, Inner

ramus With long feathered setae along outer margin

and partly terminally, remaining terminal margin

with three (LHS) or four (RMS) long spines; dorsal

inner margin with row of strong upward curved

spines for about proximal 2/3, steadily increasing

in size terminally followed by comb of smaller

spines, ceasing al first terminal spine.

Telson (Fig. Sh): triangular in shape with rounded

apex and slightly convex lateral margins; length

(excluding spines) slightly less than width; margins

with complex array of spines except for proximal

1/4, consisting of close-set fringe of short, stout

spines ventrally, 20 evenly spaced, long spines

direeted posteriorly and about 14 slightly shorter

spines directed dorsally interspersed with 1-2 fine

selae.

Colour: light tan with same specimens darker than

others, Those captured in deeper water and in total

darkness do not vary in colour Irom those eaught

near the surface. Preserved speciniens only slightly

faded to date.

Descriprion of allotype: Male measuring 14,8 mm

in total length and 2.5 mm in width, essentially

similar to holotype except for the following

differences.

Antennule (Pig. 4c): with characteristic oval-shaped

sensory Gregan arising from second segment of outer

Napelluin Basal seement of inner flagellum saucer-

shaped shielding sensary organ. Outer flagellum of

NEW SYNCARID CRUSTACEAN 67

Fig. 3. Koonunxa crenarum sp. nov. holotype ?, mouthparts. a. upper lip; b. lower lip; c. mandible RHS, aboral

view; d. mandible LHS, aboral view; e. mandible LHS, oral view; f. mandible, RHS, oral view; g. maxilla 2; h.

maxilla 1; Scale bar = 0.2 mm.

68 W. ZEIDLER

ig. 4. Koonunga crenarum sp. nov. holotype 2. a. antennule; b, antenna; c, antennule, allotype ¢; d. antenna, allotype

3) & Mxp: 1 = PI-7. Scale bar | mm.

NEW SYNCARID CRUSTACEAN 69

Gi:

Vhhd

Fig. S. Koonunga crenarum sp nov. a. spermathees, 2 (SAM C3991); b-f — PI 1-5 holotype %: g, telson, allotype

dy h. telson and uropod, holotype 2; AL cursersiantenor ot 2 specimen (4.2 mm) from Railway Drain, Bayswater,

Vie. (SAM C4016).

NEW SYNCARID CRUSTACEAN 7

TAaLe 1, Meristie dete for Koonunga crenarum sp. nov, Holotype (MH), allotype (A) and paratypes (PF), All

measurements ram dorsal aspect, Unless indicaled atherwise the number of segments refer to the flagellum.

“D" — damuged ar broken,

Character

EE DIET EES INSURES

Length (mm)

H®

C3989

C3990)

P o's

Pd's

Head 3,2 2.8 3.0 -37 2.0 - 2.4

Pereon 6.2 4.6 §.5 - 6.7 3.4 - 5.0

Pleon TT 6.2 6.7 — 8.3 44 — 5.8

Telson 1.2 1.2 1.0 - 1,5 0.7 - 1.1

Total 18,3 14.8 16.5 -20.2 10.5 -14.3

Antennule LHS (RES) 13.5 (13.6) 99D (13.1) 9 (11.6)-14.95 (13.7D) 8.4 (8.2112 (10.3D)

Antenna LHS (RHS) 8.6 19,6) 7.5 (7.1D) 7.5 (7,4) 10,5 (10,4) 5.4 (5,1)-7.5 (7.8)

Ratio Pleon/Pereon 1.24 1.35 N7- 1.25 },)6— 1.32

Breadth (min)

Head 2.8 2.3 2.3 - 3.1 1.6 - 2.1

Pereoniic 7 3.2 2.4 2,5 - 3,0 1.8 — 2.3

Pleonite 3 34 2.5 2.8 - 3.2 2.0 - 2.6

Telsor 7 {45 1.5 - 2,0 L.}o~ 4,3

Antennule (segments)

Inner ramus LHS (RHS) 16 (15) 14 (14) {5 (15)-17 (16) TD (th)-14 (14)

Outer ramus LHS (RHS) 56 (48) 39D. (59) 45 (50)-57 (57) 43 (43)-55 (43D)

Antenna (segments)

LHS (RH) 35 (35) 27 (33D) 34 (28)-40 (40) 24 (24)-33 (37)

Pereopods (segments/exopodite)

1 LHS (KES) 14 (14) 13 (13) 13 (137-15 U5) 10 (10)-12 (12)

2 LHS (RHS) 15 (14) D (15) 15 (15)-17 (18) 11 (12)-14 (14)

3 LHS (RHS) 16 (15) 16. (16) 15 (14)-18 (D) 12 (12)

4 LHS (RHS) 15 (15) 15 (D) 15 (15)-17 (18) 2 (Lt)-14 (12)

§ LHS (RHS) 15 (15) (5 (D) 14 (14)-17 (16) Tt (Ut) 14 (14)

FPleopods (seuments)

1 LHS (RES) 15 (15) 15 (15) 15 (15)-16 (16) 11 (11). 14 (13)

2 LHS (RHS) 15 (15) 15 (15) 15 (15)-16 (16) 11 ()h)-14 (14)

3 LHS (RHS) 15 (15) 15 (D) 15 (15)-16 (D) 1) ()i)-14 (4)

4 LHS (RHS) 15 (15) 1§ (D) 15 (14)-16 (D) Hi (it) 13 (14)

5 LHS (RHS) 13 (13) 13 (13) {3 (13)-14 (14) 4 (D) -12 (12)

Uroped spines

(nner ramus—inner dorsal

(HS (RHS) 12 (12) 11 (12) 13 (13) 11 (11)-12 (12)

Outer ramus—ourter dorsal

LHS (RAS) D (14) 18 (17) 15 (15)-17 (17) 13 (13) 18 (7)

59 sepments (RHS; LHS broken); inner flagellum

of 14 segments (LHS & RHS).

Atitenna (Fig, 4d); left with flagellum of 27

segments, right broken but still with 33 segments.

Pereopods: similar to holotype but all with four

dacty! claws. P1-6 without coxal lobes,

Pleopods: | and 2 with endopodites modified to

form complex copulatory styles (petasma), Petasma

(Fig, 6a) directed anteriorly against mid-ventral

surface between, and posterior to, last pair of

percopods.,

Endopodite of PI 1 (Figs 6b, c) unsegmented,

grooved for about middle half of external inner

margin to provide sleeve for endopodite of Pl 2;

cup-shaped terminally, directed inwards and towards

body. Endopodites joined about 2/3 from base by

coupling spines on extension of internal, inner

margin.

Endopodite of PI 2 (Figs 6d, v) of 2 segments of

abou( equal length, each as long as endopodite of

Pl 1, Basal segment slightly expanded proximally

with coupling spines on small pad near centre of

internal, inner margin and with small basal

projection on inner proximal corner articulating

with sternal process. Distal segment apically

pointed, hollowed out on distal, inner margin to

form coneave depression directed towards body.

Sternal process (Figs 6d, e): triangular in shape,

slightly longer than wide, slightly less than 1/2

length of basal segment of endopodite of PL 2,

filting inbetween basal |/4 of endopodites of P12.

Telson (Fig. 5g): slightly more pointed apically than

Mg. 6. Soonunge crenarum sp. nov, a, pesma, allorype 2; b, endopodite PL ventral view; c. reverse of by d, endopodite

PL 2: & reverse of dy fi

endopodites Plt (LHS) & Pl 2 (RAS) of specimens 7.9, 8.4, 9.3 & 1O4 mm length

respectively, A elirsor: 26.6 min (SAM C4016); |. endopodite Pll; k, reverse of j; L endopodite Pl 2; NM. reverse ofl.

Width PES (mm)

in type, lateral margins straight or even slightly

concave.

Etymology: From Greek mythology, “crenae” being

nymphs of springs.

Variation: The paratypes are similar to the type or

allotype. Variation in size and other meristic

characters is shown in Table 1. Not shown in Table

1 is slight variation in the number of dactyl claws

on the pereopods, the number of large terminal

spines of the inner ramus of the uropods

and the relative lengths of the pereopods and

pleopods. The pereopods usually have four dactyl

claws but sometimes in the larger specimens P4—7

may have five dactyl claws e.g. holotype and female

paratype SAM C399]. The inner ramus of the

uropods usually has three large terminal spines but

in SAM C399] and on RHS of the holotype there

are four spines. The pereopods and pleopods have

not been measured for each specimen and show

only slight variations from the holotype. In the male

SAM C3992, P2 & 7 are the longest and P4-6 are

the shortest, equal in length and slightly shorter

W. ZEIDLER

than the Mxp; the pleopods are like those of the

holotype.

The non-type material exhibits the same

variation. Specimens from various localities have

been examined in detail, and cannot be distin-

guished from those from the type locality.

Some specimens appear to be more slender than

others and at first it was thought that two species

might be present, however, the relationship between

length and breadth shown in Fig. 7 indicates that

breadth is a variable character especially in larger

specimens. The pleon/pereon ratio is also variable,

ranging from 0.99 to 1.77 but does not appear to

be related to length, breadth or sex of specimens.

Females varied in length from 5.9 to 22.6 mm

with a mean length of 13.8+0.5 mm (495% c.l.).

Males ranged in length from 7.8 to 16.2 mm with

a mean length of 10.940.5 mm (+95% cl.) which

is significantly smaller than that of the females

(P <0.01). The female sample is larger (176/60 :¢).

However, seasonal fluctuations in the proportion

of males in the population may account for some

s ¢

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30 eeeoee .

e eee | ® eee

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25 ede we ©

S@@eoe 0M oO

s A:@sghe @& 4s °

e8 eco

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20 Oe MAE A

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s@ “sie *

a ane e

15 © CBO saa

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Length (mm)

Fig. 7, Koonunga crenarum sp. nov. Relationship between length and breadth (pleanite 3). Females closed circles,

males closed triangles; open symbols refer to specimens from localities other than the type locality.

NEW SYNCARID CRUSTACEAN rx)

of the observed imbalance as 45/128 specimens were

males in July, 1984 compared to 0/11 in January,

198t) 2/17 in Mareh, 1981 and 3/28 tn March,

1982—all from the (ype locality.

Development of the petasma and sensory arken

of male

A relatively large collection of males. from the

type locally in July, 1984 (SAM C3996) included

a number of very small specimens which exhibited

various stages in the development of the petasma

and also the sensory organ of the antennule. The

development of these sexually dimorphic characters

in sysearids has nor been reported in the literature.

The endopodites of P! | and Pl 2 develop laterally

from the base of the pleopods until they overlap

and are tully developed; they are then directed

longitudinally forward but are not linked by the

coupling spines until the sternal process is also fully

developed. The earliest stage observed was al a

length of 7.9 mm (Fig. 6f. At this stage the

endopodites are only recognisable as single, small,

rounded segments and the sternal process js not

visible, In the next stage at 8.4 mm (Fig. 6g) the

endopodite of Pl 1 is beginnmg to differentiate but

the sternal process has not developed further. At

93 mm (Fig. 6h) the endopodite of Pl 1 is beginning

to form its characteristic shape; the endopodite of

P| 2 has divided into [Wo scements and (he sternal

process 1s recognisable as asmall ridge. At 10.1 mm

(Fig. 61) the endopodite of PI I is almost fully

developed with coupling spines; the endopodite of

PI 245 also almost fully developed but lacks the

concave depression at the tip and the sternal process

has not developed further. A specimen at 9.6 mim

had a fully developed petasma and sternal process

but the endopodites of Pl 2 were still not linked by

coupling spines. Other specimens al about 10 mm

exhibited fully developed male sexual

characteristics.

The sensory organ on the antennule arises from

a depression on the inner margin of the second

segment of the outer flagellum (Figs 8a, b) and

develops into a pedunculate eye-like organ with the

rounded outer surface covered with minute cup-like

structures attached to the surface by a short stalk

(Fig. 82), The basal segment of the inner flagellum

becomes modified developing a saucer-shaped lobe

on the dorsal surface which partly shields the

sensory organ (Figs 8d, ©). The earliest stage

observed was at a length of 8.4 mm (Figs 8a, b).

At this stage the sensory organ is only a small

protuberance and the basal segment of the inner

flagellum is undifferentiated. At 9.3 mm (Fig. 8c)

the sensory organ has developed into a ball-shaped

structure but the basal segment of the inner

Magellum is still unmodified. At 10.1 mm (Fig. 8d)

the sensory organ is larger and more pedunculate

and the basal segment of the inner flagellum is

beginning to differennate forming a small lobe on

the dorsal surface. A specimen at 11.3 mm (Fig. 8e)

had an almost fully developed sensory organ and

the basal segment of the inner flagellum had

become broader and saucer-shaped with a fringe

of long setae on the dorsal margin, partly shielding

the sensory organ.

Sayee (1908) in his description of K. cursor

described and illustrated the sensory organ arising

from the basal segment of the outer Magellum and

a saucer-shaped lobe shielding the sensory organ

as arising from the terminal segment of the

peduncle, An examination of some specimens of

K, cursor from Bayswater, Victoria (SAM C4016)

demonstrate that these male sexual characters

(Fig, 8f are similar to those of K. crenarum sp. nov.

and that Sayce (1908) was in error in his determnina-

tion of these features.

in view of the above it is possible that specimens

less (han 8 mm in length may be difficult to sex bur

as only 10 temales were rhs small it would make

litte difference to the proportion of males/females

observed.

Tauce 2. Distinguishing characters of Koonunga cursor and RK. crenarum sp. roy

Character A, cursor

Specimen length ¢

Antennule length

rarely exceeds 10 mn

upprox. /2 body length, when

A. crenarurn

exceeds 20 mm

approx, 14 body length, when

Head

Byes

Pelagma (d}

Darty! spines

attached barely teaches pereonite 3.

evenly pounded anteriorly, antero

laleral incision above antenna very

smal! (Fig. Si)

small but present

sternal process with large posierior

Projection (Figs 6 |, m) Note also

distinctive struclure of endopo-

ices of PL | & 2 (Figs 6}-m1)

Pt-7T.3

attached reaches past pereonite 7.

poinied anteriorly, antero-

lateral incision above antenna

distinet (Fig. 2)

absent

sternal process without posterior

projection (Figs 6d, e} Note also

distinetive structure of endoped-

ites of Pl 1 & 2 (Figs fb<1

Pi-3 = 4, P 4-7 = 45

74 W. ZEIDLER

Fig, 8. Sensory organ on antennule (LHS) of—Aoonunga crenarum sp. nov, (SAM C3996) a & b. dorsal and ventral

view, from specimen 8.4 mm; c-e = dorsal view, from specimens 9,3, 10.1 & 11.3 mm length respectively.—K, cursor

(SAM C4016) f. dorsal view, trom specimen 6.6 mm. Scale bar = 0.2 mm.

NEW SYNCARID CRUSTACEAN 15

Discussion

Aoonunga crenarum most closely resembles A.

cursor the only other described congener and can

most readily be distinguished from it by the

characters given in Table 2. In the lack of eyes it

also resembles Micraspides calmani Nicholls, 1951,

the only other member of the family Koonungidae.

K, crenarum amongst the Koonungidae

(including known undescribed species) occupies the

most subterranean habjtat and is also the largest

species, It occurs in depths exceeding 40 m and of

particular significance is its occurrence in sinkholes

that were only discovered when the root collapsed

eg. Allendale and The Shaft, All previous records

of Austrahan syncarids are from relatively shallow

depths and even the Tasmanian syncarids which

inhabit lakes (end to occur near the edges and the

deepest record is 8 m, recorded for Paranaspides

lacustris Smith, 1909 (Williams, W, D. 1974).

As sinkholes and caves are difficult and

dangerous environments to explore it is hard to

establish the relative abundance of the species with

depth although it would seem 10 be more abundant

in the surface waters, especially near vegetation at

the edges of the sinkhole or cave. Specimens from

the surface and deep water have been compared in

detail and appear to be conspecific: pigmentation

is similar but deep water specimens are slightly

darker. A. crenarum does not appear to be a strong

swimmer so it is unlikely that regular migrations

occur between the surface and the botiom of the

sinkholes. It is unlikely that the deep water

specimens represent individuals that have become

“lost” as many more specimens have been sighted

in deep water than have been collected (P. Horne,

pers. comm.). A. crenarwm therelore most probably

occurs throughout the depth of sinkholes and caves,

The occurrence of K. crenarum in several, some-

times widely separated, sinkholes and caves (Fig. 1)

suggests that these environments must have been

linked at some time in the past, An underground

connection has been a popular theory but is nor

supported by hydrological evidence (Holmes &

Waterhouse 1983) and tle animals are too large for

Interstitial travel. It is most likely that the habitats

were once connected by flood waters although they

are not located near Mats which were subject to

annual flooding before drainage (Williams, M,

1974). Lt ts possible that they were isolated before

European settlement, Whatever the connection in

the past sinkholes and caves are now isolated refuges

for a once more widespread fauna,

Acknowledgments

| am extremely grateful to Mr P. Horne who

originally discovered the synearid and collected

additional material from a Humber of sinkholes and

caves. Prof, H. kK, Schminke and an anonymous

referee arc Thanked for critical conuments.

References

De Dekkhr, P (1980) New records of Aoonunga cursor

Sayce, 1908 (Syneanda, Anaspidaceal. Trans, R. Soe,

S; Aust We4(2), 21-25.

DaumMMonn, FD A. (1959) The syncarid crustacea, a living

link with remote geological ages. Aust. Mus. Mag,, 13,

63-64

Horas, dW. & Warerrouse, J. D, (1983) Hydrology.

pp. 49-39. Jn Tyler, Mo J. Twidale ©. R., Ling, J. K.

& HWalmes, J. W-. (Eds) “Natural History of the South

East” (Koyal Society of S. Aust.: Adelaide).

Sayen, G. A. (1908). On Keonunea cursor a remarkable

new type of malacostracous crustacean. Trans. Linn.

Soc, London, 1, +6, pls 1-2.

SouMINKE, H. K (1974) Psarnrmaspides williamsi gen. n.,

sp. n., cit Vertreter ciner neuen Familie mesopsammaler

Anaspidacea (Crustacea, Syncarida). Zool Ser 3,

177-183,

-—— (1980) Zur systematik der Stygecarididac

(Crustacea, Syncanda) und beschreibung sweler neuer

arten (aivgacurella pleofelyon gen. o., sp. n. und

Srygocuris giselae sp. v.). Beayfortia 30, (49-154,

WILLIAMS, M, (1974) Draining the swamps. pp. 78-226,

Jn Williams, M. “The making of the South Australian

Landscape” (Academic Press: London and New York).

Wir tiams, W, BD, (1974) Freshwater Crustacea, pp. 63-111.

fa Williams, Wo D. (Ed.)"Biography and ecology in

Tasmania”. Monographiae biol, 25

ZrFipterR, W. (1983) Terrestrial and freshwater inverte-

brates excluding insects and arachnids. pp, 187-204, In

‘HWler, M. .. Twidale, CR, Ling, J. K. & Holmes. JL W.

(Eds) “Natural History of the South East" (Rayal

Society af S. Aust.: Adelaide),

TRIGONIOIDIDAE (MOLLUSCA: BIVALVIA) FROM THE CRETACEOUS

OF LAKE EYRE NORTH, SOUTH AUSTRALIA

BY P. A. HUTCHINGS & P. §. TURVEY

Summary

Pledgia eyrensis gen. et sp. nov., a species of bivalve mollusc apparently belonging to the

Trigonioididae, an important family of non-marine Mollusca in the Cretaceous of Eastern, Central

and South-East Asia, is described. The bivalves weather out from unconsolidated fine sand and silt

mapped as Winton Formation on the western side of Babbage Peninsula, Lake Eyre North, in the

southern part of the Great Artesian Basin. No likely non-marine ancestral forms have been found

among the Unionidae and it is possible that ancestors of Pledgia eyrensis may be found in the

Trigoniidae.

TRIGONIOIDIDAE (MOLLUSCA: BIVALVIA) FROM THE CRETACEOUS OF

LAKE EYRE NORTH, SOUTH AUSTRALIA

by N. H. LUDBROOK*

Summary

Lupnrook, N. H. (1985) Trigonioididae (Mollusca: Bivalvia) from the Cretaceous of Lake Eyre North,

South Australia, Trans. R, Soc, 5, Aust, 109(3), 77-82, 29 November, 1985.

Pledgiu evrensis gen, et sp, nay., 2 species of bivalve mulluse apparently belonging to the Trigoniaididae,

an important family of fon-marine Mollusca in the Cretaceous of Eastern, Central and South-East Asia,

is deseribed. The bivalves weather out from unconsolidated fine sand and silt mapped as Winton Formation

on the western side of Babbage Peninsula, Lake Eyre North, in the southern part of the Great Artesiatt

Basin, No likely non-marine ancestral forms have been found among the Unionidae and it is possible that

ancestors of Pleduiq evrensis may be found in the Trigoniidae.

Key Woups: Mollusea, Bivalvia, Trigonioididac, Cretaceous, Winton Formation, Lake Eyre North,

Great Artesian Basin.

Introduction

In 1982 a number of grey-coloured, thick-shelled

bivalves collected in July from an outcrop on the

western side of Babbage Peninsula, Lake Eyre

North, were shown to me by Mr Neville Pledge of

the South Australian Museum, Similar material had

been collected in 1967 by Dr B, Daily of the

University of Adelaide. Thought to be of possible

Miocene age, those collected earlier were Laken to

the National Museum of Victoria for ident|fication,

As their preservation and colour were similar to

Cretaceous marine fossils from ihe Lake Eyre region

and not to Tertiary molluscs of the area, |

considered those shown lo me lo be non-marine

Cretaceous bivalves which | had not previously seen

in the Cretaceous of the Great Artesian Basin, The

material taken to Victorgl was then returned to

augment thal in the South Australian Museum,

Since the original material was collected, the

LAKE EYRE 1:250 000 map shect has been

published (Williams 1975), Quterops af Winton

Formation (Cenomanian) are shown west of the

fault on the western side of Babbage Peninsula

(Fig. 1).

The molluses were found weathering out from

sandy silt to uriconsolidated fine sand mapped as

Winton Formation, towards che base of a low blu

4 m high, There is a possibility, nor yet

subsiantialed, that the sandy silt has been reworked

from the Winton Formation, The locality was

revisited in May 1983 by G. W. Krieg and P. A,

Ropers of the Geological Survey of South Australia

Who made further collections from (he site and

provided the composite section shown in Fig, 2, The

Winton sediments are overlain by a layer of massive

crystalline gypsum up fo 1.5 uo thick in which

*c/o Departmen! af Mines aod Energy, PO Bow 151,

Eastwond, & Aust. $063.

occasional bivalves were found rewarked, This 1s

followed by pale olive-grey gypsiferous sud and a

conipact gypsum erust,

In all, 19 more-or-less complete valves and 274

fragments were collected. Some of the specimens

were sent to the Geological Survey of Queensland

and the Queensland Museum to see whether similat

moljuses had ever been found in the Winton

Formation in Queensland, bul none were known

to have been collected here.

They seem to belong to the family

Trigonioididae, an important family of non-marine

molluses in the Cretaecous fauna ol Eastern,

Central and South-East Asia (Vane 1974),

previously not known to occur in the Southern

Hemisphere.

The family ‘lrigonioididae and ifs distribution

The family name ‘Trigomoaididae was introduced

by Cox (1952) to accommodate the genera

Trigonioides Kobayashi & Suzuki, 1936 and

Hoffetrigonia Suzuki, (940 in the Unionacea rit her

than in the Trigoniacea, Cox excluded from

consideration or did not aecepr Those genera as

belonging to the Trigeniidae where trey had

otlginally been placed. Later (1955) Cox withdrew

the family Trigonioididae, considering Trigartioides

jo be 4 pormal representative of the Unionidae.

Kobayashs (1956) restored the Trigonioididae as a

valid family which has been generally accepted since

then and classified within the Unionaces. Hiehty-

four species have been described, disinbuled among

ten genera and seven subgenera. A second family

Peregrinaconchiidae with seven species was

described by Gu, Chen & Lan (i Lan 1976), while

Martinson (1984) erected the superfamily

Trigonipidea to jnelude chree families:

78 NM. H. LUDBROOK

| apo

QUATERNARY

Fluvial deposits

Awultary sat

Brow sarc. clay and silr

with reworked gypsum —

Gypsum crust

Suond gravels

Channel sands and plays — —

TEKTIARY

ETAOUNNA FORMATION __

Thick layered alerete

CRETACEOUS

WINTON FORMATION ____

Tassil locality

Vou

RILQMETRES

P\Ae hye

i NTH

ier

| |

| ; \

| BABBAGE Ly)

PENINSULA beh

| ab tu

LAKE FYRE

NORTH

Nore Gweiteyy urfagleel (reer

CANE CYAE TT 240 CN

HEU OPO

LAKE BRE

NORTH

Fig, |. Locality plan with geology of Babbage Peninsula (lake sediments blank)

Trigonivididae, Pseudohyriidae and Sainshandiidac

(= Sainschandlidae). Martinson’s classification ts

at present available in abstract only.

The present known distribution of (the

Trigonioididac in Asian non-marine basins is

(Kobayashi 1958); Northern Asia—Lena Basin and

eastern Siberia

Fastern Asia—Amur River—Manchuria

Japan and Korea

Mongolia—norihwest China,

One species has been described from Colorado

LISA (MacNeil in Morris 1936),

The Lake Eyre bivalves

The bivalves from Lake Eyre resemble

Trigonioididae from eastern Asia chiefly in the

structure of the hinge; the sculpture differs in that

the racial or chevron paliers of the costae are al

best vestigial and the dominant sculpture is finely

concentric, The interior of [he ventral margin is not

crennlated,

According (to Kobayashi (1958), the Trigon-

imididae appeared ‘sporadically and explosively in

the maritime basins of eastern Asta at the beginning

of the Cretaceous.’ The family was considerably

augmented in the Lower and Middle Cretaceous by

the recognition of important genera such as

Plicatounio and Nipponaia (Ota 1959a, 1959b,

1963). Kobayashi attributes this sudden appearance

to some geological event which caused a change in

environment, marine bivalves being locked in inland

basins during the Sakawa cycle of orogeny.

Nevesskaya & Soloviey (1981) show the maximum

development of the Trigonioididae to be from the

Cenomanian to the Santoniafi, with minor

occurrence in the Campanian and Maastrichtian,

Martinson (1984) considers the Trigonioididae to

be characteristic of the Early Cretaceous and the

Pseudohyriidae and Sainschandiidae of the Late

Cretaceous. Reconciliation of these apparent

discrepancies will presumably be found in the full

text of Martinson’s paper when it becomes available,

The age of the Winton Formation at Lake Eyre

a considered to. be Cenomanian (Ludbrook 1978,

Moore 1982, Forbes 1982) or lalest Albian (o

Cenomuanian (Moore & Pilt 1982, Krieg 1982).

TRIGONIOIDIDAL FROM CRETACEOUS OF LAKE EYRE NORTH rh

Gypsum crust

I ry

Yee ¥ ¥ ¥)

ye. ae Noha ell \ Punrly sotted aity Hie to medium sand thornughly impreynated

; see tees | Witte finely crystalline gypsunt

Thin (ins. (in bent) yypaui-cemented quarte granules, only well sorted, grains well

rourcthes! to very well rounded, some well polished

Coarsely to miilum srystallive lairly pore gypsum wilh gonw in-eonk and arrow head

felt, Horie cosistant foyer Some parting parallel tu bedding with reprecipitatud

ypu as tne laminae parallel to partuig surfaces AY Hage of gypaum faye very thin

vand of rough, pale yellow rlay which thickwns aluny stnke and lakes plane of qypsurn

Joyel, vary slightly caltareous

c i

(Thee,

res

Ay = Slightly blearhed

Pale grey, clayey and very finely sandy il) passing down to less

olayoy, very fing poorly sorted sand, shgtlly minackaus and with

minue woody (ragments Occasional rounded quar’ granules in

tower tial af un

WINTIN FURMATION |? reworked)

Bivalve fragments ant ocuasional sigh

volve on erosion slope

6S 184 SALMT

Fig. 2. Composile section of [ossil site 6340 RS 4, Lake Eyre North.

Palynological evidence for its Cenomanian age is

given by Burger (1982), At this time, following the

disruption of Gondwana and extensive marine

Nooding during the Barly Cretaceous, there was a

change of regime in the Late Cretaceous and most

ol the area of the Great Artesian Basin was reduced

to a lacustrine remmant (Ludbrook 1978),

Systematic description

Family TRIGONIOIDIDAE Cox, 1952

Genus PLEDGIA gen. nov. (fem.)

Shell ovate in shape, elongale-ovate in the

juvenile, very thick and heavy, sculpture ol

concentric growth ridges and concentric lirae. Hinge

with anterior teeth relatively short, transversely

crenulated, posterior laterals narrow, long, net

crenulated.

Pledgia eyrensis gen. et sp, nov.

Material) A total of 19 more-or-less complete

speciinens and 274 fragments, all from the type

locality 6340 RS 4. (Fig.3), South Australian

Museum (SAM): the type series, holotype P23999a,

paratypes P23999b-g, fragments P23999h-k, all

RVs average length 33, average height 23 mm;

paratypes P24Qa-h, an ontogenetic series of 8

almost complete LVs average length 30, average

height 23 mm; P24639a-p, fragments all

approximately Jength 30, height 20 mm; P24640,

about 80 fragments.

Geological Survey of South Australia (GSSA):

paratypes 1lO184a, b, 2 RVs a, length 34, height 22,

b. length 30, height 19 mm, 10185 paratype 10185a

and 16 fragmentary RVs, largest length 40, height

fest.) 32, average of 3, length 33, height 22 m;

10186-10192 104 fragmentary RVs and 41

lragmentary LYs.

Description: Valves of medium size, transversely

subovate becoming subovate in adult specimens,

narrowly rounded anteriorly, slightly produced

posteriorly, anterior-dorsal margin well rounded;

umbo located at about the anterior one-quarter,

moderately high, prosogyrate; shell very thick,

80 N, H. LUDBROOK

Fig. 3. Pledgia eyrensis Ludbrook gen. et sp. nov. a,b. Holotype SAM P23999a RV a. exterior, b. interior; c,d. Paratype

SAM P23999b RV large specimen (incomplete) c. exterior showing thick shell with imbricating lamellae, d. interior,

worn, with crenulated 3b and deep pit; e,f. Paratype SAM P24000g LV immature specimen e. interior showing

deep muscle scars, hinge not fully developed, f. exterior showing radial ridging on both the anterior and posterior

slopes; g. Paratype SAM P24000h LV rounded form; h. Paratype SAM P23999h LV fragment showing thick posterior

area with radial sculpture; i. Paratype GSSA 10192a LV with differentiated posterior area; j. Paratype GSSA 10184a

RV, elongate-oval form with pedal retractor scar just visible under hinge; k. Paratype SAM P23999f, tilted to show

pedal retractor; |. Paratype SAM P23999g RV immature specimen, elongate-oval form with anterior cardinal tooth

not fully developed; m, Paratype SAM P23999f RV immature specimen, elongate-oval form; n. Paratype SAM

P24000b LV immature specimen with well developed posterior laterals, partly developed grooved cardinal, pedal

retractor scar visible; 0, Paratype GSSA 10185a LV with strong posterior lateral. All figures natural size.

consisting presumably of an outer organic layer or

periostracum which is not preserved, an outer very

thick (up to 5 mm) calcareous layer composed of

successive imbricating lamellae, turned upwards at

the ventral margin, which exfoliate very easily

leaving the inner layer exposed, and a crystalline

inner layer formed of very thin, smooth laminae (see

Morton 1967 Fig. 6B) not thickening at the ventral

margin. Sculpture on the flank consisting of

concentric growth ridges with microscopic

concentric lirae between them; faint vestigial radial

riblets visible particularly in the anterior part of the

flank, a small, thick, posterior area differentiated

by having thick radial sculpture. Both the outer and

TRIGONIOIDIDAL PROM CRETACEGUS Ob LAKE LYRE NORTH #l

inner layers are composed of calcite, determined by

X-tay diffractometer at the Australian Mineral

Development Laboratories (AMDEL Report

G384/85). No aragonite was detected. Interior

smooth, hinge arcuate with a dental formula of

RV 53 3 Pl PIU

LV 2 PIL PIV

5 is not always present and is créenulated or grooved

only in adult specimens, 3 is strong and high, of

moderate length and in adull specimens crenulated

on both sides with a pit for the reception of 2 in

the left valve, Pl and PLL are long and narrow,

extending the length of the dorsal margin, Pll and

PIV are fairly long and smooth. The dental formula

is somewhat similar to that of Trivanioides

(Wakinaa) Ota, 1963 as illustrated by Yang (1983),

Musele scars deep, anterior adductor sear close to

(he anterior margin and to the afilerior end of the

hinge with a small deep pedal retractor pit above

it just under the hinge; posterior adductor sear

larger, not so deep, more or less roundly quadrate,

pallial line entire, area within the pallial ling clearly

defined, ventral margin smooth, slightly bevelled

but not crenulated

Distribution: Known only tram the type locality

6340 RS 4, Babbage Peninsula, Lake Eyre North;

Winton Formation, Cenomaman,

Noamenclatures The generid name is to honour

Mr Neville Pledge of the South Australian Museum,

wha collected many of the specimens and first

brought them to my notice.

Discussion: The family Trigonioididae was based

on the species Trigonioides Koddirai now reporied

from many localities From the USSR to Japan (Yang

1974), The morphology, ontogenetic vartalion and

classification of itis and related genera and

subgenera have been extensively studied by

Martinson (1965) and by Yang (1974, 1976, 1978,

1979, |983),

Pledgia eyrensis seems to have appeared suddenty

in the Cenomanian in Australia, bur, unlike the

Tngamoididae of Eastern Asia as deserihbed by

Kobayashi (1958), it is not known to have appeared

“sporadically and explosively,” No likely non-mranne

ancestral forms have been found among the Liuion-

idav and there appears to he no close relationship

between Pledgiv evrensis and the Triassic non-

marine unronids (Ludbroak 1941) of the Leigh

Creek Coal Basin. Immature forums bear only very

slight resemblance in sculpture to the Neoconvan

Profovireus coaisi Ludbrank 1961, of which the

hinge is Hot knowa, In shape, sculplure snd (hick-

ness of shell the adult oval form seems to be mast

nearly related to the undiagnosed “Triganiid gen-

ct sp." of Skwarke (1963 pl. 6, fiz. 5) from the Birly

Cretaceous Wallumbilla Formation south-southwest

of Roma, Queensland, This rrigantid is known only

fram the single specimen figured by Skwarke, which

is embedded tn hard matrix so that the interior is

completely obscured, It does suggest, however Uae

ancestors of Pledeju eyrernsis are inost Sikely ro be

fouid in the Trigoniidae

Achuowledpinents

Tam most erateful to My Neville [ledee of Ue

South Australian Musetim for showings me the

bivalves and to Mr G, W. Krieg and Mr PAL Ropers

of the Geological Survey of South Australia lor

describing the section in whieh (ey oocur and

collecting further material. [ thank the Chief

Government Geologist and Mr P J. G. Fleoring of

the Geological Survey of Queenstand for arranging

for the examination of the specimens in Queensland

and for the loan of specimen GSQ 2559, The

figures were drafted in the Drafijng Braneh of the

South Australian Deparurent of Mines and Every,

and photographic prince assistanee wis given by

the Biostratigraphy Section of the Geoleptcal

Survey of South Australia,

References

BURGER, D (1982) Palynology Of fhe Eromanga Basin

and is applications, J Moore, PS. & Mount, 1 4

(compilers) “Eromanga Basin Symposium, summary

papers", Geol. Soc, Aust. & Pet. Explor. Soc. Allst,

Adelaide, 174-182.

Con, b. R, (1952) Notes on che Trgeniidag, with outlines

of (he family, Proc, nielec. Soe. Lond. 29283), 45-70,

ply 34,

= (1955) On fhe alfinities of the Cretaceous tamelli-

brinch wenera. Trigunioides and Hosferrianeia, tree!

Miajr 92, 345-349,

Pornes. BG. (982) Margin of the Bromangn Basin,

Sourh Australia: a review: dn Matire, PS. & Mone,

T. J. (compilers) “Eromanga Basin Symposium,

summary papers”, Geol. Sov, Aust & Pet. Explor Soc,

Aust, Adelaide, (0-125.

Kiikayasny, T1956) On the deminon of fraonporaes

andits relation to siailar peleeypad gomers, Japa v

Geol Ceaser 171), T4494, pt 4,

— (J958) ‘The Cretaceous nan-mariine pelecynusdts

tron the Nan Phin Gyn Sie ii Une nertlwasvert pare

of the Korat Plateuu, Thailand, wih u note an the

Trigdmiaididac. Cantniitions ta the s¢otopy ovat

puldecontology of Soullicast Asia MLVIT Lived

Polaconr Souridust Asia VV, W9-13K, pls SNAQXITA

& Susan Ky (938) Non-marine sfretls of rhe

Moktonw-Wakwio Series. Japon < Ceol Geos, 13,

242-257

RHIG, Ce Wo (ISR2) Stoatiomaphy aud wwotanies af the

Dalhousie qutreties, soulhwesl Eromanga Basin, Ja

Moore, POS. & Mount 1 dconmpiters) “Keomanea

Basin Symposiiin, sunniary papers”. Ciool, Soe Aust

& Per Lxplon Sow Aust) Adelie. [45-158.

42 N. H. LUDBROOK

LAN Xiu (1976) Mesozoic lamellibranch fossils from

Yunnan, China, Mem. Nanjing Inst. Geol. Palaeont.

Acad. Sinica No. 7 (In Chinese).

Lupsreok, N. H. (1961) Mesozoic non-marine Mollusca

(Pelecypoda) from the north of South Australia. Trans.

R. Soe. S. Aust. 84, 139-147.

(1978) Australia. /n Moullade, M. & Nairn, A. E.

(Eds). ‘The Phanerozoic geology of the World’ I] The

Mesozoic, A., 209-249.

Martinson, C. G, (1965) Cretaceous lamellibranchs of

the family Trigonioididae and their classification.

Paleont. Zhurn. 4, 16-25 (In Russian).

(1984) The Late Cretaceous molluscs of Mongolia,

Malac, Rey. 17, 133 (abstract).

Moore, P. S, (1982) Mesozoic geology of the Simpson

Desert Region, northern South Australia, /n Moore,

P.S. & Mount, T. J. (compilers) “Eromanga Basin

Symposium, summary papers”. Geol. Soc. Aust, & Pet.

Explor. Soc. Aust. Adelaide, 46-57.

& Pitt, G. M. (1982) Cretaceous of the

southwestern Eromanga Basin: stratigraphy, facies

variation and petroleum potential, 127-144. /n Moore,

P. S. & Mount, T. J. (compilers) “Eromanga Basin

Symposium, summary papers”. (Geol, Soc. Aust. & Pet.

Explor. Soc. Aust.: Adelaide)

Morris, F. K. (1936) Central Asia in Cretaceous time.

Bull, geol. Soc. Amer. 47, 1477-1534.

Morton, J. E. (1967) ‘Molluses’. 4th (revised) edition.

(Hutchinson & Co.: London).

NEVEesSKAYA, L. A. & Sotoviev, A. N, (Eds) (1981)

Developments and change in molluscs at the Mesozoic-

Cainozoic boundary. Academy of Science USSR,

Paleontological Institute. Science (In Russian).

Ota, Y. (19594) Plicatounio of the Wakino Formation.

Trans. Proc, Pal. Soc, Japan. 1.5. 33, 15-18, pl. 3.

(1959b) Trivonioides and its classification. Trans.

Proc. Pal. Soc. Japan, n.s, 34, 97-104,

(1963) Notes on the relationship of Trigonioides

and Plivatounio, non-marine Mesozoic Bivalvia from

Eastern Asia, Geol. Rep. Hiroshima Uniy, 12, 503-512.

Skwarko, S. K. (1963) Australian Mesozoic trigoniids,

Bull. Bur. Miner. Resour. Geol. Geophys. Aust. 67.

Suzuki, K. (1940) Non-marine molluscan faunule of the

Siragi Series in South Tyosen. Jap, J. Geol, Geogr. 17,

215-231, pls 22-24.

WILLIAMS, A. F. (Compiler) (1975) LAKE EYRE map

sheet, Geological Atlas of South Australia 1:250 000

series. (Geol. Surv. S. Aust.: Adelaide).

Yano, S. Y, (1974) Note on the genus Jrigonioides

(Bivalvia). Trans. Proc, Palaeont. Soc. Japan N.S. 95,

395-408, pls. 54, 55.

(1976) On the non-marine molluscan fauna from

the upper Mesozoic Myogog Formation in Korea. Trans.

Proc. Palaeont, Soc, Japan N.S. 102, 317-333.

(1978) Ontogenetic variation of Trigonioides (s.5.)

paucisulcatus (Cretaceous non-marine Bivalvia). Trans.

Proc. Palueont. Soc. Japan N.S. 111, 333-347.

(1979) Some new bivalye species from the Lower

Gyeongsang Group, Korea. 7rans. Proc. Palaeont. Soc.

Japan N.S. 116, 223-234,

———— (1983) On the subgenus Wakinoa (Cretaceous non-

marine Bivalyia) from the Gyeongsang Group, Korea.

Trans. Proc, Palaeont. Soc. Japan N.S, 131, 177-190,

pls 38-40.

LARVAE AND PUPAE OF LECTRIDES VARIANS MOSELY AND

LEPTORUSSA DARLINGTONI (BANKS), (TRICHOPTERA:

LEPTOCERIDAE)

BY J. E. JACKSON

Summary

Final instar larvae and pupae of two species of Leptoceridae, Lectrides varians Mosely and

Leptorussa darlingtoni (Banks) are described and figured.

LARVAE AND PUPAE OF LECTRIDES VARIANS MOSELY AND LEPTORUSSA

DARLINGTON] (BANKS), (TRICHOPTERA; LEPTOCERIDAE)

by J. E. JACKSON*

Summary

Jacksos, J E, (V98S) Larvae and pupae of Leetrides vartans Mosely and Leptorussa carlingtont (Banks),

(Trichoptera: Leptoveridae). Trans. R. Soc. §. Aust. 109(3), 83-95, 29 November, 1985.

Final instar larva¢e and pupae of two species of Leptoveridac, Lectrides verians Mosely and Lepturussa

darlingion’ (Banks), ave deseribed and figured.

Kiy Wokus: Trichoplera, Leptoceridae, |arvae. pupae.

Introduction

The taxonomy of the immatures of Australian

Trichoptera is in an early stage of development

compared with thar of the northern hemisphere

fauna, for which keys of larvae to genera and

species have been developed (Hickin 1967, Lepneva

1964, Wiggins 1977). Identilication of Australian

larvae to family is possible using the key of

Williams (1980) as revised by Drecktrah (1984), and

& key to some free-living and nel-spinning genera

has been developed by Cartwright & Dean (1982).

Few detaited descriptions of Australian larvae are

available (examples are Riek (1968), Neboiss

(1977b, 1979), and Drecktrah (1984)), and For many

species (he larvae are not known. No immatures

of Australian Lepraceridae are adequately

deseribed, despite their dominance in the Australian

fauna (Riek 1970).

The leplocerid genus Lepforussa Mosely is

monotypic: L. russata Mosely is considered by

Neboss (1977a) fo be conspecific with L.

darlingtont. The species is recorded [from

Queensland, New South Wales, Victoria, South

Australia and Tasmania (Neboiss 1983); adults of

both sexes are described by Mosely & Kimmins

(1953) and. Neboiss (1977a), Lecirides Mosely

includes ZL, varigns and the recently discovered 2.

parilis Neboiss, the larva of which is unknown; L£.

barians is recorded from South-east Queensland,

Victoria and Tasmania (Neboiss 1983). Taxonomic

descriptions of the previously undescribed final

instar larvae and pupae of Lecrrides varians Mosely

and Lepterussa darlingtoni (Banks) were

underlaken prior to a comparative study of the rwo

species’.

* Department of Zoology, University of Adelaide, GPO

Box 498, Adelaide, S. Aust. S001

eieent uddress; 2S Austral ‘Tecrace, Malvern, S. Avot,

‘Jackson, J, BE (1984). Taxonomy, brology and case

funaion of Leetrides varians Mosely and Leprortssa

dartingtoni (Banks) larvae ( Crichoptera: Leptoceridae).

Honours thesis, Department of Zoology, University of

Adelaide Unpubl.

Methods

Identification of Leecrrides variuns and

Leptorussa darlingiont was confirmed by rearing

larvae and pupae to adults.

Deseriptions of final instar larvae are based on

preserved specimens collected from a site near

Heathfield, S. Aust., (139°39'E, 35°01'S); pupal

descriptions are based on pupae reared from larvae

in the laboratory, a Few collected specimens, and

pupal cxuviae, Whole specimens were prepared for

Microscopie examination by mascerahion of soll

parts in hol KOH (5%) for 10-15 minules, rinsing

in glacial acetic acid, and cleariny m clove oil.

Specimens were then dissected and prepared as

temporary mounts in clove oll, or permanent

preparations in Depex, Drawings were made [rom

slides with the aid of a drawing tube on a Wild M20

compound microscope and a camera lucida on a

Wild M3 stereomicroscope. Head capsule width of

larvae was measured to the nearest 0.01 mm, at the

level of the eyes, using an ocular micrometer.

Morphological terminology of larvae follows that

of Wiggins (1977),

Specimens ure deposited in the Museum of

Victoria, Melbourne.

Lectrides variuns Nasely

Material examined: 25 larvae, from near

Heathfield, S. Aust., April 1984, 9 pupae from

Heathfield, April 1984, and laboratory reared

Final instar larva

Cuse (Figs 4, 2). Constructed fram roughly

quadrate leaf fragments, overlapping in a dorsal

and a ventral row; anteriorly, dorsal fragment

projects further forward than ventral fragment,

forming a tood; Filaments of plant maternal (ill

between dorsal and ventral rows, and around tlic

anterior opening.

Larva (Figs 3-19), Body length [1-13 mim; bead

capsule width 0,78-0,81 mm, lengih t=1.13 mm-

Sclerotized parts golden brown, with distinct paler

84 J. E. JACKSON

Figs 1-6. Lectrides varians final instar larva; 1, 2, case, dorsal and ventral views; 3, larva, lateral view (most setae

not shown); 4, left lateral hump, enlarged; 5, abdominal segment LX and anal legs, dorsal view: 6, thoracic pleurites.

All seale lines in mm.

TRICHOPTERAN LARVAE AND PUPAE 85

Figs 7-13. Lectrides varians final instar larva: 7-9, head, dorsal, ventral and lateral views; 10-12, pro-, meso- and

metanota, dorsal view; 13, thoracic sternites. All scale lines in mm.

86 J. E, JACKSON

markings op head and thorax, dark bands on legs,

Head (Fizs 7-9) with pale spots on parictals,

occipital margins and posterior half of genac; ovate

in dorsal view, width about 2/3 length, Antennae

about length of right mandible, short sela apically,

Frontoclypeus strongly constricted medially,

exparided posteriorly to slightly greater width than

at anterior margin; 2 shallow concavities pn

poscerolateral margins; the resulting irregularities

emphasized hy adjacent pale spors on parietals,

Eyes situated dorsolaterally, just anterior co

frontoglypeal constriciion, surrounded by pale

area.

Labrum (Fig. (5) subovate, anterior margin with

shallow indentalion, dense fringe of short hair in

indentation, ventral brushes ef Jong. fair aniere-

laterally; median paired anterior setae shon, pale.

in tront of three pairs of setae in a transverse row,

Mandibles (Pig. 16) with rounded teeth distally on

cutting edges, six on left, five on right; lel) mandible

slightly longer than right, inner surface deeply

excavated, with two brushes; right mandible only

slightly excayaled, without brushes. Maxillo-labium

(Fig, 14) cardo digiliform with proximal end

broadly truncated, bearing one medial and one

distal seta; stipes incised medially on inner edge,

a lateral and shorter medial seta on distal margin;

maxillary palp four segmented, galea lJobale, with

three finger-like sensillae apically; submertal

sclerites absent, two long sybmental setue present;

labial palps conspicuous, cylindrical, Ventral

apoatome (Fig. 8) oblong, about 1/2 length of

parietals. width about 1/3 Jengih, broades}

anteriorly, tapered to almost 1/2 width posteriorly.

Pronotum (Fig, 10). Anterola(eral angles

projected forward, acute, Shehtly upturned, anienor

margin concave, shallowly crenate with sis pairs ot

Margitial selue arising in coristirictions; lransverse

row of setae just anterior to middle of notum;

posterior margin teavily sclerotized. Mesonotum

(Fig. 11) rounded posterolaterally, setae sparse,

three setae wrouped an each anterolateral angle.

Metanotum (Pig, 12) only partially sclerotized; 2

subquadrate sclerites anterumedially and smaller

paired selerites laterally; a single seta in each medial

sclerite, three in anterolateral sclerites, ore long arid

two very shart setae posterior to cach medial

sclenite, Prosternum (Fig, (3) with dark brown

sclerotized bar on posterior margin, smaller bar on

each side, pair of mesosternal sclerites, mela-

sternum wilh §3-10 setae in two curved (ransverse

rows, Most arising from small round sclerites,

Leps (Figs 17-19), Forelees (Fig, 17) short-

scements decreasing in size distally, coma and deour

subtriangular, coxa with bard of setae offset from

posterolateral margin. Fore-trochantin horu

shaped, upturned apically, one seta on ventral

marein. Trachariter with short spur on distal angle,

distally a ventral brush of short pale serae. Femut

fenzth about 24 width; row of short pale setae

ventrally. Tibla ovlindrical; anteriorly three dark

spurs, yentrally four; row of short pale setae

ventrally. Tarsus with three spurs vencrally: tarsal

claw stou, curved, sighily shorter than tarsus, basal

spuralinost length of claw. Midleg (Fig. 18) abour

to = Teng of fereley, segments subequal.

Trochanter with ventral brush. Femur bears seven

dark spurs ventrally and eomb of minute spines

basally on distal segment. Tibia with sever spurs

on anterior face, five ventrally, Tarsus with four

spurs, now of pale setae distally; tarsal claw stout,

slightly curved, basal spur short. Hindley (Pig. 19)

about 2% fengrh of midleg. Trochanter with row

of seven small dark spurs on disial 1/3, femur witlt

seven dark spurs ventrally, twa small spurs near

division. Tibia almost 2% length of other segments,

divided subequally, basal scement with five spurs,

distal with eight, Targus with six dark spurs

ventrally; tarsal claw pobyst, basal spur short,

Abdomen (Fig. 3) dorseventrally flatrened;

lateral fringe dark, on segment VII! reduced to only

a few hairs; segment VILI with lateral row of

spicules on anterior hall; hairs ai seginents MTX,

Il, tour dorsolateral; WI-V1I, two ventrolateral;

VILL, six dorsal two ventrolateral) LX, two dorso-

lateral, Lateral humps (Fig. 4) each with clavate

stlerite and three selae; dorsal hunip conical, two

pairs of fine setae, Tergite IX (Fig. 5) rectangular;

three pairs of marginal serae postenorly, cach outel

one shorter than others, wo pairs of very short

selae offset from margin. Lateral sclerites of anal

Jezs uniformly brown. Gills single, finger-hke, in

lollowing positions: sexment 1, pasteriur dorsal;

segments H—-V1, anterior dorsal, lateral and ventral;

segment Vif, anterior dorsal and ventral (presence

variable).

Pape (Pies 20-34), Length) 11.5-12.5 miny. Lees

without apical hooks or claws, fore- and midless.

with dese hair tiiecs, Antennal bases cach with

seven erect black selac. Labrum (Fi. 23) with lateral

mnargios straight, converging anteriorly, anterior

margin rounded: five pairs of setae anterolarerally,

Mandibles (Figs 28-31) broad-based, slender

apically; inner marvin roughly serrate, with or

without a well developed projection; pairsd setae

On outer Surlace ear base, Terminalia of abdomen

(Pies 32-34) with bwo small dorsal processes, each)

bearing four strong anlerolaterally directed setae:

wale (Fig. 33) with pair of fleshy clasper-shaped

ventral protuberances with bulbous process between

bases; female (Pig. 34) with ventral hump. Anal

processes rod-like, longer than 9th abdominal

seement, apices curved dorsad and pointed, 2-3

short sefaeé arising trom inner margin Subapically,

TRICHOPTERAN LARVAE AND PUPAE 87

Figs 14-19. Lectrides varians final instar larva; 14, maxillo-labium, showing association with head capsule, ventral

view; 15, labrum, dorsal; 16, mandibles, dorsal: 17-19, fore-, mid- and hindlegs, left, All scale lines in mm.

TRICHOPTERAN LARVAL ANTT PLIPAL ba]

a 41h aboul 2/3 from basal end. Posterior angles

of abdominal segment | bear transverse oblong

plates with thickened ridges (Fig, 24); segments

l1I-V1 bear elongate anterior hookplates (Figs 25,

26) with 2-3 prominent posteriorly-directed hooks,

segment V with rectangular posterior hookplaies

(Fie 27) bearing three anteriorly-directed hooks.

Seements U-VIl with paired shaped sclerites

dorsally and ventrally, proximal bar of T reduced

in ventral selerites.

Pupal Case (Pigs 20-22), Larval case modilied

hy closure anteriorly with oblique silk membrane,

with which attached to substrale; traosverse vent

in anrertor dorsal plate, through which larval

sclerites ejected; posterior Closure perpendicular leaf

Hawineht, onentation of pupa reverse of larval

orientation.

Leprorusse darlingtont (Banks)

Material examinect 20 \arvac, from near Heathfield,

S. Aust, April, May 1984, six pupae laboratory

reared,

Final instar larva

Cuse (Pig. 35). Length 812 mim, constructed ot

saidgrains, cylindrical, tapering posteriorly, slightly

curved; anterior margin usually straight ar with

small dorsal process; posterior membrane absent.

Larvae (Figs 36-55). Body length 6-8 mm); fread

capsule Width 0.54 rom, length 0.73-0,77 mim, Head

and thorax dark brown; distinct pale markings on

head, usually pale cross on frontoelypeis, thorax

mottled. Legs golden-brown with indistinet darker

bands

Head (Figs 43-45) ovare jn dorsal view, width

aboul 2/3 length Antennae about length of right

mandible, short seta apically, Frontoclypeus sub-

pandurale, Eyes situated dorsolaterally, just anterior

to franta-elypeal constriction, surrounded by pale

area.

labrum (Piy. $2) subovate, anterior margin with

four setae in shallow indentation, one long seta

arming Crom each anterolateral raargin, medially

three payrs of setac in transverse Tow. Mandibles

(hies 54, 55) with prominent teeth distally, sin on

let{, live on right; left slightly longer than right,

inner surface deeply cacavated and bearing two

small brushes: right only sightly excavated, without

brushes, Maxillo-labium (Pig, 53): eardo blunt

proximally, attenuated laterally, bears | medial und

one distal seta; stipes with one lateral and one

shorter medial seta on distal margin; miuxillary

palps three segmented, galea lobale, a sinyle sensilla

apically: submental selerites quadrate, two long

submental setae; palpigers Z-shuped, very narrow,

Ventral apotome (Fig. 44) rectangular, width about

2/3 length, slightly narrower anteriorly,

Pronotum (Fig. 46) rectangular anterior margin

pale, with 6-7 pairs of regularly spaced setae: lateral

margins (Fig. 40) with row of about seven setae;

posterior marvin heavily sclerotized, Mesonolum

(Fig. 46) rounded posterolaterally, four pairs of

scattered setae dorsally, aboul seven setae grouped

in each aiterolatéral afale, Metanotum (Pig. 46)

mostly membranous; medially two small irregular

selerites, bearing one short seta on anteroproxima!

angle, laterally three long setae, Prosternal sclerite

absent, two small dark mesosternites (Pigs 47, 48),

metasteroum with 4-3 pairs of setae,

Legs (Figs 49-51), Anteriar face of coxae much

more densely setose than posterior face, other

seginenls wilh few setae. Forelen (Fig. 49) short and

robust. Fore-trochantin (Fiz. 40) truncated distally,

anterodorsal angle produced torward slightly, two

setae Of anterior Margin, Trochanter with two pale

spurs near distal angle, brush of short pale setae

distally. Femur width about 2/3 length, camb of

small pale spurs ventrally, Tibia dilated distally, four

spurs on ventral margin. Tarsal chameter about 1/2

that of tibia at joint, six spurs ventrally; tarsal claw

as long as tarsus, basal spur about 1/2 tenpth.

Midleg (Fie. 40) almost 2x length of foreleg, less

Tobusi, segmienis (except for short trochanter)

decreasing in sive distally. Distal seemeni of

trochanter with ventral brush of pale setae. Femur

hears comb of smal) pale spurs near base of distal

sezment. Tibia curved basally, spur ansing 2/3 from

basal end. Tarsus with ventral row of small pale

spurs on distal 2/3. Hind lep (Fig. 51) slender, alyour

2” leneth of midleg, segments subequal

‘Trochanter with comb of small pale spurs on distal

1/2 of posterior face. Femur with similar comb at

base of distal seginent. Tibia bears one spur ucas

distal end. Tarsal spur arises mediallys tarsal claw

slender, curved slightly to a fine point, length about

2/3 Jength of tarsus, basal spur about 1/4 of length.

Abdomen (Pig. 36) creamy white, shehtly darsoa-

venirally Tlattened; lateral Iringe shorl, line;

seement VITt with complete lateral raw of spicules.

lareral fringe absent: seerments VI-VIL with dorsal

hairs. V1I-?, Vil-4d, VU-6. Lareral trumps of

segment | (Fig. 37) each with @ selerotzed bar,

Flos e354, Leevides variens pupa. 20-22, case, anterior end in ventral wad dorsal views, posterior closure; 24, tabruny

and aniwelypeus, dersal. 24, riele selerorived plate of abdominal segment |; 25-27, righe abdominal hookplares,

antenor hookplate dorsal and lateral view, posterior hookplate dorsal view; 28-31. miandibtes, felt without wirier

projection (28-29) and with inner projection (40-31), 32~34, terminalia of abdomen, dorsal, male vemtrul, utd

female lateral views. All scale lines in mm-

90 J. E. JACKSON

Figs 35-42. Leptorussa darlingtoni final instar larva: 35, case, lateral view; 36, larva, lateral view (gills not shown,

most setae not shown); 37, left lateral hump, enlarged; 38, dorsal hump, dorsal view; 39, abdominal segment |

and anal legs, dorsal view; 40-42, pro-, meso- and metapleurites. All scale lines in mm.

TRICHOPTERAN LARVAE AND PUPAE 91

Figs 43-48. Leptorussa darlingtoni final instar larva: 43-45, head, dorsal, ventral and lateral views; 46, thoracic nota,

dorsal view; 47-48, meso- and metasterna, enlarged mesosternite. All scale lines in mm.

92 J, E. JACKSON

Figs 49-55, Leptorussa darlingtoni final instar larva: 49-51,fore-, mid- and hindlegs, left; 52, labrum, dorsal view;

53, maxillo-labium, showing association with head capsule, ventral view; 53, labrum, dorsal view; 54-55, mandible,

interior face. All scale lines in mm.

TRICHOPTERAN LARVAE AND PUPAE 55

Tait lL. Arrangement of the gills of L, daclingtoni.

Seement Dorsal Lateral Ventral

1 p 3br _ p 2br

i a Jor a 4br a 3tr

il a Shr a lbr a 3br

IV a 357 _ a 3br

Vv a Qbr — a /2be

Vl a tbr — a 1/2br, present/absent

Vu a 1/2br _ 1 thr, present/absent

Pe posterior, a=anterior, br=branched-

spinulose distally, a single medial scta; dorsal hump

(Fig. 38) flat, with medial transverse sclerotized

band, roughened with minute spicules anteriorly.

Tergite LX (Fig. 39) width greater than 3* length,

mostly unpigmented, with irregular dark spots;

three selae laterally on postenar margin, medial

setae much longer than others. Lateral sclerites of

anal legs mostly unpigmented in anterior 1/2, few

irregular spots, Anal claw with one small dorsal

accessory hook, Gills Hilamentous, branched,

arrangement as in Table |,

Papa (Figs 56-66). Length approximately 8 mm.

Legs without apical hooks; fore- and midlegs with

dense hair fringes, those of foreleg less dense.

Antennae long, coiled several times around end of

abdomen, antennal bases without setae, Labrum

(Fig. 61) semicircular, two short setae anteriorly,

live lony setae on each lateral margin, Mandibles

(Fig. 61) pincer-shaped, distal 1/2 of inner margin

serrate, (wo Setae On Outer Surface near base,

Terminalia of abdomen (Figs 63-66) with two small

dorsal processes, each bearing four strong antero-

Jaterally directed setae; male pupa (Fig. 63) with

three bulbous ventral protuberances, Anal processes

(Fig. 66) rod-like, longer than 9th abdominal

segment, apices curved dorsad and pointed; sparsely

covered with short pale hairs, rwo long setae arising

from inner marpins subapically, one shart seta basal

(o these and fourth about 1/5 from basal end.

Abdominal segment | bears pair of transverse

oblong plates with thickened ridges; segments

IH-V1 bear elongate anterior hookplates (Fig. 59)

with 2-3 prominent posteriorly-directed looks;

segment V with rectangular posterior hookplates

(Fiu. 60) bearing 2-3 anteriorly-directed hooks.

Segment U-VLi with paired T-shaped sclerites

dorsally and ventrally, proximal bar of T reduced

in ventral sclerites.

Pupal Case (Vigs 56-58). Larval case modified

by closure anteriorly with plate of sand grains

bound with silk just in from rim (Fig. 57):

posteriorly thick silk membrane, intact until larval

ecdvsis when trefoil-shaped vent is formed (Fiz. 58);

pupa in laryal orientation, pharale adult escapes by

opening anterior plate like trapdoor. Case attached

to substrate with stalked disks (Fig. 56) anteriorly,

or anteriorly and posteriorly.

Acknowledgments

Lam prateful to my Honours Supervisor, Alice

Wells, and to Margaret Davies tor helplul guidance

in the preparation of this manuscript

Keferences

Cartwricht, D. b & Dean J. Cc. (1982) A key to

Victorian genera of free-living and retreat-makine

Caddistly larvae (insecta, Trivhoplera). Mem. naint

Mus. bretdrta. AS, i134.

Drerekykau, G (J9R4) Deseniption of the immature

stages Of ANvecella grisea Banks (Trichoptera: Helico-

phidac) and morphological characierisics used te

distinguish between larvae of Australian Cualocicae,

Conoesucidac and Helicophidae pp. 115-122. fa 1c

Morse (Ed) ‘Proc. 4th tnternain. Symp. Trichoptera,

(utik: The Hague).

Hickin, So E. (1907) “Cauddis Larvae. Larvae of the

Hritish Trichaptera”. (Hutchinson: London).

Livny, S. Ce (1964) Fauna SSSR, Rucheimiki, val. 2,

no lt, Lichinki i kukolki pocdotryada kol’chatosh-

chupikovykh. Zoolovicheskii Institur Akademi) Nauk

SSSR, 0.5. 88. [In Russian. Translated into Eqelish as:

Fuuna of the US.S.R3 Irichoptere, vol. 2, no. L. Larvae

and Pupae of Avvulipalpia. Published by the Israel

Program for Scientiic Translauions, 1970).

Mosety, M, E. & Kinmins, D. EL (1953) “The trichoptera

(Caddisthes) of Australia and New Zealand”

pp 27-276 (British Museum (Natural History);

London).

Neboiss, A, (1977a) A taxonomic and zoogeopraphiec

study of Tasmanian caddisflies (Insevia: Trichoptera),

Mem. aaint. Mis, Fictoria, 38, 1-208.

—— (1977b) Avripleetididae, a new caddisfly family

(Trichoptera: Atriplectididac). pp. 67-23. In M. 1.

Crichton (Ed. ‘Proe. 2d Internuin, Symp. Toghaptens *

(Junk: The Tague).

(1979) A terrestrial caddisNy larva from Tasmania

(Calocidae: Trichoptera). Awst Entomol Mae. 5, 90-93.

(1984) Checklist and bibliography of the Australian

Caddis-Flies (Trichoptenn). Aust Sou Linmnot, Spec.

Publ §

Rirk, E. F. (1968) A new family al caddis flies trom

Australia (Trighopiera! Tasimiidae). 4 Aust, Ene Soe.,,

7 19-1b4.

94 J. E, JACKSON

63\,

) wa” i

TRICHOPTERAN LARVAE AND PUPAE 95

(1970) Trichoptera (Chapter 35): pp. 741-764. In WicGains, G. B. (1977) “Larvae of the North American

I. M. Mackerras (Ed.) “The Insects of Australia”. Caddisfly Genera (Trichoptera)”. (Univ. of Toronto

(Melbourne Univ. Press: Melbourne). Press: Toronto).

WILLIAMS, W. D. (1980) “Australian Freshwater Life”.

2nd edn. pp. 255-266. (Macmillan: Melbourne).

Figs 56-66. Leptorussa darlingtoni pupa: 56-58, case, stalked-disk attachment, anterior and posterior closures; 59-60,

anterior and posterior hookplates; 61, labrum and mandibles, dorsal view; 62, mandible, enlarged; 63-66, terminalia

of abdomen, male ventral, female ventral, male lateral views, enlarged anal process. All scale lines in mm.

FOUR NEW SPECIES OF HYDROPTILIDAE (TRICHOPTERA) FROM THE

ALLIGATORS RIVERS REGION, NORTHERN TERRITORY

BY A. WELLS

Summary

Four new species of micro-caddis fly (Hydroptilidae) from the Alligator Rivers region, Northern

Territory, are described in the genera Hellyethira, Tricholeiochiton, Oxyethira and Orthotrichia;

notes are given on their possible relationships. Collecting data for three species suggest that these

show strong seasonality, with two species emerging almost exclusively in the dry season (July), and

one in the wet (March).

FOUR NEW SPECIES OF HYDROPTILIDAE (TRICHOPTERA) FROM THE

ALLIGATOR RIVERS REGION, NORTHERN TERRITORY

by A. WELLS*

Summary

WELLS, A. (1985) Four new species of Hydroptilidae (Trichoptera) from the Alligator Rivers region,

Northern Territory. Trans. R. Soc. S. Aust. 109(3), 97-102, 29 November, 1985.

Four new species of micro-caddis fly (Hydroptilidae) from the Alligator Rivers region, Northern Territory,

are described in the genera Hellyethira, Tricholeiochiton, Oxyethira and Orthotrichia; notes are given on

their possible relationships, Collecting data for three species suggest that these show strong seasonality,

with two species emerging almost exclusively in the dry season (July), and one in the wet (March).

Key Worps: Taxonomy, Trichoptera, Hydroptilidae, Alligator Rivers, seasonality.

Introduction

Regular monitoring of “emergence” traps on three

natural water bodies near Jabiru in the Alligator

Rivers region, Northern Territory, yielded

information on species diversity, seasonality, and

diel activity of a number of Trichoptera (Sharley

& Malipatil 1985, and unpublished data). Amongst

the microcaddis flies (Hydroptilidae) collected (14

species in four genera), were four species which are

described here for the first time, and which are

referred to the genera Hellyethira Neboiss,

Tricholeiochiton Kloet & Hincks, Oxyethira Eaton

and Orthotrichia Eaton.

Within Australia, Tricholeiochiton is known only

from the north (Wells 1982), where five species have

now been recorded. The genus is elsewhere

represented by a single species in the Palaearctic and

two species in S.E. Asia (Marshall 1979). The sub-

genus Dampfitrichia in Oxyethira, into which one

new species is placed, has a similar northern

distribution in Australia where four species are now

known; it occurs elsewhere in S.E. Asia, the

Neotropics, the Palaearctic, and the Western Pacific

(Kelley 1984, and pers. comm.). In contrast,

Hellyethira and Orthotrichia are widespread in

Australia (Wells 1979a & b, 1983), and now total

18 and 35 known species, respectively. One

Australian species of Hellyethira is recorded from

New Caledonia (Wells unpublished data), and

Kelley (1984) has transferred a Japanese species to

this genus. Orthotrichia is absent from SW

Australia, but is especially diverse in the north.

Seven species were collected in the Alligator Rivers

study (Sharley & Malipatil unpublished data); about

54 species are known from elsewhere in the world.

Emergence data for the three seasons that were

monitored, late-dry 1982 (October/November), wet

1983 (March), and dry 1983 (July) (Supervising

* Department of Zoology, University of Adelaide, G.P.O.

Box 498, Adelaide, S. Aust. 5001.

Scientist for Alligator Rivers region 1984; Sharley

& Malipatil 1985!) give some indication of life

cycle patterns of a number of species and suggest

that three of the new species, at least, are highly

seasonal. Interestingly, each of these appears to be

more closely allied to a north-eastern than a north-

western congener.

Materials and Methods

All material was collected in the Alligator Rivers

region, Northern Territory, from three natural water

bodies, the Magela Creek (at N-T. Water Division

gauging station 821009), Georgetown Billabong and

Corndorl Billabong. Trapping methods are

described in the Alligator Rivers Region Research

Institute, Research Report 1983-84 (Supervising

Scientist for Alligator Rivers region, 1984).

Methods for preparation, drawing, and storage

of specimens follow Wells (1978). All material,

including types, is the property of the Northern

Territory Museum of Arts and Sciences, Darwin

(NTM), where it is lodged.

Hellyethira veruta sp. nov.

FIGS 1-3

Holotype: NTM I.56¢, Magela Creek, S. of Georgetown

Billabong, N-T., 11.vii.1983, A, J. Sharley.

Paratypes: NTM 1.57-63 3¢, 4? (including allotype 1.60)

collected with holotype, NTM 1.64-89 16¢, 10%, same

locality, A. J. Sharley, 1.vii.1983; NTM 1.90 19, 27.vi.1983,

NTM 1.91-94 49, 29.vi.1983, Georgetown Billabong, nr

Jabiru, N-T., A. J. Sharley; NTM 1.95 19, 30.iii.1983, NTM

1.96 19, 22.iii.1983, Corndor!l Billabong, nr Jabiru, N-T.,

A. J. Sharley.

Other material examined: 7c', 10%, 27.vi.1983, 26, 29,

29.vi.1983, 4d, 109, 3.vii.1983, Ig, 29, S.vii.1983, 4d, 99,

' Sharley, A. J. & Malipatil, M. B. (1985) Aquatic insect

emergence from waterbodies in the vicinity of Ranger

Uranium Mine, Jabiru, Northern Territory. Supervising

Scientist for the Alligator Rivers Region, Res. Rep. 1985

(unpublished).

98 A. WELLS

Figs |-S. Hellyethira veruia sp. nov. 1, 2. genitalia in dorsal and ventral views. 3. 2 terminalia, ventral view. Tricho-

leiochiton jabirella sp, nov. 4. ¢ terminalia, ventral view. Oxyethira warramunga sp. nov. 5, ¢ terminalia, ventral

view. Abbreviations: ae., aedeagus, d.pL, dorsal plate; inf.app., inferior appendage; pr., paramere; VII, VII, LX,

abdominal segments VII, VIIL, and EX, Scale bars = 0.1 mm.

FOUR NEW HYDROPTUIDAE 39

TVILIVBS, Ad, 42. YM T98S, Ut, AY, WWI 1983, 2 22,

15.vii 1983, bo, 12, (7) MORI, We WY, WYVTYRS, 1h be,

21984, Ao BviLl9BS, be, IV, WHi19B3, fy, 1,

2G 19S, 44, 22 A AGSS, UP, WAU GRS, 2G, 2KA.1YBS,

Mayela Creek, S. of Georgctown Billabong, NUL, A. J.

Shrarley,

Dyagnosis? Medium sized, with mottled fawn-cream

vestibure,

Male. Length of anlerior wing 1,5-b.7 mim,

Antenpuc 30-sepmented. Genitalia symmetrical, as

in Figs t, 2. Inferior appendages multilobed,

venttal-mos! lobe elongate, slender, bifld distally;

second lohe also slender and elongate; upper lobe

plate-like in ventral view, with apicatateral angles

attenuated. Dorsal plate membranous, with a

median serrale “rulf. Paramere slender, curved,

intersecting, medially, No subgenilal plate evident,

Female. Length of anterior wing 16-2.1 com,

Terminalia (Fig. 3) short, sternite Vil sfighrly

bilobed apicomesally, cach lobe tipped by paired

setae,

H. veruta shows close sitmilarity to AL eskensis

(Mosely) and H. sentisae Wells (Wells 1979a) which

have similar scissor-like parsmeres, The inferior

appendages more closely resemble those of A,

eskensis, although the dorsal plate of A, veruta ts

distinct rom those of the other two species, Hither-

to, | considered H, eskensis and H. seniise to be

close sister species; the precise nature of their

relationship to H. verutais unknown, bul the tire

species Clearly form a distinct tiveage within

Hellvethira,

H. #rura appears to be quite seasonal: it 1 absent

from Jate-cry season collections, rare in those from

the wet seasou, and most abundant in the dry

season Samples, Most speeimens were taken from

Mapela Creek and only few from the two lagouns.

At the sampling sile Magela Creek has a witite sand

substrate. and laoks the macrophyte community

found in (he lagoons; to the dry season the water

recedes to form a small stagnant pool in rhe streans

channel (Malipatil pers. comm.),

Trieholeloehitan jabiretla sp. ney,

FiGs 4, 6, 7

Halotype: STM L97 ft. Coridurl! Wikabons, we Jabire,

NOP, 20.717,1983, A. L Shanley.

Maratypes NTM 1.98 Ic 1.99 talletyyse) 2 collected with

Netotyye; NEM TI) te Vii 983, NTM TOL ty,

16ji,1983, NTM 1102-103 2% 18.1b.1983, Corndarl

Billabong, nr Jabiru, NT; NTM 1.104 bf) Cieargecown

Killabong, ar Jabiru, N.T., A, J, Sharley, 7), 1983.

Diagnosis: Medquri sized, with mottled wings, tips

of wings uplurmned, Ry and Ry arise indendently in

anterior wings,

Male Length of anierion Wire 2,2-2,5 mm

Antenoae 32-segmented, Genitslia as id Figs 4, 7

Segment X sheer, steenite with broact, deep, median

excavation, lateral lobes sclerotised distally. Dorsal

plate trilobed, membranous, central lobe

subquadrate. Subgenital plate membranous, broad,

narrowly cleft apicomesally. Interior appendages

stoul, slightly extended apicomesally to form

rounded, sclerotised lobes, Aedeagus with 4 black

spine arising subapically and projecting slightly

heyund apex.

Female, Length of anterior wing 2.5 mm, Antennae

32-seymenied, Terminalia (Mig, 4) short, pale;

segment VIL simple collar-like.

7, jéthirella is nvost closely allied to 7: fidelis Wells

{Wells 1982) With which it shares the general form

of ils male genitalia and wing venation However,

itis readily distinguished by the black spine on the

accleagus, broader inferior appendages, and bilobed

subgenital plate, Both species were collected from

the same localities in the emergence traps. T

Jabivella appears to be highly seasonal, being taken

oniy in the wet season (March), while 7) Jidelis was

vollected ip all seasons (Sharley & Matipatil

Unpublished data). Although congeners were

collected fram sagoons only in the Alligator Rivers

region, 7, fidelis has been taken from rivers and

streams in NE Queensland, Both Georgetown and

Corndor! billabongs have black clay substrates and

twacrophyte communities, and have slow flow only

in the wet season (Malipalil pers, comni.).

Oxyethira (Dampfitrichia) warramungs sp. nov.

FIGS 5, 8,9

Halotype: STM 1105 ¢, Goorgetawn Billabong, ar Jabirn,

21.vib.t983, A, 1 Sharley-

Paretypes: SM 1406-100 Id, 42, fineluding allotype

C107), 2Lvif,t983, NTM 1110-12 39, 27,vi.1983, NTM

(119-106 be, 47, INILIG83, NTM L.AIT IIB Id, 1%

15.94.1989, NTM $099 te, (vil 1983, NM 1120 19,

29yi,1983, NTM Li2k Id, 274i1,0983, Georgetown

Billabong, fir Jabiru, NT, A, J, Sharley, NTM L122 Is,

391.1983, NTM 1.123 12, 20.yii, 1983, Magela Creeb, 3, of

Cicorgerown Billabong, NUT, A. +. Sharley; NTM L124 [dy

Corndorl Billabong, nr Jab, NT, 238i1.1983-

Olher material exariined 19, 29vi1983, 2%, Swi IORI, 12,

$i) 1989, 22, 25.vil 1983, Georgetown Billabony, nr Jabaru,

NAT, A. 2 Sharley, NTM: 12, 262.1983, 1?) 276).19R3,

34, Ti 1983, 42, LiL LBS. 39, Si. 1983, NY. Sv 1983,

1%, Q.vil 1989, 12, 1491,9993, 22, 27.vii.1983, Magela Creek,

S of Georgetown Billabong, NTA. J. Sharley, NTM

Diagnosis: Minute caddisflies with mottled

vestiture; spur formula 0,2,4,

Male. Length of anterior wing 1.1-1.5 mm,

Aniennae 26-seemented. Genitalia as in Figs &, 9

Segment VIT1 broad, rounded, sternite widely and

shallowly excased apicomesally; tergite willy a hroad,

deep mesal concavity. Dorsal plate membranous,

rounded, broad at base, narrower distally, Aedeagus

stoyt, wifhauy titlater. Inferior appendages discrete

distally, fused at base; pajred fobes dorsal to inferior

FOUR NEW HYDROPTILIDAE

Imm

Figs 10-12, Orthvtrichia eurhinata sp. nov. &, 10. Tih antennal segment, dorsal view; [1, 12. genitalia, dorsal and

ventral views. Abbreviations: ae, aedeagus; d.pl., dorsal plate; d.prinf., dorsal process of inferior appendages; infapp.,

inferior appendages; pr, paramere; rds., right dorsal spine; sau., sensilla auricillica; sc., sensilla coeloconica; sp.,

sensilla placodea; v.h., vestitural hairy VIL, IX, abdominal segments VIII and IX. Scale bars as indicated.

appendages may represent the subgenital plate.

Female. Length of anterior wing 1.4-2.0 mm.

Antennae 20-segmented; stapes about 2~ length of

pedicel. Terminalia as in Fig. 5. Sternite VIII

triangular, apex with a narrow sclerotised band:

tergite shallowly concave apically,

O. warramunga is clearly a member of the

minima group in O. Dampfitrichia, to which two

other north Australian species, O. artuvillosus

(Wells) and Q. plumosa (Wells), belong (Kelley

1984). However it lacks their distinguishing

features—areas of androconia on abdominal tergite

VI, elongate hairs on the hind tibiae, and a patch

or border of stout black setae apicomesallly on

sternite VIII. The female terminalia of O

warramunga are distinguished only by the shallowly

concave apex of tergite VILI, compared with the

angular vertex of O. artuvillasus, In features of male

genitalia O. warramunga is probably closer to O.

plumosa than to O. artuvillosus.

Only a single specimen was collected in the late

dry season, all others were taken in the dry season

at all three localities. One male and three Females

of O. artuvillosus were also collected from Magela

Creek in the dry season of 1983 (Sharley & Malipatil

unpublished data).

Figs 6-9. Tricholeiochiton jabirella sp nov. 6, 7. 8 wenitalia, dorsal and ventral views. Oxvethira warramunga sp.

nov. &, 9, é genitalia, dorsal and ventral views. Abbreviations: ae., aedeagus: d.pl., dorsal plate: infapp.. infenor

appendage; sub.g., subgenital plare; VIII, abdominal segment VILI. Seale bars

~ Ot mm.

102 A. WELLS

Orthotrichia eurhinata sp, nov.

FIGS 10-12

Holotype: NTM 1.125 ¢, Georgetown Billabong, nr Jabiru,

17.yi.1983, A. J, Sharley.

Diagnosis: Known only from a single medium-sized

male with distinctive antennae: segments of

proximal 's of flagellum stout, broader than long,

subsequent segments becoming more elongate

towards lip; proximal segments with whorls of

vestitural hair on either side of a broad band of

sensilla auricillica (Fig. 10), distal segments with

incomplete bands or patches of auricillica.

Length of anterior wing 1.7 mm. Antennae

24-segmented; 4 terminal segments dark, preceeded

by J pale, 4 dark, 2 pale, and 13 dark segments.

Genitalia as in Figs 11, 12. Right dorsal spine only

present, elongate, tapering and curved across dorsal

plate, Dorsal plate rounded apically, wrapping

around aedeagus; a small blunt subapical process

on dorsum. Inferior appendages small, discrete,

symmetrical; dorsal process short, divided apically;

basal apodeme elongate slender. Paramere thin,

curved,

Female unknown.

O. eurhinata is in the O. adornata group in the

Australian Orthotrichia and is probably closest to

O. bullata Wells, another north Australian species

(Wells 1979b), although O. bullata has more

elaborate dorsal spines and larger, rounded interior

appendages. The form of the antennae of O.

eurhinata is quite unique, at least amongst the

Australian and New Guinean members of the genus,

Which have scattered vestitural hair on all segments

and large numbers of sensilla placodea (Wells 1984).

Acknowledgments

! wish to thank Dr Malipatil who kindly made

the Alligator Rivers material available to me

through the Northern Territory Museum of Arts

and Sciences, and who provided information on the

habitats, Mr A. J. Sharley who made the collections,

and the typing and technical staff of the Zoology

Department, University of Adelaide, for their

assistance in preparation of the manuscript.

References

Keiiey, Ro W. (1984) Phylogeny, morphology and

classification of the micro-caddis fly genus Oxyethira

(Trichoptera: Hydroptilidae). Trans. Am, Eni. Sac, 110,

435-463.

MARSHALL, J, E. (1979) A review of the Hydroptilidae

Cittchopters). Bull. Br. Mus. (Nat. Hist.) Entomol, 39,

135-239,

SUPERVISING SCIENTIST For ALLIGATOR RIVERS REGION,

(1984) “Alligator Rivers Region Research Institute,

Research Report 1983-1984". (Aust. Govt. Publ. Serv.

Canberra. 1984).

WELLS, A. (1978)[1979] A review of the Australian species

of Aydroptila Dalman (Trichoptera: Hydroptilidae)

with descriptions of new species. Aust. J. Zool. 26,

745-62.

(1979a) A review of the Australian genera

Xuthotrichia Mosely and Hellvyethira Neboiss (Trich-

optera: Hydroptilidae), with descriptions of new species,

Aust. J Zool. 27, 31-29.

(1979b) The Australian species of Ortharrichia

Eaton (Trichoptera: Hydroptilidae). Aust J. Zool. 27,

385-22.

(1982) Tricholeiechiton Kloet and Hincks and new

genera in ihe Australian Hyeroptilidae (Trichoptera).

Aust. JE Zool. W, 251-70,

(1983) New species in the Australian Hydroptilidae

(Trichoptera), with observations on relationships and

disiributions. Aust. . Zool. 31, 629-49,

(1984) Comparative studies of antennal features

of Australian Hydroptilidae (Trichoptera). Jo J. Morse

ed. “Proc. 4th Internain. Symp. Trichoptera™. (Junk:

The Hague).

A NEW SPECIES OF UPEROLEIA (ANURA: LEPTODACTYLIDAE) FROM

THE PILBARA REGION, WESTERN AUSTRALIA

BY MARGARET DAVIES, M. MAHONY & J. D. ROBERTS

Summary

Uperoleia glandulosa sp. nov., is described from the Pilbara region of Western Australia. The new

species is sympatric with U. russelli. Call information and karyotype are included in the species

description.

A NEW SPECIES OF UPEROLETA (ANURA: LEPTODACTYLIDAE) FROM THE

PILBARA REGION, WI

"STERN AUSTRALIA

hy MarGareT Davies*, M, MAHnONYT & J. D. Ropertst

Summary

Davies, M.MAvoney, M, & Roperrs, J.D. (1985) A new species of Uperoleia (Anura: Leptodactylidue)

from the Pilbara Rewion, Western Australia. Trans. R. Sow. S Aust. 1094), 103-108, 29 November, 1985.

Uperelcia glandtilosa sp. nov., is described trom the Pilbara fegion of Western Australia, The new

species issympatric with U. russe//i, Call information and karyotype are included in the species deseripuon,

KeY Worms: Uperoleta glandulosa sp, nov., advertisement calls, Anura, Leplodactylidae, karyotype,

Totroduction

In revising the genus L/peroleia Gray, Tyler,

Davies & Martin (L98la) cestricted the distribution

of the nominate spevies LL marmoratu Gray to the

type locality “Western Australia”, estimated from

Gray’s journals to be in che Prince Rezent River

areca, Later they recorded the distribution of U,

miobersi (Andersson) as confined to the Fitzroy

River drainage (Tyler ev of, 1981b). Hence records

of LU. marmarat and unsubstantiated records of

U, myobergi (as Glavertia mioberei) in the Pilbara

region of Western Australia (Mam 1965; Main &

Storr 1946) cannol be sustained,

A third species, U. russell/ (Loveridye) is recorded

from the Pilbara and js considered endemic co the

region (Main 1965; Main & Storr 1966; Tyler e/ al,

}98la; Tyler, Smith & Johnstone 1984),

During. field work following Tropical Cyclone

Jane in January 1983, two of us (J,D.R, and M.M.)

collected two species of Uperoleia near Port

Hedland. One of these species is U. russel/i and (he

other 15 conspecific with the specimen referred to

by Main (1965) and Main & Storr (1966) as U

marmarata and which is undeseribed,

Here we describe the new species and include

brie! notes on its breeding biology, call and karyo-

type

Materials and Methods

Material cited here is deposited in the Australian

Museum, Sydney (AM), the Sourh Australian

Museum, Adelaide (SAM{ and the Western

Australian Museum, Perth (WAM)-

Measurements of specimens follow Tyler ev al,

(198lap. Meastirements taken (in mm) were: eve

diameter (E); eye (o naris distance (B-N); internartal

* Department of Zoology, University of Adelaide, GPO.

Box 498, Adelaide, S. Aust. S001.

J} School of Biological Sciences, Macquarie Umivernity,

North Ryde, NSW. 2113.

j Department of Zoology, University of Western Ausiralia,

Nedlands, W.A. 6009.

span (IN); snout to vent length (S-V) and tibia

length (TL),

Osteological data were obtained from cleared and

alizarin red 5 stained specimens prepared aller the

method of Davis & Gore (1947), Qsteological

descriptions follow Trueb (1979),

Calls were recorded in the field on a Sony

TC-S10-2 reel to reel tape recorder, with Beyer MIOI

of AKG D190 microphones al a lape speed of 19 cm

s~', Six call parameters were analysed in detail: (1)

mean call period (seconds) estimated from the time

taken 10 make 20 calls (time taken with a stop watch,

halfspeed playback; (2) call duration; (3) pulse

duration; (4) pulse period; (averaged over the whole

call) (2-4, all (on milliseconds and measured from

oscillagrams made on a San-e7 5M28 direct recor-

ding oscillograph); (5) pulse number (counted

oscillograms); and (6) dommant frequency, averaged

over five calls by a Tektronix SLAN spectrum

analyser. For the first five parameters, we analysed

live ealls from each recorded frog, Pulse durations

were estimated (rom all pulses in the call, Periods

rather than repetition rales are reported as |hese are

the basic units measured; repetition rates represent

a derived unit (Gaioni & Evans 1984). Repetition

rates are 4 simple reciprocal function of periods,

Data for cach frog were averaged and the mean

Yalues were Used te calculate an overall mean, Each

individual thus contributes one datum point to the

final Figures,

Miototic chromosames were examined from

WAM R89489, using the technique described by

Mahony & Robinson (1880). Silver staining was

canned out following the procedure of Bloom &

Goodpasture (1976).

Uperoleia glandulosa sp. nay

FIGS 1-7

Uperoleia marmoran Maby 1965, Frogs of Soutl) Westert

Australia, p, 76 (parc; Maen & Store, LO66. WL Aust.

Nal. 10, p, 57; Cogger, 1983, p. 84,

Holotype: WAM RB9IS89, an adult male collected

104 M. DAVIES, M, MAHONY & J, DD ROBERTS

Fig. 1. A lateral, B dorsal view of head of Uperoleia xlandulasu sp. nov. (SAM R27081).

at Petermarer Creek, Port Hedland-Broome Rd,

W.A, (21°23'6"S, 118°48'21"E), on 10.1.1983 by M.

Mahoney and J. D, Roberts.

Definition: A moderate sized species (males

20-25 mm, female 25 mm S-V) with moderately

long limbs (TL/S-V 0,34-0.41). Maxillary teeth

absent; frontoparietal fontanelle widely exposed;

basal webbing between the toes; toes fringed;

mating call a sharp click.

Description of holotype: Maxillary teeth absent.

Vomerine teeth absent. Snout short, truncated when

viewed from above and in profile. Eye to naris

distance greater than internarial span (B-N/IN

1,36), Canthus rostralis prominent and straight

(Fig. 1A). Loreal region sloping. Tympanum not

visible externally (Fig. 1B). Fingers moderately long,

slightly fringed (Fig. 2A) with well developed sub-

articular tubercles. In order of length 3>4>2>1.

Palmar tubercles large and prominent. Super-

numerary palmar tubercles prominent. Hind limbs

long (TL/S-¥V 0.37). Toes long, fringed, with basal

webbing (Fig. 2B). In order of length

4>3>5>2>1. Large oval inner and broad outer

metatarsal tubercles. Subarticular tubercles conical.

Dorsal surface with well defined parotoid,

inguinal and coccygeal glands (Fig- 3), Dorsum

fainuy tubercular. Submandibular gland disrupted.

Ventral surface finely granular.

Male with unilobular, submandibular vocal sac.

In preservative, dorsal surface brown, Short cream,

midvertebral stripe on anterior 1/3 of body. Black,

lyrate markings on scapular region (Fig. 3) and

black patches on other portions of dorsum.

Surfaces of parotoid, inguinal and coccygeal

glands, cream. Inguinal and femoral regions cream.

Ventral surface cream with chocolate brown

suffusions. Submandibular region dark brown,

Fig, 2. A palmar view of hand, B plantar view of foot

of Uperaleia glandulosa sp. nov, (SAM R27081).

Colour in life. Main & Storr (1966) described the

colour of WAM R22921 as “Dorsal ground colour

olive-brown with black spots and blotches.

Anteriorly an indistinct vertebral series of orange

spots. Dorso-laterally a broken orange stripe from

behind orbit nearly to groin, A small orange spot

on each side of sacrum. A large orange red blotch

in groin and on outer surface of thigh. Upper

surface of limbs as on dorsum, Ventrally whitish

flecked with grey. Lower surface of limbs flesh-

coloured.”

This description conforms with our observations.

Dimensions (in mm): Snout-vent length 20.2; tibia

length 7.4; eye diameter 2.8; eye-naris distance 1.9;

internarial span 1.4,

Variation: The are seven paratypes, six adult o¢

and one adult 2. WAM R89490-2, AM R114573,

NEW UPEROLEIA FROM THE PILBARA 105

Fig. 3. Uperoleia glandulosa sp. nov. in life.

collected with the holotype; SAM R27081, SAM

R27082 (cleared and stained), 3.2 km NE

Wittenoom turnoff on Port Hedland-Broome Rd,

10.1.1983, M. Mahony, J. D. Roberts, WAM R22921,

Mundabullangana (5 km E of Hstd), 19.ii.1961,

G. M. Storr.

Males range 19.9-23.4 mm and the female is

24.6 mm S-V. All paratypes are squat with long legs

(TL/S-V 0.38-0.41) in the males: 0.34 for the

female). E-N/IN ranges 1.13-1.50. All specimens

have well developed cream glands, and basal to

slightly more than basal toe webbing. Skin texture

resembles that of the holotype. The ground colour

of the dorsum of the female is grey rather than

brown whilst the darker colour markings of all the

paratypes are brown whereas the holotype is black.

One paratype has a truncated snout when viewed

from above, as in the holotype, whereas the others

have more rounded snouts (Fig. 1B). In some of the

paratypes the midvertebral stripe is not as

conspicuous as in the holotype. Palmar tubercles

are not always as prominent as figured.

Mating call structure: The advertisement call of this

species is a sharp click: a single note formed by two

or three pulses. The pulses have a rapid rise time,

one or two oscillations, and an irregular decay to

about 1/4 of the peak amplitude before the next

pulse is initiated. Pulse periods are short (about

6 ms) and equal to pulse durations. Details of call

structure are given in Table 1. An oscillogram is

presented in Fig. 4.

Breeding biology: Most males were observed calling

around a roadside drainage channel. Males were

calling at the base of sedge clumps close to the

Tasie 1. Details of call structure of two Uperoleia glandulosa sp. nov. Values are means with standard deviation

and range in brackets,

‘Temperature Pulse Pulse Pulses Call Call Calls Frequency

Tec Number Duration sec! Duration period min~! (Hz)

(ms) (ms) (s)

25 2.3 63 160 14.2 0.74 81.1 2875

(Air dry) = (0.4, 2.0-2.6) (0.8, 5.7-6.8) (0.6, 13.4-14.6) (0.08, .69-.8) (35, 2850-2900)

106 M. DAVIES, Mo. MAHONY & J. D. ROBERTS

water's edge or in a similar position actually in the

water, Males with a similar call were also heard in

the Petermarer Creek and around a flooded claypan

about 3 km southwest of the Turner River on the

Northwest Coastal Highway, in the Turner River but

hig. 4, Oscillographs of advertisement call oF Uperoleia

vlandilosa sp. nev.

not on the Yule River. Males were calling on the

third night after two days and two nights of heavy

rains (114 mm at Port Hedland).

Uperoleia russelli was calling at the same sites,

Osteology: Skull poorly ossified, sloping

anteroventrally, Small portion of sphenethmoid

ossified posteriorly to medial extremities of

palatines with ossified portion extending about 1/3

of the length of the orbit in ventral view. Medial

ossification of sphenethmoid absent dorsally and

ventrally. Prootic not fused with exoccipital.

Exoccipital not ossified either dorso- or ventro

medially. Crista parotica short, stocky, not overlain

by otic ramus of squamosal. Carotid canal groove

not developed on frontoparietals. Frontopanctal

fontanelle extensively exposed, overlapped laterally

by frontoparietal elements extending narrowly for

9/10 length of orbit, Anterior margin of fontanelle

formed by sphenethmoid and medial cartilaginous

elements, at level of anterior rantus of plerygoid,

Posterior margin formed by articulating postero-

medial extremities of frontoparietals (Fig, 5A),

Nasals moderately ossified, approximately

(riangular with maxillary processes directed more

ventrally than laterally; widely separated medially,

Maxillary processes not in bony contact with well

developed preorbital processes of very shallow pars

facialis of maxillary.

Palatines robust, reduced laterally, abutting and

just overlying anterior extremities of sphenethmoid

ventrally. Parasphenoid robust, cultriform process

elongate, extending anteriorly to ossified portion

of sphenethmoid in ventral view (Pig, 5B), Alary

processes expanded laterally, at right angles to

Kia. 5. A dorsal, B ventral view of skull of Upereleta glandulosa sp. nov, (SAM R270K2}. Seale bar

5 mm.

NEW UPLROLELA PROM THE PILBARA 107

cultriform process extending almost (to level of

anterior extremities of medial rami of pterygoids,

Medial ramus of pterygoid moderately long,

acuminate, not in bony contact with prootic region.

Posterior ramus moderately Jong and broad.

Anterior ramus in long contact with moderately

developed pterygoid process of palatal shelf of

maxillary. Cartilaginous quadrate present between

base of squamosal and quadratojugal. Quadrato-

jugal robust, in Firm contact with maxillary,

Squamosal shaft robust; tiny zygomatic ramus,

elongale unexpanded otic ramus lying alonyside

lateral extremities of crista parotica.

Maxillary and premaxillary edentate. Palatal shelf

moderately deep with well developed palatine

processes not abulting medially, and moderately

developed pterygoid processes, Alary processes of

premaxillaries moderately broad, bifureate and

perpendicular. Vomers absent. Columella bony,

Pectoral girdle arciferal and robust. Omosternum

and xiphisternum present, Sternum cartilaginous,

Clavicles slender, curved, poorly separated meclially.

Seapula bicapilate; about same length as clavicles.

Suprascapula about 2 ossified,

Eight non-imbricate presacral vertebrae. Sacral

diapophyses poorly expanded, Relative widths of

Iransverse processes Il>sacrum>1V>ll>V=

VI>VIT=VILL.

Nig. 6.

slandulosa sp, vay. (SAM R27082),

Lateral view of pelvic girdle of Cfpercletu

4 \\

ee oe + eer

aN Ob ak a

Bicondylar, sacrococeygeal articulation, Urostyle

with crest extending 34 its length. Ilia extend

anleriorly to sacrum. Hial erest absent. Large

papillate, dorsal protuberance on small dorsal

proniinence (Fig, 6), Humerus with strongly deve-

loped anteroproximal crest, Phalangeal formula of

hand 2,233. Distal tips of terminal phalanges

knobbed. Small bony prepollex and palmar sesa-

moid, Phalangeal formula af foot 2,2,3,4.3. Well

developed bony prehallux.

Karyolype: The karyotype of Uperoleia glandulosa

15 Shows in Fig. 7) the diploid number is 2u = 24.

The chromosomes are arranged in two groups on

the basis of size. Pairs 1 to 6 are large with relative

lengths (R.L.) ranging from 15% to 9%; pairs 7 Lo

12 are small with R.L. ranging from 6% to 2%.

Chromosome pairs 3, 4, 5 and 8 are submetacentric,

with the remaining chromosomes being metacentrie.

A prominent secondary constriction which is silver

stain positive, and thus considered to be the

nucleolar organiser region, occurs subterminally on

the short arm of chromosome 4. A distinctive

secondary constriction mid-way along the short arm

of chromosome & was not strongly silver positive

and is not considered Lo be a nucleolar organiser

region,

Etymology: The specific epithel is from the Latin

glanduia in reference to the prominent parotoid,

inguinal and coceygeal glands in this species.

Comparison with other species. Uperoleia glan-

dulosa is distinguished from many congeners by the

presence of a widely exposed frontoparietal

fontunelle, This feature is shared by C1 arenicola,

U. borealis, U. inundatu, LU. orientalis, U, russelli

and U, talpa. All of these species, except for U.

orientalis for which the data are not known, are

“long call" species whereas U, glandulosa has a very

short call, U! glandulosa has a slronger back pattern

and less extensive webbing than U. orientalis:

From the three “short call” species in Western

Australia, Uf lithomoda, LU minima, and U aspera, U,

hth

xs “om

Vip. 7. Karyotype of Upervlera glandulosa sp. nov. Arrows indicate rhe location of the Hucleolar Oreaniser rewion

of chromosome 4; inset shows (his region With silver sigiting. The location of a secondary constriction on Chromosome

& is not silver positive and is indicated by an arrowhead.

108 M, DAVIES, M, MAHONY & J, D. ROBERTS

glandulosa is distinguished by its greater toe

webbing and greater trontoparietal fontanelle

exposure,

Discussion

It would appear that the hew species that we

describe here has been encountered previously al

Mundabullangana but misidentified as U,

marmorata by Main (1965) and Main & Storr

(1966), These authors also questioned the identity

of some of the material that they assigned to Ui

russelli (as Glauertia russell’), considering, that it

might represent UL. myobergi.

No material is available to confirm the existence

of a third species in the Pilbara, but during the

course of the fieldwork by M.M, and J,D,R,, it was

thought that a third call type was detected, Hence,

although we can now establish that two species are

present in the Pilbara (U. glandilosa and U.

russelli), further field work is required to establish

if a third species is present.

Although the diploid number, 2n = 24, of several

species of Uperoleia has been reported (see

Morescalchi, 1979 for a summary), to date no

karyotype has been figured and chromosome mor-

phology has not been described for any species of

the genus. Karyotypie examination of 10 species of

Uperoleia (Mahony & Robinson in press), revealed

a suiking conservatism of chromosome morpho-

logy and, in particular, location of the nucleolat

organiser region. While this conservation means

that species of Upervleia cannot be distinguished

from one another chromosomally, the consistent

location of the nucleolar organiser region is a

distinctive feature of the karyotypes of Uperoleia

species, enabling them to be distinguished from all

other myobatrachine genera.

Acknowledgments

Field work was supported by an Australian

Biological Resources Study grant to M.M, and

funds from the Zoology Department, University of

W.A. to J.D.R. We thank Steven Donellan for Fig. 3.

Laboratory studies were funded by an Australian

Research Grant Scheme grant to M. J. Tyler and

M.D.

M. J. Tyler critically read the manuscript and js

thanked for helpful discussion.

References

Buoom, 5. BE. & Gooorasrurr, C, (1976) An improved

technique for selective silver staining at nucleolar

organiser regions in humin chromosomes. Hum. Genel,

34, 199-206,

Cocarr, H, G. (1983) Reptiles and Amphibians of

Australia. 3rd Edin. (Reed: Sydney),

Davis, BD, D. & Gorn, V, R. (1947) Clearing and staining

skeletons of small vertebrates. Fie/diana Tech, 4, 1-16,

Gatonr, 8. 1. & Evans, C. S. (984) The use of rate or

period to describe temporally patterned stimuli, Ami,

Behay, 32, 940-941,

Manony, M. J. & RoBinson, E, 8, (1980) Polyploidy in

the Australian leptodactylid frog genus Neobatrachus.

Chrontasania (Berl.) 81, 199-212.

& (in press) Nucleolar organiser region

(NOR) location in karyotypes of Australian ground

frogs (family Myobatrachidae). Genetica

Main, A. R, (1965) Frogs of Southern Western Australia.

(Western Australian Naturalists Club: Perth),

& Storr, G. M. (1966) Rate extensions and

notes on the biology of frogs from the Pilbara region,

Western Australia. West, Aust. Nat. 10, 53-61.

MORESCALCHI, A, (1979) New developments in vertebrate

cytotaxonomy, I. Cytotuxonomy of the amphibians.

Genetica 50, 179-193.

Trure, L. (1979) Leptodactylid frogs of the genus

Telmatobius in Ecuador with the description of a new

species, Copeia 1979(3), 714-733,

Tyior, M. J. Davies, M. & Martin, A. A, (1981a)

Australian frogs of the leptodactylid genus Uperoleia

Gray. Aust. J. Zool. Suppl. Ser. 79, 1-64.

, & (981b) New and rediscovered

species of frogs from the Derby-Broome area, Western

Australia, Ree. West. Aust. Mus. 9(2), 147-172.

SMITH, Lb. A. & Jounstonre, RoE, (1984) “Frogs

of Western Australia” OWA, Muse: Pert)

THE AUSTRALIAN NAUCORIDAE (INSECTA, HEMIPTERA-

HETEROPTERA) WITH DESCRIPTION OF A NEW SPECIES

BY IVOR LANSBURY

Summary

The Naucoridae of Australia are redescribed and keys to genera and species are provided. A new

species, Naucoris subaureus sp. nov. is described from Western Australia. Lectotypes are

designated for Naucoris australicus Stal and N. subopacus Montandon. Distributional and

ecological notes are given for all six species.

THE AUSTRALIAN NAUCORIDAE (INSECTA, HEMIPTERA-HETEROPTERA)

WITH DESCRIPTION OF A

NEW SPECIES

by Ivor LANSbURY*

Summary

LANSBURY, [

new species, Trans. R. Soc. 8.

. (19K5) The Australian Naucoridae (Insecta, Hemiptera-Heteroptera) with description of a

Aust, 109(3), 109-119, 29 November, 1985,

The Naucoridae ol Austealia ave redescribed and keys lo zenera and species are provided, A new species,

Naueoris subaureus sp. nov. is deseribed from Western Australia, Lectotypes are designated for Nawecorts

ausindicns Stand No Subopacus Montandon, Distibutional and ecoloyical notes are eiven for all six species.

Kiy WoRbS: Naucoridae, Naucoris, Aphelocheirus, distribution.

fntroduection

Phe deseribed Australian naucorid fauna of live

species (Nuucoris ausiralicus Stal (1876), N.

congres Sal (1876) and Aphelocheirus australions

Usinwer (1936) from Queensland, N, subopacus

Monlandon (1913) and MN. rhizomatus Polhemus

(M984) from the Northertt Territory) 16 relatively

depuuperate compared with that of New Guinea,

La Rivers (1971) lists 20 endemic species in cight

endemic genera, from thal area.

Naucoris Fabricius is an ‘Old World’ genus with

ed. Mie spceies recorded from the Oriental Region,

Tt has not been recorded from New Guinea or New

Zealand,

The tnelusion of Aphelocheirus Westwood inthe

Naucoridae is debatable. China & Miller (1959) and

Popov (1970) included the genus in the Naucoridae

subfamily Aphelocheirinae. However, Poisson

(1957) considered (he genus merited family stacus

and iis omission by La Rivers (1971) [rom his world

checklist of Naucoridae implies acceptance of this

contention, A recent review by Hoberlandt & Stys

(1979), retained lphelocheirus ina subfamily of the

Naucoridae,

Polhemus (1984) briefly reviewed the Naucorinae

of Australia. In this paper, | redefine the Australian

species of the Naucoridae, including v1, wustralicus,

and describe a new species of Nowcoris tram

Wester Austrtlia,

All ineasurenrents xiven are in mm.

Kev jo Vustralian genera af Naucoricdee

Vertex even? rounded between Mie eyes (Pres |, 7k RD.

fore femur (merissare (Pig. 55) Rostrum short, not

reaching The lore coxae Antennae short, three

stemented and not visible fran above —-.....

sete . Navvoriy babricius 1775

Vertex nitduded herween the eyes (Pig, 62), Pure fernur

HOt jerussate (Pies 63 & 64). Rostrom long, redehine

* Hope Entomolpgical Collections, University Museuin,

Oxtord, Wik.

the mid-coxae, Antennae long, foursegmenied, often

visible from above ~ Aphelucheirus Westwood 833

Naucoris Fabricius

Body variably llattened dorso-ventrally. lore

femur broadly triangular and inerassate, fore tibia

folding tito a shallow sulcus along femoral margin.

Mid and hind legs “cursorial’. Antennae three-

segmented lying beneath eyes on underside of head,

Rostrum three-segmented, two scements visible, not

reaching fore coxae Fore legs separated by a

prominent pilose ridge produced cephalad. Between

mid and hind coxae, small plate-like keel, not readily

visible when viewed from side. Head and pronorum

variously punctate light and dark browns anterior

margin of pronotum more or less straighr, Clavius

and corium unicalonous or bicolored; membrune

hot always clearly differentiated from corium and

emboliar fracture nol always entire, Underside of

abdomen variably pubesvent. Ventral latero-tergites

shining, width variable, always becoming obsole-

scent at apical margin of third sternite, posteriorly

reaching seventh sternite. Sternites divided by fold

or suture either side of mid-line (Fig. 2) Male

genital capsule “boat-stiaped", laterally heavily

selerotised, apically membranous. Parameres

asymmetrical. Male fifth stermue medianly

asymmetrical. Fernale operculium slightly rounded

and distally emarginate. Wilhin Australia, Maucoriy

is Of a larly uniform appearance.

Kev lo the Ausiralian species of Naucoris

! Postener jurgin of pronoun oot produced

candad at humeral angles (Figs 1, 7,48), Venter

ciiher appearing bare or with fine short hairs 2

Posterior margin of preanotum produced candad

al Miuineral aiwles (hig. 41) Al) abdominal

veritrites clothed in fine Holden pubescence

No subaureus sp. nov,

5 sutellum and most of embolium pale yellow

(Fig, by ee ee eee JN. ausrralicus Stal

Setitellim Unicolorous dark reddish brown or

black, emboljum basally variably pale yellow to

10 1. LANSBURY

Figs |-6, Nauwcoris australicus Stal; 1, dorsum and

yellowish brown, arise? ways dark brown to

black (Fig. 7) . ~#h5 ¢ 3

Posterior aneles of connexiva 4-6 sharply

produced (Fig. 10), Shining area of 3rd ventral

latero-tergite basally broader than anterior width

of 4th and broadly infuseated (Fig. 39) .,,...

oo... N rhizomatus Polhemus

Posterior angles of connexiva 4-6 not sharply

produced (Pig. 9). Shining area of 3rd ventral

latero-tergite not basally broader than anterior

width of 4th and mot broadly infuscated

(EIBE ASHABY. Fee h e Lee adic Ae aie 4

Dorsum of head and pronotum rugulose and

rather dull. Shining area of ventral latero-tergites

narrow (Pigs 9 & 38) N. subapacus Montandon

Dorsum of head and pronotum smooth and

usually shining. Shining area of ventral latero.

tergites broad (Figs 35-37) ...N. cangrex Stal

4. (4)

Naucoris australicus Stal

FIGS 1-6, 26, 32-34, 51

Naucoris australicus Stal, 1876, p. 145; Lundblad, 1933,

p. 62; La Rivers, 1971, p. 71; Polhemus, 1984,

pp. 157-158.

Lectotype: Female, Austral boreal, Thorey; | ¢ and

2, venter of Lectotype 9. 3, genital capsule and 4, 5, left and right

parameres; 6, aedaegus of d, Eidsvold, Qld; ms, median sternite; op, operculum; ps, parasternites; vil, ventral

latero-tergites; it- vil sternites.

1 ¢ paralectotypes Australia Orient med. Thorey in

the Riksmuseum, Stockholm, vid.

Distribution: Queensland.

Male §8.0-9.6 long, width 5.3-5.7. Female

(lectotype) 9.5 long, width 5.7.

Calour: Head, pronotum, scutellum and most of

embolim pale yellow, Head shining with a triangular

group of shallow brown punctures. Pronotum

shining, medianly faintly and irregularly

transversely striated; anterior margin dark brown,

groups of brown punctures converging from inner

margin of eye towards the median line posteriorly.

Between these groups of punctures, third group of

larger darker punctures; posterior margin slightly

darker with short irregular rows of shallow

contiguous brown punctures. Clavus, conum, apical

region of embolium and membrane brown with

obsolete narrow, yellowish-brown stripes extending

from between the embolium and claval suture

almost to membrane. Connexiva pale yellowish

brown, distal angles faintly infuscated with irregular

prominent brown spots. Underside and legs pale

yellow.

AUSTRALIAN NAUCORIDAE

bigs 7-10, Naucoris spp. 7) No congrex Stal & dorsum, Mogeill Farin, Qld, § N. swhopacus Montandon § dorsum,

Foge Dam, N14 9, No subopaces Montandon ¥ abdomen, Adelaide River, NT, 10, N. ritizomiatus Polhemus 2

abdomen, Adelaide River, NT.

Structure: Anterior interocular space almast the

same as posterior Pronotal humeral width 2,3-2.4 «

median length, rhe later between 1.4-1.8+ head

length. Pronotal lateral margins. slightly convex.

Seutellum between 15-18% broader than long,

lateral marvins Simuale. Labrum LS-1.7% broader

than Jong. Mesosternum vonspicuous, slightly

produced cephalad, Postero-lateral angles of the

fourth and fifth connexiva forming an approximate +

90° allele, sixth and seventh slightly produced, the

sixth more promigent ut the female (Fig. 1).

Depression oF seventh parasternite conspicuous,

almost reaching outer margin of sternite, Male fifth

sternite (Pig, 26), Shiming lateral margin of third

ventral latero-terite narrow (Pigs 32-34). Female

seventh stermile about 2/3 median length of sixth

(Vig, 2), Operculum (Fig. 31) about 1.3% broader

than long. Male genitalia (Pigs 3-6),

Material examined: Lectotype female, Austral

boreal, Thorey (Stockholm). Gne ef, Queensland,

Bisvold; one ¢ ‘Queensland’ (British Museum,

Natural History),

Polhemus (1984) sives data for a single male from

Qld, 14 miles NW of Aye, 20 m, 14.41.1962, B.S

Ross, E.G. Cavagnaro. This specimen in the J. T.

Polhemus eollecdou.

Stal's 1876 account of Naweoris is confusing

Where it refers to Australian species. He

distinguished N. ausiralicus from N, congrex

because the former had a pale sculellum (“scutello

pallido”). In the original deseription he refers to a

male. The type series received from Stockholm has

a female labelled TYPUS, the other twa speeimens

have dark brown seutella and are referable to N.

congrex. The female labelled TYPUS is hereby

designated Lectotype and is labelled as such as it

is the only specimen which agrees with Stal’s

description. The remaining male and lemiale

labelled “Australia Oriemt and Thorey” are hereby

designated paralectotypes ol N. australicus

although they are in faet N, congren.

The slender data available sugwest thal AN,

australicus is contined to Old.

Naucoris australicus is similar in peneral

appearance lo N, congrex. Ibis easily distinguished

by the pale yellow scutellum and embolium.. Across

the widest part of the body it has three prominent

pale spots, shared by no other Ausuralian nauvorid.

Woodward ef uf (1970) Fig. 26, 72e, figured N.

australicus, not N. congrex as stared.

Naucoris conzrex Stal

FIGS 7, 11-25, 27-28, 35-37 & 50

Naucoris congrex Stal, 1876, p. 145; Lundblad, 1933, p. 62;

La Rivers, 1971, p. 71: Polhemus, 1984, pp. 157-159,

Holotype: Pemile, Moreton Bay in the

Riksmuseum, Stockholm, vid,

2 1, LANSBURY

Figs 11-25, Naucoris congrex Stal male genitalia; 11, 13, 16, 17, 20 & 21, Tasmania, Tomahawk River; 12, 14, 18 &

22, Victoria, Yea River; 15, 19, South Australia, Piccaninie Blue ponds; 23-25, “austrelicus” part of type series

~ congrex: (11-12) genital capsules; 13-16 & 23, left paramere, 16, different aspect; 17-20 & 24, right parameres,

20, different aspect; 21, 22 & 25, aedeagus.

Distribution: Tas., Vic, S. Aust., N.SW, and Qld.

Males 7.7-8.7 long, width 4,8-5.3. Females 8.5-9

long, width 5.4-6,

Colour; Occurs in two forms as follows. Dark form,

head and pronotum yellowish-brown and shining.

Head with a broad band of brown punctures

tapering or converging towards anterior margin.

Pronotum anteriorly, irregularly, transversely striate

with shallow brown punctures; centre of disc with

scattered larger punctures; posterior 1/3 finely

striate with short rows of punctures coalescing into

brown lines; lateral margins smooth, Scutellum

varying reddish-brown and black; inner lateral

margins of clavus pale. Embolium basally pale

yellow, apically verging towards corial colour.

Membrane dark brown to black with many greyish

punctures. Connexiva pale yellow, slightly

infuseated posteriorly with dark brown punctures.

Pro- meso and metasternum mostly black; lateral

margins Or mesosternal ridge yellow. Ventrites black

with silvery hairs. Ventral parasternites dark brown

with crescentic pale yellow indentations across

ventrites. Legs pale yellaw.

Pale form: Head and pronotum pale yellow back-

ground, brown punctures on head reduced in

density and coverage, wholly confined to basal half

of head. Pronotal pattern reduced; brown stripes

on posterior 1/3 almost obsolete. Clavus and

corium brown, membrane black. Sculellum as in

dark form, Prosternum straw-coloured; coxal plates

slighly pigmented. Sternites dark yellowish-brown.

Structure; Anterior interocular space slightly

AUSTRALIAN NAUCORIDAE 13

narrower than postenor space; inner lateral margin

of eyes almost straight, Head width to length ratio:

males 3.1-4.1« yreater than length, average 3.4x.

Females 2.4-4™, average 3.5«. Pronotal humeral

width 2.2-2.5% median length, average 2.4; latter

about twice median length. Pronotal lateral margins

evenly curved. Scutellum between L.S-1.7% broader

than long, lateral margins slightly sinuate, Labrum

between 1,3-1.8»~ broader than long, average 1.5%.

Mesosternum similar to N. awstralicus. Postero-

lateral angles of connexiva forming 90° angles; sixth

connexiva slightly produced in female (Fig. 7).

Depression of seventh parasternite large reaching

inner margin of ventral latero-lergite. Male fifth

sternite (Figs 27 & 28), little variation between Tas.

und Qld forms. Shining lateral margin of third

latero-tersite broad (Figs 35-37), Female fifth and

sixth sternites same length, seventh sternite 2/3

length sixth sterntte. Opereulum basally irregularly

emarginate (Fig. 50). Male genitalia (Figs 11-25),

imm

Slight variation between Tas. and mainland forms.

Material examined: Holotype female, Moreton Bay

(Queensland) Riksmuseum, Stockholm. Tas.: Georgetown,

G. E. Cole; Launceston; Wedge Bay, C, H. Hardy, South

Australian Museum. Pawleena Lagoon, Coal River; Pond

at Karanja; Blackmans Lagoon; Tomahawk River, Cape

Portland Lagoon; Icena Lagoon; Moriarty; Newnham; St

Bernard’s Creek; Flinders Island, Lagoon at Emita; lagoon

near Robinson's Farm; North East coastal lagoon; King

Island, Lake Flanagan; the preceding via Dr P. S. Lake,

and now in the Australian National Insect Collection

(ANIC) Canberra.

Vic.: Yallock Creek near Melbourne; farm Dam,

Hammano road near Neerim (some adults teneral also

immatures); farm dam near Bunyip River, Labertouche

North (immature); Lake Purumbete; Eleocharis swamp

about 8 km from Colac; farm dam about 2 km from

Barwon Downs (immature); billabongs along margins of

Barwon River at Inverleigh; “Sheepwash” lagoon between

Cathkin and west of Molesworth, Yea Rd; backwater of

Yea River at Yea (25-29.iii.1979, I. Lansbury) (ANIC,

Canberra); Noble Park, 6.iv.1918, F. E. Wilson (SAM).

S. Aust.: Adelaide, H. M. Hale; River Torrens, Murray

Bridge, H.M.H.; Bridgewater, H.M.H.; Lucindale, A. M.

Fips 26-40, Naucoris spp, male 5th sternite of 26, N. australicus Sval, Qld; 27, N. congrex Stal, Tomahawk River,

Tas; 28, N. compres Stal, Moggill Farm, Qld; 29, N. subaureus sp, nov. Millstream, Qld; 30, N. rhizomatus Polhemus,

Coomatlie Creek, N.T; 31, N. subopacus Montandon, Fogg Dam, N:T. 3rd ventral laterotergite of 32, N. australicus

Stal, Lectotype fernile; 33, N. australicus Stal, male, Eidsvold, Qld; 34, M. australicus Stal, male, Qld; 36-37, Naucaris

congrex Stal; 35, Holotype female; 36, male, Tomahawk River, Tas; 37, male, Moggill Farm, Qld; 38, NM. subopacus

Montindon, male, Forg Dam, NPs 39, NV. ratzomarits Polhemus female, Adelaide River, N-T.; 40, N. subaureus

sp. oy, Holotype male, Millstream, Old. ‘Terminology p protonum; iii ps 3rd parasternite.

114 |

Lea; Roanka Sta., Blanchetown, in shallow water along

edge of Cumbunga Creek, 12.v,1973, R, (ons (SAM), Mt

Gainbier region, Piccaninie Blue ponds near Glenelg River,

20.iy.1979, 1, Lansbury; “Ewens" ponds near Mt Gambier,

21. iv.1979, LL; Naracoorte region, Hack’s Swamp near

Bool Lagoon, 23 iv.1979, 1.L.; same data, Bool Lagoon,

23,iv 1979, TL Kangaroo Island, river just beyond

Karratla on road lo Kingscote, 9,iv.1979, LP, (ANIC,

Canberra).

Qld; Cunnamulla, H, Hardeastle (SAM); Atherton Table-

land, S km from Mareeba on Molloy-Mareeba road,

22,1979, LL; Brisbane, Moggill Farm dam, 6.iv.1979

(some \mmature) U1, (ANIC, Canberra).

A male from Tasmania, Tomahawk River,

2.x1.1972, B. Knott has a pair of tunnel-like

structures made with detritus and whal seem to be

short naucorid hairs lying along the 2nd/3rd

episternal suture where the middle coxae articulate

within the thoracic cavity. These structures are

almost certainly phoretic in origin; no remains of

any organism were found,

The broad shining lateral margin of the third

ventral latero-tergite im both sexes and the

conspicuously asymmetrical fifth sternite of the

inale are diagnostic.

Naucoris subopacus Mentaaden

FIGS 8, 9, 31, 38, 53, 7, 56-61

Nauvoris subopacus Montandan, 1913, pp. 223-224;

Lundblad, 1933, p. 62

1984, p. 160,

; La Rivers, 1971, p. 71, Polhemus,

LANSRURY

Leclo/ype; Female and three $4 paralectotypes,

Northern Territory, Adelaide River, in the British

Museum (Natural History) vid, Two female

paralectotypes, same locality SAM, vid.

Distribution: W.A,, NT, and Qld.

Males 6,6-6,9 long, width 4-4.2, Females 6.6-7.6

long, width 4,3-4.6.

Colour; Dark form; head dark yellow with brown

punctures, finely rugalose appearing dull, in

artificial light rather more shining. Pronotur

medianly rugulose; most of anterior 1/3 and middle

1/3 covered with brown punctures; anteriorly

divided with pale triangular area; within paler area,

group of shallow brown punctures With many

longitudinal brown stripes posteriorly. Lateral

margins smooth yellowish-brown, Scutellum pitehy,

reddish-brown to black with faint reddish-brown

area medianly, Clavus and corium reddish-brown;

dark specimens from Fogg Dam, N-T. have obsolete

yellowish-brown patches. Embolium apically yellow

merging into corial colour, Membrane blotchy

yellowish-brown, membrane of left hemielytron

smokey-brown. Connexiva yellowish-brown,

posteriorly lightly infuscated. Mesosternum laterally

yellow, otherwise dark brown to black. Meta.

sternum dark reddish-brown. Median sternites

reddish-brown, ventral latero-tergiles sulfused with

—

yer raers

eee ~

\ oF

Vigy 41-49. Nawcoris sabaurens sp. noy, Molutype male, Millstream, Qht4l, dorsum; 42, genital capsule; 43-44, fer

paramere; 45 46, right paramere; 47, apex of Fig. 45; 48-49, aedeapus.

AUSTRALIAN NAUCORIDAF Is

crescenuic yellow indentations. Legs yellowish-

brown,

Pale dorm: Vesd and prouotim pale yellawish-greys

brown spots on head not scattered byt in groups

either side of midline. Punctate area of pronotum

not so clearly difVerendated from background

colour; pattern similar but alrmtost obsalere.

Scutellum paler, Prregular yellowish markings on

vlavus aud corium obsolete, Connexiva grevish-

vellow with faint infuseation along posterior

inareins. Underside paler; mesosternum almost

completely pale yellow, metasternum tinged with

reddish-brown along keel. Pigmentation of dorsum

decidedly fuvitive, fading rather quickly; Dark areas

of liewd and prefiotun fade rapidly Jeaving

sculpluring alld rugdlose areas. Confiexiva lend 16

fade to unilorin erevish-yellow and scatellum lases

its ulmost blach colouring post-mortem.

Structures Anterior interocular distance always less

jhan posterion mflerocular space Inner margin of

eyes Convelzing anteriorly (Fig, 8), Greatest width

of head 4od.c media length; variation in part due

to position of head post-mortem; card-mounted

specimens have head pushed forward in front of

pronurum. Vresh or “wer” specimens tend to have

head “dipping” below level of pronotum. Pronotal

frumeral width between 2.25-2,5% median length.

lalter beine ca, 2» median lenath Pranotun

elevated or “hump-bavked”, lateral margins diverge

id, Somenmes sthonyly rugulose with conspicugus

inregular seulprnng over most of disc. Seutellam

finely punctate; male about 1.7 « broader than long.

female 16-19. Pasterp-lateral angles of connexiva

hroadly exposed torniog approximate 90° anule,

Labrum 14.07. broader tian long. Mesosternal

nidee prominent, clearly produced cephalad. Male

fifth srernite slightly asymmetrical. Stemites five

and six of male with long hairs; remainder of

stermites dark, tomentose. Pitth female median

sternite slightly shorter than sixth, seventh nor as

lonp as filth (Pig. 9), Operculuna (Fig. 53), Shinn

area of third ventral larero-tereite narrow (Pig. 38),

Male genitalia (Pips 36-61),

Lemotype designation” The type series ol Naweors.

subapaeks comprises bwo spouts: NV. sabapens anc

N. rhisematus. Montandon did nor desienate any

specunen as Type, A female labelled North

Australia, Adelaide River, Adeluide River Station,

July 14-18)h, 1891, J, Walker is hereby designated

and labelled Lectotype. Three females with identical

data are designated paraleciotypes. All the

foregoing we in the Brirish Museum Natural

History, A further tetiale paraleccorype in the

South Australian Museunt hus been designated

which was orpinally prt of Montaidan’s syntypic

series, A axl tetnale from North Australis,

Adelaide River, Adelaide River Station, 8-131h

August, 1890, J, J. Walker is labelled and designated

paralectorype of N. subopacus al(houeh it is in fact

N. rhizamatus. All the preceding material was

collected during the Voyage of H.M.S. Penguin

1890-1893 and presented to the British Museum by

the Lords of the Admiralty, 1896,

Mutertal examined: M1. Koalpanyah, GF Hilly 30 miles

gust of Darwin, G.RH., SAM. MeMinns lagoan neal

Darwin, 4-16.,1979, 1, Lansbury; Foge Dam near Darwin,

460.1070 LL billabong near Nourkiiie Rock, Kakadu

Navonal Park, 401979, 11.) Arntiem Highway, Mary

River pauls, 17v1979 EL. ANIC, Canberra. Jabjlokse

area, Magela Creck floodplidin, Buffalo billabong,

73.4,1979 (immature) RK. Tai) same dita, 20.90,1979;

Jahiluka Billaboni, (7.411979 (invature); same dita,

1541979 (immature); Nankeen Billabong, 1941979)

Winmurra Billabong, Mhty.197%, Ry Tair, ANIC Canberra.

Old: Split Rock, 14 km seuth of Laura, 23-26yi.1975,

G, B. Monteith OM, Brishane. -

W.A5 Beverley Spings, 1iy,1969, DD. Criuliani;y WAM,

Perth.

Naucoris subapacus 1s ia small robust species; the

broadly exposed conneaiva and narrow shining third.

ventral larero-tergite distinguish NV. subapadens from.

the rest of the genus within Australia,

Nauvoris rhizomatus Polhemus

FIGS 10, 30, 39 & 52

Naucory snhopaeus Mowandoan, Wk, pp, 223-224

(ptirunt.)

Nanecors rlizaimatus Polhemps, 984, pp. 187-158,

Flalotype: Male, NT, near Darwin, Coonialie

Creek, U.8i.1977, 1 1. Polhemus, im (he ANIC,

Canberra, Paratype male and leniale originally

deposed in Oxford (Polhemus, 1984: 157) is now

in ANIC, Canberra,

Superfictally this naucorid resembles A.

subopacus. The latter has narrow shining latero-

tergites (Fig. 9); those af N, rhizemaras are broad,

distally infuseuted. The lateral margin of third

latero-rergire basally much broader (han apex of

fourth (ig. 39), this featare comman to both sexes.

Seutellum relatively shorter thaw in. subopacis,

always 2 broader diai long: latcral margins more

rounded and apex far less acuminate than in N

subopoens. Nlesosternal ridge more produced

cephalad, Male tilth sternite (Fig, 30). Female

operculum (Pig. 52).

Muferial evarrineds One 2 paraioetatype of NV. sufiopdens,

N, Austcuia, Adelaide River, Adeluide River Staten,

B-13th August, R80, 1. J Walker SAM,

Sal; Jabiluka pegion, Mayela floodplam, Bultalo

Billabong, 227.1979, R. Tail, 0 ingmatinc, connessval

markings prominent; same data, 2011974, | ot liken “ith

ft ON saidraparews,

Polhemis (984) describes the habiat where MV.

rhizomatus was lound jn Australia as a deep water

creek, the bugs beitig found amongst the tatled

rools of Pandanus. Neucoris subopocus tends to

be found in shallow weedy habitats, The slender

lo 1. LANSBURY

Imm BD

Figs 50-61. Female operculum of 50, Naueoris cengrex Stal, Holotype; $1, N. ausiralicus Stal, Lectotype:

52, \. rhizamaius Polhemus Adelaide River, N.Ty 53, N. subopacus Montandon, Adclaide River, NT. 54, NV,

subaureus sp. noy. paratype, Millstream, Qld, $5, M. sudaureus sp. nov. fore leg, Holotype male, Millstream, Qld;

N, subopacus Moniandon, male genitalia, 56, 57, 59 & 61 Fogg Dam, NT; 5% & 60, 30 miles east of Darwin,

NT. 56, genital capsule; 57, 58, right paramere; 59, 60, left paramere; 61, acdeavus.

evidence suggests that oceasionally N. subopacus

and N. rhizemutus occur in the same habitat,

Naucoris subaureus sp. nov.

FIGS 29, 40-49, 54 & 55

Holotype: Male and paratype female, W.A.,

Millstream, 23,vii,1958, R. P, McMillan in WAM,

Perth. Two ?% paratypes (one teneral without dorsal

pigmentation), W.A., Drysdale River, 18-21.viii.

1975, (14°39'S, 126°57'E), 1. FB. Common & M, S,

Upton, One ? paratype, W.A., Drysdale River,

3-8yii. 1975, (15°02'S, 126°55'E), LEBC. and

M.S.U, ANIC, Canberra,

Distribution; W.A,

Male 7.0 long, width 4.6. Females 7.8 long, width

4.5-4,9,

Colour; Head yellowish-brown, medianly a

longitudinal stripe of contiguous brown spots,

constricted midway along length. Pronotum

medianly shining, lateral margins dull yellowish-

brown. From anterior margin two broad brown

bands almost reaching posterior 1/3; anteriorly with

two secondary groups of brown punctures directed

towards middle of disc, Pronotum lightly striated

with inverted brown “V" shaped pattern between

secondary groups of brown sculpturing; much of

disc covered with shallow pits; middle of posterior

margin with short brown bar directed cephalad.

Scutellur pale with broad brown band from base

to apex. Embolium basally yellowish-brown apically

dark brown outlined in pale yellow. Clayus and

corium dark brown with confused pattern of

yellowish-brown lines, Membrane dark brown with

small yellowish nodules. Connexiva anteriorly

yellowish, posteriorly dark brown to black,

Underside shining pale yellow other than dark

brown embolium. Ventral latera-tergites basally

dark brown. Legs pale yellow.

Structure: Posterior interocular space greater than

anterior width. Inner lateral eye margins more or

less straight, converging anteriorly. Greatest width

of head 3 median head length and just over half

of pronotum. Pronotal humeral width about 2.5 =

median length, lateral margins slightly convex;

humeral angles directed caudad. Scutellum

punctate, just under 2 broader than long,

Emboliar fracture anteriorly prominent, posteriorly

obsolete. Clavus and corium not differentiated;

membrane continuous with corium and coriaceous

(Fig. 41). Postero-lateral angle of sixth connexiva

slightly produced in female; third-sixth postero-

lateral angles of male connexiva forming an

approximate 90° angle. Labrum about 1.5 broader

than long. Mesosternal ridge elevated posteriorly,

AUSTRALIAN NAUCORIDAE Wy

crest covered with shiming yellow. hairs, Mesasternal

carina vestigial, Sternites covered in thick shining

golden pubescence. Ventral latero-tergifes narrow

(Viv 40), Female sixth sternite 2% longer than fifth

and (28~ lotger than seventh, Opereulum

(Fig. 54), Seventh parasternite with vestizal

depression. Male fifth sternite (Fig. 29), Front leg

(Vig. 55). Male genitalia (igs 42-49), capsule

scleratised, not membranous apically, paramerces

long.

Naucons subaureus is easily reeagnised by the

Uvick golden pubescence covering the median

stemites, produced pronatal angles and striking

paltero on the clavus and vorium,

Comments on distribution end bidlagy uf

Australian Naucoris

Within Australia [here ape two species pairs

N, ausioalicus/congrex and N, subapacus/rhizo-

matus. The fifth species \. subaurens is anomalous.

Nanecoris unsiralicus/congrex; These are super-

fically Sinlar wl size and general coloranan and

in having an “easterly” distribution. There are

insufficient data to comment any further on N.

austraticus. The inmost itortherly record ol N.

vongren is Marecba, Atherton Tablelavid, Qld, Its

oocurrence so lar ori aay be as a result of a

“Chinahe’ anomaly caused by the “high” plateau

wilh a more temperate climate compared with the

tropical climate of the surmounding areas. Mv

coneren is. common tty Tas, atid data sugpest that

at the southern end of the range, 1s normally un-

yolting, Oceasionally a partial second genention

may dverwinter in the immature stages. Data bused

on samples collected 1972-1973 from Tas, (small

lake, Pawleena; Coal River 4 miles parth af

Richmond; small lake, Upper reaches Sorell River

and Blackmans Lagoon) lend to support this hypo-

thesis. Mature adults are present trom

Mareh=Qetober, but by then numbers appear to be

Jow, Patring and oviposition presumably take place

jo August-Sepreniber, by Moyerbet, 3rd instar

nymphs are present, finmalure slages continue Lo

be present umil Vebruary, Teneral adults start

appearing im January or earlier (ne dala are

available Jor December) By March adults are

mature and the immature stages ane nor usually

presen!. Immatures were colleefed in May trom

Blackmians Lagoon possibly representing an

overwintering population af immature NV. carigrex,

In Tas. N comgrex has 4 Jaw index speeres

diversity, for caample 1M. corderex lo common) It

OL competes Orlier waterbugs, OP saine 64 halvilats

sampled MN. caverex Was found in ren, elalir of

Which have tive other species Ob aguahe and sea-

aquatic Hereraprera, Greavese diversity was in

Blackmans Lagoon (nine species total) which was

sampled quite eaterisively. On che inamlaad, NV,

congrex is Tound most commonly in Vie, This

probably js due, in parr, to the greater number af

freshwater habitats in Vig, compared with’S, Aust,

and also reflects lack of collecting in N|SW, Ip Vie,

N. conerex was found in nine out ol 37 habitats;

preatest species diversity was 18, the lowese chree,

ol the latter, V. congrex was rhe comimonest species.

Iynmature stages were collecied fram Mareh—-hine

in Vic and Old,

Nanceris subopucus/rhizomaius: These two

Species are remarkably similar in size, coufiguration

and coloration. Both have a “northerly” distribution

extending from NW, Australia, across. the NT. to

Old near Laura, Mast af the habilats sampled by

the author in 1979 where N. subopacus was found

also had Diplonvelins (Belostomatidac) presen.

The belostamatid populations were always much

ereater jiunierieally than those ol No subopacus,

Naucoris sibaureus. Does Hol belo to either

o! the species pairs and is unlike other Naucoris

species fram SE Asia. The Millstream Oasis sourh

of the Great Sandy Desert if noted lor its distinetive

fauna, especially Odonata (Watson 1981), Data on

waler-bugs are scanty, The occurrence oF WN.

subaureus in the Drysdale River area of NW

Australia sugeests that the Sandy Desert does not

impede the dispersal al water-buys between the

Millstream Oasis and the Kimberley,

Aphelocheirus australicus singer

FIGS 62-69

Aphetlocheirus ausiralious Usinger, 1937, pp. 341 342.

4 prepa truss Woodward ef af, 1970, p. 456 (richtioned

only),

Useful diagnestie leatures ares Head, pronotum,

scitelliim and most of clavus more or less shining,

Coridm and embelium dull, Underside appearing

smooth and shining,

Body dorso-ventrally compressed. Head longer

than broad, antennae long, four-segmented, usually

visible from above, Rostrum reaching mid-coxae,

Provotuin transverse, lateral margins exphinate;

median Igneth clearly shorter than median fead

tenelh, postenor marvin roundly emarginate in

front Gf seutellum, Embolium basally broad,

tapering apically along margin of coriunm,

Membrane distinel from corium. Connesivum

broadly exposed, (Usinger’s fiyire shows first visible

coniextval sepment infuscated_) Speeuneny fram N.

Qld have this sepmvnt pule yellow, nor broadly

infuseured, as are remainder (Fig. 62). Fernara of

all Jegs dorso-ventrally flattened (Pigs 63, 64), Pare

and mid legs alike Male genitalia (Figs 65-69);

within capsule, pair of processes attached to lateral

plates, distally pedeesses heavily sclerotised with

dark brown spines apically (Pig, 6) These

8 1. LANSBURY

\ ke

SOY vi TTI

ea a

‘i.

TET ly

Imm

| aan

\ << 67 Imm,

Figs 62-69, Aphelocheirus australicus Usinger, male, Cape York, Lockerbie, Qld. 62, dorsum; 63, fore leg; 64, mid

leg: 65, genital capsule; 66, detail of spinose processes on lateral plates; 67, aedeagus; 68, 69, left and right parameres.

structures not found in Naucoris, Aedeagus more

robust (Fig. 67). Parameres elongate, symmetrical

with long spines distally (Figs 68, 69).

Hoberlandt & Stys (1979) comment on the “appa-

rent” venation of the membrane of some Apehlo-

cheirines. The female from “Captain Billy Creek”

appears to have vestigial venation. By examination

under a strong light, it is possible to detect slight

folds in the texture of the membrane, the folds

forming two irregular cells with unconnected

brachial pattern. A male from Lockerbie does not

have a cellular pattern but more a confused anasto-

mosing system, Originally described from Cairns,

Qld the holotype female is said to be in the

California Academy of Sciences,

Material examined: (all macropterous), Qld, 142°45'E,

11°40'S, Divicing Range, 15 km west of Captain Billy

Creek, Cape York Peninsula, 5-12.11.1976, G. B. Monteith;

Upper Qld, Lockerbie area, Cape York, 13-27.iv.1973,

G.B.M. QM, Brisbane.

Aphelocheirus is normally thought to be

restricted to well-oxygenated water; the bugs are

found crawling about beneath rocks and stones,

Typical Aphelocheirus habitats are the Boulders at

Babinda near Cairns where the creek flows very

rapidly over and under granitic boulders. The

habitats in Cape York are described by Monteith

as swampy with sluggish streams. All Cape York

material was collected at light (T. EB. Woodward in

litt, 4.iii,1977). Hoberlandt & Stys (1979)

commented on flight in the Aphelocheirinae and

all the known material of their new taxa were taken

at light.

In the Palaearctic region the presence of

Aphelocheirus is taken as a reliable indicator of

water purity. The species A. aestivalis has an

efficient plastron allowing gas exchange to take

place without the bug having to surface, in contrast

to Naucoris. The Cairns habitat tends to suggest

AUSTRALIAN NAUCORIDAT 19

that Australian A phelocheirus has a similar plastron

respiration system, However, the presence of the

species in Cape York occurring in swamp sluggish

creeks need nol be considered unusual as Thorpe

& Crisp (1947) list a wide variety of habitats where

Aphelocheirus aestivalis (F.) has been found in

Europe. Within the Palaearctic region,

Aphelocheirus is dimorphic but it is not known if

it is so in Australia,

Acknowledgments

| wish to thank Dr P.S. Lake, Monash University

and Dr A, Neboiss, National Museum of Victoria

for their unstinted assistance whilst | was in

Victoria. Dr G. Gross, South Australian Museum,

Adelaide who enabled me to visit the Mt Gambier

region; Prof. W. D. Williams of Adelaide University

who arranged the trip to Kangaroo Island, Mr

Graham Griffin of C.S.LR.0., Alice Springs and

Dr Gary Fitt (then in Darwin) for their valuable

assistance in the Northern Territory. Mr C. Pedersen

of Noranda Mining for his extended hospitality at

Koongarra. Mr Walford-Huggins for assistance in

the Molloy region of Queensland, Dr T, E,

Woodward, Dr G. B. Monteith and Dr R. Kitching

for theie great help in southern Queensland, Dr

C.N. Smithers, Australian Museum, Sydney for his

assistance, The work was commenced during the

tenure of a grant from the Leverhulme Trust

(London) and grants from A,B.R.S. and C.S.1.R.0.,

Canberra. Finally thanks to Dr W. Dolling, British

Museum (Natural History) and Dr Lindskog, Riks-

museum, Stockholm for the loan of critital types

and to Sandra Lawson of the Zoology Dept, Univer-

sity of Adelaide for typing the final draft of the

maruseript.

References

China, W, b. & Mthink, N.C. b. (1959) Checklist and

keys to the families and sublamilies of Hemiptera-

Heteroptera, Bull, Be Mus. nal. Hist. Entumeotlouy 8,

Homprtanpr, b. & Spys, B. (1979) Tampocoris asiaticus

Gen, and sp. t.—A new uphelocheirine from Vietham

and further studies on Naucoridae (Heteroptera), Sp.

nur Mus. Prage 33(B), 1-20.

La Rivers, [, (971) Studies of Naucoridae (Hemiptera),

Bial. Soc. Nevada Memoire UW, 1-120.

Lusparan, O, (1933). Zur Kenntnis der aquitlen und

semi-uquatilen Heniipteren you Sumatra, Juva und Bali.

Arch. Hydrobiol. Suppl. 12. (Tropische Binnengewasser

iv), 1-489,

MomTanbon, AL 1, (1913) Hémipteres aquatiques. Notes

et descriptions de deux espéces nouvelles. Aull. Sect.

sci, Acad, rowm, 1, 219-224,

Porson, KR, (1957) Faune de France fase. 6). Hetéroptéres

Aquatiques (Paul Lechevalier; Paris).

Pounrwus, J.T. (984) A review of the Naucorinae of

Australia (Heteroptera: Nauecoridae), .. dust ent See.

23, 157-160,

Poroy, Y. A, (1970) Notes on the classification of the

Recent Naucoridae (Heteroptera, Nepomorpha). Bull.

Acad, pol, Sci, Ser Sei Ch V1, 16, 93-98,

Sta..-C. (1876) Enumeratio Hemipterorum, A. svenska

Vetensklkad. Hand. 144), 1-162.

Trhorer, WH, & Crisp, D. J. (1947). Studies on plastron

respiration, 1, The biology of .phelocheirus (Hemip-

tera, Aphelocheiridae) and the mechanism of plastron

retention. . exp. Biol. 24, 227-269,

Usinuer, R. Lb. (1937) A new species of Aphelocheirus

from Australia (Hemiptera, Naucoridae). Ausr. Zeal.

SB, 341-342,

Wa4rson, J A, L, (1981) Odonata (dragontlies and

damselflies). /a A. Keast (Ed.) Ecological Biogeography

of Australia; Vol, 2, 141-1164. (W. Junk: The Hague).

Woonwarp, T. B,, Evans, JW, & Eastop, Vo F (1970)

Hemiptera. Jn CSIRO “The Insects of Australia”

(Metbourne University Press: Victoria),

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 109. PART 4

COASTAL SEDIMENTARY FACIES AND FORAMINIFERAL BIOFACIES

OF THE ST KILDA FORMATION AT PORT GAWLER, SOUTH

AUSTRALIA

BY J. H. CANN & V. A. GOSTIN

Summary

The St Kilda Formation, within Gulf St Vincent and surrounding coast lands is redefined to include

all Holocene sediments deposited under the influence of marine processes. Upper surfaces of the

formation are not confined, but rather may be surfaces of active sedimentation.

COASTAL SEDIMENTARY FACIES AND FORAMINIFERAL BIOFACIES

OF THE ST KILDA FORMATION AT PORT GAWLER, SOUTH AUSTRALIA

by JH. Cann* & Vo AL GOSTIN|

Summary

Cann, J. & Gostin, V. A, (1985) Coastal sedimuntary facies anc foraminiferal biofacies of the St Kilda

Formation at Port Gawler, South Australia. Trans. R. Soe, 8. Aust. 104904), 121-142, 29 November, 1985.

The St Kilda Formation, within Gull St Vuicent ane surpoundine coastal fads ts redefined to include

all Holocene sediments deposited under the influenes of marine processes, Upper surfaces of the formation

are not coulined, but rather inay be surlaees ef wcuve sedimentanion.

Port Gawler is an area where marginal marine sedimenuis are accumulating under (he balling, frapping

and binding actions of scuzrasses, cyanobacterial mals, muilwroves, samphires and sallhush. These plant

COMMINTTNeS Geeur i essentially discrete vores, successively adjacent and panulel to the lidal shore lines.

Their influence causes progradarion and aggradation of bioclastic curbonate-quariz sands and muds. Bivalyes,

gastropods and foraminifera are abundant aod contribute significantly fo the carbonate conmiponent of

sediment. Distribution of molluses is closely reluled to plant conununities, sediment type and period of

dal inundation, Thus a series of laterally adacent sedimentary favws are recognised 10 be associated wilh

the following sub-envirorments: subtidal and inlerlulal seagrass (meadows; mangrove woodland; tidal

distributaries; cyanobacterial mars of rhe weer sand Mat; high tide beavh; supratidal lagoons or sabkhas,

dunes and storm ridges; the estuary and wssecited backwaters of rhe Gawler River

Foraminiferal piofacies, based on selected sieved fractions of dense-liquid Motation conventrates are

established for (he various sedimentary onvirouments. tn (wo vibrooores, down-hole assemblages of

foraminifera are similar to those of modern Port Gawler enwronments. Palucoenyironmental interpretation

of the cores, based on bork lithelowy and assemblies ol toraminitera, shows plat the earliest sediments

of the Holocene transgression were evsentially samphire muds and shelly storm debris. Later sedimentation,

after sea level stabilised, aceurred through processes Of progradation and iggradation, inextricably related

to communities Of seugrissés, Mangroves, cyanobacterial mats and samphires,

Key Worbs: Port Gawler, Gulf St Vineent, Hloloceng, St Kilda Formation, loraminilera, molluses,

coastal sediments,

Introduction

The eastern coast of norjhern Gulf St Vincent

is normally subject to a low everey wave regime.

This fact, together with northward lonygshore driv,

ensures that itis an area of active sedimenrarion.

Low topoeraphic relief of the coastal areas, and

extensively developed sand and mud ITals, result in

broad intertidal areas. High tides associaned with

storm Surges cuHuse inundauian of ooriutally

supratidal environments.

From seaward of Ihe low lide shore line tr ite

supratidal areas there is a pronounced zonarion at

plant communities, These ronge from subridal and

interlidal seayrass meadows, through mangrove

woodlands aud cyanobavtenal macs, to supraddel

samphires and saltbush. In various ways the plants

baffle, trap and bind sediment! causing queradauon

and progradation of the sand Mats, Molluses and

foraminifera are abundant and their shells ane fests

contribute significantly to the aecumulatine

sediment. Also, species of these fauna are

*Schoal of Pure and Applied Sciences, SAC ALL,

Smiths Rad, Salisbury Bast, S. Aust, 3109,

} Department of Geology, Linwersity of Adelaide, Bex

498.0, PLO., Adelaide, 5. Auae. S001

distinetively distributed through the zoned plant

communities and there are several consistent

plant/ammal assacianons.

This paper documents relationships belween

plant comoiuinities and sedimentary processes at

Port Gawler, northern Gulf St Vincent. A method

of foraminiferal analysis, involving dense-lquid

floration concentrates, and selected sieved fractions,

is deseribed und used to establish foraminiferal

biofames for the various sedimentary environments

These data are used in palaeoenvironmental analysis

of twa cores of Holovene sediment.

For Por Ciawler and surrounding coastal areas,

if is proposed that all Holocene sediments,

deposited under inarine influenee, rightly belong

to the St Kilda Formation. This usage includes those

sediments, subtidal, intertidal and supratidal,

forming al present.

The St Kilda Formation

{uo Sauth Austratia considerable confusion has

prevailed in the interpretation of the St Kilda

Farmiation since Firman (1966) first proposed this

and other Holocene stratigraphic units.

On the one hand, Firman (1966) defined (he

formation as “varjous lithologies deliniied for

122 | H CANN & Vo AL GISTIN

mapping purposes by the law lying swampy tract

which extends a ule or more inlane near St Kilda

and elsewhere along the modern coast." That is, te

included the madern supratical samphire sediments

us a facies of the St Kilda Murmaion. This

interpretation was followed by Thompson (1969)

who referred “light-grey shelly strasided beach live

deposits and shelly silts and sand overlain in places

by modern intertidal aud swanip depasits” to the

Holocene St Kilda Formation, and showed these

sediments to be distributed along the eastern coast

of Gulf St Viligent, extending inland aboul 5 km-

On the other hand, Firman’s definition relared

sediments of the St Kilda Formation uy che “raised

sca-bed” of Tate (1879, p. 69) and to the marine

sands and clays of the Osborne fiigh sea level

(Aitchison, Sprige & Cochrane 1954), Daily ef af

(1976) elaboraled on this interpretation, clearly

relating the formatiou ro “a high stand of the

Flandrian sea when shelly sand . , , was deposited,”

Their evidence for 4 Supposed higher Holocene sea

level, about 1.5 metres above modern sua level,

consisted largely of stranded beach rides, and they

cite dates of 3800 + SOO and 1120. 75

radiocarbon years BP. respectively lor (he base ane

top of the St Kilda Formation, By implication,

marginal marine sedimeucs younger than

1120 + 75 radiocarbon years BP. were therelore

excluded from the St Kilda Pormation, This

interpretation was recently followed by Tosetl (1983)

in the Draft Management Plan for Lhe Port Giawler

Conservation Park. A figure illustrating bore hole

stratigraphy, after Lindsay (1945). refers to 1.5

metres Of modern sediment (black sandy clay and

mud with plane fibres) overlyiug 4.6 metres of Se

Kilda Formation (sandy clay with stiell fragments),

Thus aribiguity persists in ihe wee of the

formation name, One interpretation includes the

modern marginal marie sediments togerher with

those of earlier Holocene age, The other, based on

the premise of a Higher than present stand of jhe

Holocene sea, excludes thrse sediments younger

than 1120 + 75 radiocarbon years BLP.

Belperio ef a/, (1983) reviewed evidetice relating

to Holocene sea levely in South Ausiralin and

concluded that there was lil (le evidence to support

the notion of higher Halucene sew levels in the

Adelaide region. They empliasised phe Un portance

of processes of coastal propradanion ane inerticlsl

and supratidal deposition, and cvtsiderad shel!

ridges ti rate poorly as evidenee lor higher sem

levels, In this paper we follow Belperyo eh ah (E483),

‘Lindsay, of M. (1965) Sttarigraphy und

inicropalaeantolagy of three deep Gores, hundred af Port

Gawler S&S Aus Depr Milles Repl Book ASvS1

(Unpublished),

Consequently we see ne reason (o invoke any reeen|

lowering of sea fevel and we are unable to

siratieraptically differentiare the iasine Holocene

sediments. Within Gulf St Yineent and surrounding

coastal lands We Uerefore recognise the St Kilda

Formation to include all Holocene sediments

depusiled jinder the influence of marine provesses,

including estuatine deposits, coastal dunes .and

storm ridges, As such, the St Kilda Formation is

a readily (nappable unit, consisting internally of

yarous sedimentary facies, deposited under a

variety of marine processes. Sediments of the

Lipson Formation and Semaphore Sand (Firman

1966) muy thus be interpreted as facies of the St

Kilda Formation,

The base of the Sc Kilda Vormation is well

defined. Vhe unit distonformably overlies

Pleistocene sediments of the marine Glanyille

Formation (Flymah 1966; Cann (978) and the

alluvial Pooraka Formarion (Firman 1966), Earliest

depositor ja the Adelaide region has been dated

alo 44) | 90 radiocarbon years B.P; (Belperio e/

al, (983). In our preset usage, upper surfaces of

the St Kilda Formation are wet contined, bur rather

may be surfaces of active sedimentation, The

Germein Bay Formation of Spencer Gulf is

equivalent! (Hails ef ah 1984).

The study ares

‘The study ares ts sittiated about 12 kin northwest

of St Kilda, the type location for the St Kilda

Formation (Fig. (). The area ts almost fopographi-

cally Mat wilh samphire swampland to the east,

passing threugh mangrove woodland and open

sundflats to scagrass Meadows wesiwards.

liiMedialely eastwards are salt comcentration

evaporation ponds, che most northerly of a syste

ledding 10 salt extraction at Dry Creck. The area

inclides the estuary of the Gawler River, site of the

now disused Port Gawler {Lisbon Whart, Fig. 2),

Gawler River has its source in the Para Faull

Block, appyOxiately 40 km east of Port Gawler

{os an Intermittent stivam, penerally confined to

a varpow, Meander channel as it traverses the

northern Adetaide Plain. At times of heavy rainfall,

fine graiued wlastle sediments are transported inte

the estuary, from where they are redistributed

threnughouwe the sttidy afea by coastal marine

processes,

Phe rezion of South Arstmlia which fnetudes

Port Gawler expenences a pronounced Mediter-

Huean type of elite Susumers are mostly hot and

ity With Miaxumumn ternperatunes greater than 40°C

Hu UuecoUtOn. Winters are generally mild, night

tempura ures rarely felling to 0'C” Average annual

rainfall of 420 pay al Port Gawler occurs mostly

ST KILDA FORMATION 123

Gowler mf —

ihn ©

, .

PURT GAWLER

STUY oFEb

f * Port adeinney”

] i kien Mateotingl

| =a 1b

Praraki Sunnie SOtTH H

| AUN TRAUS |

Ba is Older rocks moely Joeudeun i

ay wrtiony semen

4 |

0 5 iu WEST

Hig. L Losation map showing surface distribulion of (he

Halocene St Kilda and Pooruka Formations, os used

in (hits paper and other simplified geology.

Pig. 2. Ruins of Lishon Whit, Port Gawler, This facility

with Historivally import for the expart of grain and

wool Wile years 1839 ta 1914. ¢Toteff er ad. 1944),

during, the cooler months, though oceasional

summer (hundersronms produce heavy falls of rain,

Fivaporation bas been estimated at 1800 mm

aumually (Towel! LIBS)

Nhe areas influeneed hy normal tidal changes of

sew level ut Port Gawler are extensive; The gradient!

seawards from the samphite salt marsl is almost

{tiperceplible and maximum astronomical tidal

variitins are about 2.5 m. Armaximum high tide,

sea water covers the sand flats and floor of the

mungrove woodland, and flows through tidal

distributaries to the samphire marshland beyond,

Al minimum low lide, (he sea relreats to the extreme

seaward margin of the sand [lat.

Superimposed on normal tidal (luctuations are

the longer term effects of surges with periods of

i to 20 days, and thought lo relate to the passage

of weather systems (Provis & Radok 1979). Kelly

(1984) has reported storm surge sea level

Muetuations of 0.5 m along the southern Australian

coast, When a storm surge coincides with a spring,

high lide, unusually high tides result. Abnormally

large volumes of water move across the sand flats

and through the mangrove tidal creeks, flooding the

samphire marshland and adjacent (normally)

Supralidal areas, When these conditions are further

accompanied by local winter storm activity, with

strong south westerly winds, sea water is driven ever

more deeply jito the coastal environments, The

combined eflects of tidal currents and storm waves

in these circumstances can cause cousiderable

erosion und redistribution of sediment,

The wave regime of Gulf St Vincent has been

discussed briefly by Bye (1976) and, more

comprehensively and with particular reference to

the Adelaide beaches, by Wynne et al, (1984), The

configuration of the Gulf is such that, along the

eastern coasl, prevailing south westerly winds

generale waves of maximum fetch south of

Adelaide, Adelaide beaches are subject to a lower

wave energy regime which generates significant

alongshore (ransport of sand northwards towards

the sludy area, Except under storm conditions, wave

energy from Port Gawler to the top af the Gulf is

very low.

Vevelation

The role of yegetalion in determining the nature

and cisiribution of sediments at Pom Gawler is

fundamental and is discussed in greater detail below.

In general the Various cyanobacteria, algae,

seagrasses, Mangroves and other plants pertorn) two

significant functions, Firstly, they are the autotroplis

in food chains leading 160 organisms which secrete

shells or tests of calcium carbonate. These shells

and tests, cither intact or comminuted, ultimately

become part of the sediment. Secondly, in a vuricty

of ways, they actually cause deposition of sediment,

trapping grains of calcium carbonate, quarks and

other mineral matter.

Posidonta australis, the broad leal species, is a

seagrass commonly known as “tape-weed”, It is

common in Gulf St Vincent, living from low ude

level to depths of about 10 m. Posidertia hus tough,

roughly textured leaves that host a variety of

124 JH. CANN & V4, GOSTIN

epiphytes (Womersley 1956). The fat ventral surface:

of the foraninlfer Nahecvlaria lucifuga, abundant

at Port Guwler, often shows surtae-like imprints of

these leaves. Womersley & Thomas (1976)

bonsidergd that few animals feed directly on the

Posidenia leaves. The rhizome/root system and leaf

sheaths of this plait are dense, fibrous and resistarn

to organic decomposition, Pasidenia uuvrilis leaves

can survive only brief periods of emergence at low

uides. At Port Gawler it grows in (he owlermosl areas

of the sand hat ist the lower intertidal zone, and

seawards,

Zostera ertuellerd is another scagrass, of narrower

leaf, commonly referred to as “eel grass". Like

Pasidoriia, this plant Has a rhizome/rout system and

hosts epiphyte algae and animals (Womersiey &

Thomas 1976), At Port Gawler, Zostera, lovether

with species of Heterazosrern, Lepilaend and

Ruppia (as described by Ruhertsan (84), forms

extensive, dense, seagrass meadows seaward of the

bare sand flat, and seaward ol, and adjwoent lo the

inangrove woodland. These seagrasses oan survive

longer periods of emergence at low tide

Avicennia australis var resinifera is the ouly

species of mangrove Found m South Austra and

neeurs ac Por Gawler in bork marure woodland and

ws younger colonising plants (Butler ev al. LT;

Harbison (98); Burton 19ST; L982; Talbar (982;

‘Yotcff 1983), Tidal disthibutaries form a dendritic

patceru through the mature woodland, Sea witer

flushes accumulated salt from around the mangrove

roots (Futler es af 1977) and distributes algal,

sengrass and other organic flotsam. Much of this

organic debris probably provides muirient for the

mangroves and is a source of food for a variety of

gastropods, Mangroves ure confined in their

distribution seuwands by the need tov their vertically

promruding prevmatuphores to be periodically

exposed to rhe air, aod landwards by the need for

repular Mushing of accumulated salt from the roots

by sea water at high ride (Chapman 1975). Juvenile

mangroves are colonising the Zasfera meadow

seawards Of the bare sand Flat at Port Gawler.

Durlug the past decadle we have observed their

establishmen| progressively southwards over several

hundred metres from the older trees

Mats of cyanohacteria (= blue-green alpac)

colonise both the floor of the mangrove woudiind

and the mid-tide bare sand lar They are dull preen

in colour tind slippery. Mats wre constructed art the

sediment/water interface by one or more species of

cyanobacteria and may also contain & Viriery of true

bacteria and other algae. Cvanebacteral (mats are

among the measl productive of aquatic

pholosynthelip systems (Bawld 1981), AC Hort

Gawler the mars are apparently erazed by a variety

of gastropods. Their distibuctor onthe open sand

ut appears delimined by grazing gastropods

seawards, and by the infrequency and short duradon

of high tds) iundation of the sediments closest

to the beach.

Halosarciu helocnernoldes and Sarcecornia

guingueflora are dominant plants of the saniphire

salt marsh community. These are low-growing,

Ueshy plants which have considerable salt tolerance

and eccupy the zone inimediately landwards of (he

mangroves,

Sedimentation at Port (cawler

1. Posidonia-Pinna Facres/Zone

M low Lide iL 1s possible to observe the seaward

marein of the sand flai, Pesidenia australis

vrows in patches 20-30 oe in area. Pinna

biculer(. P dolabrata and Subitopinna virgate)

(Butler & Brewster 1979), known locally us

“razor fish”, is conspicuously abundant, growing

in among the seaurass. The sediment is coarse,

poorly sorted, shelly sand. Carbonate content

is greater than 90%, much coming from lime

secreting organisms hosted by the Pesidania,

Voraminifera are particwarly plentiful among

the smaller grains; among the larger shelly

organisms, & bicolor may grow to a length of

20 cm ina single year (Butler & Brewster 1979),

Patches of Posidania on the outermost areas

can be observed im various stages of burial by

the mobile sand (ig. JA), ‘The baffling action

of the seagrass apparently traps the sediment,

Studics in the Bahames by Scoftin (1970) and

Neuman ef al (1970) show that seagrasses can

reduce water velocity from 30 em see~!

(sufficient to anspert loose sand grains alone

the bare sca floor) down to wera al the

sediment/water interface As the sediment

accumulates, Pasidania grows upwards, but is

constrained by its inability to survive low tide

emergence and jt ullimately dies. Large areas of

the remains of recently dead seagrass together

with numerous emply Pinna shells, valves gaping

and in life position, may be observed

immediately lindwards of the present day low

water shore line (Fig 3B),

Carbonate sand is therefore accumulating uf

to jow tide level, causing the outcr margin af the

sand flat to prograde seawards, In some areas,

after death of Posidomid, the uppernast

sediment! remains bare of vexetation and

consequently subject to tidal and storm. waye

transport. Such surfaces become scoured,

channelled and rippled as the shelly sand is

redistributed accordine to enerey ecouwdilons.

Mostly, however, the sediment mass remaitis

essentially coherent, reinforced by seagrass Ohi,

ST KILDA FORMATION 125

Fig, 3A, Patch of partly buried, living Posidonia, adjacent to remains of recently dead Posidania; outermost sandfat,

B. Dead Posidonia remains with dead Pinna, valves gaping, in life position; outermost sandflat, C. Remains of

agglutinated worm lubes, approximately 2 cm diameter. These tubes are constructed of carbonate sand grains and

orientated vertically in the uppermost 0.3-0,5 m of sediment, Scouring by waves or tidal currents cause them to

fragment, Outermost sandflat. D. Dense, fibrous remains of dead seagrass (Zostera) alter death and loss of foliage;

section exposed in tidal channel; wrist watch for scale is 22 em long. E. Recently dead Katelysia (x24), valves attached,

convex upwards; bare surface adjacent to Zostera. F. Zostera in the seagrass meadow.

J. HL CANN & VA, GOSTIN

hy large vertically onented Pitta shells, and by

Numercis Worn Mibes of agelutinated stiell geil,

Algal prowtlr on (he substrate helps to minimise

scouring by tidaleurendrs aud storm waves. We

have Guserved considerable provracation of the

sind Mat during Me past decade,

. Zostera-Katelysia Focies Zone

In the mid tidal area, the sand flat is colonized

by n scagntss meadow, consisting of sevens!

specs, in which Zostere vel? predominates

(Mg. 3) The dominant Myalve mollusc ts

Kutelysia sp, (Fig, 2E). &. scataring has been

recorded from Port Crawler hy Toceff (1983), bol

thrve species of Aotelvyar may be present

(hudbriek 1984). Disarticulated valves of

Aatelysia sp. have been widely redistributed

throughout (he study area,

Like Pasidenia, Zostera balfles and traps

sediment. ‘lhe miel tidal area is less inflvenced

by Wave action and tidal currents, $0 grain shee

16 currespondingly smaller and there is a

siynificant mud component, Sediment is rich in

uranic arte, supporting bacterial reduction

of sulphate ions from sea water, Below the

sediment/ water iterface it is black and as a

strong sulphide odour (Pig, 44), Burrowing by

sinall crabs tends to extend to the underlying,

arain supported sand of the Pasidonia-Pinka

lanes. Bioturbation tends (0, 1 part, oxidise and

homogenise the sediment,

Ageradation of sediment in the Zasters

nicadow coyitinues fo a level determined by the

penod of low tide emergence, Ultimately the

plants on the landward side of the meadow an:

inadequately watered ac high tides and they che.

On the seaward side, the Meadow advances over

Hie Ol) Posidoniae Pinn substrate, ane (he liner

sediment facies thes progrades.

. Mangrove Facies/ Zane

At Port Gawler we have observed the

colunising advance ol juvenile mangroves,

Aptoennie australis var resin(fera, across portant

of the Zostera meadaw (Fig. 4B),

Iris evident that the finer, muddier and

orpanic-rich sediment trapped by this seagrass

is an ideal substrate for niangrove colonmation.

The mangroves therefore represent a third majar

stage Of plawc/sedimene succession

Young mingrave inees quickly establish a

lateral roda system, each moot bedring a Series

of vertically protruding pneumatophores. Each

Plat is suproliided by Fadlating rows of these

spike-like siructupes, 20-3) cai leh. Fora tiie

both mangrives wnet seavrass co-exist, bul as the

density of pncumatophores increases, so toa

does the amount of organic flotsam that is

trapped by, and adheres to, these subaerial roots.

The Zostera thus dies under a blanket of

accumulating marine compost,

Ping sediment continues to accumulate in this

facies as the mangrove woodland grows to

maturity. Algal and seagrass debris bring

epiphytic carbonate organisms which remain as

grains in the sediment. Thus aggradation

proceeds to levels where the floor of the mature

mangrove woodland (Fig. 4C) is inundated for

only brief intervals during high tides. The

sediment/waler injerface is then colonised and

stabilised by a cyanobacterial mat which

introduces new depositional processes (discussed

below). The substrate is intensively burrowed by

the small mud e¢rab, Helice haswellianus,

facilitating sea water permeability and aeration,

The mangrove sediment thus becomes oxidised

and homogenised.

Throughout this episode of deposition a

dendritic patiern of tidal distributaries develops

and is maintained.

. Vidal distributary facies

Tidal water movernent is concentrated alony

distributary channels which are best developed

within the mangrove woodland. As a

consequence of the dendritic pattern of

channels, current speeds are variable and

sediment type varies accordingly. Major, shallow

channels have coarse, shelly debris, rich in valves

of Katelysia and Anapella (Fig. AF),

disarticulated and convex upwards, Deeper

backwater channels have a high component of

black sulphide mud.

A major component of the shelly fauna within

the mangrove tidal distribularies is the small

turriform gastropod Batillaria (Zeacurnantus)

diemenensis (Ludbrook 1984), Where channels

meer the open sand flat, shells of (his gastropod

are numerous and current aligned (Fig, 4B),

They have been widely redistributed throughout

the Study area.

. Cyanobacterial Mat-Sand Flat Pacies/Zone

Bauld (1981) has illustrated the ability of

cyanobacterial mats to fix sediment in Spencer

Gull, He describes how entangled trichomes of

Microcoleus sp., and their enveloping

muciliginous sheaths, trap and bind sediment

in a thin, coherent layer. In addition, photo:

synthesis at the mat surface removes carbon

djoxide (rom séa water, increasing pH and

favouring precipitation of calcium carbonate.

ST KILDA FORMATION 127

Fig. 4A, Sediment of the Zostera meadow, shelly sand, rich in sulphide and mud, B. Juvenile colonizing mangroves

in the Zostera meadow; algal and other plant debris adheres to pneumatophores and smaller plants. C. Mature

mangrove trees adjacent to a tidal distributary. D. Tidal distributary with mature mangrove woodland, left, and

inner sandflat with cyanobacterial mat, right. E. Gastropod shells, mostly Bari/laria, are a major component of

the sediment in the mangrove tidal distributary (Figs. 4C-D). F. Sediment of tidal distributary which separates

the Zostera meadow from the inner sandflat, Bivalves, mostly Aafelvsia and Anapella are disarticulated and convex

upwards,

1H, CANN & ¥. A. GOSTIN

Cyanobacterial mars cover a large part of the

otherwise unvegetated sand fal, the floor of the

mangrove woodland and same of the supraudal

arcas, These are therefore sites af sediment

aggradation. Sediments bound by eyanobacteriul

mats-are known to withsland curren! velocities

up to five times higher than those required to

erode matefree sediment (Neumann ef a/, 1970).

Sections cut through the active mat (Fig, $C)

reveal a thin layerol oxygenated sediment, below

which is black, sulphide-rich, laminated, silty

sahd (Fig, 5D). Anacrobic bacteria, nourished

by fhe organic matter of previous layers of the

mat, reduce sea waler sulphate ions. Metals,

particularly iron, are fixed in this way. Black

sediment turns to a pale rust colour on

prolonged expasure to the air, Calcium

carbonate content may be as high as 70%.

The cyanobacterial mats support a dense

population of grazing gastropods, notably

Salinator sp, On the seaward! edge of the bare

sand flat, persistent grazing appears to inhibi

establishment of cyanobacteria (Fig. 5B). la this

area, large numbers of the small bivalve

Anapella sp., probably A. eyeladea (Ludbrook

1984), live clustered together immediately below

the sand surtace (Fig. SA), Disarticulated yalves

of Anapella are widely redistributed in sediments

of che study area.

The bare sand flat is sharply separated trom

the Zostera meadow to the west, and the

imangrove woodland to the south (Pig, 4D), by

intertidal distributarics. These ddal channels

have remained essentially unchanged for the past

decade.

Beach facies

Hivh tide beach sediment at Port Gawler has

a valchum carbonare content of abour BOM, Whe

texture is predominantly sandy, but shells

origidating in all of the other fiers oecur as

constituents, Wide variation in grain size, [rom

shell gravel to find sand, may be reluted to

variable wave energy, Which is a function of

water depth and wind sirengih, Sections cul

through tke beach reveal horizontal 1 gently

tilted, well-sorred laminae, and, rarely, higher

anvle cross beds.

Tlotsam seagrass debris appears to signi-

ficantly assist stabilization of beach sediment tr

seaward, While salrbush and other plants are

established in, and fix, the upper beach sands

(Pig. SE), The tigh tide beach is therefore a mass

of accumulating sediment, progradiny seawards

geross the janer Saud flat.

—

Dune facies

Windblown carbonate sunds occur some

50-100 m inland of the beach. This area has

been considerably modified by hurman wetivily,

bul the low dune forms, stabilized by sa}thusls

and other vegetation, are clearly idenfiliable The

sediment is well sorted and consists of fine

broken shell and entire stnall gastrapods and

foraminifera, Sections cul through the dunes

reveal coarser, underlying sediments,

Storm ridge facies

Storm ridges are Formed ut times of extremely

high tides and local storm activity Shefly

material is dpyven shorewards under these

conditions, and may pile up ina ridge of coarse,

poorly soried shell debris, somewhat paralleling

the shore line, but seawards of the beach, If the

ridge is continuous, that part of rhe sand flat

between the newly formed storm ridge and the

pre-existing beach is isolated from further wave

action, A new heach facies is established on the

seaward side of the storm ridge, and a new dune

facies fo landwards.

At northern Pon Gawler, extensive storm

ridges unve existed in the area between che sale

evaporation pans and the present high tide beach

and dune facies, Shell-grit mining has-effectively

removed most of this sediment, the location of

ibe ridges now being marked by a number of

parallel, elongate, shallow depressions.

Immediately south of the Gawler River, a

storm ridge some 1,5 km long and up to 2 im

high lies within the mangrove woodland (Pig, 6).

lis situated several hundred metres landwards

of, and parallel to, the seaward margin of the

mangroves, The seaward side of the ridge 1s

straight and steep, suevesting the form of «

stranded beach (lotelf 1983), On this surface

Harbison (pers. comm.) has observed numerous

large, square-cut pieces of timber, sinilar to

(hase used inv the corstruction of early Saurls

Australian jewes. These tenis of driftwood were

evidently emplaced at the tinte of, or shortly

following, formation of the storm ridge.

Along the coastal plain, north of Pert Gawler,

numerous more ar less parallel storm neges may

be observed.

Supratical lagoon facies

Areas of sand flats that have been isolated

from open marine influence may still reveive sea

warer via fidal distributaries. Alternatively, water

may be supplied by seasonal heavy ram or rising

ground wate, On the floors of these suprajidal

lazoons, or sabkties, Gyametuctenal mats alten

flourish. They maiotain low diversily

ST KILDA FORMATION 129

Fig. 5A. Dead Anapella bivalves (« 1) in life position, immediately beneath surface of inner sandflat; section exposed

by mangrove tidal distributary. B. Grazing gastropods (x 1) limit growth of the cyanobacterial mat on the seaward

areas of the inner sandflat. C. Exposure of sediment beneath the cyanobacterial mat; dark horizon is sulphide

rich, D. Detail of sediment slab, Fig 5C. Uppermost laminae are flushed with photosynthetic oxygen; dark lower

laminae contain sulphide from bacterial reduction of seawater sulphate. E. Inner sandflat and vegetated high tide

beach. F. Mature mangrove trees overhang the Gawler River estuary near the ruins of Lisbon Wharf.

130 J. H. CANN & V. A. GOSTIN

Fig. 6. Aerial photograph of the Gawler River estuary showing the locations of Lisbon Wharf ruins and the site

of vibrocore VC 138. A storm ridge and sabkha, south of the estuary, are indicated.

populations of carbonate fixing organisms such

as the gastropod Salinator sp. and the

foraminifer E/phidium sp. (Cann & De Deckker

1981). Photosynthesis favours chemical

precipitation of calcium carbonate and fine,

wind blown sand is trapped by the mats.

Gypsum crystals may form during summer

evaporation. Samphire plants grow around the

margins of these lagoons, where they trap

supratidal seagrass flotsam and sediment. In this

way they gradually encroach on the area and

contribute to its shallow infilling. Supratidal

lagoons are therefore sites of active sediment

accumulation,

Supratidal lagoon sediment, stabilized by

samphire plant growth, occurs adjacent to the

storm ridge south of the Gawler River (Fig. 6).

To the north of Port Gawler, extensive supratidal

lagoon systems occur landwards of the storm

ridges. Within the study area this facies has

effectively been replaced by salt evaporation

ponds.

10, Estuarine facies

Large quantities of seagrass, algal and other

plant flotsam may be observed on the surface

waters of the Gawler River estuary, transported

back and forth by the tides. Some of this

material becomes entangled in the

pneumatophores of the mangroves that

overhang the water way (Fig. 5F). Other plant

debris accumulates on the muddy bed of the

estuary, particularly in backwaters. The resulting

sediments are therefore peats and organic rich

muds. Sediments of this facies were encountered

in vibrocore VC 138 and are discussed further

below.

Distribution of the various facies described

above is shown by the block diagram Fig. 7.

Foraminiferal biofacies at Port Gawler

Sediment samples were collected, on two

approximately straight line transects, from the

environments described above. The first transect

extended from high tide beach to the edge of the

Posidonia meadow, approximately along the line of

break in block diagram, Fig. 7, the second from a

small sabkha within the samphire zone to a

mangrove tidal distributary. Samples of

approximately 200 ml were taken from the surface

to a depth of about 5 cm. Most samples contained

appreciable amounts of plant debris.

SY KILDA FORMALION 131

STORM RIDGES

HIGH TIDE BEACH

———

JUVENILE

ACCESS

os” ROAD

ev y- SAMPHIRE

YY y- ZONE

41.

} -

ate

iu Ay

Mangroves

Zostera

Posidonia

fa any |

aes Sandflot

Terrestrial sand and clay

iin

Fig. 7. Schematic block diagram illustrating modern depositional environmenis and related sedimentary facies of

the St Kilda Formation at Port Gawler. Sites of vibrocores VC 13@ and VC 138 are indicated.

PORT GAWLER

AREA

TABLE 1. Summary af the sedimentar\ facies of Port Gawler

Facies Environment Sediment Lype

Posidonia—Pinne — subtidal, outermost sand flat carbonate yuartz shelly sands

Zosteru—Katelysia — intertidal sand flat muddy carbonate quariz sands

Mangrove intertidal mangrove woodland carbonate quartz silts, muds and sands with organic debris

anc cyanobacterial mats

Tidal distributary throughout the tidal range channel deposits with convex upwards disarticulated bivalves

and current aligned gastropod shells

Cyanobacterial high lide inner sand flat line carbonale quartz sand bound by cyanobacterial mats

sand flat

Beach high tide shelly carbonate quartz sands and shell gravels

Dune supratidal, back of beach well sorted carbonate sand, reworked from beach wand otter

sediments

Storm ridge supraticdal coarse comminuted shell; lag deposits of this facies underlic

present day sand flat sediments

Supratidal lagoon supratidal, samphire sypsiferous carbonate quartz muds, silts and sands

or sabkha

Estuarine Gawler River estuary and peat, organic mud and related sediments

hackwaters

132 J. CANN & ¥, A. GOSTIN

In processing the laboratary samples, the high

organic content made u difficult te determine the

number of foran\initera with respect to a dry weight

of sediment and this line of investigation was not

pursued. However, all samples contained sulficiens

numbers of foraminifera for relative percentage

analysis.

After boiling in frestr water for about an hour

to break down the organic debris, samples were wet

sieved and sediment of sand size (2.00-0.062 mm)

was retained. Foraminifera were concentrated from

these sand fractions by Jlotation on

tetrabromoethane, Residues were inspected Io

ensure that most tests had been separated. The dred

foraminiferal concentrates were further sieved! inte

phi grain size fractions for microscopic

examination, For each sample, percentace

abundances of foraminifera species were determined

for the fractions 1.00-0,50 mm and 0,50-0,25 mim-

Analysis of these fractions only, rather than the

entire sample, has a number of important

advantages.

Ellison (1951) has commented on the time

consuming nature of qitantitative foraminiferil

analysis in palaeoecalegical studies. The work

reported here was part of a more extensive

investigation involving several hundred surface

sediment and core samples.” It was importan) that

these samples be processed efficsently, Coarser

fractions require lower powers of magnification,

ereatly facilitating identification and separation of

individual specimens,

In finer fractions, juvenile individuals constitute

a high percentage of the foraminifera. Species

identification is more difficult for juveniles,

particularly for miliolid genera, Schnitker (1967)

observed thal tests of juvenile THlocu/ita lnmeiara,

asexually produced in laboratory cultures, differed

significantly in form from that of the parent. It is

well Known that some species of Triloeteline may

exhibit quinqueloculine coiling as juveniles,

becoming triloculine only as adults (e.g, Loeblich

& Tappan (964), Such changes in form ane further

complicated by microspheric/megalospheric¢

dimorphism,

(n natural sedimentary systems, (ests of smaller

forantinifera are easily winnowed tram their

environments of origin to be deposiied elsewhere

Also, during intraenvirotimerntal Lansport, smaller

Mofe fragile species aré more prone to atritigd and

mechanical destruction than larger, more robust

forms, Finally, when considenfg core samples, pose

diagenesis preservation will favaur larer forms, less

prone to solution, In this study, specles of smaller

genera, such as Bulindnoides, wre considered (or

*Cann unpublished data

serenilly tave less value as environmental indicators

than those af larger genera, such as Pencroplis.

In the studied size fractions twenty-eight species

of foraminifera were observed, The percentage

numerical distributions of the more common

species, with reference tu the sedimentary

environments of Port Gawles, are shown in Tig. §.

These species are illustratedt by scanning electron

photomicrographs in Figs 9 and 10,

Systematic notes on selected species

of Foraminifera

Nubecularia Jacifuge i extremely variable in

morphojogy, its shape often influenced by the

object to which it adheres, Some forms ure plano-

convex, the Mat surface usually incomplete and

revealing a planispinel arrangement of chambers.

A ereat many individuals have globular, twisted,

tube-like tests, with multiple apertures and were

presumably unattached. Both attached and

umiattachel forms are recownised here as the same

species.

Peneroplis planaius forms thin, translucent,

planispiral tests in deeper water, but in warm,

tntertidal areas, where salinity rises with high rates

of evaporation, tests are thick and often aberrant.

The range a! morphology that may be observed in

such populations of Peneropliy has been well

illustrated by Sellier de Civricux (1970) for the

Mediterranean and by Hughes-Clarke & Keij (1973)

for the Persian Gull. Similar fornis occur in Port

Gawler sediment and are considered here is a single

Species.

Spiroloculina spp, here includes. §. aatdarum and

also forms havitig chambers of more quadrate

section, lacking the numerous oblique costae of A

anrifuram, but bearing distincs longitucinal ridges

wt the edges oof the chambers, and often a third,

parallel, and in the centre of the flat peripheral

switace, Such forms are referrable to S. tricasta

Cusiiman & Todd, 1944, Some individuals have

been observed to have features intermediate between

those definitive of these two species,

Disrorhis dimidiaras is here used essentially in

the sense of Hedley ef af. (1967). hey recognised

that forms having grearly varying morphology, such

ws keeled of lobate periphery, high or low spire, large

or small ventral Maps, correctly belong (o a single

specres, AID of their ifustrated forms have been

recognised in vie Port Gawler material, and when

Many speamens are examined ac one time, it is

apparent that vanalion i conlinuos.

Alphidian macefiiforme is medium to large for

The genus, distinguished by its tiuck, lens-tike forms,

Wi) aunicrous Involage, aon-lnflated chambers.

Sutures are raised, curved and limbate, joined by

ST KILDA FORMATION 133

ELPHIDIOM_MACELLIFORME —

ELPHIDIUM CRISPUM

DISCORBIS DIMIDIATUS

PENEROPLIS PLANATUS | a

OO MILIOLINELLA LABIOSA=—————————— SH aA

———___TRILOCULINA, INFLATA——————

——— ———SPIROLOCULINA SPP SS > —>

CRIBROBULIMINA MIXTA 7

SIEVED SAMPLES: FRACTION SIZE 0.50-0.25mm

1,00-0.50mm

PENEROPLIS PLANATUS

NUBECULARIA LUCIFUGA

SIEVED SAMPLES: FRACTION SIZE

CRIBROBULIMINA MIXTA

<= SEAWARDS LANDWARDS =>

am) z =o

q iw rs)

Pr 5 :

= - Fic

b =z at o

= a= e2hg We ae Lu lu

> aa <t 2g 52 a, iid

z2>= or r= So on Zu =) tea

oo ex iu eC rst rama 2 — rr

20 go fra wero Or Ta ox

ot —— os FWe 25 az a =O

ow ><a ow asia an =a 2 aa

a= on n= 620 =a =H x ann

ADJACENT SEDIMENTARY FACIES (NOT TO SCALE)

Fig. 8. Foraminiferal biofacies: relative abundance of selected species through modern, adjacent sedimentary

environments, Port Gawler, South Australia.

134 LH. CANN & VA. GOSTEN

many slender retral processes. There is a prominent!

imperforate umbilical boss which is neither raised

nor depressed, The interiomurginal aperture is

Obseuted by pustular carbonate low on the apertural

face, The periphery bears a low narrow keel. The

surface is uniformly white and glossy,

This species, which is common in South

Australian gulf waters, has only recently been

described (McCulloch I981). fl is likely thar in

previous records of Australian Holocene

foraminifera, it has been identified as ElpAidiuen

advenum Cushman 1922, Cushman (1922)

originally described 2. odvenc as having a depressed

umbilical region, and lacking any significant boss

or similar material, However, in subsequent work

(1939) he referred forms having a rhomboid sectlon

and (lush umbilical plug (ee. Brady 1884, PL CX,

figs. a, b) to the species, Austrahian workers (eg,

Chapman 1941; Parr 1945; Collins 1974) hive

followed this later interpretation,

Vella (1957) maintaiied a detinetion between

forms having a depressed sembilicus with i small

central boss of clear shell ayaterial, and thease horns

having the umbilicus covered by Mattencd, glossy

plugs not protruding beyond the outline of the stietl,

For the latter form Vella (1957) erected

Elphidionortian eharlovensis, which appears elusely

similar to the species figured by Brady, cited above,

McCulloch (1981) also believed (hat Custinan’s

later imerpretation ol Alpfidiwe agvvenun inchided

more Than Owe species and accordingly established

Elphidiuem macelliferine, Uescrited above, Wer

descripljons aad figures agtee With species occuring

in the Holocene sediments of Morl Gawler.

Apthiorpe's Evmdidivee sroce//uene (1980, pl. 26,

fig. Halse apnedtrs to belongs cof mtaeedlliforen

and her & advend (1980, pl, 26, fie. 10) may be a

juventle Of Glee species,

Port Gawler vilrocores VC 156 andl VO 13%

Vibtoocores were taken at the sites indicated yn

Fig. 7, VC 136 on the open inner sind Hat and VC

138 in the mangrove wordland, adjacent to the

Gawler River estuary. Both cores penerrated almost

3 merres of sediment.

In the laboratory, the cores were split lengthwise

and lithological logs prepared. These are presented

as Flies. tl and 12. From one half, samples of

approximately LOO ml of sediment Were taken at

24 cm intervals and processed as described above.

For cach sample, percentage abandances of

foraminifera were decermines! for the sive frietions

Fig. 9. Trochienmina teffata (Montagus 1808 a- © 120; d,

1,00-0.50 mm and 0.50-0,25 mm, The percentage

distributions of the more cormmon species were

determined. These dara are shown down core for

the productive intervals, here considered as St Kilda

Formation, In Pigs. 13 and 14.

Each core represents a sequence of near shore

marine environments, recording both the initial

Holocene transgression and some of the later

episodes of sedimentation that followed sea level

stability, The down core foraminiferal assemblages

are, at least in part, indicative of those

environments, and may be compared with the

fOramingleral data obtalned for the various facies

deseribed nbows,

The reliability of the data was initially tested by

comparing two samples from the cyanobacterial

mal facies. the first from the surface transect, the

second from the top of VC 136, The sample

localities were about 500 metres distant from cach

other on the mid-inner open sand flat,

Pig. 15 illustrates this comparison in the form of

histograms. The data for the size tractions

0.50-0.25 mm are remarkably similar. That for the

coarser fractions are less convincing, perhaps

reflecting the Senaller number of specimens counted

for thal size range. The number of |ndividuals

recovered from che fraction 1,00-0.50 mim were 80

and 123 for the transect and top of VC 136

respectively; equivalent numbers for the finer

fraction were 211 and 717.

For the sediment size fraction 0.50-0.25 mm,

where the number of individuals comprising the

fraction is approximately 200 or greater, the

percentage abundance of foraminifera species ts

considered 10 be a reliable envirommental indicator

Results and interpretation of cores

bre I36

The earliest sediments of the St Kilda Formation

corded in Yibrocore VC 136 represent the

supratidal facies of the samphire zone. Fig. 16

compares the percentage abundance of species of

foraminifera from the modern samphire with that

observed at 50 omin VC 136. The overlying coarse,

shelly sand, sampled at 25 cm, is interpreted as

remnant, storm driven shell debris. Although oo

foraminiferal data were determined for an actual

storm rider, the percentage abundance of species

obtained for the high tide beach, developed on the

seaward side of a low storm ridge, is essentially

similar to thal al 25 cm in VC 136. The top saniple

of VO 136 has been discussed above,

~ 660. Colhratulinina mixta Cushotan 1927 ef, 40;

gp, «O60. Noliwularie (veiw Defrance 1825 hj. «40. Syrroloenling antillerven VOrbigny 139 bol 950,

Spirolocuiina trieasa Cushman & Todd 1944m on, a). Trilovuliny inflata UOrbigny UR2b o-G) 9 60, Miliotinella

fotesa WUrbiginy) TARY ret, «ROL Peneroplis plonotiey (Fieltcl & Moll) 1798 u-v, » 30.

J. H. CANN & V. A. GOSTIN

ST KILDA FORMATION 127

VIBROGORE VC 136

with some line quartz sand.

Depth in centimetres

Silty, very tine, quartz sand

POORAKA FMN, ST- KILDA FMN.

Dark yellowish brown clay.

Penetrated 294 cm

Recovered 272 cm

Description

Yellowish grey, mixed quartz-carbonate, well sorted, fine sand.

Raddish brown, stiff, sandy mud

Environment

Cyanobacterial sandflat

Greyish yellow, poorly sorted, coarse shell deors

Storm debris shoreface

Dark greenish grey bioturbated mud with poorly sorted,

muddy, shell dabris and fine quartz sand

Samphire

Bark yellowish brown, indistinotly mottled, micaceous, sandy silt

Alluvial flood plain

Mean grain size Incréases down core to

very coarse , angular, quartz sand with quartz pebbles

River channel

Alluvial Hood pigin

Fig, 1, Descriptive and interpretive log of Vibracore VO 136,

Vibrocore VC 136 thus records the Holocene

transgression as initially marginal marine supratidaf

sediments, overlain by storm debris, remnant of the

episodes of storm ridge formation, The uppermost

sediments are the result of aggradation on

cyanobacterial mats.

VC 138

The proximity of the site of vibrocore VC 138 to

the Gawler River estuary suggests that sediments

penelraled by this core would have been deposited

under condilions of fluctuating salinily, However,

tidal channels 2 Km and 2.5 km south-east of Port

Gawler probably indicate former sites of

debouchment of the Gawler River, The present

course of the estuary has apparently developed only

in later Holocene.

Although only 57 individuals are recorded at

75cm in VC 138, for the size fraction

030-0,25 mm, the percentage distribuhon of

foraminifera species agrees closely with dara

established for supratidal samphire sediments

(Fig. 16), The distribution of species at 125 cm is

very similar to that observed for the high tide beach

atid apparently indicates storm sediment. Thus the

transpressive sequences of VC 138 and VC 136 are

essentially the same

From 100 em to 75 cm the increase in numbers

of Spiraloculina spp,, Miliolinella labiosa, Discorbis

dimidialus and Elphidiuni crispurn, with decreasiny:

numbers of Peneraplis planatus, is consistent with

the development of seagrasses (Fig. 17). The

increase in Ammonia beecarii, a noted polyhaline

species (eg, Murray 1971), suggests increased

influence of tresh water from the Gawler River,

A dramatic change in the foraminiferal

assemblage occurs at 50 um, as Trhochameina

inflata numbers increase trom almost Zera (6 over

70% and most other species disappear,

Trochumminu inflata is well established in the

literature as an estuarine dweller (eu. Collins 1974).

Apthorpe (1980) reported the species fron low

salinity waters of the Gippsland Lakes, Victoria.

Cano (1984) has found it to be a significant

component of the fauna of the upper Onkaparinga

River estuary, south of Adelaide, South Australia,

The abundance of Trochummiina inflata, veaching

LOO Of the fraction 0,50-0,.25 mm at 25 cm,

marks the development of the Gawler River estuary

al ils present site,

Finally, reappearance of most species in the

uppermost sediment of the estuarine manzrove

woodland suggests a return to conditions of

tolerable salinity. If this is so, there may have been

hip. 10. Discorhis dimudiatus (Parker & Jones) 1862 ac, «35; d, «350. Ammonia heecarii (Linne) 1758 ep, = 120;

= 70. Blphidiumn macelliforme MeCulloch 1981 iso, » 60, ThichoAvalus trapicus (Collins) 1958 p-r, <5: 5,

S5fl.

{Although 7 fropicus abundance is less Haan 3% (royehout bor) VO 146 and VC 134, iF is present in significant

Tumbers in the surface sediment of the estuarine maunerove woodlaad Cann & De Deckker (981) reported f)is species

trom a saline lake.

138 J. H. CANN & ¥. A, GOSTIN

VIBROCORE VC 138

Description

Disarticulated Katelysia.

Live mangrove roots.

Carbonate lodules,

very fine, silty, quartz sand.

Carbonate nodules and ?burrows.

Brownish grey to reddish brown,

stiff, sandy mud

POORAKA FORMATION

Penetrated 308 cm

Recovered 274 om

Medium grey, poorly sorted, coarse shell and shell sand.

Carbonate cemented, fine quartz sand.

Yellowish grey, moderately sorted, medium, quartz sand-

Environment

Estuarine mangrove woodland.

Estuary-

Zostera

Seagrass meadow sandilat.

Posidonia

9 Yellowish brawn, mottled, organic rich mud. Rare shells.

= = Dark grey to black muddy peat with live mangrove roots.

= Light grey, moderately sorted, coarse to very coarse,

= gastropod sand, with minor larger «hells.

Zz Live mangrove roots and ?sea grass roots

iva]

Storm debris shoretace.

Sampnire.

Medium grey, bloturbated (?crab burrows), fine, sandy, quartz sill,

Alluvial Nood plain.

Light olive grey, micaceous, poorly sorted,

River channel,

Alluvial Hood plain.

Fig. 12. Descriptive and interpretive log of Vibrocore VC 138.

a real decrease in fresh water debouchment due to

increasing climatic aridity, or due to human

modification of fresh water drainage since

European settlement of the area. Alternatively, the

reappearance of these species may simply be due

to the trapping of sea grass debris, with associated

foraminifera tests, by the pneumatophores of

recently grown mangroves.

Conclusions: six thousand years

of coastal accretion

Following transgression of the post glacial

Holocene sea to its present level, an ordered

yonation of plant and animal communities became

established. Favourable conditions ensured vigorous

organic growth and rapid production of bioclastic

sediment. An originally alluvial landscape was

rapidly modified as storm shell ridges were thrown

up along most of the coast. Saline marshlands

formed landward of the ridges. The sediment

redistributing effects of high tide waves and storms

persisted to varying degrees along different parts

of the coastline.

Around sites of debouchment of the Gawler

River, deposition of estuarine muds favoured the

development of seagrass, mangrove and samphire

plant communities. In contrast, in the northern part

of the study area, prolonged wave-dominant

sedimentation gave rise to a succession of storm

shell grit ridges.

The northern area is today little influenced by

wave action. Only minor amounts of new sediment

are being added to the high tide beach, and young

mangrove trees are rapidly colonizing the Zostera

seagrass meadow. As these new Lrees coalesce with

the existing mangrove woodland, and with

ST KILDA FORMATION 139

SIZE FRACTION 1,00 - 0,50 mm SIZE FRACTION 0.50 - 0.25 mm

® of — —T _

a || fe : z

E =

£25

o <x

0 QO

7 a

= ss =

50, x

£ H

o w

O75

e & 0 20 40 60 80 100

Ny uw z

nt oO =

ro nr} 4 ve

= a x 3 3 = fa 3 g S Fs oly

Fa = w & £ = = w Sle

o = ca S c 2S 2 2 3 2 t/a

c ce =. = i — £& —

5 = = o a 2 S cc = 2 E €

a“ & 5 3 = Q 2 = & ‘a 5 35

= 9° rs) 2 e ° 3 2 a Ss

° oO x 3 o @ re 3 o a Q 2 \JH.

Oo re oO Zz a Q oO Zz ao a w wi i198

Fig. 13. Percentage distribution of selected species of foraminifera down core #VC 136, open inner sand flat, Port

Gawler, South Australia.

SIZE FRACTION 1.00 = 0.50 mm SIZE FRACTION 0.50 - 6.25 mm

of ; ¥ ———— = i

a j | jf NN i] T |

= J vA \ f \

wo f ye ‘

E 25 g . < Y j

= s oh ‘nls \ f f

c | ~. } vA ‘ \ ff i i"

6 i ie , i \ | \ / | | | \2

« 50 J \ \ ta ( \¢ » . XN vw nN | \ in)

emi bt Ny \ Jf ; 7 / \ [=

E hoo \ \ ff *, 7 / | |<

ars / \ ) (4 ) / \ | | a

| \ / AN iy fi \ re

f ‘ é \ <

400 ( ) \ | " | f o

| | | =

| | i *“

125 | | | ag ) :

} | \ / }

| \ | | \ /

VSO; \ ) | \ } } if

\ , | . | } | / | |

\ / 1] \ fot | ; | |

75 Insufficient samole | | \ } } —

\/ ree VA NV Ny i

200 Vv 4 J \

c = _ Eee

5 a 20 40 60 680 100 a

wm % 5

= uu 2

= 4 oo

7 E

E = 3 2 = ' 2 = 3 =

wv. “C1 E 2 3 & = = =a a = s 2 E& Sia

ra < 5 = a = S = = Se — o so Slua

5 ey : = = Ff) + g 722 8 £3 & tis

= ‘Sy 5 = = 2 : E tz 3 e me HY =

BS 3 E 3 Ee 4g 222 8 s 2 5 5/5

o ES 3 3 a 5 5 S a S - ze =/e

5 5 6 = 5 5 8 2 85 2 5 2 Fs 6/*

oo 5 = w a « zZz5 = oa Go < t &

Fig. 14. Percentage distribution of selected species of foraminifera down core #VC 138, estuarine mangrove woodland,

Port Gawler, South Australia,

140 J. H. CANN & VY. A. GOSTIN

N=717 N=123

VIBROCORE VC 136,

O =-2.-6M

£53283 352 £ 5|3]2 2

x Ze 2% Boas x Z)als 5

EGa 2 Ssz2e E Sisils 2

eo 2fE-ZSERVE % Pee a oe

= 8 5327 3 E SO = S|,/5 0

Ezgee2F,28 | =a B-3 o

Sa 2 ce 253 E = slal@e

oa 85 ¢ fie 5 a 21els

co at8 88 5 25 e oisis §

saga°3R cats S 2}/s/2 3

6 Zz an) 4 6 z/*/o =

uw ir

N=211 N=80

TRANSECT SAMPLE,

MID-INNER OPEN SAND FLAT.

0.50 - 0.25 mm JH.C, 1985 1.00 - 0.50 mm

Fig. 15. Comparative percentage distributions of species

of foraminifera for two samples from the inner open

sandflat. N refers to the number of individuals

constituting the size fraction.

continued sedimentation, the northern area will

come to resemble the rest of the tide channelled

samphire and mangrove coastal zone.

Thus it may be concluded that the segment of

coastline discussed in this paper has evolved from

saline marshlands and storm shell ridges, to a

seagrass-sand flat-high tide beach zonation, formed

under vigorous activity of waves (such a condition

still prevails along the Adelaide metropolitan

beaches), and finally to an environment of tide

dominant sedimentation involving mangroves.

The following conclusions refer to the Holocene

St Kilda Formation at Port Gawler.

1. The Formation consists of a number of marginal

marine sedimentary facies which are closely

related to different plant/animal communities.

2. The earliest sediments were transgressive,

consisting predominantly of samphire muds and

shelly storm debris.

3. Later sediments were formed by processes of

progradation and aggradation at present day sea

level. These processes are inextricably associated

with various plant communities.

4. The sediments of most facies are rich in calcium

carbonate, up to 90% for the shelly sands of the

Posidonia seagrass facies.

—s Ss

Xx 3Gs322E

esgeiee2s

o 2Z2f-3s2685

c Os AE o

FS Wie eS Eo

f ie} Oo. Ss =) oO

SSG FE RMSE

ae Bee Ve

° im ai us” a. rs a 5

» @oeee g = 2

2 5 =oy,t ov

5 7 a

4 VIBROCORE VC 138,

175 - 177cm

N=57

N= 522 VIBROCORE VC 136,

50 — 52cm

)

N= 289 TRANSECT SAMPLE,

SAMPHIRE ZONE.

J.H.C. 1985

0.50 - 0.25 mm

Fig. 16. Percentage distributions of species of foraminifera

for samples from vibrocores VC 136 and VC 138,

compared with a surface sample from the samphire

zone. Only the size fraction 0.50-0.25 mm is considered.

N refers to the number of individuals constituting the

size fraction. The similarity of the histogram for the

samphire zone and the mid-inner open sandflat (Fig.

13) reflects the landward sites of deposition and

similarity of sedimentary processes.

5. Distinctive, though sometimes subtle, changes

in the percentage abundance of foraminiferal

species occur from one facies to another.

6. Reliable comparisons of foraminiferal biofacies

can be achieved using the size fraction

ST KILDA FORMATION ial

N=675

VIBROCORE VC |38,

75-77 om

TRANSECT SAMPLE,

ZOSTERA MEADOW

= |S lee!

5 %

Fggzgsigleg

S232 2 Flaja —

Epme § Elsie 2

Src SS waiele se

e2ew S\|E|% F

ea GE ,|B/E 8 25

Et. Ss @ 2 =a w

SFxePBoe alfl|s E N=1307

BpPeES Selielece

2s BREE 18/2 5

5.2. % = aje|4s

= a =

5 = = 4a

at TRANSECT SAMPLE,

POSIDONIA MEADOW

jones) 9 SeHbth” ot wae

050 —-0.25 mm

Fig. (7. Percentage distribution of species of foramiiitlera

for sutface samples from sea grass meadows with a

saniple from vibracore VO 138. Only the size fraction

0.50-0.25 mim is considered, N refers bo the number of

individuals constimting the size fraction,

0.50-0.25 mm of floated foraminiferal

concentrates,

7. Down core documentation of relative

abundances of foraminiferal species, in

conjunction with lithological studies, can be

used to determine palaeoenvironments and thus

the sedimentary history,

Acknowledgments

The Marine Sciences and Technologies Grants

Scheme provided financial support for the coring

programme. The Council of the South Australian

College of Advanced Education granted

professional leave to John Cann lor the preparation

of Lhe paper, Sharon Proferes drafted figures 1, 7

and 8; Francis Gorostiaga assisted technically in

recovering the vibrocores; Brenton Bowman

processed samples for foraminiferal analysis;

Richard Bacrett prepared final photographic plates;

Chris Moore typed the final manuseript, Some of

the research reported here was completed by John

Cann as part of a Ph.D. programme at the

Universily of Adelaide under the supervision of Dr

B, McGowran, We thank Dr B. McGowran, Dr

A. P. Belperio, Mc M. Lindsay and Professor

H. B.S. Woinersley for critically reading the

manuscript,

References

Al@Hison, G. D., Spatag, R. C., & Cocuramr, G. W.

(1954) The soils and geology of Adelaide and suburbs,

Bull, weal, Surv, 5, Aust. 32, ch. 4, 19-28,

Aprooree, M. (1980) Foraminiferat distribution in the

estuarine Gippsland Lakes system, Victoria. Proc, R.

Sae. Vie. 91, 207-232.

Bavlb, J. (981) Geobjological role of cyanobacterial

mats in sedimentary environments: production and

preservation of organic matter, RMR LD Ausn Geol.

Geophys. 6, 307-317,

BALPORIO, A, By HATLS, JR & Gostin, ¥, A. (1983) A

review Of Holocene sea levels in South Australia. /a D.

Hopley (Ed.), “Australian sea levels in the last 15 (00

years, a review.” Dep, Geogr, Jumes Cook Uniy, North

Queensland Moenegr, Ser, Qcoas. Pap, 3, 47-47,

Meany, HB. (1884) Report on the foraminifera dredwed

by H.M.S, Challenger, during the years 1873-76. Rept

Sel, Results Baplor Voyage H.M.S, Cliallenger, Zool,

Pr. 22, 9, 1-814.

Bukron, Tb. (9829) Mangrove changes recorded north

of Adelaide, Safic 6, 8-12,

(1982b) Mangrove development north of Adelaide,

1935-1982. Trans. R. Soc. 8. Aust. 106, 183-189.

BuTier, A. J, Detrtks, A. M., McKiniue, S.C. &

Tomas, D, P, (1977) Distribution and sediments of

manyrove forests ia South Avstralia, (bj WOT, 35-44,

& Brewster, by J, (1979) Size distributions of the

fan-shell Pinna bicolor Gmelin (Matluseas

Eulamellibranchia) in South Australia, ar J Mar:

breshwat Res, Mh 245-44,

Byi, J. A. T. (1976) Physical oceanography of Gult Si

Twidale, M. J, Tyler & B. P. Webb (Eds.) “Natural

History of the Adelaide Region,” (R, Soe, S. Aust.

Adelaide.)

CANN, J, H. (1978) An exposed reference section lor the

Saaayile Formation. Q. geol. Notes, zeal. Surv. 8. Aust,

65, 2-4,

(1984) Quaternary foraminifera, environments and

palacoenviranments of Spencer Gull, 8, Alusy, Afar

Seminar Ser. Abstracts.

& Dr Deckkrr, P. (1981) Fossil Quaternary and

living foraminifera from athalassie (nen-marine) saline

lakes, southern Australis, < Puleont, 55, 660-670.

CHarman, F. (1941) Report on foraminiferal soundings

and dredgings of FS. Endeavour along the continental

shelf ot the south-east coast of Australia. Trans. R, Soe,

S, 46s) 5, 145-211,

CHAPMAN, V. J. (1975) Mangrove Vegetation. (J, Cramer;

Germany.)

Coxrtims, A. C. (1974) Port Phillip survey 1957-63.

Foraminiferida. Mem. Nain. Mus. Wie. 35, 1-61.

CUSHMAN, J. A. (1922) Shallow-water toramunifera ot the

Tortugas region, Carnegie Inst, Washington, Publ 31.

Dept, Mar, Biol, Papers V7, \-85.

(1939) A monograph of (he foraminiferal family

Nonionidae US. Geol Sure, Prof’ Paper 194, 1-100.

142 J. H. CANN & V. A. GOSTIN

DAILy, B. FIRMAN, J. B., ForBEs, B. G. & Linpsay, J. M.

(1976) Geology. pp. 5-42. In C. R. Twidale, M. J. Tyler

& B. P. Webb (Eds.) “Natural History of the Adelaide

Region.” (R. Soc. S. Aust.; Adelaide.)

ELLison, S. P. (1951) Microfossils as environment

indicators in marine shales. J. Sed. Petr, 21, 214-225.

FirMAN, J. B. (1966) Stratigraphic units of Late

Cainozoic age in the Adelaide Plains Basin, South

Australia. Q. geol. Notes, geol. Surv. S. Aust. 17, 6-9.

Halts, J. R., BELPERIO, A. P., Gostin, V. A. & SARGENT,

G. E. G. (1984) The submarine Quaternary stratigraphy

of northern Spencer Gulf, South Australia. /n J. R.

Hails & V. A. Gostin (Eds.) “The Spencer Gulf Region.”

Mar. Geol. 61, 345-372.

HARBISON, I. P. (1981) The case for the protection of

mangrove swamps: geochemical considerations. Search.

12, 273-276.

Hep ey, R. H., HurRDLE, C. M. & BurpettT, I. D. J.

(1967) The marine fauna of New Zealand: intertidal

foraminifera of the Corallina officinalis zone. N.Z.

Dept. Sci. Ind. Res. Bull. 180.

HuGHES-CLARKE, M. W. & Kew, A. J. (1973) Organisms

as producers of carbonate sediment and indicators of

environment in the southern Persian Gulf. Jn B. H.

Purser (Ed.) “The Persian Gulf, Holocene sedimentation

and diagenesis in a shallow epicontinental sea.”

(Springer-Verlag: Berlin.)

KELLY, L. R. (1984) Characteristics of the southern

Australian sea level signal. S. Aust. Mar. Seminar Ser.

Abstracts.

LogEBLicH, A. R. & TAPPAN, H. (1964) “Treatise on

invertebrate paleontology.” Jn R. C. Moore (Ed.) Part

C, Protista 2, Sarcodina (chiefly Thecamoebians and

Foraminifera). (Geol. Soc. Amer, and Uniy, Kansas

Press: Lawrence.)

Lupprook, N. H. (1984) Quaternary Molluscs of South

Australia. Dept. Mines & Energy, Aust. (S.A. Govt.

Printer: Adelaide.)

McCuLLocu, I. (1981) Qualitative observations on Recent

foraminiferal tests, part IV, with emphasis on the Allan

Hancock Atlantic expedition collections. (University of

Southern California: Los Angeles.)

Murray, J. W. (1971) “An atlas

foraminiferids.” (Heinmann: London.)

of British

NEUMANN, A. C., GEBELEIN, C, D. & SCOFFIN, T. P.

(1970) The composition, structure and erodability of

subtidal mats, Abaco, Bahamas. J. Sed. Petrol. 40,

274-297.

Parr, W. J. (1945) Recent foraminifera from Barwon

Heads, Victoria. Proc. R. Soc. Vict. 56, 189-218.

Provis, D. S. & RADOK, R. (1979) Sea level oscillations

along the Australian coast. Aust. J. Mar. Freshwat, Res.

30, 295-301.

RopBerTSON, E. L. (1984) Seagrasses. Jn H. B. S.

Womersley (Ed.) “The Marine Benthic Flora of

Southern Australia. Part 1.” ch. 6. (S.A. Govt. Printer:

Adelaide.)

SCHNITKER, D, (1967) Variation in test morphology of

Triloculina linneiana d’Orbigny in laboratory cultures.

Contr. Cushman Fdn. 18, 84-86.

Scorrin, T. P. (1970) The trapping and binding of

subtidal carbonate sediments by marine vegetation in

Bimini Lagoon, Bahamas. J. Sed. Petrol. 40, 249-273.

SELLIER DE CivriEUX, J. M. (1970) Mutaciones recientes

del género Peneroplis y relaciones filogénicas con otros

Soritidae. Rev. Espan. Micropal. 2, 5-12.

TALgor, U, (1982) Environmental change in the Port

Adelaide-St Kilda area. Taminga 19, 22-39.

Tate, R. (1879) The anniversary address of the president.

Phil. Soc. Adelaide (R. Soc. S, Aust.) 1878-79.

THompson, B. P. (1969) ADELAIDE map sheet,

Geological Atlas of South Australia, 1:250 000 series.

(Geol. Surv. S. Aust.: Adelaide.)

TorerF, S. (1983) Draft management plan for Port

Gawler Conservation Park, Gulf St. Vincent, South

Australia. Dept. Env. & Planning, Adelaide. (S.A. Govt

Printer: Adelaide.)

VELLA, P. (1957) Studies in New Zealand foraminifera.

N.Z. Geol. Surv. Pal. Bull. 28, 1-64.

WomERSLEY, H. B. S. (1956) The marine algae of

Kangaroo Island. IV. The algal ecology of American

River inlet. Aust. J. Mar. Freshwat. Res. 7, 64-87.

& THOMAS, I. M. (1976) Intertidal ecology. pp.

175-185. In C. R. Twidale, M. J. Tyler and B. P. Webb

(Eds.) “Natural History of the Adelaide Region.” (R.

Soc. S. Aust.: Adelaide.)

Wynne, A. A., FOTHERINGHAM, D. G., FREEMAN, R.,

Mou Lps, B., PENNEY, S. W., PETRUSEVICS, P., TUCKER,

R. & Extis, D. P. (1984) Adelaide coast protection

strategy review. (Dept, Eny, & Planning: Adelaide.)

REDESCRIPTION OF TWO TRICHURID NEMATODE PARASITES OF

VERTEBRATES IN AUSTRALIA AND PAPUA NEW GUINEA

BY DAVID M. SPRATT

Summary

Anderson & Bain, 1982 placed Paratrichosoma Ashford & Muller, 1978 as a synonym of Capillaria

(sensu latu) Zeder, 1800. The reasons for this synonymy are discussed herein and Capillaria

crocodilus (Ashford & Muller, 1978) comb. nov. from tunnels in the epidermis of Crocodylus

novaeguineae Schmidt in Papua New Guinea is redescribed from co-type specimens.

REDESCRIPTION OF TWO TRICHURID NEMATODE PARASITES OF VERTEBRATES

IN AUSTRALIA AND PAPUA NEW GUINEA

by DAVID M, Spratr*

Summary

Sprarr, DM. (1985) Redescriplion of lwo tfichiitid fHematode parasites of Vertebrates in Australia and

Papua New Guinea, Trans, &, Soe. & Aust. U4), 143-150, 29 November. 1985-

Anderson & Buin, 1982 placed Perarrichosoma Ashford & Muller, 1978 as a synonym of Cupiffaria

(sensu lain) Zeder, ROW The reasons for this synemymy are discussed herein and Capilaria eracodilus

{Asitford & Muller, 1978} comb: nov. [rom (unhels in the epidermis of Crocodylus navarguineae novtegwinede

Sehandt in Papua New Guinea is redescribed from co-type specuinens,

Vrichosomoides nusulls Bioewa & Aurizi, 1961 is described and illustrated (rom the nasal cavity of

Rattus fuscipes (Watertouse) in southeaster) New South Wales, This represents the first record wf ihe parasite

in Australia and &. fuseipes is (he only murid fost known to date

Key Wornos, Peretrichosomu, Cupillaria, WHelinsameides. Nemavoda, distribution, husis, morphology,

Muridac,

Introduction

The Trichuridae is a cosmopolitan family of

parasitic nematodes which are poorly understood,

both morphologically and biologically, During

study of the Wrichurids of Australasian vertebrates

the need arose to re-examine two unusual species,

one described originally as Paretrichosama

crocodilus Ashtord & Muller, 1978 from Crocudylus

novaeguineae novaeguineae Schmidt at Moitaka

crocodile farm, Port Moresby, and 7richosomoides

nasalis Biowca & Aurizi, 1961 first reported fom

Rattus norvegicus (Berkenhout) in Rome (Aurizi,

1958) and not known to occur in rats in Australia.

Redescription al’ these species ancl consideration of

their taxonomic placement form the basis of this

report.

Materials and Methods

Nematodes were fixed in hot, 10% neutral

buffered formalin and cleared in lactophenol.

Co-types of 7, nasalis were kindly loaned to ure

by Professor E. Biocca of the Istituto di

Parassitologia dell’ Universita di Roma. Loan of

male and female paratypes of Paratrichosome

crocodilus was kindly arranged by Mrs B. A. Hareis

of the British Museum.

Measurements were made with the aid of an

ocular (iicrometer, drawing tube, and measuring

wheel, and are presented in micrometers unless

otherwise stated. Where possible, the range of

measurements 1s followed by the inean, in paren-

theses, Ulustrations were made with the aid of a

drawing tube,

Type specimens haye been returned to their

respective insiiiwions. Representative specimens of

* Division of Wildlife & Rangelamds Research, CSTRO,

BO. Box 84, Lyneham, A.CT, 2602.

7. nasalis lrom R, fuseipes in Australia have been

deposited in the British Museum (Natural History)

(B.M.(NH) No. 1981/3537-3540), the [stituto di

Parassitologia dell’ Universita di Roma, the

Muséum national d'Histoie naturelle, Paris (MN

476 HB), the South Australian Museum (V3244),

the Australian Melminthological Collection (13855),

the United States National Museum Helmintho-

logical Collection No, 77454 and the helminth

collection of the Division of Wildlife and

Ratigelands Research, CSIRO (N498, 717, 882),

Capillaria crocodiius (Ashford & Muller, 1978)

comb. nov,

FIGS !-7

Material examined: fram Crocodylus noavaegwineve

novwerninege Moitaka cracadile farm, Pon Mateshy, Id,

IY paratypes, BM(NH) coll. No. 197&/917-920.

Redeseription

Long thin nematodes with attemuated anterior

and blunt posterior extremities. Cuticle

exceptionally thick, cuticular striae not observed.

Iwo exceptionally broad lateral and one narrow

ventral bacillary bands, Hypodermal gland cells of

hands papilla-like in appearance due to necessity

fer neck of cell to traverse (hick cuticle to external

pore opening. Lateral alae absent, Cephalic

extremity minute, with minute dome possibly

consisting of two lips; stylet and buceal capsule not

observed, Oesophagus commencing as farrow

muscular tube, broadening posterior to nerve ring,

narrowing before reaching stichosome, passing

through stichosome on dorsal or lateral surface but

exiling trom it on ventral surface as narrow

muscular duct, Stichosome not extending as far as

Intestine. Narrow muscular oesophagus forming

unusual opaque junction with inlestine, consisung

of two rather thar three lissue segments and not

D. M. SPRATT

: e &

A : SNA

= — es

A —— hE Ano

a a Ae

Renna :

TRICHLRIDS OF VERTEBRATES 145

appearing as true valve. Two mescnchyinal cells not

present at oesophago-inteslinal sunchion,

Stichosome composed of approximately 36 stichn-

cytes, Much longer than Witle, large nucleus Wor

observed in each stichocyfe. Iilestie large and

conspicuous. ‘Tail broad and bhunt, without papillae

in female and without alae in male Anus and clones

terminal in female and male respectively. Male with

shor ejaculatory duct, long cloaca lined with

cutiele, lung robust non-sclerotised spicule, and

spicule sheath (=cirrus sense Anderson & Barn,

1982) with robust blunt spines or scales. Fentale with

vulva slightly posterior lo qeésephagorintestinal

jinction, vagina vera lined with cuncular spines oF

scales, Epgs moderately thick-shelled, umtanned,

shell without ornamentation, eaes iy discal uterus

not embryonared.

Male

Length 22.7 mm. Width at nerve ring 10, at

ovsophago-intestinal juncyion 37; maximum

width 66. Length of muscular oesophagus 30k total

length of oesophagus 13.5 mm. Length of

stichosome 13.2 mm} stichocytes 33 in number:

length of stichocytes 260-330. Nerve ring SO from

cephalic extremiry. Total length of cloaca 930; length

of spiny spicule sheath 150; length of cjaculatory

duct 100; spicule pouch entering clowet 213 from

caudal extremity, Spicule present, not sclerotisedt,

450 in length, & in maximum width, capicolan with

characteristic “bubble” shape, spicule broad

proximally, with broad but taperiig point distally,

Junction of intestine and eyiculatory duct simple,

both ducts same width at junction, Spicule sheath

with robust but generally blunt spines or scale-like

ornamentation (not everted jn this specitwen}.

Spicule retractor muscle iisertitig 150 anterior to

sphincter muscle delineating modification of distal

vas deferens into ejaculatory duct.

Lateral and ventral bucillary bards commencing

in region of nerve ring; lateral bands 2-3 cells wide,

cells regular in distribution; yeorral band one cel)

wide, cells irregular in distribution. Lateral bands

broadening rapidly, 1-143 cells in) width

approximately § mm posterior ta cephalic end and

practically encircling worm; bands narrowing

abruptly near tail tip, 7 cells in width ar level of

entry of spicules pouch lo cloaea. Ventral bacillary

band with single column of cells spaced irregularly

along body length,

Female

Length §7.5 mun. Width at verve ring 70, al vulva

113; maximum width 135. Leagih of muscular

oesophagus. 270; total length of oesophagus

19.2 mm. Length of stichosome 18.7 mm, sticho-

cytes 37 in number; length of stichocytes 300-580,

Nerve ring 60 from cephalic extremity. Vulva

2.2 mmm from anterior extremity, Vagina vera 109

tong, fined with thick cuticular scales similar in

morphology (0 those on male cirrus, Rectum 201

long, lined with thick cuticle, Anus terminal.

Lateral and ventral bacillary bands commencing

in region of nerve ring. Lateral bands 4—6 cells wide,

cells regular in disposition; ventral band one cell

wide, cells irregular in disposition. Lateral bands

broadening rapidly, 10«13 cells in width approxi-

mately 5 mm posterior to cephalic extremity,

peactivally epcircling worm; bands narrowing

abruptly stpproximately 0.5 mm from tail tip.

Yentral bacillary band with single column of cells

spaced irregularly along body length.

Discussion

My olservatioms differ markedly from those of

Ashford & Muller (1978), This trichurid nematode

species frou epidermal tunnels in the abdominal

shin of crocodiles possesses {i) conspicuous

bacillary bands comprised Of papilla-like hypo-

dermal! gland cells duc to the neck of the gland cell

having ta traverse the thick body cuticle to the

extemal pore opening, (ii) a stichosome comprised

of 32-37 elongate Michocytes arranged in a single

column, (ii) a long broad non-sclerotised spicule,

(iv) a Spieule sheath with robust blunt spines and

scale like ornamentation, (y) a long muscular cloaca

with thick cuticular lining and (vi) unembryonated

eggs in the distal uterus of the female, none of

which were reported by Ashford & Muller (1978).

The caption to their Fig. $ contradicts their state-

ment on p. 216, “Eggs unembryonated when laid”

and should read, “Egg from skin, containing &

larva”,

‘The-abovementioned features are characteristic

of the trichurid nematode subfamily Capillariinae

Figs 1-7, Copillavie erosodiive (Ashford & Muller, L978) comb. nov. 1. Male posterior end showing junction of intestine

and ejaculatory duct with distal Clo#es. 2, Male posterior end showing capitulum of spicile, spicule retractor and

protractor muscles, and cloaca. 3, Male caydal end showing cloaca, spiny spicule sheath and distal end of spicule.

4, Blum spines und scube- like ormamentaliot on spicute sheath of mate. +. Female anterior end showing oesophago-

intestiqal junction, gutva and vapina with seale-lihe ofmamentation, ventral view. 6. Female caudal end, ventral

view. 7, Lateral baculary band of fenmale stiowing neck of plird o¢fls rakersing thick cuticle to exterior pore openings.

Scale lines: Figs 4, 7, Wh arns Pip, 2) 20 wm; Ves 1,2, 5, 50 9m, Abbreviations: ci—spoule sheath (cirrus sensu

Anderson & Brain, 1982), cl—cloaca, ed—ejncularory duct, ® intestine, o—oesophagus, par—protractor muscle

of spirale, recut. rim—rerracton muscle of spicule spe=spicule, st—stichverts, va—vagina, vo—vulva.

146

(sensu Anderson & Bain, 1982) and differ from the

Trichosomoidinae to which Ashford & Muller

allocated their genus. In contrast, members of the

Trichosomoidinae possess a primitive oesophagus

with 60 to 150 stichocytes sometimes arranged in

two or three columns, males have a short cloaca but

lack both a spicule and a spicule sheath, and eggs

are embryonated when laid by female worms.

Anderson & Bain, 1982 placed Paratrichosoma

Ashford & Muller, 1978 as a synonym of Capillaria

(sensu latu) Zeder, 1800. The reasons for this

synonymy are presented above and the crocodile

parasite is formally recognised as Capillaria

crocodilus (Ashford & Muller, 1978) comb. nov.

Despite the foregoing, C. crocodilus possesses

several features which distinguish it from many

other members and which may warrant

consideration in any future comprehensive revision

of the genus Capillaria. These are as follows: (i)

exceptionally thick cuticle, (ii) exceptionally broad

lateral bacillary bands, (iii) anterior muscular

oesophagus with median swelling similar to that

occurring in first-stage larvae of the Metastrongy-

loidea, (iv) exceptionally long stichocytes, (v)

stichosome terminating well anterior to oesophago-

intestinal junction (vi) form of oesophago-intestinal

junction, (vii) absence of two mesenchymal cells at

oesophago-intestinal junction, (viii) absence of alae

or papillae on male and female caudal extremities,

(ix) vagina vera lined with cuticular scales similar

to those on spicule sheath of male and (x) insertion

of spicule retractor muscle well anterior to (rather

than at level of) sphincter muscle delineating

modification of distal vas deferens into ejaculatory

duct.

Ashford & Muller (1978) reported that two

crocodiles, C. novaeguineae novaeguineae and C.

porosus Schneider were susceptible to infection with

Capillaria crocodilus, that the parasite was rare or

absent in crocodiles from areas of saline water and

thal transmission of the parasite does not occur

under some rearing conditions on crocodile farms.

Undulating nematode worm trails, probably

attributable to C. crocodilus, have been reported

from C. acutus Cuvier (Garrick in Webb & Manolis,

1983), C. intermedius Graves (King & Brazaitis,

1971), C. johnstoni (King & Brazaitis, 1971; Webb

& Manolis, 1983) C. moreletii Duméril, Bibron and

D. M. SPRATT

Dumeril (King & Brazaitis, 1971), C niloticus

Laurenti (King & Brazaitis, 1971) and C. porosus

(King & Brazaitis, 1971; Webb & Messel, 1977).

In the Northern Territory of Australia worm trails

are more common in larger specimens of both C

Johnstoni and C. porosus (Webb & Messel, 1977;

Webb & Manolis, 1983). In Queensland, worm trails

occur in C. johnstoni 3 years of age and older, and

there is an increasing prevalence and density of trails

with age. (K. R. McDonald, pers. comm.).

Other species of Capillaria (sensu latu)

(Capillariinae) occur in the skin both of cold-

blooded and of warm-blooded vertebrate hosts

(Gallego & Mas-Coma, 1975; Moravec & Cosgrove,

1982; Wade, 1982). In addition, Lyne & Sommerville

(1965) reported a species of Capillaria in skin

sections of the lip and scrotum of the marsupial

bandicoot, Perameles nasuta Geoffroy in Australia.

I have recovered adults of species of Capillaria from

these sites in R nasuta and Perameles gunnii Gray,

and from epithelial tunnels in the tongue of R

gunnii, P- nasuta and the small dasyurid marsupials

Antechinus stuartii Macleay and A. swainsonii

(Waterhouse). The genera Anatrichosoma and

Trichosomoides (Trichosomoidinae) also contain

species which occur in tunnels or burrows in the

epithelial tissues of their mammalian hosts (see

Discussion in Spratt, 1982 and this paper).

Trichosomoides nasalis Biocca & Aurizi, 1961

FIGS 8-21

Specimens of Trichosomoides nasalis were found

in the mucosa of the distal nasal cavity of 5 of 232

Rattus fuscipes examined from forested coastal and

montane habitats (sea level to 1220 metres) in

southeastern New South Wales. The parasite was

not found in 77 R. lutreolus (Gray), 13 R. rattus

L., 97 Mus musculus L. and 8 Mastacomys fuscus

Thomas examined from these same regions.

Numbers of female nematodes recovered from

individual rats were 4—34 (X=12), those of males

living in the uteri of females 2-8 (x=5),

Material examined: from Rattus (=Epimys) norvegicus,

Rome, Italy, T. Crapulli, co-type 2 containing 2 co-type

do in utero, fragments 2 co-type ??, 1958, in Istituto di

Parassitologia dell’ Universita di Roma.

Figs 8-21. Trichosomoides nasalis Biocca & Aurizi, 1961 from Rattus fuscipes. 8. Anterior end, female, dorsal view.

9. Oesophago-intestinal junction and vulva of female, lateral view. 10. Caudal end of gravid female, lateral view.

11, Caudal end of non-gravid female, lateral view. 12. Cephalic end female, en face view. 13. Egg from distal vagina

of gravid female. 14. Egg adjacent to that in Fig. 13, opposite profile, 15. Anterior end male, lateral view of cuticular

ornamentation—pore openings of hypodermal gland cells interrupted by partial but conspicuous transverse striae.

16. Anterior end male, lateral view. 17, Oesophago-intestinal junction, male, lateral view. 18. Caudal end male,

lateral view. 19. Elevated hypodermal gland cells of bacillary band in posterior half of female, lateral view. 20.

Lateral bacillary band, mid-body region of female. 21. Lateral bacillary band 0.5 mm from caudal end of female.

Scale lines: Fig. 12, 10 wm; Figs 13, 14, 20 um; Figs 8, 10, 11, 13, 21, 50 pm; Fig. 9, 100 um.

TRICHURIDS OF VERTEBRATES

_—

—

148 D. M. SPRATT

From R. fuscipes, Lee’s Creek, A.C:T., P. Haycock and

D. M. Spratt, 1599, 66¢¢ , 22.i1.1979; Bundarra Creek,

Snowy Plains, N.SW., D. M. Spratt, 3499, 91d¢d,

22.11.1978; Emu Creek, Bondo State Forest, near Bondo

N.SW., P. Haycock and D. M. Spratt, 629, 21dd,

24.xi.1978; Nadgee State Forest, near Eden N.SW., P.

Haycock and D. M. Spratt, 799, 23¢¢ , 12.xii.1978; E.

Walter and P, Haycock, 599, 286d , 1.xii.1982.

Redescription

Female relatively long and narrow, with slightly

attenuated anterior and blunt posterior extremities.

Male minute, occurring in uterus and vagina of

female.

Female (measurements of 18 specimens)

Length 6.5-15.7 (11.9) mm. Width at nerve ring

30-34 (32), at oesophago-intestinal junction 60-100

(84); maximum width 90-200 (150). Cuticle thin,

with conspicuous, closely-spaced transverse striae

anteriorly, less conspicuous in midbody, ‘barely

discernible posteriorly. Cephalic end narrow bearing

single stylet, able to be retracted into minute buccal

capsule, Oral opening dorso-ventrally elongate, with

6 minute lips each bearing single papilla. Amphids

relatively large, papilliform. Oesophagus

commencing as narrow muscular tube, broadening

posterior to nerve ring, narrowing as it enters

stichosome, passing laterally through stichocytes.

Length of muscular oesophagus 120-200 (160); total

length of oesophagus 1.38-2.41 (1.80) mm. Length

of stichosome 1.20-2.21 (1.64) mm, Number of

stichocytes 69-84 (74), anterior stichocytes regularly

aligned in single column, posterior ones irregularly

aligned in 1, 2 or 3 columns; cells approximately

square, becoming more triangular in posterior half

of stichosome, all with single large nucleus. Nerve

ring 18-50 (26) from cephalic extremity.

Oesophago-intestinal junction conspicuous, with

two small mesenchymal cells, 1 dorsal, 1 ventral,

each with single large nucleus. Vulva 1.40-2.45

(1.83) mm from anterior extremity, with small lip

on each of anterior and posterior margins, with

exceptionally thin cuticular lining. Vagina vera

132-167 (150), muscular, with exceptionally thin

cuticular lining. Posterior intestine with narrow

lumen and relatively thick muscular walls in

immature females, with wide lumen and thin

muscular walls in gravid females. Rectum with thick

cuticular lining. Anus terminal, without lips or

swelling. Tail blunt, often twisted or distorted in

gravid females.

Lateral bacillary bands commencing in region of

nerve ring as column 1-2 cells wide, broadening to

3-4 cells width at posterior end of stichosome, cells

arranged irregularly. Bands becoming much wider

just anterior to or level with vulva, extremely wide

from here to posterior end and leaving only narrow

dorsal and ventral columns of cuticle unornamented

except for diminishing transverse striae. In this

region hypodermal gland cells projecting above level

of body cuticle, papilliform, with single pore

Opening at apex, most pronounced in posterior half

of body in gravid females. Lateral alae not observed.

Eggs in distal uterus 70-80 (75) long, 34-76 (40)

wide, variable in shape, thin in one profile thick in

opposite, plugs not protruding, shell dark amber

brown, smooth; eggs containing larvae; larvae

hatching in distal uterus in some specimens.

Females containing 2-8 (5) males, usually in

distal uterus or muscular vagina of non-gravid

females, usually in posterior uterus of gravid

females. One female with anterior half of male

protruding from vulva.

Male (measurements of 10 specimens)

Length 1.25-1.65 (1.38) mm. Width at nerve ring

22-32 (28), broadening in posterior half; maximum

width 40-70 (54). Cuticle with conspicuous

transverse striae throughout. Cephalic end with two

minute lateral papilliform structures, presumably

amphids. Buccal capsule minute, 4-6 (5) long, stylet

not observed. Oesophagus commencing as broad

muscular tube, narrowing posteriorly as it enters

stichosome, passing laterally through stichocytes.

Length of muscular oesophagus 60-110 (87); total

length of oesophagus 530-630 (582). Length of

stichosome 440-530 (493). Stichocytes as described

in female, cell walls often indistinct making

counting difficult; approximate number stichocytes

65-74. Nerve ring 25-34 (28) from cephalic

extremity. Oesophago-intestinal junction indistinct.

Intestine broad, thick-walled. Intestine and vas

deferens uniting to form short muscular cloaca,

70-80 (76), without cuticular lining anteriorly.

Cloaca modified distally into short thick-walled

cuticular duct without musculature, 30-37 (34).

Spicule absent. Spicule sheath absent. Tail blunt,

cloacal opening terminal.

Lateral bacillary bands similar to those in female,

not as broad, with fewer gland cells. Bands

commencing as column of single cells posterior to

buccal capsule, broadening to column 2 cells wide

posterior to nerve ring, continuing throughout body

length as column 2-3 cells wide. Gland cells

becoming papilliform near posterior end of

stichosome, projecting above level of body cuticle,

each with single pore opening at apex. Gland cell

openings separated or grouped in lots of 2 or 3 by

short conspicuous transverse striae.

Discussion

Members of the genus Trichosomoides are

parasites of the mucosal surfaces of murid rodents,

Only two species are known; T. crassicauda

TRICHURIDS OF VERTEBRATES 149

(Bellingham, 1865) Railliet, 1895 from the urinary

tract, generally the bladder, of wild and laboratory

rodents throughout the world, and 7. nasalis Biocca

& Aurizi, 1961 from the nasal cavity of R.

norvegicus in Rome. This species was recorded in

the wild for only the second time by Cross ef al.

(1970) who reported a prevalence of 4.2% in R.

exulans (Peale) in Central Java, Indonesia. Later,

Cross & Santana (1975) reported T: nasalis in 24%

of R. coxingi Swinhoe examined on Taiwan.

Number of worms per infected host ranged from

1-15 with a mean number of 4.5 nematodes per

animal. Bernard (1964) described T. gerbillis from

the stomach of Gerbillus pyramidum_ hirtipes

Lataste in Tunisia but this was later transferred to

Anatrichosoma (Pence & Little, 1972), Males of the

genus 7richosomoides are unique among parasitic

nematodes for their habit of dwelling in the uterus

and vagina of the female worm.

Morphological differences between 7. nasalis

from R. norvegicus in Rome and the specimens

from R. fuscipes in southeastern Australia are slight.

Biocca & Aurizi (1961) illustrated 59 stichocytes in

the stichosome of the female. A variable number

of stichocytes occur in male and female nematodes

from R. fuscipes and I observed 70 stichocytes in

a co-type female fragment (non-gravid) from R.

norvegicus. Measurements of morphological

features of both male and female 7) nasalis from

R. norvegicus reported by Biocca and Aurizi (1961)

and observed by me are slightly greater than those

of specimens from R. fuscipes. These differences

are viewed as insignificant, possibly host-induced

and certainly not warranting separate specific status

for the material from R. fuscipes in Australia, which

is here recognised as T. nasalis.

A characteristic feature of male and female 7:

nasalis is the form of the lateral bacillary bands,

particularly in the posterior half of the body. In this

region the hypodermal gland cells project above the

body cuticle, are papilliform or dome-shaped and

bear a single pore opening at their apex. This feature

was observed by Biocca & Aurizi (1961)—“...

papilla-like cuticular elevations nearly 10 »m in

diameter.”—and may be seen in their illustration

of the female nematode. These authors’ observation

of the conspicuous nature of the elevations on the

ventral surface of female worms is misleading. The

bacillary bands originate on the lateral surfaces but

become extremely wide posterior to the vulva,

leaving only narrow dorsal and ventral columns of

cuticle without gland cell pore openings, and thus

unornamented, except for weak transverse striae.

Spratt (1982) commented on the similarity in form

of the hypodermal gland cells in species of

Trichosomoides and three members of the genus

Anatrichosoma.

The records of 7. nasalis in R. exulans in

Indonesia, R. coxingi on Taiwan and now in

indigenous R. fuscipes in Australia add weight to

the suggestion that this nematode species may be

widely distributed throughout the world (Cross ef

al., 1970; Cross & Santana, 1975).

Acknowledgments

The author is indebted to Professor E. Biocca and

to Mrs E. A. Harris for loan of type material, to

Mr P. Haycock and Mrs E. Walter for collection

of specimens, to Drs I. Beveridge and J. Calaby and

Mrs E, Walter for constructive criticism of an earlier

draft of the manuscript, and to Mrs J. Rudd for

typing the manuscript.

References

ANDERSON, R. C. & BAIN, O. (1982) CIH Keys to the

Nematode Parasites of Vertebrates No. 9, Keys to genera

of the superfamilies Rhabditoidea, Dioctophymatoidea,

Trichinelloidea and Muspiceoidea. Commonwealth

Agricultural Bureaux, Farnham Royal, England, 26 pp.

ASHFORD, R. W. & MULLER, R. (1978) Paratrichosoma

crocodilus n. gen. n. sp. (Nematoda: Trichosomoididae)

from the skin of the New Guinea crocodile. J/.

Helminthol., 52, 215-20.

Aurizi, A. (1958) Su un nematode del genere

Trichosomoides parassita delle cavita nasali dei ratti di

fogna (Epimys norvegicus) di Roma. Nuovi. Ann. Ig.

Micr. 9, 264-266.

BERNARD, J. (1964) Trichosomoides gerbillis n. sp.

parasite stomacal d’une gerbille d’Afrique du Nord.

Arch, Inst, Past. Tunis, 41, 33-38.

Biocca, E. & Aurizi, A. (1961) On a new parasitic

nematode Trichosomoides nasalis n. sp., from the nasal

cavities of Epimys norvegicus: and considerations on

the family Trichosomoididae Yorke and Maplestone,

1926. J. Helmirtaol., R, T. Leiper Suppl., 5-8.

Cross, J. H & SANTANA, F. J, (1975) Trichosomoides

nasalis in the nasal chamber of Rattus coxinga on

Taiwan. Chin. J. Microbiol. 8, 183-184.

—., Kunoin, W. D., Jones, G. S., Gapa, A. M.,

JOSOSOEDARNO, S. & GUNAWAN, S. (1970)

Trichosomoides in the nasal cavity of Java rats. J.

Parasitol. 56, 566.

GALLEGO, J. & Mas-coma, S. (1975) Capillaria garfiai

n. sp. (Nematoda: Trichuridae), parasito de la mucosa

lingual del jabali, Sus scrufa Linnaeus, 1758

(Mammalia: Artiodactyla). Vie Milieu 25, 237-48.

Kina, F. W. & Brazaitis, P. (1971) Species identification

of commercial crocodilian skins. Zoologica 56, 15-70.

Lyne, A. G. & SOMMERVILLE, R. I. (1956) A nematode

parasite, Capillaria sp., in the skin of the marsupial

bandicoot, Perameles nasuta. Aust. J. Sci. 28, 205-6.

Moravec, FL. & CosGrove, G. E. (1982) Pseudo-

capillaroides xenopi gen. et sp. noy. from the skin of

the South African clawed frog, Xenopus laevis Daud.

(Nematoda: Capillariidae). Rev. Zool. afr. 96, 129-37.

150 D. M. SPRATT

Pence, D. B. & LitTLe, M. D. (1972) Anatrichosoma

buccalis sp. n. (Nematoda: Trichosomoididae) from the

buccal mucosa of the common opossum, Didelphis

marsupialis L. J. Parasitol. 58, 767-773.

Spratt, D. M. (1982) Anatrichosoma haycocki sp. n.

(Nematoda: Trichuridae) from the paracloacal glands

of Antechinus spp., with notes on Skrjabinocapillaria

Skarbilovitsch. Ann. Parasitol. 57, 63-71.

Wane, S. E. (1982) Capillaria xenopodis sp. n.

(Nematoda: Trichuroidea) from the epidermis of the

South African clawed frog (Xenopus laevis Daudin).

Proc. Helminthol. Soc. Wash. 49, 86-92.

Wess, G. J. W. & MESSEL, H. (1977) Abnormalities and

injuries in the estuarine crocodile, Crocodylus porosus.

Aust. Wildl. Res. 4, 311-19.

—— & MANOLIS, S. C. (1983) Crocodylus johnstoni in

the McKinlay River area, N.T. V. Abnormalities and

injuries. [bid., 10, 407-20.

AUSTRALIAN AND NEW ZEALAND SPECIES OF ELACHISTA AND

HALOTHRIX (ELACHISTACEAE, PHAEOPHYTA)

BY S. SKINNER

Summary

Three species of Elachista Duby, E. claytoniae sp. nov., E. australis J. Ag. (Syn. Portphillipia

australis (J. Ag.) and E. orbicularis (Ohta) Skinner and one species of Halothrix Reinke, H.

ephemeralis sp. nov. are described from southern Australia. E. australis and E. lindaueri Chapman

are described from New Zealand, and a key is given to the Australia and New Zealand taxa of

Elachista. The systematics of the genus Elachista are discussed.

AUSTRALIAN AND NEW ZEALAND SPECIES OF ELACHISTA AND HALOTHRIX

(ELACHISTACEAE, PHAEOPHYTA)

by S, SKINNER*

Summary

SKINNER, S (1945) Australian and New Zealand species of Elachisra and AHalothrix (Elachistaceae,

Phacophiyta). Trans. &. Soc. 8. Aust, 1094), 151-160, 29 November, L98s.

Three species of Flachista Duby, A. clayranie sp. nov., & australis J. Ag. (Syn, Portphillipia australis

(1, Ag.) Silva) and A. arbiewduris (Olitay Skinner and one species al Haloshriv Reinke, A ephemeralis sp

mov, are deseribed from southern Australia. E. australis and £. lindavert Chapinan are described from

New Zealand, und a key is given to the Australia und New Zealand taxa of Flachistd. The systematics of

the penus Bluchista ave discussed,

Key Worns: Elachista: E clavtoniae sp. nav, E australiss E orhieulariss FO lindaueri; Halothtix;

H. ephemeralis sp. nay, Pertohillipia: Elachistavcac; Chordariales; Phacophyta; southern Australia.

Introduction

The family Flachistaceae includes three genera

and about 30 species, and is Usually placed in the

order Chordariales. The absence of any hair-like

structures and the presence of determinate cortical

assimilatory filaments (“paraphyses’) in

combination with long, indeterminate agsinilatory

filaments clearly separate this family trom other

families in the order. The pulvinate thallus is

composed of a lilarnentous medullary system which

radiates from a basal layer and supports a cortex

of both determinate axsimilatory filaments and

reproductive structures, beyond which project the

long assinvilatory filaments, The cells of

assimilatory filaments of all three genera contain

a jarge central nucleus, numerous discoid

phacoplasts with many species having prominent

pyrenoids projeeing from the phaewplasts, and

some physodes, All are epiphytes,

The venera included in the family are Malarhrix

Reinke, epiphytic on seagrasses, Blachisia Duby

(including Symphoricacces Reinke and

Partphillipia Silva), epiphytic of larger alpae, and

Herpodiscus South (1974), epiphytic on Durvillaea

Elachiste species are found as epiphytes on various

Phacophyta and Rhodophyta. with one species (

zosterge Noda in Noda & Kitami 1971) oma manue

angiosperm. Plurilocular sporangia, which ure

produced by most species, are Uniseriate, filiform

and are borne on sporangiophores among the

cortical assumilators.

Haloririx difters from Elgehiste by having

iutercalary plurilocular sporangia Whieh develop, in

i sorus, in cells within the long assimilatory

filaments, It also shows relatively fewer phacoplasts

in the photosyntheti¢ cells, and, at least in 4.

*Departmem of Botany, University at Aclelnide, GPO,

Box 498, Adelaide, S. Aust, 5001

lumbricatis (Kiitz.) Reinke, a greater development

of chizoidal filaments buttressing the medulla—

both of these differences are ones of degree rather

(han contrast wilh Elachisia, The hosts af Halothrix

are usually marine anglosperms (Zostera,

Heterozostera and Phyllaspadix),

Elachista (= Portphillipia) australis J, Agardh

has been recorded from Victoria and Tasmania

{Wemersley 1967) and & orbicularis (Ohta) Skier

from Western Australia and South Australia

(Skiner 1983). The present study shows that there

is also one further species of Elachisia and at least

one of Halothrix in southern Australian waters. In

New Zealand, two species of Klachista have been

recorded, F, australis in the south, and the endemic

E. lindaueri Chapman,

A subanraretic species of &/aehista, E. antaretica

Skotesberg 1953, is also Included in the key beeause

of its recent discovery on Macquarie Island (Ricker

pers. comm,)

Key to the Genera

|. Plant pulvinate to spreading, epiphytic on larger algae;

colls of long assimilatory (laments will) ounerous

pavked phacoplusts; plirilocular sporangia, when

presenl, uniseriate and Formed on sporangiophores

arising from medidlary fluments and within the

long sssimilatary Thamenis with numerous but

scattered phacoplasts; pluriloealar sporangia

intercalary mm sor) on lony assimilatory filaments,

J, Agardh 1848: 7) ISRK2; 9.

Chapman 196): 16, Haye 1995 117, Harvey (846: 99,

Kiellman (897; 220. Kuekuck 1929) 2L figs l0-[4

Lindauer, Chapman & Aiken 1961 214. Rosenyvingc

19345: 19. Suuvagean 194m: 139. Skottsberg 1907: 54-

1053-537, Seedelins 191) 162. Takamatsu (dae 145

Yamada 19285 1).

152 S SKINNER

Syimphoricoceus Reinke (89) 17, Kuckuck 1929: 32,

Kjellinan 1897) 219.

Poriphillipta Silva 70: 9da.

Thallus epiphytic, pulvinale tu globose,

hemispherical of spreading, usually 2-20 mim high

and aggregating in considerable numbers on the

host.

Basal layer of small isodiametric¢ cells, forming

a plate on the host surface, often with short pegs

is extensions of the lower cell wall. Medulla a

filamentous cushion of usually elusely packed cells

supporting and completely sheathed by the cortex.

Medullary filaments of non-pigmented, cylindrical

to clongate-ovoid or subglobose wells with

branching from most vells. Rhizeidal filaments

develop from cells of the lower medulla in many

species, and two species (2. australis and &,

lindaver() form interweaving, transverse supporting

filaments [Klammernrgane (Kuckuck 1929, p. 199],

clampirons formed laterally from cells of lower

inedulla.

Cortex of short assimilatory filaments and

reproductive organs terminal to the medullary

filaments, Cortical assimilatory Tlilaments

determinate (associated with sporangiophores),

filiform Lo clayale, straight to arcuate, with 5-20

eylindrical to inflated cells, containing fewer

phaeoplasts than cells. of long assimilators. Long

ussimilatory filaments (sometimes referred (7 as

assimilatory haivs) indeferminate (except in EL

sledlaris Areschoug ‘and Gonodia marchentae

Setchell & Gardner), protruding well beyond the rest

of the thallus wih a meristem of short cylindrical

cells, ind above with usually elongate, cylindrical

pigmented cells with thick walls.

Uniloculat sporangia in (he cortex, on short

pedicels, on terminal medullary cells or from the

base of assimilatory filaments, ovoid or pyrtfarm,

with a terminal pore,

Plurjlocular sporangia Witiseriate, filiform. with

8-48 loculi, cither in specialized, corymbose lateral

branches from the outer medulla or laterally from

long assimilatory filaments.

Tipe species; Elachista seutulata (Smith) Aresch,

A genus of at least twenty species, found on a

Wide variety of hosts and known from temperate

and colder seas. Three species occur in Australia,

and two in New Zealand. Sexuality has never been

clearly demonstrated In the life evele, which, at least

in Some Species, appears to be direct.

Key (0 (ite species of Elachista in Australia and New

Zealand

!, Medullary Flamemts of one kind only, unitocular

sporangia present; plurilocular sporanuia, wher

present, with 8-16 locull Th rook rm cz

Medullary filaments of two kinds, unbranched

filaments which bear the long assimilmory filaments,

and subdichotaious Mlainents which bear the cortex;

unilacular sporangia absent, plusilocular sporangia

with 16-24(—48) loculi_..... Elachista orbleularis

2. Medulla compact, filaments branching from most

cells, no transverse supparting, Filaments presen

cells Of long assimilatory filaments at least as wide

“wy medullary celly oo... — "|

Medulla loose, with sparsely branched filaments

producing transverse supporting filaments below:

eclls Of long assimilaiory filaments narrower than

theditlary vells. —..-_- - . eae |

3. Bath wnilocular and plurilocular sporangia (aay be

Present on the one plant, plutilocular sporangia usually

more common; cortical pssiimilatary Silenents

cylindrical and plentiful, , Elachista clayloniae

Unilneutar sporangia omly present) cortical assimilatory

filaments clavate bur sparse —_ Blachista anterctica'

4 Cells of medullary Mlaments cylindrical, 7-10 pm

mm diameter; cortigal assimilarocy Cilansents tiliforn,

wavy, cells narrow, cylindrical and uniform

wn) Flachisra dustrais

Cells of medullary tlaments Ovaigd-cylindrical,

2030 um in diameter. cortival assimilatory

HHaments clivale and arcuate, upper cells larerally

ini (Taateal

me veecveeey Blachista lindaueri

'Elochista antarctica Skottsberg 1953 has recently been

collected from Macquarie Island (Ricker, pers, comm.)

and may occur On other subantarctic islands south of

Australia und New Zealand. Skottsbere (1953) considered

that this taxon did not produce cortical assimilators

Cortical assimilators are present but uncommon on

material from Mucquarie Island (Ricker MA-24,

BiLIS78; ADU AS53082),

a ee a a

Fig. |. A-K, Blachista elawaniae sp. nov, (ADU, AS50809, Holotype}. A. Habir, on trond Of Sargassum sp. B. Upper

sponmeia. D. Filament showing long assimilator and sec\iid cortical branches. EL, Upper medulla supporting cortes

and long asstmilatary filament Lower inedijila and base. G-E. Elachista australis. Ag. (ALL A34809), G. Habit,

on Viptiogihore. . Filament showing narrow (ausverse filaments from lower nodes, medullary and long assimilatory

filament dnd cortical branches With cor(\gal asvimilalors. [. Cortex with untocular sporangia among cortical

assimilators. <M Alaloririy ephemeratis sp, nov. (ADU, A32664, TYPE), 1. Habit on Mererezestera tasmtanica.

K 1&2. Filament showing upper medulla, cortical bianehes with woilocular sporangia, meristem and cells of long

assiilatory Hamernt with sorus of plurilocwluc sporangia, &. Individual cortics! assimilator. M. Medulla and base

of long assimilatory filament.

ELACHISTA AND HALOTHRIX IN AUSTRALASIA

153

154 S. SKINNER

Elachista claytoniae sp. nov.

FIG. 1 A-F

Thallus dark brown, epiphytic on the margins

and lamina of the lower leaves of Sargassum

(Fig. 1A), pulvinate, hemispherical, 0.5—5 mm high.

Basal layer (Fig. 1F) of irregular, isodiametric

cells 8-15 wm in diameter, each bearing one

medullary cell. Medullary filaments (Fig. 1 D—F)

closely packed, subdichotomous, cells pyriform,

below grading to subglobose above, non-pigmented,

20-45 gm in diameter, L/B 2.5-4. Cortical

assimilatory filaments (Fig. 1B) straight or slightly

curved, filiform, with 15-25 pigmented, cylindrical

cells 4-6 (-8) in diameter, L/B about (1-) 2-3,

borne terminally on branches of medullary

filaments together with reproductive structures.

Long assimilatory filaments (Fig. 1D) arising from

terminal medullary cells and extending to two or

three times the length of the rest of the thallus;

meristematic zone indistinct; assimilatory cells

cylindrical. 12-20 »m in diameter, L/B 1.5-2.5, with

numerous discoid phaeoplasts.

Unilocular sporangia ovoid to obpyriform,

23-28 wm in diameter, L/B 2-3, occurring among

the plurilocular sporangia (Fig. 1C). Stalked

spherical cells, among the young plurilocular

sporangia, have been observed in the isotype MELU

20520, and the holotype ADUAS0809, but zooid

masses were not observed in such organs (Fig. 1B),

and their function is unknown.

Plurilocular sporangia (Fig. 1D, E) are grouped

on corymbose branches on medullary filaments or

immediately below the meristematic region of long

assimilatory filaments, filiform, uniseriate, with

occasional oblique cross walls and 12-16(-—24)

loculi, 5-8 »m in diameter, opening by a terminal

pore, outer wall persistent.

Diagnosis; Thallus pulvinatus, 0.5-5 mm alt; fila

medullae conferta, cum cellulis aut pyriformibus

inferne aut subglobosis superne; fila assimilata

corticata aut stricta aut leviter arcuata, filiforma

cum 15-20 cellulis; fila assimilata longissima cum

cellulis teretibus, 15-20 »m diametro, L/B 1.5-2.5;

sporangia unilocularia ob pyriforma; sporangia

plurilocularia uniseriata, filiforma in sporangio-

phoribus corymbosis aut ex medulla superne aut

ex filis longissimis.

Holotype: ADU, AS50809 (Clayton, 21.ix.1970).

Isotype MELU, 20520.

Type locality; Ocean Beach, Sorrento, Victoria.

Distribution: Port Phillip Heads, Vic.

Hosts: Sargassum spp.

Specimens examined: Queenscliff, Vic. (Clayton,

6.ix. 1969; ADU, A50331); Sorrento, Vic. (Clayton,

4.ix.1971; MELU, 21207); Point Nepean, Vic.

(Clayton, 4N.1969; ADU, A50332).

The specific epithet, “claytoniae”, is chosen to

honour the collector Dr Margaret Clayton, and her

distinguished work on Phaeophyceae in southern

Australia.

Elachista mollis Takamatsu 1938a Fig. 5 is similar

in form to E. claytoniae in that the cortical branches

support the sporangiophores, but it has clavate

rather than filiform cortical assimilatory filaments.

E. globosa Takamatsu 1938a and E. zosterae Noda

& Kitami 1971 also are similar to E. secundata in

general morphology. E. globosa has globose cells

throughout the medulla, and E. zosterae is

somewhat like Halothrix species below the cortex.

Elachista australis J. Agardh 1882: 13.

De Toni 1895: 440. Guiler 1952: 78. Lucas 1909: 19;

1913: 58; 1929: 14. Philippia australis (J. Ag.) Kuckuck

ex Oltmanns 1922: 34, fig. 327. Kuckuck 1929: 19, fig.

8. (Non Philippia Klotzsch 1834: 354), Philippiella

australis (J. Ag.) Silva 1959: 63. Lindauer, Chapman

& Aiken 1961: 215, fig. 42. Womersley 1967: 229. (Non

Philippiella Spegazzini 1896: 566). Portphillipia

australis (J. Ag.) Silva 1970: 944,

FIG. 1, G-I

Thallus medium brown, epiphytic on upper

ramuli of species of Xiphophora (Fig. 1G),

compact, globose to pulvinate, 0.5-5.0 mm high.

Basal layer restricted in area, with a plate of small

isodiametric cells forming pegs into host surface

and supporting the medullary filaments. Medulla

filamentous (Fig. 1H), cells cylindrical, weakly

pigmented, 7-15 »m in diameter, L/B 5-8.

Branching of medullary filaments at intervals, with

five or more cells in each internode, and often two

laterals arising at each node; in lower medulla one

or more lateral filaments may form interweaving

transverse supporting filaments, while in the upper

medulla lateral branches usually give rise to cortical

assimilatory filaments and associated unilocular

sporangia, with the main axis of the medullary

filament continuing as a long assimilatory filament,

Cells of transverse supporting filaments 4.5-5.5 »«m

in diameter, L/B 5-7.

Cortical assimilatory filaments (Fig. 1 H—I)

flexible, sinuous, of 25-30 cells at maturity, 4-7 «4m

in diameter, L/B 3-7. Long assimilatory filaments

broader and extending beyond the thallus by 2-5

times; cells short with numerous small phaeoplasts,

10-12 um in diameter, L/B 1-2.

Unilocular sporangia (Fig. 11) borne on a one

celled pedicel or sessile on lower cells of cortex,

ovoid to cylindrical with a terminal pore, 45-60 »m

long and 20-50 «m in diameter.

Plurilocular sporangia unknown.

Type locality: Port Phillip Heads, Victoria (Harvey,

Alg. Aust. Exsicc. 101 as Leathesia sp. nov.?).

Type: Herb. Agardh, LD (45972),

GLACHISTA AND HALOTHRLY IN AUSTRALASIA iss

DistriPutions Mpallo Bay to Port Phillip Heads,

Victoria, the north and east coasts of Tasmania, and

Stewart [,, New Zealand,

Host range: Niphophora chondrophylla and X.

gladiata.

Specimens examined; Port Phillip Fleads, Vic (Marvey,

Alg. Aust, Exsice, 101; ADL, APRSS1); Apollo Bay, Vic.

(Homersley, 10.x11.1969; ADT, A34d809): Point Lonsdale,

Vie (Skinner, 47,1978; ADU, AS9067% Lilverstone, ‘Tas,

(Gordan, 1851966; ADU, A30107); Bellerive, Derwent R.,

Tas. (Radway, Qet. 1909; ADU, ASSIS): Boat Harbour

Point, S ‘las. (Horiersiev, 16.x1982; ADU, ASSTSD);

Tessetated Pavenients, Eaglehawk Neck, tis (Parsons &

Womersley WAAHT, ADU, ASIVKE CHR, 399502);

Ringaringa, Stewart 1, New Zealand { Womersley, 41,1966,

ADU, A29SI0),

Clachisre-australis and &. lindduert Chapman are

distinctive species, Both show a loose and extensive

medulla with transverse supporting filaments and

the development of a subeortex by concentration

of later] branches in the upper medulla to support

the cortical assimilators. & Jindaueri (see below)

has larger cells Chroughout; the medullary cells are

inflated rather than cylindrical as in & euyrealis,

avd the vortival assimilators are arcuate rather than

sinuotis, Wiih the upper ceils ivilated laterally, The:

presence of the Lransverse supporting filaments was

used by Kuckuck (Kuckuck in Oltmanns 1922,

Kuckuck 1929) (o distinguish the genus Pii/ippue

from Llachista, There are several other species of

Alachisia e.g. &. mollis, E secundata and & globosa

Takamatsu 1[998a, which show a coucentration of

branching in the upper medulla, although one of

these taxa prodiice transverse supporting filaments,

The relentian of the genus Portpiillipia Silva (=

Philippia Karckuck non Kbotzsch) with the inclusion

ol &, australis and E, tindaveri, does nol seem

warranted, and # australis is here returned to

Elachista,

Flachista Jindawer! Chapanan Wel; 6, tig. 2.

Lindaucr, Chapman and Alken $961; 215.

HIG, 2 A-1D

‘Thallus yellow-brown, globose to pulvinate,

epiphytic on Viphaphare clondrapkylla,

Basal layer contravted to a very smal! disk.

Medullary filaments (Pig, 24, H) very lone and

sparsely branched, cells inflated-cylindmeal, 20-30

envin diameter, L/B 2-3; lower laterals forming

thizoid-like, hranghed, transverse supporting

filaments with often more than one from (he same

node! |~3 Upper branches (Pig. 2A, C, DO) from one

node, forming a subsorte, supporting Lhe cortical

assimtlators wnd unilocular sporangra Cortical

assimilatory filaments sligtily elavate, arcuate, che

upper cells literally expanded and deli. Long

ussimilatory [anvents (hig. 2A) usually terminal

io the main axis of a medullary flamer, will) a

meristem at or just above the level of the cortex;

mature cells cylindrical, 12-20 pm in diameter, L/B

I-1,5,

Unilocular sporangia sessile ovoid to pyriforen,

30-50 am in length and (2-18 »m in diameter, L./B

3-6, borne laterally on lower cortical cells,

Plurilocular sporangia unknown?

Tepe Jocality: Bay of Islands, New Zealand

(Lindauer, 6,xi.1937).

Tipe: AKU (Herb, Lindauer No, 29).

Spevimen examined: lsotype (ADU, A4573) and

Lang Beach, Russel, New Zealand, lower culittoral

pools (Warersley, 28.%,1969% ADL, A35069),

Disiribution; throughout New Zealand.

Chapman's (1961) description makes no mention

of the transverse supporting filaments, and is

incomplete in the description of branching in the

medulla) His deseription of “plurilocular

gametaugia fortied by modilicauon of ends of

assimilarars", without a figure, is unsubstantiated

In the specimens examined,

Flachiste orbicvlaris (Olua) Skinner 1983; 98,

Pips 1-3.

Genodia urbicularis Ota 1973: 21, Cie 1). Noda 1975:

tu Nig, 12.

No further dala have been obtained to add to the

species descriplion af Skinner (1983),

Tepe Locality: Tappi, Aomori Pret, Japan (Ohta,

11.1970),

Type (Ohia 133) in Herbarium, Niigata University.

Distributions Japan, and southern Australia (from

Rattnest Island, W. Aust, to Port Hacking, New

South Wales), This species is present on its hosr

throughout the year.

Hosts: Unduria species in Japan, Evklonia radiata

(C. Avardh) J, Agardh im Australia.

The two most closely related taxa to Elachista

orbiculuns are &. nigra ‘Wakamatsu 1938a, and

Gonodia fuxifermis Noda in Honda 1969 nom.

nud, (a species of Elachista) bath endenite (o Japan,

While (he other species show the formation of some

laterals from those medullary filaments which

support the long assimildtery filaments, &

orbieularis develops such laterals ouoly very

ovcasionally, and then from the lower-most cells,

Genus HALOTHREN Reinke [884: 49,

Hamel 1995; 126. Kuckuck 1929: A&, figs 15-17.

Rosenvinge 1935: 37. Takumutsd 1948b: ISL Yamada 92H:

313. Verde 1909; 123

Thallus pulvinate, hemispherneal, loasely

compacted, up to 2 en high, epiphytic on

Zosteracede. lodividual plants may be derisely

clustered, giving the appearance of one plant.

Basal layer a central disc of isodianietric ells

With, al the margins, a rhizoidal system from small

isodiametic cells in the lowermost medulla,

ELACHISTA AND HALUTHRIX IN AUSTRALASIA al

Medullary filaments of pyritorm or subglobose non-

pigmented cells, With branching conlined mainly to

upper vells. Cortical assimilarary filaments clavate,

arcuate or straigl, of a lew ovoid or eylindncal cells,

formed immediately below the meristematic zone of

the log assimilatory filaments, on Iaferal medullary

cells. Lony assimilarory filaments narrower hasally,

extending in lenath many times beyond the rest of

the thallus, with short cells (L/B about U with

Numerous scalfered phaeoplasis,

Unilocular sporangia arising [ram = upper

medullary cells, ovoid-eylindrieal or ureeolate, with

a terminal pore and plug.

Plurilocular sporangia in sort on anticlinally

divided evoups of cells 1 upper paris of long

assiimibatory Caments

pe species: H. lumbricalis (Kine) Reinke.

A venus of five described species, A. lumtbricalis

from the north Atlantic and Baltic, ahd four species

from Japan, An additional new species is here

described from southern Australia. The principal

difference between this venus and &achisto is the

speciilized plurilocular sporangia on the surface of

the long assivulatory Hlamens in Aalosiriv, While

i. slellaris, forms pluciloeular organs on fong

assimilatoey Filaments, they do not involve the prior

subdivision of the whole mother cell as in Mulethrix,

Pedersen (1974) fas shown thal Halothric lembricalis

his a simple direct lite cycle bn a recent paper

Pedersen (1984) has sugpested the inclusion of

Halothrix in the Giraudiacesxe on the basis al

similarities in morphogenesis, However, the

concemration of branches in the medulla

immediately below the meristem in Halarhriv is also

seen in various species of Elachisia, nolably in E

mollis, [. claytoniae, FE globosa and &. zasterae.

More importantly the development of the

photosynthedc part of the Gtraweia thallus, above

the meristem, invelves an orderly sequence al

periclinal divisions of (he cells of each whorl or rer

(Skinner & Womersley, 1984), which i not seen if

Heloihriy. Myriotrichia species show possibly closer

parallels to Giravelia, as the axis of the thallus in that

menus widensues periclinal division of assimilatory

cells belore the production of unilocular and

pluriloculur sporangia (Kuckick, 1899),

Halotheix ephemeralis sp. nov.

HG. 1 J-M

Thallus brown, epiphytic on Mererozastern

tasmanica teal blades (Fig, 11), offen so nuimeratis

as to cover Taost of the blades: pulvinale, loosely

compacted, 5-15 maim, high,

Basal plate of small isodiametric cells (0-13 aim

in diameter, adnate to host surface. Medullary

filaments (Fig. 1 -M) with 3 ar 4 lower tapering

cells 20-25 py in diameter, L/B 25-4, infrequently

branching, and 2 Lo 4 upper subglobose cells 20-30

em in diameter L/B about J, branching

subdichatonously, Cortical assinvilatory filuments

straight, Of 8-15 glightly inflated cells, eradine

upwards fron cvlindrival to subglobose, 10-15 jm

in diameter, L/B I-LS (Pig. 1 Ke, L). Long

assinilatory filaments (Fig, Kiyo, M} many fimes

longer than the rest of the thallus, cells increasing

in diameter upwards, shart but broad, upper

photosynthetic cells eylindrical, 30-30 am in

diameter, L/B 05-2

Unilocular sporingia (Fig. 1K.) cylindrical, with

a terminal damed plug and pore and tapering

basally, 15-30 ym in diameter, usually paired on

lateral medullary branches arising immediately

below the meristem ola tong assimilatory filament

together with vertical axsimilatory filaments,

maturing sequentially,

Plurtlocular sperangia (Fig. 1K,) in iutercalary

son. barne on 2-6 madified cells of tong

assimilatory filaments, the soral mother cell

undergoing Ewo or three anticlinal divisions, with

individual sporangia (some with a basal cell) arising

from periclinal divisions of these secondary cells;

often present on plants before unilocular sporangra

develop.

Diagnosis: Thallus usque ad Ha/othrix ambieua,

sed filis assimilatis corticais suictis Fillformibusque

nec arcuatis cunt cellolis inflars, uque sporangiis

uniloculariis pyriformibus, sporangus plurilowlaris

intercalaribus ab cellitis lili longissimi,

Thpe Jocalitv: Aldinga rect, S. Aust, in shallow

pools and gutters especially near the southern

seaward margin of the platform,

Ficlaivpe; ADU, 32664 (Womersley, 29 viii.1968).

Distribution: Only known from rype locality

(Aldinga reel, 8, Aust) (Skinner 14.18.1977; ADL,

A48249 and Ad825); 18_i0.1978: ADU, Ad9553).

There are two speeumens, MELU, 21072 and 21143,

on Hererezesiera from localities in the Port Phillip

area of Victoria, which are labelled NM. /umhricalis,

but the plants are very small, mov very fertile and

Hot idencifable hevond wenus,

Host: Helerazslera tasmanica

Specimens examined; As above,

The macrothallial stage is present fora very short

season, hence the specific epithet, IL has been found

only during August and September, fora period of

less rhan six weeks. This species dillers from éd.

Fig. 2 Floehore Jide’ Chapman (ADU, A35069), A to D all part of one filamentous system. A Lipper medulla

WIM Cortes and itil Jung assimiaior B. Lower medulis with transverse Hlaments C\D. Medullary and corlical side-

branch systenys

158 S. SKINNER

ambigua Yamada (based on a specimen from

Muroran, Hokkaido, Japan (Kurogi, 8.v.1978; ADU,

A49376) by possessing straight, not curved, slightly

medianly inflated cortical filaments, and an almost

cylindrical, not ovoid, unilocular sporangium. The

differences between this species and H. /umbricalis

are the possession of unilocular sporangia, the

absence of inflated cells towards the top of the

cortical filaments, and the discrete basal layer rather

than a rhizoidal system. H. ephemeralis is also

similar to H. tortuosa Takamatsu, which also has

straight cortical assimilatory filaments but a less

clearly defined branch system at the cortical-

medullary boundary than H. ephemeralis.

Discussion

The southern Australian and New Zealand

regions have four species of Elachista and one of

Halothrix. One other species, E. antarctica, is

confined to subantarctic islands such as Macquarie

Island. &. claytoniae from Port Phillip Heads,

Victoria, and the single species of Halothrix, H.

emphemeralis from Aldinga reef in South Australia,

appear to have restricted distributions, although the

plants may be quite common within that

distribution for short periods of the year. Elachista

australis, although sometimes locally common in

the summer months, is not often collected, and does

not appear to be as widespread as its two host

plants, Xiphophora chondrophylla and X. gladiata,

the former confined to south eastern Australia and

the latter to Tasmania.

On the other hand, Elachista orbicularis has a

wide distribution in Australia—from Rottnest

Island in W.A. to Sydney in N.S W.—and occurs at

most times of the year, usually in large numbers on

the host. As the first record of this alga in Australia

came from Port Noarlunga, S.A. in 1975, it is quite

possible that the species is a recent introduction

from Japan. E. orbicularis is sometimes associated

with epiphytic ectocarpoids which are frequently

seen on senescent plants of Eck/onia radiata, and

may thus have been overlooked. The recorded

localities are rather disjunct; there are no records

for Victoria, and although the host is usually found

as a major component of the sublittoral on rocky’

coasts throughout South Australia, plants of

Ecklonia radiata infected with Elachista orbicularis

have not been found in South Australia outside of

the rather narrow range of Gulf St Vincent and

Encounter Bay.

Portphillipia is here synonymised with Elachista.

While £. australis and E. lindaueri are clearly highly

specialized members of the genus, the production

of transverse supporting filaments hardly justifies

the separation of the two species into a distinct

genus. These two species are the most structurally

evolved of one developmental line within Elachista.

No such generic separation has been suggested for

the other divergent group of species which includes

FE. intermedia Crouan & Crouan, FE. nigra and E.

orbicularis, although they also show specialization

of form and function of cells and filaments of the

medullary tissues.

Interspecific relationships in Elachista

On the basis of relative complexity of vegetative

morphology and characteristics of sporangia, the

genus Elachista may be separated into three

sections. The comparison of species presented here

has been made largely on published descriptions

and illustrations together with examination of

herbarium material of a few of the species.

As will be seen from the lists of taxa included

in the sections of the genus, no attempt is made to

change the nomenclature of taxa not recorded in

southern Australia. The separation of Elachista

Duby (which produces two kinds of assimilatory

filaments) from Myriactula Kuntze (= Myriactis

sensu Kiitzing, = Gonodia Nieuwland 1917), which

produces one determinate kind of assimilatory

filament and indeterminate phaeophycean hairs,

dates back to Kjellman (1890). Pedersen (1979)

interpreted the hair-like filaments produced by

crowded plants of E. /ubrica as modified long

assimilators, not as hairs as Jaasund (1960) had in

separating E. /ubrica from Elachista and including

it in Myriactula. Many of the northern Pacific taxa

which have been included in Elachista, or excluded

from it, on the basis of descriptions and illustrations

noting presence or absence of hairs, should have

their nomenclatural status reviewed in the light of

the two opinions as to the nature of those hairs.

Section J; Plants partly endophytic, with a basal

system of rhizoids, penetrating beyond the host

epidermis; medulla compact, filaments diffusely

branched; cortical assimilatory filaments absent,

but long assimilatory filaments present; unilocular

sporangia present; plurilocular sporangia both in

corymbose groups at the base of long assimilators

and laterally from cells of long assimilators.

Taxa included: Elachista stellaris Areschoug.

Wanders ef al. 1972.

Section 2; Plants epiphytic, with discrete basal layer

often with peg-like projections from basal cells and

rhizoids from lower medullary cells; medulla

filamentous, with one kind of filament, branching

irregular; cortical assimilators (paraphyses) present

in most species; unilocular sporangia usually

present; plurilocular sporangia, when present, in

corymbose groups in the cortex, individual

sporangia with (6-)8-16 loculi (E. scutulata may

have 16-24 loculi), In E. /ubrica Ruprecht,

ELACIUSTA AND HALULHRIX IN AUSTRALASIA 180

plurilocular sporangia may also form laterally to

cells of long assimilators (allhough the origin and

tuncoon of these structures is interpreted otherwise

by Pedersen (1979)). Pedersen (1984) provides strong

evidence of conspecilily of &. /ubrica and E.

Jucicola. There are 1wo subsections,

Subsection 2a: Medullary filaments close packed,

branching from most cells, or with shor! internodes

of 10 or Fewer vells, with concentration of branching

in upper medullary filaments (to Form a subvortex)

in Some species} plucilocular sporangia present in

many species. Taxa included: Gonodia marchaniae

Setehell & Gardner 1924; LJuchistu covcapharae

Takamatsu 1938a, Noda 1969; &, antarctice

Skottsber, 1953; £. /ubrica Ruprecht, Kuckuck

1929; & fucivolu (Velley) Areschoug, Hauck 1885;

&, meridionalis Skotisberg 1907; EB. seutilare

(Smith) Areschaug, Kjellmanp 1897, Hamel 1935; £

flaccida (Dillwyn) Areschoug, Hamel 1935,

Takamatsu 1938a; £. glubosa Takamatsu 1998,

Noda 1969; E. clavioniae Skinner sp. nov., Garadia

jobnstonii Setchell & Gardner 1924; £. zasterae

Nodain Noda & Kitami 1974; EB me//is Takamatsu

1938a.

Subsection 2h: Medullary filaments loose,

branching at iniervals of (S—)10—15 cells, with upper

branches forming a subcortex to support the cortical

assimilators and unilocular sporangia, and lower

branches of the medullary filaments modified to

interweaving (ransverse supporting filaments:

plurilocular sporangia unknown,

Tae included: Elachisia lindaueri Chapman (96;

#. australis J. Agardh 1882, Kuckuek 1929,

Section 3: Plants epiphytic with discrete basal layer

often with peglike projections from basal cells and

rhizoids from lower medullary laments; two

morphologically distinct Kinds of medullary

filaments, One with broader cylindrical to ovoid

cells, supporting the long assimilatory filaments,

and the other of narrower clongate-ovoid cells,

branching throughoub supporting fhe cortical

ussimilatory filaments and reproductive organs;

unilocular sporangia present jn Some species;

plurilocular sporangia im corymbose graups,

individual sporangia with (16-24-48 local, Me

second kind of medullary filament may arise

laterally from the firs) kind of medullary filament

or separately from ihe basal cells.

Taxa included: Elachista nigra Takamatsu 1934a; E-

intermedia Crouan & Crovuan (= £&. neplecta

Kuckuck 1929) Sauvageau 1936; &. fenvis Yamada

1928, Takamatsu 19388 (incl, f pacifiea), Noda

1969, 1974; Mvriacsula sarzassi (Yendo) Feldmann

1943, Noda 19642; &, orbicilaris (Ohta) Skinner

1983, Noda 1975, Ohia 1973; Gonaeia Jusiformis

Noda in Honda 1969, nom- nud-

Taxa excluded from Elachista

Elachista ramoso Skotisherg 1907 = Gononeme

ramase (Skortsberg) Skotisberg (92; &, masarioides

Skousberg 1921 = Mvriacnila rosarioides

(Skortsberg) Skottsberg 1953; & pusilla Skortsberg

1921 = Myriaciula pusilla (Skottsberg) Skottsberg

1953. The following six taxa are described or figured

as having true phacophyvean hairs and should thus

be |ucluded in genera in the Learhesiaceac: &.

minulissina Taylor 928, 1960, E crassa Takamatsu

1998a; &. taentaefarmis Yarnada 1928; & vellasa

Takamatsu 1938a; & sadoensis Noda 1969; £,

sargussicola Noda 969.

Acknowledgments

Much of the present work Was Undertaken as part

of the author’s doctoral research, supported by a

University Research Gram at the Universiiy of

Adelaide. The work was completed while Lhe author

was a research associate with Prof. HW, AS.

Womersley with support from (he Marine Sciences

and Technologies Grants Scheme, Much thanks are

due to Dr M, Kurogi of Mokkaido Liniversity,

Japan, Dr M,N, Clayton of Monash Lniversity,

Victoria, and Dr R. J. King of the University of New

South Wales for the loan of specimens. Partjoular

thanks are duc to Prof. H. B.S. Womiersley, ancl

also Dr M, J. Parsons of DSR, Christchurch,

New Zealand, for reading and discussion of the

Mmapuseript

2¥endo (1420) describes this axon as having occasianal

hairs, bul Noda’s (1964) description and ilhastrarions are

without hairs, There may be two (axa involved here

References

Acai, 1 G. (848) “Species, genera ef ordines

Alyarum.” Vol. 1, (Cileerup: Lured.)

(1882) Till algernes systematik, Mya tidrag. Aleta

Univ, Land 17, 1136, pls 1-3

Cuarmas, V1. (1960) New entilies and combitanans tn

ihe Phacophycene of New Zealand. Bull, Res, Cour

Israel, sect 13, WD, 18-28,

Or Ton, | B. (1895) “Sylloge Alparutn onium

Hhucusque cognirarum.” Vol. 3 (Padua.)

Duy, 1. E. (1830) “Botaincon palliciuim” tf) (Pars |

Froonann, J (6943) Une mouvelic espéoe de Miadae tite

parasiie diy Greeilaria armate I. Ag. Moll, Sac, Hist,

Nat IAfriew de Nord 34, 222-229,

160 S. SKINNER

GUILER, E. R, (1952) The marine algae of Tasmania.

Checklist with localities. Pap. Proc. R. Soc. Tas, 86,

71-106.

HamEL, G. (1935) “Phéophycées de France” II, 81-176,

(Paris.)

Harvey, W. H. (1846) “Phycologia Britannica” Vol. I.

(Reeve: London.)

Hauck, F. (1885) Die Meeresalgen. Vol. II. Jn L.

Rabenhorse, “Kryptogamen—Flora von Deutschland,

Oesterreich und der Schweiz”. (Kummer: Leipzig.)

Honpa, S, (1969) On the marine algae of Toyama Bay in

the Japan Sea. Bull. J. Soc, Phycol, 17(3), 104-108.

JAASUND, E. (1960) Elachista lubrica Ruprecht and

Elachista fucicola (Velley) Areschoug. Bot. Mar. 1,

101-107.

KJELLMAN, F. R. (1890) “Handbook i Skandinaviens

hafsalgflora’, Vol. I. Fucoideae. (O. L. Lamms:

Stockholm.)

(1897) Phaeophyceae. Jn A. Engler & K. Prantl

“Die Naturlichen Pflanzen familien” 1(2). (Englemann:

Leipzig.)

Kuorzscu, J. F. (1834) Ericearum cel. Adelberto de

Chamisso descriptarum. Pars Addita. Linnaea 9,

350-367.

Kuckuck, P. (1899) Beitrage zur Kenntnis der

Meeresalgen, 5-9 Wiss. Meeresutersuch., Abt,

Helgoland, N.F, 3(\): 13-82.

(1929) Fragmente einer Monographie der

Phaeosporeen, Wiss. meeresunters. N.F. 17(4), 1-93.

LINDAUER, V. W., CHAPMAN, V. J. & AIKEN, M, (1961)

The marine Algae of New Zealand, I]: Phaeophyceae.

Nova Hedw, 3, 129-350, pls 57-97.

Lucas, A. H. S. (1909) Revised list of the Fucoideae and

Florideae of Australia. Proc. Linn. Soc. N.S.W. 34 9-60.

(1913) Notes on Australian marine algae, I. Ibid,

38, 49-60 pls 1-5.

(1929) The marine algae of Tasmania. Pap. Proc.

R. Soc. Tas. 62, 6-27.

NIEUWLAND, J, A, (1917) Critical notes on new and old

genera of Plants, IX. Amer, Midland Nat. 5, 30.

Nopa, M. (1964) Marine algae in the vicinity of the

Shioyazaki Cape, Fukushima Prefecture. J. Fac. Sci.

Niigata Univ. ser. 2, 4, 33-75.

(1969) The species of Phaeophyta from Sado

Island in the Japan Sea. /bid. ser. D, 6, 1-64.

(1974) Some marine algae collected on the coast

of Kashiwazaki Province facing the Japan Sea. [bid,

ser. D, 11, 65-74.

(1975) Some marine algae further added to the

marine flora of Sado Island in the Japan Sea. Jbid,

ser. D, 12, 91-107.

& Kitami, T. (1971) Some species of Marine

Algae from Echigo Province facing the Japan Sea. Ibid,

ser. D, 8, 35-52.

OutTA, T. (1973) Some new and rare marine algae from

Tsugara straits between Honshii and Hokkaido. /bid.

ser. D, 10, 11-28.

OLTMANNS, F. (1922) “Morphologie und Biologie der

Algen”. Vol. Il. Phaeophyceae-Rhodophyceae. (Fischer:

Jena.)

PEDERSEN, P. M. (1979) Culture studies on the brown

algae Holothrix lumbricalis and Elachista fucicola

(Elachistaceae). Bot, Not. 132, 151-159.

(1984) Studies on primitive brown algae

(Fucophyceae). Opera Bot, 74, 5-76.

REINKE, J. (1889) Algenflora der westlichen Ostsee

deutscher Anthiels. Bericht Komm. wiss Unters. deutsh.

Meere Kiel. 6.

ROSENVINGE, L. K. (1935) On some Danish

Phaeophyceae. Mem. Acad. R. Sci. Lett. Dan.

Copenhague, Sect. Sci. 9, 6(3), 40 pp.

SAUVAGEAU, C, (1936) Second Mémoire sur les Alques

Phésporées. Bull. Stat. Biol. d’Arcachon 33, 1-204.

SETCHELL, W. A. & GARDNER, N. L, (1924) Phycological

Contributions. VII. Univ. Calif. Publ. Bot. 13, 1-13.

Sitva, P. C, (1959) Remarks on algal nomenclature, II.

Taxon 8, 60-64.

(1970) Remarks on algal nomenclature, IV.

Tbid.94]1 19, 941-945,

SKINNER, S. (1983) The life-history of Elachista

orbicularis (Ohta) comb, nov. (Elachistaceae,

Phaeophyta) in southern Australia. Br. phycol. J. 18,

97-104.

& WomersLey, H, B. S. (1984) Southern

Australian taxa of Giraudiaceae (Dictyosiphonales,

Phaeophyta). Phycologia 23(2), 161-181.

SKOTTSBERG, C. (1907) Zur Kenntnis der Subantark-

tischen und Antarktischen Meeresalgen, 1.

Phaeophyceen. pp. 1-172, pls 1-10. Jn Wissenschaftliche

Ergebnisse der Schwedischen Siidpolar-expedition

1901-1903. Vol. 4,

(1921) Botanische ergebnisse der Schwedischen

Expedition nach Patagonien und dem Feuerlande

1907-1909. VIII Marine Algae. I. Phaeophyceae. K.

svenska Vetensk-Acad Handi. 61(11), 1-56.

(1953) On two collections of Antarctic marine

algae. Arkiv for Botanik ser. 2, 2(7), 531-566,

SouTH, G, R. (1974) Herpodiscus gen. noy. and

Herpodiscus durvilleae (Lindauer) comb. nov., a

parasite of Durvillea antarctica (Chamisso) Hariot

endemic to New Zealand. .. R. Soc. N.Z. 4(4), 455-461.

SPEGAZZINI, C. (1897) Plantae Patagoniae australis. Rev.

Fac. Agron. y Vet. La Plata 3, 485-589.

SVEDELIUS, N. (1911) Elachistaceae, p. 162. In A. Engler

& K. Prantl “Die natirlichen Pflanzenfamilien nebst

ihren Gattungen und wichtigeren Arten” 1(2).

(Engelmann: Leipzig.)

TAKAMATSU, M. (1938a) Elachista aus dem nordéstlichen

Honshii, Japan. Saito Ho-on Kai Museum, Res. Bull.

14, 145-176.

(1938b) Halothrix aus dem nordoéstlichen Honshi,

Japan. Ibid. 14, 181-192,

Tay or, W. R. (1928) The marine algae of Florida, with

special reference to the Dry Tortugas. Carnegie Inst.

Washington Publication 379, 1-219.

(1960) “Marine algae of the Eastern Tropical and

Subtropical coasts of the Americas”. (University of

Michigan Press: Ann Arbor.)

Wanpers, J. B. W., Hoek, C. VAN DEN, & SCHILLERN-

VAN Nes, E. N. (1972) Observations on the life-history

of Elachista stellaris (Phaeophyceae) in culture. Neth.

J. Sea. Res. 5(4), 458-491,

WomersLeyY, H. B. S, (1967) A critical survey of the

marine algae of southern Australia. II. Phaeophyta.

Aust. J. Bot. 15, 189-270.

YAMADA, Y. (1928) Report of the Biological Survey of

Mutsu Bay, 9. Marine algae of Mutsu Bay and adjacent

waters, II. Sci. Rep. Tohoku Imp. Univ. ser. 4, 3(4),

497-534.

YENDO, K. (1909) Notes on algae new to Japan. Bot.

Mag. Tokyo 23, 117-133.

(1920) Novae Algae Japoniae. Decas 1-11. bid.

34, 1-12.

THE SPATANGOID ECHINOID LINTHIA FROM THE LATE EOCENE OF

SOUTHERN AUSTRALIA

BY K. J. MCNAMARA

Summary

Five specimens, one from the late Eocene Tortachilla Limestone in South Australia, and four from

the Pallinup Siltstone in south-western Australia, are described as a new species, Linthia pulchra

sp.nov. The absence of a lateroanal fasciole is not thought to be of sufficient importance to warrant

placing this species in any other genus other than Linthia. The concept of the genus Linthia is

emended to include forms both with and without a lateroanal fasciole.

THE SPATANGOLD ECHINOID LINTHIA FROM TILE LATER EOCENE OF

SOLTHERN AUSTRALIA

by K. J. MCNAMARA*®

Summary

McNamara, K, J, (985) The spatangoid echinoid Linthia tram the Late Eocene of southeri Australia,

Trans, R. Soe. 8. Aust, W%4), 141-165, 29 November, 1985,

Five specimens, one from the Late Focene Turtuchilla Lijnestone iy South Australia, and four fram

(he Pallinup Siltstone in south-western Australia, are described as a new species, Linthin pulehra sp. nov.

The absence of u lateroanul fasciole is nar thous ht to be of sallicient importance to warrant placing this

species in any yeous other than (deride ‘Che concepi of the genes LinMiie ws crended to include forms

both with and withoul a Jaleraaual fascjole,

Key Worps: Spatangoid echinaid, Linda Eocene, new species, southerrt Australia.

Introduction

Within the ‘lertiaty sequence of southern

Anstralia the rich spatangoid echinoid fauna has

yielded a number of forms which, at sometime or

other, have been assigned to the genus Lindhva. Tate

(1885) described Linthia antiaustralis from the Barly

Miocene of the Murray River cliffs; H. L. Clark

(1946) placed Megalasrer compressa in Linthia: and

Pritchard (1908) placed two of McCoy's (L882)

species, Pericosmus nelsoni and Pericosnius viyas

in Linthia, along with a new species, &,

mooraboolensis, However, as discussed below, none

of these species belong in the genus Lintila.

Consequently, this common, cosmopolitan, Tertiary

genus has not previously been described from

Australia.

Iris the ain of this paper to deseribe what is not

only the largest Known spatangold echinoid from

the Late Eocene Tortachilla Limestone in South

Australia and the Pallinup Siltstone in Western

Australia, but also to record the presence of f inthia

in the Australian Tertiary.

The material wpon which (his paper is based

consists of a single specimen from the Tortachilla

Limestone in South Australia and four from the

Pallinup Siltstone in Western Australia, Although

the Tortachilla Limestone specimen does nel have

its adoral surface preserved, much of its aboral

surface is in an excellent state of preservation. ‘Uhre

of the specimens from the Pallinup Siltstone are

preserved only a8 interial moulds, but with bowk

aboral and adoral surfaces preserved. ‘The other

specimen is poorly preserved and consists of jm

external and internal mould. Sufficient details are

preserved in the five specimens (o be contident that

they are conspecific and represent a hitherto

undescribed taxon,

Measurements on the specimens were made with

a vernier calliper to an accuracy of O.1 mm. A

* Wester Australian Muselm, tranecis Street, Perth,

Western Australia @000,

Wutnber of parameters are expressed as percentages

of maxiniuim test length (VL), Specimens are

housed in thé collections of the Museum of Victoria

(NMYV); Western Australian Museum (WAM) and

Cieological Survey of Western Australia (CiSWA).

Stratigraphy

The Tortachilla Limestone is a yellow-brown,

ercen and grey bioclastic limestone which in oulerop

in the Willunga Embayment of the St Vincent

Basin, South Australia, attains a maximum

thickness of 2m in cliffs at the south end of Maslin

Beach (Reynolds 1953; Cooper 1979). tt is early Late

Eocene tii age (Ludbriok 1963; McGowran 1978;

Lindsay 1985), It contains a rich marine invertebrate

fauna, predominant amongst which are the

echinoids. Most forms from this unt have been

described, including the tollowing: Srereocidaris

eudmore’ Philip, 1964; 5. /osteri Philip, 1964; 8S,

inermis Philip, 1964; S. hispida Philip, 1964; S,

infricale Philip, (964; Salenidia tertiaria (Tate, 1877)

(see Philip 1965); Orrholophus bittneri Philip, 1969;

Tatechinus audus Philip, 1969; Fibularia gregata

fate, 1885; Behinolampas posterocrassa Gregory,

1890 (see McNamara & Philip 1980a); Apatopygus

vincednaus (late, 1891); Australanthus longianus

(Gregory, 1800); Pisolenipas concinna Philip, 1963;

Gireliaster tertiarius (Gregory, 1890): G_ bellissae

Foster & Philip, (978; Gillechinus cudmnrei Fell,

1964 (see McNamara e7 a/. in press), Hemlaster

(Bolbaster) sp. soy. (MeNamara, in press);

Prenaster @ldingensis Hall, 1907; Protenaster

preaustralis McNamara, 1985; Schizaster (Paraster)

tate) McNamara & Philip, 1980b.

‘The Pallinup Siltstone, a member of the Werrilup

Formation within the Plantagevet Group, ts a silt-

stone of spongolite which occurs discontinuously

along the southern coast of Western Australia (see

Datrash & Kendrick 1980, Fig. Ll) This richly

fossilifermus uml is comadered to have been

deposited ina shallow shelf cavironment (Darragh

162 K. J. MCNAMARA

& Kendrick 1980). Echinoids form a major part

of the marine invertebrate fauna, but remain

essentially undescribed. Gillechinus cudmorei has

been recorded from the unit (McNamara ef a/. in

press), Other forms present include Schizaster

(Paraster) sp. nov.; Prenaster aldingensis; Giraliaster

bellissae; and ?Pericosmus sp. Locally spines of

regular echinoids are common. On the basis of the

bivalve fauna, Darragh & Kendrick (1980) con-

sidered that the Pallinup Siltstone correlates with

the Blanche Point Formation, Tortachilla Limestone

and Browns Creek Clay. These units cover the

planktonic foraminiferal zones P15-16 (Ludbrook

1973; McGowran 1978).

SYSTEMATIC PALAEONTOLOGY

Order Spatangoida Claus, 1876

Family Schizasteridae Lambert, 1905

Genus Linthia Desor, 1853

Emended diagnosis: Test commonly small, heart-

shaped, anterior ambulacrum not petaloid,

depressed with groove at margin; apical system

ethmolytic, subcentral with 4 genital pores; petals

of almost equal length; peripetalous fasciole sharply

indented between petals; lateroanal fasciole present

or absent.

Remarks; Kier (1984) has recently provided a

diagnosis of Linthia. The southern Australian Late

Eocene form accords well with this diagnosis in its

possession of a depressed ambulacrum III with

prominent anterior notch; ethmolytic apical system

with four genital pores; petals of similar length; and

indented peripetalous fasciole. However, Kier (1984)

diagnosed the genus as possessing a lateroanal fas-

ciole. Although the Schizasteridae are generally

typified by their possession of both a peripetalous

and lateroanal fasciole (Fischer 1966), in some

genera the lateroanal fasciole is not always entire

or present in all species or in all specimens of a

single species (Mortensen 1951).

McNamara & Philip (1980b) have discussed the

problems involved in using the characters of the

fascioles in the supraspecific classification of the

Schizi.steridae. For example, the lateroanal fasciole

may be present or absent, as in Paraster; only

present in juveniles, as in Abatus; reduced or lost,

as in Brisaster; incomplete, as in Tripy/us and Para-

brissus; or absent altogether, as in Kina.

Although species of Linthia typically possess a

lateroanal fasciole, examples have been described

in which it may be incomplete in some individuals,

or missing entirely from an entire population.

Gregory (1906) described Linthia oblonga

(d’Orbigny, 1854) from the Cenomanian-Turonian

of Sinai and Egypt. In this species the peripetalous

fasciole is invariably preserved, but the lateroanal

fasciole is either absent or present only as obscure

traces. This absence is not a function of preservation

as the tubercles and peripetalous fascioles are well

preserved. Newton (1904) similarly observed this

variability in L. oblonga from Sinai.

A similar situation is found in a form described

by Bather (1904) from north-west Nigeria as

Hemiaster sudanensis. This species is ethmolytic

and in appearance belongs more appropriately in

Linthia. This was also the view of Lambert & Thiery

(1925). Bather noted how specimens from certain

localities show the lateroanal fasciole (for example

Bather 1904, P1. 11, Fig. 13), whilst from the other

localities it is clearly absent (Bather 1904, Pl. 11,

Fig. 7).

Consequently the revised diagnosis of Linthia

presented above takes into account the ephemeral

nature of the lateroanal fasciole. Although much

of the test of the Tortachilla Limestone specimen

described herein is very well preserved, the lateral

parts of the test unfortunately suffer from extensive

post-mortem encrustations of bryozoans. Con-

sequently only one small portion of the lateral part

of the test, where the lateroanal fasciole would be

expected to be, is well enough preserved to indicate

whether or not this fasciole is present. Close

examination in this region near to the junction with

the peripetalous fasciole, has revealed the absence

of a lateroanal fasciole. The only specimen from

the Pallinup Siltstone which shows part of the

external surface (WAM 66.637) is not well enough

preserved to provide corroborative evidence one way

or the other. In all other aspects this species

resembles a typical Linthia, within which genus it

is therefore placed.

Linthia pulchra sp. nov,

FIG, 1

Material: Holotype, NMV P20455, from the Late

Eocene Tortachilla Limestone, southern Maslin

Beach, South Australia. Paratypes GSWA F5828,

from the Late Eocene Pallinup Siltstone, at Stokes

Brick Pit, Albany, Western Australia; WAM 66.637

from the Pallinup Siltstone at Bremer Bay, Western

Australia; WAM 85.710, 85.711 from the Pallinup

Siltstone near Albany, Western Australia.

Diagnosis; Test relatively narrow, maximum width

anterior of centre; apical system slightly anterior

of centre; ambulacrum III long, moderately depres-

sed; anterior notch well impressed and broad; petals

long and relatively narrow.

Description: Test reaching a maximum length of

80 mm; relatively narrow, width being 90.5% TL

in both undistorted specimens (NMV P20455 and

GSWA F5828); maximum width about one-third TL

from anterior margin; height 50-54% TL, highest

point slightly posterior of centre in weakly

developed keel; posteriorly truncated. Anterior

* SOUTHERN AUSTRALIAN EOCENE ECHINOID L/NTHIA

163

Fig. 1. Linthia pulchra sp. nov.; A, NMV P20455, holotype, aboral view, from the Late Eocene Tortachilla Limestone

at Maslin Beach, South Australia; B, WAM 66.637, paratype, aboral view of latex cast of external mould, from

the late Eocene Pallinup Siltstone, Bremer Bay, Western Australia; GSWA F5828, paratype, C, lateral view, D, aboral

view, E, adoral view, from the Late Eocene Pallinup Siltstone, Albany, Western Australia; all x1.

notch deep (8.7% TL) and broad (11.3% TL).

Apical system ethmolytic with four genital pores,

anterior pair being smaller than posterior pair;

moderately depressed and situated 39% TL from

anterior ambitus; madreporite long. Ambulacrum

111 depressed and broad; bearing large number (42)

of pore pairs; within each pair pores aligned about

15° to transverse line and separated from each other

by a prominent, swollen interporal partition. Flanks

of ambulacrum III bear larger tubercles than found

anywhere else on test. Floor of ambulacrum III

covered by dense array of miliary tubercles. Petals

deep; anterior pair diverge at about 130° and are

41% TL in length, bearing up to 44 slightly

conjugate pore pairs, outer pore of which is slit-

like and inner nearly circular. Posterior petals

164 K. J. MCNAMARA

diverge at about 60° and are 31% TL long in

holotype, 28% TL long in paratype, bearing up to

38 pore pairs which are similar in form to those of

the anterior petals. Posterior petals slightly narrower

than anterior pair, being 7% TL compared with 8%

TL. Peripetalous fasciole slightly indented in

interambulacra 2, 3, and 5; strongly indented in

interambulacra 1 and 4.

Adoral surface slightly convex. Peristome anter-

iorly situated, posterior margin being situated 20%

TL from anterior ambitus; semicircular in form,

width 16.5% TL; slightly sunken. Labrum does not

project anteriorly; constricted at one-quarter length

from anterior, flaring strongly posteriorly. Plastron

nearly flat; length 56% TL; width 40% TL. Peri-

proct slightly sunken; dimensions unknown.

Discussion: Although the specimen from the

Tortachilla Limestone assigned to L. pulchra was

collected some 1800 km from those in the Pallinup

Siltstone there can be little doubt that specimens

from the two regions are conspecific, even though

the Pallinup Siltstone specimens are largely

preserved as internal moulds. The only differences

between specimens from the two regions lies in the

slightly flatter test and the more sunken periproct

in the only complete uncrushed Pallinup Siltstone

specimen.

Such a wide distribution for an echinoid taxon

is not unusual in the living Australian fauna.

For instance, Breynia desorii has a range of nearly

4000 km along the Western Australian coast (Mc-

Namara 1982). Protenaster australis has a similar

range along the southern Australian coast

(McNamara 1985). In addition to the occurrence

of L. pulchra in both the Tortachilla Limestone and

Pallinup Siltstone, Gillechinus cudmorei is also

found in both units (McNamara ef al. in press), as

is Prenaster aldingensis A single incomplete

specimen of Giraliaster bellissae is known from the

Pallinup Siltstone. Foster & Philip (1978, p. 814)

questioningly referred a specimen from the Torta-

chilla Limestone to this species. They also record

it from the Late Eocene Wilson Bluff Formation

near the South Australia/ Western Australian border.

They further note the similarity in age (planktic

foraminiferal zones P14-15) between this unit and

the Tortachilla Limestone (P15). The Pallinup

Siltstone, as noted, lies somewhere within zones

P15-16.

A number of species from the Australian Tertiary,

as noted above, have been placed in Linthia.

However, none belong in the genus. Linthia

antiaustralis (Tate, 1885) has recently been placed

in Protenaster (McNamara 1985). Megalaster

compressa (Duncan, 1877) was placed in Linthia

by Clark (1946), but is now regarded as belonging

in Pericosmus (McNamara & Philip 1984).

Pericosmus nelsoni (McCoy, 1882) placed in Linthia

by Pritchard (1908), has recently been made type

species of a new genus, Waurnia (McNamara &

Philip 1984). Pericosmus gigas (McCoy, 1882) was

also regarded as belonging in Linthia by Pritchard

(1908). It is now considered, along with Linthia

mooraboolensis (Pritchard, 1908) to belong in

Victoriaster. Consequently, the Late Eocene form

described herein and placed in Linthia, is the only

unequivocal record of the genus from the Australian

Tertiary.

The only Australian spatangoid which could

possibly be confused with L. pulchra is Protenaster

philipi McNamara, 1985 from the Late Oligocene

Waurn Ponds Limestone in Victoria. L. pul/chra can

be distinguished by its lower test, more centrally

located apical system, deeper ambulacrum III,

broader petals and semicircular peristome. Pro-

tenaster preaustralis McNamara, 1985 which

coexists with L. pulchra in the Tortachilla Lime-

stone, is also superficially similar, but differs in its

possession of a more anteriorly situated apical

system; barely sunken ambulacrum III; much

shallower anterior notch; longer, narrower petals,

the anterior pair running almost transversely across

the test.

Linthia pulchra differs from the type species L.

insignis (Desor, 1853; see de Loriol 1876, P1.6,

Fig. 1; P1.7; Figs 1-2) from the Eocene of Switzer-

land in its broader anterior petals; more anteriorly

situated apical system; lower test with truncate

posterior margin; and less anteriorly projecting

labrum. The Middle Eocene L. wilmingtonensis

(W. B. Clark, 1915; see Cooke 1959, P1.29, Figs 1-4;

Kier 1980, P1.17, Figs 3-6) from North and South

Carolina has, like £. pulchra, a deep ambulacrum

II bearing many pore pairs. The two species can

be distinguished by the narrower petals of L.

pulchra and narrower ambulacrum III.

Linthia sindensis (Duncan & Sladen, 1882-6; see

Lambert 1933, P1l.4, Fig. 16) from the Eocene of

Pakistan and Madagascar compares with L.

pulchra. However, the Australian species can be

distinguished by its narrower petals and deeper

ambulacrum III.

Jeannet & Martin (1937) illustrated a specimen

from the Late Miocene of Java which they called

Schizaster spec. aff. subrhomboidalis Herklots

(Jeannet & Martin 1937, Fig. 61). This rather worn

specimen differs from S. subrhomboidalis (see

Gerth 1922, Pl.62, Fig. 3) in possessing a deeper

anterior notch; more anteriorly eccentric apical

system; and longer posterior petals. In these respects

this specimen bears some resemblance to L. pulchra.

However, its slightly shorter posterior petals and

broader ambulacrum III preclude placing the two

forms in the same species.

165

SOLMTEERN ALISTRALIAN EOCENE ECHINOID LINT ARIA

Acknowledgments

Professor G. M. Philip and Mr J, M. Lindsay are

thanked for reading the paper und offering

suggestions fur ifs improvement,

References

Barvee, FA. (1904) Eacene echinaids from Soknte.

Geol. Mag. Wee 3(1), 7927-304,

TEARK, HE. (1946) The cominodenn faut) Austrahia.

Ths composition and ils origin, Publ, Carnes. Inst. 566,

1-57

Cooke, C. W. (1956) Cenovoie echinoids of Eastern

United States. Araf Pap. US, Geol. Surv 321, 1-106

Coorna, Bt. (1979) Bacerie lo Miocene straleriphy of

the Willugga binbay ment, Rep, Dept Mines Enerny 8.

Aust, 50, 1-89

Darnaun, T. AL & Kespeick, G. We (i980) Bocene

bivalves from the Pallinup Siltstode pear Walpale,

Wester Australia, A Roy, Sov, Wo Aust. 63, 5-20,

Feit, H. B. (1964a) New genera of Tertiary echinoids

Repel ispeiceay Australia. Mem. nail Mus. Vite 2t,

Misco, AG. (966) /# Moore RoC (Bd) “Tiearise on

tnveriebrate Palaeontology, Pl U, Echinodermata J,

Asterozoa-Eehimozaa.” Goal, Soc. Amer. (Linig Kansas

Press: Lawrence),

Foster, R. J. & PHiuie, GM. (1978) Tertiury holastersicl.

echingids fran Australian and New Zealand

Palaeontology 21, TH-822

GERTH, H. (1922) Echineiden Jn Dic fossilicn yon Jaya,

K, Martin. Sament, Gea, Retolsmus, Leiden (4),

497-520,

GREGORY,..L. W. (1890) Some additions to the Australian

Tertiary Echinoida, Geol, Mae, 27, 481-492,

(1906) Fossil Echinvidea fram peas and Egypt.

Geol, Mag. Dec §, 3, 216-227 & 246-25

Hari, ToS, (1907) Four new cchiaoids from the

Australian Tertiary. Proc. Ro Sac. Vie. 19, 47-53.

JEANNEL, A, & Marin, A, (1937) Ueber neozoisehe

Eohinoidea aus dem Niederlaendisch-Indisclen Arch-

ipel, Lend, Creal, Meded, 8, 215-308_

Kier. P.M. (980) The echinvids of the Middle Eocene

Warley Hill Formation, Santee Limestone, and Castle

Hayne Limestone of North av South Carolina,

Smithson, Conte, Puleabiol, 39, 1-102,

(1984) Possil spatangoid echinoids of Cuba,

Smithsan, Cane, Paleabiol, 88, '-336,

Lammogt, 4 1994) Gohinides de Madagascar communi

yes par M_ PL Besaine. Ann, geal, Surv. Mines Mada.

vasvar dt, IAD

& Tripey, P. 1909-1925. Essai de nomenclatiore

raisonnde des Behinides (Ferriére: Chsumont).

Lintisay, I. M-. (1985) Aspects of Sourh Australian

lertiary foraminiferal biestratigraply, witht emphasis on

studies of Massilina and Subboting. fir MeL indsay

(Ed. ), Stratigraphy, palacontology, malacology; Papers

in honour of Dr Nell Ladbroak. Spee. Publ. Dept Mines

Lnerey S. bust. 3, 18-241.

Lona, PB. de (1976) Descriptions des éelinldes Ternaries

de la Suisse Mem, Soc peléant Suisse 2. 85-142.

Luperook, N. H. (1963) Correlation of the Tertiary racks

of South Australia, 7rans. &, Soc So Aisi, 87, $-15,

(1973) Distribulion and stratigraphic wdility uf

Cenozoic molluscan Rrunas in southern Ausiraha.

Tohoku Uni. Sei. Dep, 2nel ser. (Geol), Spec. Val. No.

6. 141-261.

McCoy, # (1882) Prodromus of the palaeonrology of

Victoria (Decade VII), Melbourne.

MatGowkan, B. (1978) Early Tertiary foraminiferal bio-

stratigraphy in southern Australia: a progress report.

nD. J. Belford and V. Scheibnerova (Eds), The Crespan

Volume: Essays in honour of Irene Crespin. Bull, Bur,

Miner Resour Geol, Geaphys, Aus(, 192, 83-45.

McNAMARKA, K. J. (1982) Taxonomy and evolution of

living Specics ol Breyaiy (Echinoidea: Spatangoida)

from Australia. Ree. West. Aust. Mus. 10, 167-197,

(1985) Taxonomy and evolution of the Cenozoic

spatangoid echinoid Profenuster, Palaeon(ology 28,

311-330,

— in press, Taxonomy and evolution of ‘Tertiary

species Ol Hentiaster (Spatangoida: Herniusteridae)

from Australia. Pulaeontology.

& Pouitip, Ce M, (19808) Tertiary species of

tchinolampas (Echinoidea) from southern Australia,

Men, nar, Mus Vie. 44, 1-14

& (L980b) Australian Tertiary schizasterid

echinoids. Alcheringa 4, 47-65,

A 41984) A revision of the spatangoid

echinoid Pericusmus (rom [he Tertiary of Australia. Ree.

Mes Aust, Mus. 11, 319-356,

& KRUSE, PD, in press. Tertiary brissid

échinoids of southern Australia, Alcheringa.

Mortensen, T. (14S), A monograph of the Echinoidea

3f2), Spatangoidea i] (Reitzel:; Copenhagen). 593 pp.

Newtom, R. B. (1904) Linthia oblonga (Orbigny) from

Senai, ‘Geol, Mag, dec, 5, vol 1, 441-445.

PuHicie, G. M. (1963) Two Australian Tertiary

neolampadids, and the Classification of gassiduloid

echinoids, Palaeurtology 6, 718-726,

= (1964) The Tertiary echinoids of soutt-eastern

Australia, HW. Cidaridae (2). Prove. R. Soe. Wie. 77,

433-474,

(1965) The Tertiary echinoids of south-eastern

Australia, IIL, Stirodonta, Aulodonja, and

Camiurodonta (1). Prac. R, Sac, Vie, TR, IAM,

(1969) The Tertiary echinoids of south-eastern

vastralia. 1V. Camarodonta (2). Proc. KR. Soe, Vie, 82,

234.275.

PRITCHARD, G. B, (1908) On the occurrence of (he genus

Linthia in Victoria, with description of a new species

Pree. Ro Soe. Vie. 14, 21, 302-400,

Reraaps, M. A. (1983) The Cainozoie succession of

Maslin and Aldinga Bays, Sout Australia, Trans, R,

Soc. &. Aust. 76, 114-140.

Tare. RK. U877) On new species Of Belemnites and

Salenia tram the Middle Tertiary of South Australia.

O. dL geol. Soc, bawidon. 33, 256-258,

(1885) Miscellaneous contributions to the

paleontology of (he older rocks of Australia. Sef, Sey,

Rec N.S. 1, 1-5,

~ F891) & biblicgraphy and revised list of the

described cehingids of the Australian Booene. with des

ri eal of some tww species. Thats. R. Sree. 3. Aaavi.

. 270-282.

SPATIAL DISTRIBUTIONS OF STOCKING INTENSITY PRODUCED BY

SHEEPFLOCKS GRAZING AUSTRALIAN CHENOPOD SHRUBLANDS

BY ROBERT T. LANGE

Summary

The distribution of estimated sheep stocking intensity (ha sheep ') is described from within each of

several paddocks (large wire-fenced subdivisions) of arid zone stations (ranches) in Australia. The

estimation method is based on demonstrated proportionality between time spent and egesta

deposited by sheep in the localities they visit. Egesta deposition measurements of which the

intensity estimates are transformations are shown experimentally to correlate with immediate sheep

effects upon the vegetation.

SPATIAL DISTRIBUTIONS OF STOCKING INTENSITY PRODUCED HY

SHEEPFLOCKS GRAZING AUSTRALIAN CHENOPOD SHRUBLANDS

by ROBERT T. LANGE*

Summary

Lance, R. T. (1985) Spatial distributions of stacking (ntensity produced by sheepflocks grazing Australian

chenopod shrublands. Trans. R. Soc. S. Aust, W9(4), 167-174, 29 November, 1985.

The distribution of estimated sheep stocking intensity (ha sheep” ') is deseribed fram within each of

yeveral paddocks UVarzge wire-fenced subdivisions) of arid zone stations (ranches) in Australia. The estimation

mettind 15 based on demonstrated proportionali(y bebween (ime spent and evesta depusited hy sheep in

the localities they visit, Egesta deposition measurements of which the intensity estimates are transformations

are shown experimentally to correlate with immediate sheep effects upon the vegetauon.

The magnitudes of estimated sheep stocking intensity are shown to vary greatly from place 1p place

within the one paddock, In the studied region about one third of each paddock’s area experienced intensities

higher than the paddock average, eradityg upwards to about six times higher, while about two (hinds

experienced lower intensities grading dawn to zeta. Implications for perennial vegetation in the paddocks

ure diseased,

Kry Worps: Chenopod shrubland, sheep praying, stocking pressure varialioo,

Introduction

Sheep pastoralism in the Australian anid zone is

notable for its great weol production but also tor

the damage which the sheep do to the perennial

vevetation. There is a general belief that there are

sale slocking intensities which do not harm the

vezetation and excessive intensities (“overstocking”)

which do, Many authors have used smatl-paddock

experimental trials (of arca one ha or 50) to

demonstrate the relevant relationships between

stocking intensity and vegetation reactions

(Cunningham & Walker 1973; Hamilton & Bath

1970; Harrington 1979; Leigh & Mulham 1966;

Leigh & Wilson 1969; Leigh, Wilson & Mulham

1968, 1969; Trumble & Woodroffe 1954; ‘Tupper

1978: Williams 1970; Wilson 1974, 1976; Wilson &

CGractz 1980, Wilson, Leigh & Mulham 1969;

Wilson, Mulham & Leth 1976),

The application of their results to [ull scale

pastoral paddoeks i5 complicated because flocks do

not spend their time evenly throughout the large

areas in which they wander (Waite 1896; Osborn,

Wood & Paltridse 1932; Valentine 1947; McBride,

Arnold, Alexander & Lynch 1967; Barker 1979;

Squires 1970, 1974, Whalley, Robmson & Taylor

1978), So although the paddock average stocking

intensity may be known (PSI = enclosure area

divided by flocksize), the intensity on any actual

part (SIP — average stocking intensity of the part)

remains unknown, Without some means of

estimating SIP at given points within paddocks,

there js no connection with the experimental-trial

literature. The difficulties of this situation were

* Depariment of Botany, University of Adelaide, Box 498,

G.P.O, Adelaide, S. Aust. S001,

pointed out by Rawes & Welch (1966, 1969), Ares

& Leon (1972) and Anderson & Currier (1973)

An approximate solution to these difficulties

follows demonstrations by Rawes & Welch (1966,

1969), Lange (1969), Ares & Leon (1972), Squires

(1974), Lange & Willcocks (1978) and others, that

quantilative relationships exist between flock use

of pasture localities and the amounts of egesta

which tall in them, From these relationships SIP

can be estimated. A particular basis demonstrated

by Lange (1983) is used here to examine the spahial

and temporal distribunon of estimated SIP in arid

yone paddocks of the Whyalla region, South

Australia, during the period 1969-1982.

Methods

Computations and Graphical Summaries

As im the study of Hilder (1964) the enclosure

(paddock) is imagined to be divided into many

equal parts from each of which egesta accumula-

tions can be measured. Over any observational

timespan the exact average Stocking intensity ou

each part (Lange 1983) then is:

area of the part (ha)

ee = SIP (ha sheep ')

enclosure » fraction of total

Hlocksize flocktime spent

in the part (=F)

The use of ha sheep '| rather than its reciprocal

follows Harrington (1979), Cunningham & Walker

(1973), Rattray (1960), Goodall (1971) and Vesey-

Fitzgerald (1974) because in the arid zope jt mostly

has values >1. Except in small-scale calibration

experiments (Lange & Willcocks 1978; Lanve 1983)

thete was no practicable means of obiaining F

168 R. T. LANGE

directly because canopies of the western myall trees

(Acacia papyrocarpa Benth) prevented surveillance

even from a tower. Egesta accumulations in contrast

were easily measured and had the special advantage

of reflecting total flock activity at all times in all

places.

An approximate SIP was obtained for each part

by substituting the fraction of enclosure total egesta

deposition on the part as a surrogate for F (the

fraction of total flocktime spent in the part). In this

study that substitution rested on 3 experimental

demonstrations in the Whyalla region, that

sheeptime spent on a part was roughly proportional

to egesta recovered from it (7 = 0.92 to 0.98,

p <0.001). This proportionality was subject to the

requirement that the size of the part, the length of

the accumulation span and the size of the egesta

sample were all relatively large (Lange 1983).

As in similar studies (Rawes & Welch 1966, 1969;

Ares & Leon 1972; Squires 1974; Hilder 1964),

egesta on the part was sampled, in this case from

240 m? of ground surface per part. Over

protracted periods it was recovered, oven dried,

weighed, volumed and stored every few weeks to

avoid field decay losses.

Cumulative Paddock Area

fe) 0-5 1

LiLtitiiiiitt

7 127sheep ha!

~ 56

= r

2 2

5 1 Go

<4 ©

a. =

7p) O5 I

0-25

2-5 sheep ha | 0125

‘

Fig. 1. The cumulative distribution of log, SIP (stocking

intensity of the part) for the 30 parts of the sheep-

stocked enclosure described by Hilder (1964), The right-

hand scale expresses SIP variation in terms of PSI

(paddock average stocking intensity). In this case sheep

ha ! rather than its reciprocal is used because of the

very high stocking intensity.

The first form of graphical summary was the

paddock cumulative distribution of log, SIP as

shown in Fig. 1 which was prepared as an example

from published data of Hilder (1964, his Fig. 1).

This summary is directly comparable in preparation

and applications to the cumulative probability

distribution P, of a normal distribution (Smith

1954, his p. 581) which in most cases it

approximates. Log x is used to compress and

normalize SIP scores which vary greatly even from

very small heavily stocked enclosures (Hilder set 20

sheep in just over | ha). Log, is used because

published data about the sensitivity of vegetation

to SIP are from factorial experiments involving an

SIP doubling scale (e.g. Wilson, Leigh & Mulham

1969; Graetz & Wilson 1979).

The second form of graphical summary was a

3-dimensional graph of which the base plane

represented the paddock surface and the vertical

axis was scaled in log, SIP. The surface then was

contoured in intervals of one log, cycle with PSI

as datum, thus exhibiting successive doublings

(upwards) and halvings (downwards) of SIP from

PSI. Rotation of these graphs allows perspectives

of the spatial distribution of SIP as in Fig. 2 which

was prepared as an example from the same data of

Hilder (1964) as was Fig. 1.

Fig. 2. Perspective view of a 3-dimensional graph showing

the distribution of SIP values in the enclosure described

by Hilder (1964). Values are grouped in class intervals

of one log, SIP cycle with PSI as datum. The scale

expresses the SIP variation in terms of PSI.

Study Locality

The studies were conducted on Middleback and

Nonowie stations near Whyalla, South Australia,

an environmental context which already has been

thoroughly described by Barker (1979), Jessup &

Wright (1971), Rogers & Lange (1971) and Noble

(1979) with maps and descriptions of most of the

paddocks mentioned here. It is arid chenopod

shrubland of A/ériplex and Maireana with variable

low woodland of Acacia, on undulating desert

loams. Rainfall is very unpredictable and averages

about 220 mm yr! (1940-1970). The whole region

is fenced into paddocks approximately 1200-2000

ha which are stocked continuously with merino

STOCKING INTENSITY IN ARID SHRUBLANDS 169

Tait L.. List of studies undertaken in the Whyalla region, South Austrilia, to determine the magnitudes dnd

distributions of SIP (stocking intensity of the part of the paddock in hea sheep~') in each case. PSI ~ paddock

averuge Stocking intensity.

Study Paddock Area PSI Dates Number

No. name (ha) (ha sheep |) of paris

I, Wanga 1153 5.8 Aug. 1969-40

Feb, 1970

inc.

2. Wertige 2280 6.5 Oct, 1971- 37

Muy 1972

ine.

3. Wernzo 2280 65 dune 1972- 37

Sept. 1973

ing,

4. Overland 1290 5.5 June L978- 50

Jan. (979

ie,

5, Twor Mile 1936 67 Sept, IYRO- = 52

Nov, 1980

ine,

6, Two-Mile )936 6.7 tan, (9RI- 52

March 1981

ine,

% Purpunda 1145 5,4 June 1982 59

Sept, 182

ithe,

sheep at PSI of about 6 ha sheep ' (Lange,

Nicolson & Nicolson 1984),

Validation Experiments

Validation that egesta accumulation was roughly

proportional to flocktime spent by sheep in parts

of the discussed paddocks was published by Lange

& Willcocks (1978) and Lange (1983).

In the work of Ares & Leon (1972), validation

of egesta accumulation as a useful variable to

measure in pastoral research rested on the

persuasiveness of its correlations with stock effects

upon pasture plants. The sane applied where

distance from water was used as the stocking

variable (Osborn, Wood & Paltridge 1932; Valentine

1947; Lange 1969; Barker & Lange 1969; Squires

1974, Graetz & Ludwig 1978; Barker 1979; Fatchen

& Lange 1979). To provide an equivalent sort of

validation that egesta accumulations (and hence SIP

estimates) correlate wilh concomitant effeets on

vegetation in the Whyalla region, several

experiments can be cited.

Lange (1984) showed that the observed proba-

bility of random ourer shoots of saltbush (Atriplex

vesicuria Heward ex Benth.) being grazed by sheep

was largely accounted for by using concomitant

sheep egesta accumulation as the independent

variable in regression equations, Data were from

bushes at arbitrary localities within 5000 ha of

Whyalla shrubland ranged by sheep. The regression

equation was

y = 1.80 4 017K PF = 0.78, p <0.01

where » was percent loss of marked shoots over.a

6 week period afd +. was concomitant sheep egesta

accumulation in kg dry wt, on plots 60 = 40 m.

Across the different parts of small experimental

paddocks stocked heavily, the relationship on plots

10 » 10m was much clearer, namely

y = 0416 + 0.004x, 7° = 0.91, p <0.001.

One further validation experiment* is reported here

to extend demonstrations of the usefulness of egesta

accumulation as an index of SIP, at least in some

contexts. Stipa nitida Summerh. & C.b. Hubbard

and Danthania caespitosa Gaud. in Freye, are

prominent tuft grasses occurring between bushes

in mixed chenopod shrubland of 2-Mile paddock

(map and description in Barker 1979). A tract of

this vegetation 30 « 200m was fenced to include

a drinking trough and was surveyed into 10 equal

parts.

In cach part all grass tufts >5 cm tall ona plot

20 » 0.2 m were estimated for biomass (x. dry wt.,

technique of Andrew, Noble & Lange 1979) and

measured for height (cm). The enclosure was then

stocked with 10 merino sheep lor 6 days, the grasses

then remeasured and egesta dropped by the sheep

was collected from plots 28 ~ 2 m superimposed

over the grass plots. The prediction was thal egesia

” This experiment was performed by Chester .. Merrick,

an Honours student in the authors programme.

170 R_T. LANGE

accumulations would index reductions in grass

height and biomass via regression equations.

Fullscale Siudies of Paddock SIP

Table J lists studies which were undertaken

between 1969 and 1982 (see Barker 1979, Noble 1979

and Lange, Nicolson & Nicolson 1984 for paddock

maps), Wanga paddock lies east of Wertigo and

Overland lies north of 1-Mile.

Results

kalidation Experiment

The experiment yielded the very highly significant

mulliple regressions:

v = 36.99 — 0.47x + 0.932, Fr = 0.99,

Pp <0,001

where » = final mean grass height (cm),

x = epesta dry wi (kg)

and <=. = initial mean grass height (cm)

and y-~ 141) O15y « 0.952, r2 =0,99,

p <0,001

where » « [inal grass biomass (g),

xX ~ epesta dry wt (Kz)

and z = inihal grass biomass (g).

Full Seale Studies

Data from (he seven Separate paddocks were all

\he same in principle, differing only in detail of

Cumulative Paddock Area

is) O5 1

Lit pp pd ay

19)

wo it 'a

7 ©

Ia 2 a

2? z=

A 12

£ 3 2

a 0-5 1)

al 4

~ 0-25

01425

0-06

\ 0-03

Not Visited —

Vig. 3, The cumulative distribution of log. SIP for (he

50 purts of Overland Paddock June 1978-January 1979

inclusive (Study 4),

spatial paltern and particular history, A selection

of data is presented here as a basis for discussion.

None of the omitted cases tells a different stary or

contradicts the examples presented,

Overland Paddock (Study 4)

Fig. 3 shows the cumulative distribution of los,

SIP for the $0 equal parts of Overland Paddock

for the period June 1978-January 1979 jnclusive,

Four parts accumulated no egesta during this period

so SIP for them was oo ba sheep~!, Over the used

parts of the paddock SIP values graded smoothly

upwards from PSI, over about one third of the area,

to a highest intensity about 2.5 cycles (6«) above

PSL. Over about two thirds of the used area they

graded smoothly downwards from PSI to lowest

measurable intensities about 4 cycles (16) below

PSI, Only | of the 50 parts actually had SLP — PSL.

Fig. 4. Perspective view of a 3-dimensional graph showing

the distribution of SEP values in Overland Paddock

(Study 4). Swamp Dam is the southern warer poi

Fiz, 4 shows the corresponding SIP surface of

the paddock with log, PSI as datum and with

class-intervals of | log, cycle, This spatial pattern

had the following weather and flock management

history. In June 1978 drought had restricted

drinking sources for sheep in Overland paddock ty

Swamp Dam in the south part. The flock was

obseryed to depend on Swamp Dam uoril it dried

oul, Then a temporary source of water was

introduced (by pipeline) to a point at the wesr end

of the north boundary af the paddock, to which

the sheep were introduced.

Two-Mile Paddock (Studies 5 and 6)

Fig. 5 shows the cumulative distributions of log,

SIP for spring and for summer 1980-81,

respectively, in 2-Mile paddock, Fig, 6 shows (hese

distributions compared with a theoretical normal

cumtlative probability distribution for the 52 parts

of the paddock. The diagonals represent required

lines for perfeet fit and the dotted lines show

observed fit,

Rank order correlation of spring and summer

SIP scores across the 52 parts of 2-Mile paddock

was nol significant, implying that the Mock shilted

STOCKING INTENSITY IN ARID SHRUBLANDS I

Cumulative Paddock Area

; (a) 0:5 1 “16

Litt} jt | | jf |

1 r

ex ®

3 2 ° 5

5 1o*

os 3 On

~ iT

o 4 0-5

2&5 O-

0125

. | 0-06

Visited L

Not

Mig. 5. The cumulative distribution of log, SIP for the

52 parts of 2-Mile Paddock in spring (fine line) and

summer (heavy line) 980-81.

its pattern of paddock use from spring lo summer,

1980-81, Hence the resemblance of spring and

summer cumulative distributions is demonstrated

to be independent of spatial shifting of use pattern.

Fig. 7 shows the SIP surface of 2-Mile paddock

for the combined spring and summer periods. Peak

stocking pressure was located not at the watering

points bul against the northern fence in a drainage

line.

Wertigo Paddock (Studies 2 and 3).

Fig, 8 shows the SIP surface of Wertigo paddock

for the period October 1971-May 1972 inclusive

with log, PSI as datum and SIP jn class-intervals

of I log, cycle. The flock was observed throughout

this period to use only the southwestern water point

and 1o graze away from it in two directions, namely

southeastwards into the southeast corner of the

paddock and north along the western boundary.

One part of the paddock was never visited,

Fig. 9 shows the corresponding surface for the

period June 1972-September 1973. The flock was

observed throughout this period to use Wertigo

paddock in approximately the same pattern as

previously, There was a highly significant rank

correlation ol SIP scores between the two periods

(r — 0,846, #7 < 0.001), indicating relative stability

Observed (Empirical) Cumulative Probability.

05 j

Theoretical Cumulative Probability

(Normal Deviate )

i=)

Observed (Empirical) Cumulative Probability

0-5

Theoretical Cumulative Probability

(Normal Deviate)

Fig. 6, The cumulative distributions of loz, SIP for

spring and summer I980-81 in 2-Mile” Paddock

compared with theoretical normal cumulative

probability distributions. The diagonals represent

required lines for perfect tit and (he doued lines show

observed lit. Spring above, summer below.

Fig, 7. Perspective yiew of a 3-dimensional graph showing

the distribution of SIP values jn 2-Mile Paddock for

the combined period Spring and Summer 1980-81

(Studies 5 and 6).

72 RK. T. LANGE

Pig. 8. Perspective view of a 3-dinrensiinal wniph showing

(he distribution of SIP values in Wertiga Padilock fev

the period Oct, 1Y71-May 1972 ine fStudy 2).

Fig... Perspective view of a 3-dimensional graph shows

the distribution af SIP values in Wertige Paddock for

jhe period June 1Y72-Sepr 1974 ine. (Suady 3).

ol use pattern, Kangaroo egesta mcovered from the

sume places also correlated between periods (r

0.39, p < 0,05),

Discussion

The demonstiations by Rawes & Welch (1966,

1969) and by Ares & Leu (1972) that measurements

of evesta deposition in open paddock situations

jndex stocking intensities of sheep, are extended by

(he presene data. That extension is nit merely tu

the Australian and zone context, but also towards

a wore quanritlive footing.

Data jeported here show very signitlcane

quauutative relaponships beiween sheep epesta

deposinors and simulaneous effects on plants at

localities Chtoughout the range of the freely-roam ine

sheep flock, Furthermore those relauionships have

been demonstrated over time intervals so brief as

to admit no scope for interpretations other than uf

cgesta deposition as al index of simultaneous

stocking or gmiving pressure “The only logical

conclusion From the regression equations of the

validation experiment is that egesta accumulation

on plots indexed the simultaneous loss Of pritss

height and biamass caused by the sheep, and from

the equation in Lange (1984) that it indexed the

SuNUlANcous cating Of salehush shouts by che flock.

lo the extene thar rhe SEP esrimiates reported here

are direcc transformations of phe egesta deposition

data, (he SIP estimates also relate to effeets of stork

on vegetation, Ti is reasonable from the presest

evidence to suppose, if plants are being uffected by

stock in their padduck, thal the plants are likely to

be affected most where SIP estimates are highest

and Jeast where they are lowest. But the particular

advance introduced here is the capacity to place

quantitative estimates (ha sheep ') upon the SIP

at particular places within the ranges of flocks, and

thus to link those places with the exlensive small-

plot experimental literature mentioned in the

Introduction. The results of examining those links

will be discussed elsewhere; the aint here is Co discuss

intrapaddock SIP for its own sake,

Moch evidence indicates fundamental similarities

in the relationships of SIP in all paddocks examined

from the Whyalla region and extending, so far as

data exist, 10 published cases from elsewhere lit

Australia, Thus the logarithms of SIP values in each

ol the examined paddocks were approximately

normally distributed, Something intrinsic in the

behaviour of sheep flocks is suggested by that fact.

constraining them to distribute theur flocktime very

unevenly, on average, throughout their range, Lest

that be attributed simply to the vast range area

available to relatively small flocks in (he Whyalla

region, or to the piosphere effeet (Barker 1979), il

should be recalled that the same applied even to

Hilder’s data concerning 20 sheep penned inside |

ha,

That inflexibility has important practical

implications, The first is that the more even spread

of flocktime [hroughout each paddock, desired by

station managers (Lange, Nicolson & Nicolson

1984), seems less likely to be achieved simply by

further fencing of paddocks turo smaller areas, than

was earlier thought.

The second is in relation Lo absolute flocksize anc

paddock sive, and PSI. Ie is traditional in the

Australian sheep industry to make stocking

comparisons just on the basis of PSI (in ha

sheep ‘or, in wetter more productive places, sheep

ha |). But as shown in Fig. 5 of Lange, Nicolson

& Nicolson (1984), any particular PSI (say, 6 hit

sheep |) is set throughout the industry by widely

different combinations of flocksize and paddock

size, Present results show that even i the relatively

small paddocks of the Whyalla region, SEP values

reflect the limits to which flocks can or will range

away from their water-point. There is no evidence

to suggest phat those limits are different in big

paddocks,

It ought to be evident from this, and from the

inexorable tendency of the flock to exert SLP values

grealer (han PSI over about one third of the Mock’s

reabzed range, ital PSL becomes progressively more

srossly misicading as an Indes of SIP, us paddocks

STOCKING INTENSITY IN ARID SHRUBLANIS W3

and locksizes increase at he one PST. Thatis unless

the larger paddocks have correspondingly more

Waterpoints, which in many Austrahan cases they

do not.

There ape other applications of the present

approach which are independent of consideralions

of wool production and pasture. Over 60 species

of rare and threatened Australian endemic flora are

to be found scattered within the sheep paddocks

ial enmesh their remnant distribiniens in this

region. Clearly thei fate is closely related to the SIP

they experience, whether high or low, Lis iiportiua

to establish what their situation is, by means of the

present Jechnique. as part of developing

conservation measures for them.

Acknowledgments

The author most gratefully acknowledges the

financial support of this work provided by the

Australian Research Grants Scheme, ragether with

supporr from the Broken Hill Proprietary Co. and

Mitsubishi Motors (Australia) Ltd.

References

ANDERSON, E,W. & Coneitr, Wo b (1U74) Bealuatines

zones OF dihzation. J Aarige Mune. 26, 87-9],

AsbRUW, M, BL, NobLe, TR, & Lancet, RT 978) A

none desiructive method for estimating the waht of

forage on shrubs. dust ba sep AV, 225-231.

Ags, JO. & Lrosx, RB. C_ (1972) An ecoloxical

Asseysment of the influence of grazing on plant

community structure, A Leak 0, 325-331,

BAKKER, S. (1979) Shrub population dynamics under

praving-—within paddock siudies. J Ro Gracty & K-

Howes (eds) Studies of te Australian And Zone IV.

Chenopod Shraiblands. C.S,1,R.0, (Aust) Diy, of Land

Res. Manag, pp. 83-100.

& LaAnot, BK, 1. (1969) Effecrs of inoderate sheep

Stocking on plint populitions of a blackoak-hluchush

association. lust «Bor V7, 827-537

CUNNINGHAM, GML & Walker, Po, 1973) Growth and

survival of trulew (lege anew Moot @& Bent)

in western, New Soutlr Wales, Trop. Grass/ 7. 69-77.

Farenen, TIL & Lance, R. TT. (1979) Piosphere patent

and dynaimivs in achenopod pastire sraved by calle.

Prk. Gimeta & K. Howes (eds) Studies of (he Australian

And Zone Vo Chenopod Shrublans. C.S.LR8,0, (Aust)

Div, of Land Res. Manag. pp. 160-169,

Gompaxir, DW. (1971) Extensive grazing systemis. Ch. 9,

pp, 173-187. In J. B. Dent & f. R. Anderson (Eds)

SYSIINS analysis 1 Uprichitural management, (Wiley

New York),

GkAtty, B,D, & Lopwie, JA. (1978) A method far che

analysis Of piosphere data applicuble ta range

assessment luvs Ruagel J 1, 126-136.

o% Wisos, AOD. (1979) An asseysiment of

herbivore diets in the chenopod shrublands tn R

Cirnety & kh. Howes (eds) Studies of (he Australian Arid

Zone LY. Chenopod Shrublands, CS A.R.0) (Ansty Dy

OF Land Res Manas. pp 44-159

TEAMit tom, 2, & Uavres, 1. Cy, (1970) Performance of sheep

and caltle pritzed Separately and wogether. least, A fay

Aert. and An ftast. 10, 19-26,

HARRINGTON. Ci oN, (LUT) The effvers af eral puts and

sliwep on The strub populations ina semi arid woodlind.

lust, Rangel iv, B44-445,

Hitbbre, Lo (1964) The distribution of plant autrient:

by Sheep a pasture, Proe, Aust Se Ar Pred 5,

24 124k:

Jessth, RoW. & Wario, M. 2 971. Cengzorn

sediments, soils and climates ar Whyalla, Seurh

Australia, Geoderma 6, 275-308

Lancr, Ro 1.1969) The wlospher sheep inick wad dupe

pullers. Rartige Manag 22, 396-400.

(1983) Earimanion ol ahcep stocking infensily at

any locapon in anit cay paddoeks, Jrwas. KR. See 5,

Aust UT, 137

(1984) Leaf marking mh ranecland araving ariidies.

Troms. Be Sow S. Aust (8, 42,

Nicarson, AD. & Nicotsow. BA W984

Vegetatign management Of chenopod rangelands in

South Australia. Ws. Rangel. Jf, 4h-Sa

~— & WILLCOCKS, M_C. (1978) The relation between

sheep-time spent and egesta accumulated within an arid

gone paddock, dusn Ah Aap. Agric Ane Hush, 1,

To4-767

LEIGH, J. Ho& Muiiiam, W, E. (1966) Selection af diet

by sheep graving Semraricl pastures on the Riverine

Plain. 1. A bladder salrbush (triples vesicaria)—cotrony

hash (Adcdia dptiiia) community lust 2 Bogol Aer

An. Hush. 6, 460-467

& Wilsons, A, DL (1969) Utilization ot eliyryplex

species by sheep, /h Re Joes (ed) ‘The Hiolopy of

Atriplex. C.5.L600, (AgsL.) Diy ot Plain Wid. pp. 97-104

' & MULiAM. WoL. (968) A study wt

merino sheep graging & votton- bush (Kochia aphyita)

grassland (Stipa veribilis—-Donthoria caespiiasa)

community on the Riverine Plaig, dust J Agric Res.

19, 947-96).

a et (1979) A study OF sheen gana

i Bielah (Cusuaritu oristatiy) Rosewou ul (Helerodendrum

ol foliam) slinib woodland in Weslern New Seurk

Wales, «lust J) Axrie, Res, 30, 1223-1236,

Meribel, O., ARNOLD, Co OW, ALEAAnDER®, G. &

Loyach, J, J, 0967) Beological aspects of the behaviene

of domesticanmals, Prow. ecot Sur Ausn 2) (F165

Nopieé, | R, W979) Computer simulation models of

extensive chenapod shrubland eruzing systems, dn R

Graere and K. Howes feds) Srugies af the Aubtratian

And Zone. ['V. Chenopad Shrublands. C.S4.8.0. (Aus)

Div. of Land Res. Manag pp. 125-144.

Crnoke. "TG. B., Woon, JOG. & Pau kinoe T. 932)

Choy the prowel aiid eae lin Lo ary ii OF Te perennial

saltbush (4ryploe veyjewri—un ceolouicnt siindy of

see biog factor, Pre. finn Sue NS Wales 37,

7-42

ee THY | ML (1960) The grass cover of drica. DAW,

Agnc Studies No, 49,

Rowhs, M & Wriri D. (1966) Purchor studies ost sheep

grain the nerthecn Penioines 1 Br Gresxtand Sec

Zi, 56-4

Semeed 5 (1969) Dypland productivily of wereiation

and sheep at Moar House Naliooal Mature Reserve,

Westinoriand, Piglind. Owes Supplemwenium tb, 72,

Roores, RW. & Lanen, RT (S71) Lichen populations

vn arid soil enushs aroun Sheep watering places in Sends

Australia. Orkus 22, 83 LOG.

& Oo, (Lond. 3rd Edn, V2 pp

174 R. T. LANGE

Sauires, V. R. (1970) Growth of lambs in a semi-arid

region as influenced by distance walked to water. Proc,

Aust, Soc. An. Prod. 8, 219-225.

(1974) Grazing distribution and activity patterns

of merino sheep on a saltbush community in southeast

Australia. Appl. An. Ethol. 1, 17-30.

Trumpce, H. C. & Wooprorre, kK. (1954) The influence

of climatic factors on the reaction of desert shrubs to

grazing by sheep. Jn J. CloudsleyThompson (Ed.)

Biology of Deserts’ Symp. 3., Inst. of Biol. (Lond.).

pp. 129-147,

Tupper, G. J. (1978) Sheep production on Danthonia

caespitosa—Stipa variabilis grassland in response to

fertilizers and deferred grazing. Aus. J. Expt. Agric.

An. Hush, 18, 210-217.

VALENTINE, R. A. (1947) Distance from water as a factor

in the grazing capacity of rangeland. J. For. 45, 749-754,

VESEY-FITZGERALD, D. F. (1974) Utilization of the

grazing resources by buffaloes in the Arusha National

Park, Tanzania. E&. Afr. Wildl. J. 12, 107-134.

Waite, P. (1896) Pastoral development—past and

present. The Adelaide Observer (S. Australia). Sept.

26, 1896.

WHALLEY, R. D, B., Robinson, G. G. & Tayitor, J. A.

(1978) General effects of management and grazing by

domestic livestock on the rangelands of the northern

tablelands of New South Wales. Aust. Rangel. J. 2,

179-190.

Wittiams, O. B. (1970) Population dynamics of two

perennial grasses in Australian semi-arid grassland. ./.

Ecol. 58, 869-875.

Witson, A. D. (1974) Water consumption and water

turnover of sheep grazing semi-arid pasture

communities in New South Wales. Aust. J. Agric. Res,

25, 339-347,

(1976) Comparison of sheep and cattle grazing on

a semi-arid grassland. Aust. J. Agric. Res, 27, 155-162.

& Graerz, R,. D, (1980) Cattle and sheep

production on an Afriplex vesicaria (saltbush)

community. Aust, J. Agric. Res. 31, 369-378.

, LeiGu, J. H., & MULHAM, W. E. (1969) A study

of Merino sheep grazing a Bladder Saltbush (Atriplex

vesicaria)—Cotton-Bush (Kochia aphylla) community

on the Riverine Plain. Aust. J Agr, Res. 20, 1123-36.

, MuLHAM, W. E., & LeiGH, J. H. (1976) A note

on the effects of browsing by feral goats on a belah

(Casuarina cristata) rosewood (Heterodendrum

oleifolium) woodland. Aust. Rangel. J. 1, 7-12.

THE FOSSIL SEA URCHIN FELLASTER INCISA — AN EXTENSION OF

RANGE

BY T. SADLER AND N. §. PLEDGE

Summary

The fossil clypeasteroid sea urchin, Arachnoides incisa, was originally described by Tate from three

specimens collected from so-called Miocene beds at Red Bluff, Lake Tyers, east Gippsland. This

unit is now recognised as the Lower Pliocene Jemmys Point Formation. Foster & Philip revised the

generic allocation and indicated that the species also occurs in the Pleistocene of the Parish of

Werrikoo, Victoria.

BRIEF COMMUNICATION

THE FOSSIL SEA URCHIN FELLASTER INCISA—AN EXTENSION OF RANGE

The fossil clypeaslernd sea urchin, Arachnoides incisa,

was Originally described by Tate! from three specimens

collected from so-called Miocene beds at Red Blull, Lake

Tyers, east Gippsland. This unit is now recognised as the

Lower Pliocene Jemmys Point Formation. Foster &

Philip® revised the generic allocation? and indicated that

the species also occurs in the Plerstogeng of the Parish

of Werrikoo, Victoria.

Recently, one of us (7.S.) discovered specimens at

Willowhank, near Murray Bridge in the Murray Basin of

South Australia, some 850 kim roi the type locality in

the Gippsland Basin of Victoria, Subsequent invesiyation

and collecting has produced numerous specimens. This

note records the mew occurrence and offers additional

information on the species.

Sysremtattes

Class Echinoidea

Order Clypeasteroidea A, Agassiz

Family Arachhoididae Duncan

Genus Fellasier Durhum

Fellaster incisa (Tate 1893)

Arachnoides incisd Tate, 1893: 192, pl. 13, fig, 3;

Fellaster inctsa (late), hoster & Philip, 1980: 156, pl. 20,

fig. 4, 6—7., text fig. 1.

Amended diagnosis; Test very flat, rising slightly

towards apical disc; almost circular, transverse diameter

slightly erearer than longitudiial diameter. Apical disc

slightly anterior of ceatre. Ambitus sharp, incised at end

of each ambulacral groove. between which tl is undulose

Ambulacra inflared with median depression. and abruptly

doclivous al the sides; declivily decreases beyond end of

petals. Petals reach almost two-thirds the way 19 ambitus,

converging slightly al their ends on each ambulacrumn-

Amibulacra slightly more thar twice width of

inlerambulacra. Ambulacral plates obliquely combed,

forming five chevron patiern pointing away from apical

disc. Ornamentation minutely granular, Most apparent at

ambitus. Periproct supramarginal with shallow depression

between it and ambitus. nu definite anal notch, Peristome

centrally placed and round: Actinal area slightly concave

with ifterambulacral bulges near peristome (Pip. 1).

Fig. |, Fellaster incisa, (8) P24857, adoral, (b) P24856,

adapical,

Material: Seven complete specimens and several nearly

complete measurable specimens have been collected

(Table 1), together with numerous fragments, some of

which show the plate pattern clearly.

Taste |. Measurements. S, collection number, South

Australian Museum; L, longjtudinal diameter (from

frontal lobe); T, transverse diameter; H, heizht; A, distance

fron centre of apical dise ta posterior margin; P, distance

from centre of periproct to postenor margin. All

Medsurements in millimetres.

N i T H A P

P24857 59.9 64.5 30,3

P24658 59.8 64.3 5.1 3) 2 38

P2459 55.4 60.2 6.9 29.3 2.8

P24860 62.8 29.2 31

P24861 $3.0 60,6 6.2 27.5 45

P24862 A7.3 4.6 2,7

P24ko4 33.1 40,8 5.1 18,4 2.6

P2485 43.5 47.2 o.4 24.2 3.4

P2466 346 7B 4.4 18.0 2.7

P24867 33.2 35,5 4.0 17.5 2.6

P24869 54,5 61,2 6.2 28.7 3

P 24870 45,2 BA 32,8 3.9

P2487 44.0) 49.6 5.8 23.7 3.1

Occurrence; Most specimens came from outcrops

interpreted as Norwest Bend Formation, at Willowbank,

& km north ot Murray Bridge, and others near Sunnyside

Lookout, 3 km upstream, opposite Mypolonga. Specimens

occur throughout the lower 10 metres of exposed Norwest

Bend Formation at Willowbank (Fig. 2). Fragments were

(raced as far us Ponde, 5 km south of Mannum,

Ruagy calédresus sandsione

Highes! Pellasier found

Massive cAlcarcous saoodhonm HIGH I eat

Current bedded gravel

Curfent-hedded CHICA AGUA BANdatONe

wilh Follaster ingis4 common

Curtent-hefided gravel

Owes! Fellaster tound

Fine grhinfd MichGhols, KAIK mi mon

SaANGston® wilh thin Gedang

Worm Tubes COMNOn ema! too

FORMATION

NORWEST BEND

Miahire, Bulb 1a yellow chlcwrenie

with Mumeraus lreeguiee Mai dgroonds

apd marl bands. Richly tnssrlitoernas

with Loveniy lorbest. Mangetyeria

Qusicniis, Divaives, Dryorosas end

burrowed horizons

MANNUM FORMATION

Reserve Dom ramp

Fiz, 2. Stratigraphic section at Willowbank.

176

Abundance and completeness of preservation decreased

markedly towards this northerly location.

Remarks: In the area of interest, the Norwest Bend

Formation (Fig. 2) consists of current bedded, flat-

laminated and massive, medium to very coarse, micaceous

sands, with some gravel bands, biogenic clasts, and well-

rounded pebbles and cobbles of igneous and metamorphic

rock: quartz, Kanmantoo gneisses and schists, and

probable Murray Bridge Granite. Cementation varies

vertically; some beds are barely cemented while others

form very hard ledges. Some intervals show extensive

burrowing by “worms” forming more or less vertical tubes.

Fossils are not common, and at Willowbank, specimens

of Fellaster incisa apparently outnumber all others, which

comprise mostly scallops (Chlamys cf. antiaustralis), a

few fragments of Ostrea sp. and bryozoa. Only one other

species has been found; a single specimen of a small

unidentified sand dollar urchin. Some specimens of

Fellaster show evidence of a round hole bored into the

adapical surface, presumably by a carnivorous gastropod.

However, no fossil gastropods were found, presumably

'Tate, R. (1893) J. Proc. R. Soc. N.SW. 27, 167-197, pls

10-13.

Foster, R. J. & Philip, G. M. (1980) Proc. R. Soc. Vict.

91(2), 155-160, pls 19-20.

because the porous nature of the sediment would allow

their aragonite shells to be dissolved away rapidly.

Molluscan fossils gradually increase in number as Fellaster

decreases towards the northern limit of its observed range.

The largest specimens of Fe/laster seem to occur at

Willowbank while smaller ones were predominant near

Sunnyside Lookout. Most specimens were found in upright

horizontal position. A few, however, were overturned and

one was observed in almost vertical orientation. No

preferred lithology could be discerned, although better

specimens seem more common in the flat laminated to

massive finer sands. As would be expected, only broken

fragments were found in the gravel horizons.

Conclusions: Fellaster incisa (Tate) occurred widely in

southeastern Australia during the Pliocene, from the

eastern Gippsland Basin to the western part of the Murray

Basin. It appears to have preferred sandy bottom

conditions. For reasons unknown it was one of the few

animals to inhabit the Murray Bridge-Mannum area

during the deposition of the Norwest Bend Formation.

3Durham, J. W. (1966) Clypeasteroids. Jn Moore, R. C.

(ed.) Treatise on Invertebrate Paleontology,

Echinodermata. 3(2), Geol. Soc. Amer. (Lawrence:

Kansas).

T. SADLER and N. S. PLEDGE, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000.

AGE AND GROWTH OF THE BLUE-THROATED WRASSE

PSEUDOLABRUS TETRICUS

BY S. A. SHEPHERD AND LISA J. HOBBS

Summary

The blue-throated wrasse Pseudolabrus tetricus (Richardson) is one of the most common fishes of

inshore rocky reefs on moderate to high energy coasts of southern Australia. Yet except for

taxonomic and habitat notes, nothing has been published on its biology. This note describes the age

dependent growth and length-weight relationships of P. tetricus.

BRIEL COMMUNICATION

AGE AND GROWTH OF THE BLUE-THROATED WRASSE

PSEUDOLABRUS TETRICUS

The blue-(hroared wraske Pseudolabrus tetricus

(Richardson) is one of the most common fishes of inshore

rocky reefs on moderate lo high enerny coasts of sonthiern

Aysiralie. Yet except for taxonomic and habitat notes!”

nothing has heen published on its biology. This note

describes the age dependent growth ana length-weight

telalionships of A fefricus

This study iy bused on the analysis of scales of 58

individuals vaprured by speuring a! West Island and

udjacent areas i Encounter Bay, South Australia,

supplemented by small samples of large individuals from

Cupe Jervis, Yankalilla and Cileasons Landing, Yorke

Peninsula, The data tor another 17 individuals were

rejected because the seule circuli were foo indistinct to read

aecutarely, Three scales were laken [rom behind (he left

pectoral fin of each lish, cleaned in sodium hydroxide

slition and mounted between microscope slides.

Standard techniques’ were used for reading (he scales and

for back-valéulation of growth, Scales Were exiumined for

growth cheeks or closely spaced circuli by using a

microfiche reader, Photocopies of the projected seale

images were made and mean distances trom the focus

alone two diagonal posterior radii (where cireuli are most

clearly lefined) to successive cireuli and to (he margin were

taken for each sample, The sewe circull have been found

Io be annular for other (émperate labrids'* and are

therefore assumed to be anmnuli for this species. All lengths

are expressed as total length.

The relationship between seale raditis (R) in mim and

rotal length (L) in ent was found to be Tinear, and the

following equation was fitted ro ihe data by the leas!

squares method (Pig. 1).

Rk 0.0045 | O405 L tr 0.95)

This equation was then used to back calculate the length

or fish at the tine of formation of each annulus.

Because sex inversion [ram female to male is sizerelated

and occurs at a length of about 35 cm (Shepherd &

~ afl ’

PF,

me ee

i) ‘ *

try

- "

~ * ‘

. ‘ a ‘

E oy

E :

we -

2/4 ‘

-] ' ‘

w “

a pie

a ~ ‘

. *. c

L —————— ——— $$$ $4

in wef w 0 an

FOTAL LENGTH \cmi

Tin, t Plo of scale radii Versus total length for P

relricus,

Clarkson, in prep.) data for females and males were

combined for the purpose of the calculations,

The number of annuli increased with fish size, and

annuli maintained the same position on the scales of fish

of different ages. Subject to minor deviations discussed

below mean back-calculated lenyths agree with mean

observed lengths at each ave. Hence the back-calculations

validiate the lechnique at least for the first five age ckasses,

Older ave classes would need validation by an independent

method."

The means of back-calculated lengths at age Were

plotted by using the graphics facility of the Lotus 1-2-3

microcomputer package and a von Berlalan!'fy growth

curve was then fitted jnteractively by eye. The von

Bertalantl'y equation for the length L, in cm at age | in

years is:

L, 46.0 [I-exp(—0.22 (L + 0.5)

Table | presents the mean length at capture (those in

the i+ age group are aged between i and i+ years), the

back-caleulated length, their means and the fitted von

Bertalanffy points. The data and the fitted curve are shown

in Fiz. 2.

40 ae.

— +

= 30 a

E co ™

2 Pa

z "

2 20 y

=| rf. =

7 aé

7} 4 —

“10

ie} | A. 4 + 4 4 — ——

2 4 6 8

AGE CLASS (years)

Fig. 2. Plots of true total length at age (a dash indicates

the age class), lengths at age from the back-calculations

(crosses), and (he fitted von Berralantfy curve for P.

TPIFICUS.

The length weight relationship, determined by regression

aualysis Wak W 0.052 L270" (Re OBS; No» 61)

where W is the weight in g and L the total length in em,

The mean back-calculated lengths up to age 4 for fish

aved >4 years ure vrealer than the true length (Table 1).

This effect, if true and not an artifact of sample size, 1s

the reverse of Lee's phenomen’’*, Possible causes are:

(1) Hon-random. or biased sampling in. which slower

zrowing fish are under-represented; this may arise because

of (he gear used or the habits of the fish; and (2) selective

natural or fishing mortality’. P rerricus is not sought by

commercial or recreational fishermen (although ibis olen

a subsiantial part of the catch taken incidentally by

reereational anglers”), and the sample was uiken by

178

spearing; hence is unlikely that selective fishing mortality

occurs. However, selective natural mortality in which faster

growing fish survive better could occur. Alternatively

biased sampling may have occurred because the largest

samples were from catches of spear fishermen who selected

the largest individuals for the purpose of spearfishing

competitions.

Some labrids show a point of inflexion on the growth

curve upon the transition from female to male!!"!, This.

growth spurt is thought to occur because energy hitherto

used for egg production can be invested in growth. The

plot of true length at age for P tetricus (Fig. 2) shows

a slight inflexion at about 5 years of age, when sex

inversion occurs, but this needs verification by tagging

studies.

P tetricus appears to have a similar longevity to its warm

temperate congener of northern New Zealand P celidotus

(Bloch & Schneider)*.

We are grateful to J. E. Johnson and P. S. Clarkson

for collecting the samples and to the latter for their

preparation. Dr P. R. Sluczanowski fitted the von

Bertalanffy curve and with Dr G. K. Jones criticised the

manuscript.

TABLE 1. Mean back calculated total lengths (mm) at age for P. tetricus. N is sample size, mean length at capture

(mm) is the true length and calculated length (mm) is derived Jrom the fitted von Bertalanffy growth curve.

Mean total Von Bert. Annulus

Age N length at calculated total

capture (mm) length (mm) 1 2 3 4 5 6 7 8

1+ 2 125.0 129.3 102.3

2+ 4 166.0 194.6 101.1 158.6

3+ 10 224.3 247.0 120.8 181.3 222.0

44+ 8 282.4 289.1 136.6 208.2 227.2 265.9

5+ 7 354.0 322.8 167.7 214.8 254.3 293.3 327.4

6+ 13 376.2 349.9 146.5 196.1 246.4 294.6 334.5 371.3

7+ 6 377.7 371.7 163.5 212.6 243.7 283.0 304.7 332.6 356.1

8+ 8 388.8 389.1 154.9 203.0 231.6 275.1 304.9 338.2 366.4 390.6

Unweighted Mean 137 196 238 282 318 347 361 391

Weighted Mean 142, 198 237 284 321 353 362 391

'Last, P. R., Scott, E. O. G. & Talbot, F. (1984) “Fishes

of Tasmania.” (Govt. Printer: Hobart).

Scott, T. D., Glover, C. J. M. & Southcott, R. V. (1974)

“The marine and freshwater fishes of South Australia.”

(Govt. Printer: Adelaide).

*Tesch, F. W. (1971) in Ricker, W. E. (ed.). “Methods of

Assessment of Fish Production in Fresh Water.” IBP

Handbook No. 3 (Blackwell: Oxford).

‘Jones, G. P. (1980) Copeia (4), 660-675.

*Quignard, J. P. (1966). Naturalia monspel. ser. Zool. 5,

1-247.

Beamish, R. J. & McFarlane, G. A. (1983) Trans. Am.

Fish. Soc. 112, 735-742.

ses, R. (1958) Int Comm. NW. Atlantic Fish. Spec.

ubl. 1.

8Ricker, W. E. (1975) Bull. Fish Res. Bd Canada 191,

215-217.

°*Jones, G. K. (1983) SAFIC 7 (4), 9-18.

'OWarner, R. R. (1975) Fishery Bulletin 73, 262-283.

"Dipper, F. A., Bridges, C. R. & Menz, A. (1977) J. Fish.

Biol. 11, 105-120.

S. A. SHEPHERD and LISA J. HOBBS, Department of Fisheries, 135 Pirie Street, Adelaide, S. Aust. 5000.

RECORDS OF ROTIFERS EPIZOIC ON CLADOCERANS FROM SOUTH

AUSTRALIA

BY RUSSELL J. SHIEL AND WALTER KOSTE

Summary

In a review of the rotifer genus Brachionus (Rotifer: Monogononta), it was noted that some species

typically were collected as epizoites on planktonic crustaceans. For example, Brachionus rubens

and less commonly B. variabilis occurred on Daphnia and Ceriodaphnia species. Brachonius charini

was reported from eastern Europe as a commensal in the branchial chamber of Coenestheria davidi,

a phyllopod. B. sessilis was recorded as an epizoite of Diaphanosoma brachyurum, and the known

hosts of B. rubens and B. variabilis were listed as Daphnia magna, D. pulex, D. longispina, Moina

rectirostris, Ceriodaphnia sp., Polyphemus pediculus and several macroinvertebrates.

BRIEF COMMUNICATION

179

RECORDS OF ROTIFERS EPIZOIC ON CLADOCERANS FROM SOUTH AUSTRALIA

In a review of the rotifer genus Brachionus (Rotifera:

Monogononta), it was noted that some species typically

were collected as epizoites on planktonic crustaceans!.

For example, Brachionus rubens and less commonly B.

vartabilis occurred on Daphnia and Ceriodaphnia species.

Brachionus charini was reported from eastern Europe as

a commensal in the branchial chamber of Coenestheria

davidi, a phyllopod?. B. sessilis was recorded as an

epizoite of Diaphanosoma brachyurum, and the known

hosts of B. rubens and B. variabilis were listed as Daphnia

magna, D, pulex, D. longispina, Moina rectirostris,

Ceriodaphnia sp., Polyphemus pediculus and several

macroinvertebrates?.

Although B. rubens, B. sessilis and B. variabilis are

known from Australian waters‘, all were collected free-

swimming from the open water of billabongs (Magela Ck,

N.T., and Murray-Darling River system, Vic., N.S.W.).

Significantly, the host species listed by Koste? do not

occur here’. We document here the first record of a

facultatively epizoic brachionid from Australia, and add

to both the known epizoic taxa and their “host” species.

Fig. 1. LP SEM view (1.01 * 10? x) of B

novaezealandiae epizoic on the head (dorsal) of

Pseudomoina lemnae. Staurastrum (alga) and attached

ciliate protozoans also can be seen.

{In a collection for phytoplankton taken from a farm

dam at Hahndorf, S, Aust. (27.vii.83) a rich population

of planktonic cladocerans hosted a population of

brachionids. The predominant cladoceran was

Pseudomoina lemnae, with Daphnia carinata s.l. and

Ceriodaphnia sp. also abundant. Most larger individuals

carried from 5-40 female rotifers attached by sticky foot-

gland secretions to the hosts’ carapace; most of the rotifers

were ovigerous, with 1-2 large amictic (parthenogenetic)

eggs.

Some animals with attached rotifers were preserved in

4% formalin and prepared for light- and scanning electron

microscopy (SEM)*. Representative SEM micrographs

are shown in Figs | and 2. The rotifers have contracted

into their loricas in response to the preservative, and some

distortion of the lorica is evident, a result of the desiccation

process for SEM. However, the taxonomically significant

ventral margin is clear on the micrographs. The caudal

lorica margin and foot-opening were examined in PVA-

mounted preparations. The elongated anteromedian spines

and absence of a flap over the foot-opening distinguish

the specimens as Brachionus novaezealandiae (Morris),

which, despite its name, is widely distributed in sodium-

dominated waters in the southern hemisphere and may

be cosmopolitan in this biotope’.

B. novaezealandiae is not listed as epizoic by Koste?,

however Morris’ original description (= B. variabilis yar.

novaezealandiae) notes that his material came from a pond

at Totara, N.Z. “where they were in numbers, parasitic on

Daphnia thomsoni’*. The type slide was found in the

collection of the British Museum (Natural History), and

although the coverslip was broken and the specimen

somewhat desiccated and distorted through loss of

mountant, it was identifiable clearly as the specimen from

which Morris’ figure was drawn. The slide is labelled

“Proposed sp. nov. Brachionus novae-zealandiae C. B.

Morris 1917.1.1 (all Parasitic on Daphnia) Totara, N.

Otago. Formalin 25.10.11”. Comparison of the South

Fig. 2 Anterior view (1.86 x 10° x) of two B

novaezealandiae, same host. Animals are contracted.

Single parthenogenetic eggs carried. Scalar 100 um.

180

Australian epizoic form and the type material indicates

that the pronounced caudal spines are reduced in the S.

Aust. specimens, but are within the range of variability

of this species reported from R. Murray B.

novaezealandiae?.

Undoubtedly rotifer-cladoceran associations will be

found to be more widespread among the Australian

Rotifera with further study. A diverse assemblage of

phytoplankton and protozoans, some of which can be seer:

'AhIstrom, E. H. (1940) Bull. Am. Mus. Nat. Hist. 77,

143-184.

2Kutikova, L. A., Kosova, A. A. & Khodorevsky, O. A.

(1976) Zool. Zh. 55, 763-767. [Russian].

3Koste, W. (1978) Rotatoria. Die Radertiere Mitteleuropas

(Uberordnung Monogononta). 2 vol. revision of M. Voigt

(1956). (Gebr. Borntraeger: Stuttgart).

4Shiel, R. J. (1983) Proc. R. Soc. Vict. 95, 33-37.

in Figs 1 and 2, also occurred on the three cladoceran

genera examined. Such epizoite communities are virtually

unstudied in Australian inland waters.

We thank Brendan Atkins, then a Botany undergraduate

student, University of Adelaide, for bringing this material

to our attention, and Charles Hussey, British Museum

(Natural History) for his hospitality and assistance during

a visit by one of us (RJS) to examine the BMNH rotifer

collection.

5Smirnov, N. N. & Timms, B. V. (1983) Rec. Aust. Mus.

Suppl. 1, 1-132.

6Koste, W. & Shiel, R. J. (in press) Aust. J. Mar.

Freshwater. Res.

7Pejler, B. (1977) Arch. Hydrobiol. Suppl. 53, 255-306.

8Morris, C. B. (1913) Trans. Proc. N.Z. Inst. 45, 163-167.

9Koste, W. (1979) Aust. J. Mar. Freshwater Res. 30,

237-253.

RUSSELL J. SHIEL, Dept of Botany, University of Adelaide, G.P.O. Box 498 Adelaide, S. Aust. 5001 and WALTER

KOSTE, Ludwig Brill Strasse 5, Quakenbriick D-4570, Federal Republic of Germany.