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VOL. 109, PARTS 1 & 2 
28 JUNE, 1985 

Transactions of the 

Royal Society of South 



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

Australian waters - 1 

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

(Hymenoptera: Formicidae) in the Mt Lofty Ranges, South Australia 17 

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

from small Dasyuridae (Marsupialia) - 25 

Barnett, B. & Schwaner, T. D. Growth in captive born tiger snakes (Notechis ater 
serventyi) from Chappell Island: implications for field and laboratory 
studies 31 

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

Leptodactylidae) 37 

Edmonds, S. J. A new species of Phascolosoma (Sipuncula) from Australia - 43 

Tyler, M. J. & Miller, C A. Surface architecture of the dorsal epidermis in Australian frogs 45 

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

Australia ----------- 49 

Brief Communications: 

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

Scincidae) in South Australia ------- 55 

Smith, J. A. Tides of the Onkaparinga estuary, South Australia 57 

Glover, C J. M. Additions to the fish fauna of South Australia - 59 

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 ------ 61 






VOL. 109, PART I 


CONTENTS, VOL. 109, 1985 

PARTS 1 & 2, 28 June 

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

Australian waters 1 

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

(Hymenoptera: Formicidae) in the Mt Lofty Ranges, South Australia 17 

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

from small Dasyuridae (Marsupialia) ------ 25 

Barnett, B. & Schwaner, T. D. Growth in captive born tiger snakes (Notechis ater 
serventyi) from Chappell Island: implications for field and laboratory 
studies - 31 

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

Leptodactylidae) - 37 

Edmonds, S. J. A new species of Phascolosoma (Sipuncula) from Australia - - 43 

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

frogs ------- 45 

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

Australia ------ 49 

Brief Communications: 

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

Scincidae) in South Australia ------- 55 

Smith, J. A. Tides of the Onkaparinga estuary, South Australia 57 

Glover, C J. M. Additions to the fish fauna of South Australia - 59 

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 ----------- 61 

PARTS 3 & 4, 29 November 

/tidier, 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 

darlingtont (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 factes 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 

McNamara, 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 -----------177 

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


Insert to Transactions of the Royal Society of South Australia, Vol. 109, Parts 3 & 4, 29 November, 1935 



by Russell 7. Shiel & Walter Koste 


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. 



by Ri'SM i i J. Shu i '* & Wai if k Kosti - + 


Sililt , R. X. iV Ki'Mi , \\ (I9X5| New species and new reeouK n| Kolilera f AschHuimrlt.-,) |t -m VuwftJlSh 

mpefn. nuns. i-t. Sm s. Ann, uw), i i\ :k itmc, was, 

One hunuicd;t! ; e sIm\. seven laxaaic;tdded roilie fWrilM i^conlc-d lumi Ah-.mjVui w.ikts Nlitfiu 
1 lie total lomci (.(Hi. hfaft tac;i described and figured ore? U'ftiuIflUi fwtelUt vai nov., h'<<wr/i>s>mmtr,-iT 
M>, iios., /. tttimtuutti v»r. nm., Asplan<-)unt btigktwKU asvftit/n'irnu Ssp, inn., and fcmulbifitit} AutSi Vi 
Sift nov. hi addition, Hvxitniuv tVQ/tirti iSernov) U ivdescrihed and figured and its svnmmm il.swi-.t-d 

Ki | Wi.kU 1 -- KiniK'i i, mlv vntficics. new tafttiJV. XiMnlu 


Although knowledge of the systemnties SHEd 
ecology cT Australian Rolifern has improved 
considerably in recent years, il remains frat'meiuary. 
Ineluninarx laxonomie works (Shicl & Kosic 1979, 
Kosic & Slnel I9K081, b) stemmed liom a suidy of 
the /ooplankton of the Murray-Darling basin, From 
samples collected alone the Mauela Creek, 
Northern Icmlotv (Koste IVS1, Kosie&Shiel 1983, 
fail ef a/. I9S4) and in southwest Western Australia 
(Brptk&Sfiid 1983. kosic tv«/. II&3J. The rotifer 
fauna of most ol Australia remains unstudied. 
While this ean be attributed in part lo ;i .shortage 
ol Wfift*ro|Oj5&t$J rather lhan of rotifer-, (ef. 
Oumoiu J9X3), there are undoubtedly more 
collectors of /ooplankton in Australia than the 
rotifer records indicate Systematic difficulties 
appear to be a major obstacle. 

We noted earlier (kosic & Slnel i9N0a) rhnl major 
.systematic works deal with the northern hemisphere 
fauna; there is limited miormano n on the Rotilcra 
of ihc southern continents. Indeed, a global revision 
ol i he ClaSS ii desirable, particularly since the 
recognition of ecotypic variation in response to 
habitat variability. In ihc hraehionids, for example, 
extensive intraspcvilic variation in si^c. lorica 
moiplmloev or spine length has been demonstrated 
in response io habitat temperature, pi I. salinily Or 
predators, (c.e. Gilbert 1967. C.icen 1977, Colbert 
& Stembereer 1984). I his raises the difficulty o\' 
specific determination when the exient of inlra 
specific variability is not known lor most rotifer 
venera, and accounts in part for the proliferation 
of "species" based on vcrv limited matciial. 

h> help overcome laxonomie ptoblems we 
commenced a comprehensive illustrated key lo Lhc 

IXp.niinenf of Houinv, I. ol Adelaide, Bw> 4*>>:. 
w I'd,. AdclayJe, s. Ana. sum 
Uuhvip Hull sn.tssf s t Ottiiknibr&tk, t) 4?7n. laio-;il 
Kl | Mihlic (»l Cit-rmatry, 

planklomc monotronom Rotilcia i.coi.t t d trimi 
Austialian waters. In view oi complex loliler 
assemblages, pauieulailv in tropical Australia, \m* 
have tound it necessary to extend the ke\ IU ioclutlc 
facullaiively or occasionally planklomc m\.i ( 
including Bdelloidca. The first diagnostic |<Cft have 
been prepared (Kosic iV ^hid in press a,bl. bur an 
entire ivvision may take several veais. Ileie we 
updale the species list and describe new ia\a fn»m 
our (and other) collcciionv 

In the fust checklist, Slnel & r\o>ie (19791 
recorded 331 ra\a, in 73 genera. A further 106 lasa 
were listed b\ Kosic l\: Slue! (l9S()a). One of these, 
cited as lilinia fiof/tnatim (stvi il, hoftHa/tr, 
Kosic. 19X0). was an cimi: it i- lt l-inoixan species. 
To the 43f» known ta\a ue add hett a lorthe/ \(r, 
collated Irom the literature since I9S0, rn>H1 
material made available \£> us by others, and ftotn 
our collections. An importani '.onuilmlion nas 
made by Iktvins (19N2), who prodded li livi of 
rotifers identified from collections made in 
Australian waters oxer a petioci ol 30 years. 
o\' these a'e first recoids i'ov the continent, and a 
numbei of species are new. However several species 
deseitbed as new appear to be synonymy .jttelacs 
o\ preservation oi misidentificuiiuns. Thc-c aie 
discussed m Ihetext. Inl'rasiibtpced'ie tavi hiiherio 
described as f. or var. Mrc? dcsciibed in ivialion tu 
the nominate s>p. >imp!v Co dueomcul ^an.ibduy 
ami are not named. Mole detailed ceoloiiiLa! d.u.i 
aud/ot material will lacrhtale a later icvision. 

The format follows thai of Koste t V Sliiel (lySOhl 
/\!1 records are listed -systematically rbtlowiflg 
Dormer (1965) for the lidelloidea and Kosic (J97xb) 
for the Vlonocronon!,.. Lucahly (o\ neaiest named 
feature) and date o\' collection arc eicen Rartg.e> 
of water quality are pven wtica available HublLslicd 
records art cited bv aulluu, unpublished records by 
colleeior, (Jnlv new uisa are filmed I..;^ and 
paraiypes are todeed with ibe Snnih \usfi, t i, i 
Museum tv\M,i. 




Family Habrotrochidae 

Habrotrocha bidens (Gosse), 1851 
Loc: Bombala, N.S.W., in tree stump mould; 

28.xii.1949, 8.1.1950. 
Lit: Berzins (1982). 

H. cucutlata Murray, 1911 
Loc: Bombala, N.S.W., creek moss; 8. i. 1950. 
Lit: Berzins (1982). 

H. fusca (Bryce), 1894 
Loc: Bombala, N.S.W., tree stump mould; 

Lit: Berzins (1982). 

H. ligula Bryce, 1913 
Loc: Bombala, N.S.W., tree stump mould; 

Lit: Berzins (1982). 

H. microcephala (Murray), 1906 
Loc: Melbourne, Vic; moss and lichens; l.ii.1951. 
Lit: Berzins (1982). =77. constricta according to 

Dormer (1965). The latter is known from N.S.W. 

(Murray 1911). 

H. tranquil/a Milne, 1916 
Loc: Bombala, N.S.W., tree stump mould; 

Lit: Berzins (1982). 

H. tripus (Murray), 1907 
Loc: Yarra R., Ivanhoe, Vie., in terrestrial lichens; 

Lit: Berzins (1982). 

Family Philodinidae Bryce, 1910 

Embala 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.S.W.; 11. xi. 1953. 
Lit: Berzins (1982). 

Pleuretra brycei (Weber), 1898 
Loc: Geraldton, W.A., moss/humus; undated, 

Lit: Berzins (1982). 

Macrotrachela aculeata (Milne), 1886 
Loc: Bombala, N.S.W. , tree stump mould; 

Lit: Berzins (1982). 

M. angusta (Bryce), 1894 
Loc: Bombala, N.S.W, tree stump mould; 

Lit: Berzins (1982). 

M. concinna (Bryce), 1912 
Loc: Geraldton, W.A., moss/humus; undated 

Lit: Berzins (1982). 

M. gunningi (Murray), 1911 
Loc: Melbourne, Vic, moss/lichens; l.ii.1951. 
Lit: Berzins (1982). 

M. nana (Bryce), 1912 
Loc: Yarra R., Ivanhoe, Vic, moss/lichens; Li. 1951, 

Lit: Berzins (1982). 

M smithi Milne, 1916 
Loc: Bombala, N.S.W., moss; 8. i. 1950. 
Lit: Berzins (1982). 

M. speciosa (Murray). 1907 
Loc: Melbourne, Vic, moss/lichens; l.ii.1951. 
Lit: Berzins (1982). 

Mniobia circinata (Murray), 1908 
Loc: Melbourne, Vic, moss/lichens; l.ii.1951. 
Lit: Berzins (1982). 

M. conarus Berzins, 1982 
Loc: Heyington, Melbourne, Vic, moss/lichens: 

Lit: Berzins (1982). 

M. I i neat a Rahm, 1932 
Loc: Heyington, and Yarra R., Ivanhoe, Vic, moss; 

14.1.1951, 14.iii.1976. 
Lit: Berzins (1982). 

M. magna (Plate), 1889 
Loc: Bombala, N.S.W., moss/lichens; 8.1.1950. 
Lit: Berzins (1982). 

M. ocypetes Berzins, 1982 
Loc: Melbourne, Vic, moss/lichens; l.ii.1951. 
Lit: Berzins (1982). 

M. scarlatina (Ehrenberg), 1853 
Loc: Melbourne, Vic, moss/lichens; l.ii.1951. 
Lit: Berzins (1982). 

Family Adenitidae Bryce, 1910 

Adineta cuneata Milne, 1916 
Loc: Bombala, N.S.W., tree stump mould; 

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 ai (1981), Koste et al (1983). 

AIM KM | \\ K(»||| 1 KA 

family Brachiohidae Wesenbei^UittUi |889 
Bmchkmm twndQius per&orwius \hlstrom, (940 

I oc: VViiunuria BillahoiK-, Ma.ccla c 'k , near 

.labilnka, NT,; l5.iv.IV8G. 

pH 5.4-4. 28.5°( , 6.: m^l ' G„ 23 ^s an ' 
LH: Kostc (1981). 

ft flilcatux I. reduoius Kosie & Shiel, 1983 
l oca WTnmtirra Billaborig, near Jabilidra, Manela 

c k. NT., 4.\ii.iyso. 
I -ii: Kosic & Slue! (t^MJ, 

IS: tatftf Shid. p>s3 

I oc: Shccpuash Billaboiig, Yea, Vic; l9.t.M$Z 
pll 7.0-7.7, 24 X \ vat O,, < 50-200/iS cm ', 
sol'i macmesitmi bicarbonate wale™, high Iram- 

parencv < ■ i nth). 
I It: Shiel (*983>, 

S pinneenaua Kosic & shiel. |&j3 
Loc- Cue, sonihwesi W.A.J viii.1978. 
I .il : Geddes ?/ <//. (I9S1). 

Ii mnhlndetuutasuncylowuHiws (Sclimarda), 1859 
Loc: I.. Bullcnmerri. Vic; 22.ivl98.3 
(oil: I, |. Pouliny. 

Ii tltiudtult'Htuttis rlwtnwns (Lmilcrborn), lg93 
Loc: I . Bnllcnmeiii. Vic; 22.-ix.I983 
(oil: I, J. Knvlitig. 

ft QwdridentQlu% minor Kostc & Shiel, 1983 
I oc: BilluhOrife» ) Ma.uela Ck, NT,! l3.vl,l979; I ake 

Grace area, SOUtftWCM W.A.; \.i 1981 
Lit: kostc vt a!. (1983), 

keruK'ila tfltadwa (lisfK'fsa (C'arlin). 1943 
Loc: Vina K. ncai I van hoc, Vic; I4.hi.l97& 
I it: Hci/iiis (I9S2). 

Laitnly Luchlaiiidae Marios, 1059 
I . pyrijormis Gosse, 1S5I 
Loc; Moorabooi «.. Hallan. Vic,S ft.vii.1954, 
I it: Her/ins (1982). 

OlptGHOftfatm ptcg&m <Y\ ier/cjski). 1X93 
loc l , ( afani, Mi Buffalo, Vic ( -P. fui/utitisu 

Mauci. 193d); 25.ii.!95S. 
J ii: Berlins (1982). 

Q propaitth mtwrodaety/a (Rawer), 1965 

I .a. Maccla ( k. N.I., 15. i\. 19X0, Yuira K. cast of 

Warbnrton. Vic; 19. Hi. 1976 
I it: Kostc (1981), Ber/ins (19X2). 

\ltui}n<hum vudactylotuut (Gossc), 1X86 
Lou: Magela < k., N.L. r5.iiv.198d; lake Gracc- 
Noftetnan area, sOUifivvtwl W.A.; viii.1978, \.I9XI. 
I It! KOStC (1981), Kostc cf ai ('1983). 

Lamilv Mytilinidac Barto>. 1959 

Myiifuia Uiunthaphoru Haiici, 1938 

Loc; Solomon Dam, Lake Moondarra, Old. 

Coll: P. Hawkins, [. On. 

A/, hisukata (Lucks), 1912 
loc: Yarra R.. DigJu'S l-alls. Vic; I2.iii.l97d 
I il: Bcr/nis (1982). 

M vcntralis t. tonzirfaciylu Wullcrt, 1965 
I OC! Mat-cla (k near Jablluka, N.L; 
Ill: Kostc (19X1). 

Lophovhmu vurv&ta Beralns, 1982 
LOCI Kiii^ Patrol Ck, Kinglake, Vic; 18.\.]953. 
Lit: Ucr/ins (1982). Inadcciuately figured ami 
desctibed. May be a formalin artefact, 

/.. oxY\!vrnum (Gossc), 1851 
Loc Jackson's Creek, Moorabooi K.. \'ic; &KU96J 

I it: Bci/ins (t9S2). 

Lamilv Tt ictioniidac Barlos, |9S9 

Trichoma retmctti var. imtlis tsicnroos), iS9s 
loc: Magela Ck near labiluka. N.I.; l5.i\.19So. 
Lit; Kostc (1981), 

Miicmclwetus uliumirui (Arevaln). I9IS 
Loc: Riddel's ( k. Sunbury, Vic; 06.hi.l9M. 
Lit; Bcr/ins (19X2) (s>n: A/, austral tens is Bet/ins), 

M. dannaell Kostc $ Shiel, 198 3 
I oc Ltnltalo Billabony, MagelaCk, NX; IO.\i.l9SO. 

pH 53S, 30.9 (, 2 4.15 ppm, 29 MS cm '. 
Lit: Koslc& Shiel (1983). 

Family ( olurcllidae Bartos, 1959 

( t'htidla vohtnis (bhrenberg), 1830 
Loc; Mooiabool R (1 Yarra R., Mt Donna Buanc, 

Vic; 9-.vii.1954, 25. i\. 1962. I2.iii.l976, 
Lil: 13c. /ins (1982). 

C H'sst'Uitu ((.lascott). 1983 
Loc: C0gh)lP£ Ck, Vic; I1.\i.(953. 
1 ii. Bcnil'OA (1982), 

C. huiispirtam utuituMci (Llnenbcru), 1832 
Lot*: widespread. Victoria, perennial. 
Lil: Ber/ins (1982), 

/ t'ljudelta acuminata si?\a>\nua (BartOiJ, 1955 
Loc: Couhill's C'k; I . C Mi BulTalo, Vic; 

ILvi.1953, 25.ii.1955. 
I it: Bcr/ins (1982). 

L. acuiufint/a M'p/e/ucoMunt Bet/in-., 1982 
loc: L. Catain, Ml Bnl'lalo, Vic; 25.n.l9_vv 
I it: Bcr/ins (1982). 


L. amphitropis Hairing, 1916 
Loc: Barker's Ck, Castlemaine; Yarra R., Vic; 

21.iv.1953, 19.iii.1976. 
Lit: Berlins (1982). 

L. amphitropis victohensis Berzins, 1982 
Loc: Middle Tarwin R., Vic; Ol.v.1953. 
Lit: Berzins (1982). 

L. apsicia Harring, 1916 
Loc: L. Euramoo, Qld; viii.1978. Magela Ck, NT.; 

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. 11, Fig. 5) and we consider 
L. buangensis a synonym. 

L. chorea Berzins, 1982 
Loc: Hunter R. near Gundy, N.S.W. (?ll.xi.l953). 
Lit: Berzins (1982). Also very poorly figured, and 
possibly a formalin artefact. 

L. decora Berzins, 1982 
Loc: Loddon R., Vic; 
Lit: Berzins (1982). 

L. latusinus f. mucronata Koste, 
Loc: Magela Ck near Jabiluka, NT; 
Lit: Koste (1981). 

L. lindaui Koste, 1981 
Loc: Winmurra Billabong, Magela 

pH 5.44, 28.5 °C, 6.2 mg X > ; 
Lit: Koste (1981). 

L. minorui Koste, 1981 
Loc: Leichhardt Billabong, Magela Ck, NT; 

pH 5.55, 30.1°C, 5.53 mg X ' 2 , 58 jt*S cm" 1 . 
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, NT; 

pH 5.47, 29.9°C, 5.45 mgV 1 2 , 44 ^S cm 1 . 
Lit: Koste (1981). 

Lepadella patella patella (Muller), 1786 n. van 

FIG. la, b 
Loc: Buffalo billabong, Magela Ck, NT; 8.xii.l980. 
Coll: R. D. Tail. 
SAM V3941 

In this sample were four Lepadella with a circular 
lorica which resembled L. patella (Muller) described 


Ck, NT; 
23/xS cm '. 

Fig. 1. Lepadella patella (O. K Muller) 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 NT. 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 Jim, lorica width 62^im and toes 21 ftm, whereas 
the Buffalo Billabong form is 98 fun long, 92 ^m 
wide, with toes of 24 fan. 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. 

At S1K.M IAN KUilt IK \ 

/. patella hihhu (Hauor), 1958 
Loc: Yarnup Swamp, near I . LJnicup. soulhwesi 

V\.A.; 8 v.1981. 

17.0 ( , 1600 /XS em '. 
lit; KflsW el til. (1983). 

/- tpnntrtcarinala iSxauoos), 1898 
loc Coglnll's Ck, Vic; 1953. 
I. it: Dffryjns (1982). 

/.. i/thtdnvunrtuta \e\earinma Klcnicni. 1959 
Loc: CoghillN Ck. Vic; 1953. 
I h: Bei/ins (I982L 

/- ftawtibitrui (Lucks), 1912 
Loc RiddtTsCk near Sunburv, Vic; A.; 6. in. 1961. 
I ii: Bei/ins (1982). 

/.. //y/w M>ei\ 1934 
loc: Siralhpine pool, Brisbane, Qld; 2l.iv,l950. 

I to. pecans (I9S2). 

I leterolepafletlu heterorfuelyla Ladccw, 1925 
L0£ Ma,HL'la C k near Jabihika, N.L; 15 iv. 1980 
I II: Kosle (1981). 

L.imilv Lecauidac Battos, 1959 

/.. (Hviitimonostvla) inopinata I. s w/w/t/ (Llauei ), 

I oc: Can ns, N.Qld 

Coll: R, Hamond (1X*I. /. wmpodu bv C. II. 

L. hifatxtt en tome Bcr/ins. 1982 
loc Creswick Ck, ( lunes, Vic; ILiv.1953. 
I il: Bcr/ms (1982). 

/- India xoniuid (Marring & Myers), 1926 
Luc Jackson's Ck, Sunbm v, OYuii 61; Moorabool 

R-, Ballan, Vic; 9.vii.l954. 
Lit; Bcr/ins (WM'|, 

/_. capeis (Warring & Myers), 

loc: Nana R., near Yairagleu, V-c; 18. Hi. 1976. 
I n Ber/ins (I9S2). 

I . deeipiens (Murray), 1913 
I or: ( aims, QUI. 
Cull: \>, \ Tvlci (Del. C M leuiando). 

/. elaelris Hairing & Myers, 1926 
loc Mine Valley Billaboim near Jabihika, NT; 
Cull: R. I. Shiel. 

£ T laniellakt thalera (Hairing & Myers), 1926 
Loc lorcklale Lagoon neai JVrlh, \\'.\. 
C oil: J. van Alphcit. 

/. lunaris ausl raits Ber/iih. 1982 
loc I . Catam, Ml Buffalo, Vic; 25 h. 1955. 
1 il; Bei/ins (1982). Doubifnl, poorly figured. 
?lormalin artelael. 

/-- lunaris consthcta (Murray), 1913 
Loc: R. Murray, Blaneheiown, S.A. 7.V.I9S2. 
Coll: R. J Shiel. 

/ . lunaris perplexa Ahlsiroin. 1938 
Luc: Vana R., Yarru Cileu, Warburion. Vic 14, Is 

Lit: Bei/ins (1982). 

/.. monosty/a (Dueiay), 1897 
loc: I. I uramoo, Qld; viii.1978. 
1 il: Green (1981), 

/.. ohtttsa (Murray), 1913 
I oc L. Uuramoo, Qld, viii.1978; CoghilPs Ck, Vic; 

I il: Green (1981), Ber/ins (1982). 

/_. opias (Harring & Myers), 1926 
Loc: Nana R., McMahon's Ck, Vic; 19.m. 197ft. 
Lit: Ber/.ins (1982). 

L. perpusilla (Haucr), 1929 
Loc: Yarra R., near Ivanhoe, Vic; Riii.1976. 
I it: Bcr/ins (1982). 

/. pyhjormis (Daday), 1905 
Loc: MagclaCk near Jabiluka. NT., !5a\.l9SO; I . 
Luraniuo, Bi oin field Swamp, Atheiton 
Tableland, Qld, viii.1978; souiliern Vic. 
ILiv.1955, 1953. 6.iii.l96l. 
Lit: Koste (1981), Green (I9SI), Bei/ins (1982). 

/. scutata [Httrring cV Myers), 1926 
Loc: Magela Ck I'loudplain, NTh l5.iv.|9S0. 
I u : Kosle (I98l). 

/. teihis (Harring & Myers), I926 
loc: I . liiiramuo, Alherion tableland, Qld; 

I .it: Green (I98I). 

A. (M.) uagaltata ladecw, 1 925 var. nov. 

Mc. 2a, b 

l oe: Billabon^ o\' Coleman R.. Cape Yurke. Old 
Coll: B. V. Timnis. 
S\M V >944 

A MUijIe individual of (bis species was identified 
only from a ligure^iven by Hauer (I938, Liy. 7la, b) 
(see also Kosle I978. I ig. 80, 6a-cJ. Tbe species is 
rare, but seems to be cosmopolitan. Wulfeit (1966) 
described variable ecotypes from the Sokolu R., W. 
Afne;i. and from Indian waters. The characteristic 
anterior projections of the lorica were not visible 
or were absent on the Cape Yorke variain 
Measurements: Venlral lorica length HW/xm, ventral 
lorica width 83 /xm, dorsal loiica length 94 /nn, 
dOTsal lorica width 72/xm. iocs inch claws40^m, 
claws I2^m. 


Fig. 2. Lecane (Monostyia) unguitata Fadeew, var. nov. a, ventral, h. dorsal 

Lecane (s. sir.) 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.ii.1955. 
Lit: Berzins (1982). 

L. boorali Koste & Shiel, 1983 
Loc: Meekatharra, W.A.; 20.viii.1978. 
Lit: Koste el al (1983). 

L. clara (Bryce), 1892 
Loc: Moorabool R., Ballan, Vic; 9.vii.l954. 
Lit: Berzins (1982). 

L. curvicornis nitida (Murray), 1913 
Loc: Magela Creek floodplain near Jabiluka, NX; 

Lit: Koste (1981). 

L. formosa Harring & Myers, 1926 
Loc: Hunter R. near Gundy, N.S.W.; 11. xi. 1953. 
Lit: Berzins (1982). 

L. hastata (Murray), 1913 
Loc: Lake Grace-Meekatharra area, southwest 

W.A.; xi.1981. 
Lit: Koste et al. (1983). 

L. inermis (Brycc), 1892 
Loc: Creswick Ck, Moorabool R., Serpentine Ck, 

Vic; ll.iv.1953, 9.vii.l954, 22.iv.1953. 
Lit: Berzins (1982). 

L. levistyla (Olofsson), 1917 
Loc: Magela Ck floodplain near Jabiluka, NT.; 

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; ll.iv.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.iii.l961. 
Lit: Green (1981), Berzins (1982). 

L. pyrrha Harring & Myers, 1926 
Loc: Dam, Beaufort, Vic; 04.xi.1953. 
Lit: Berzins (1982). 


/.. rhyiida Harring & Myers, 1926 
I.oc: Yatra R., McMahon's Ck, Vic; 19.iii.I976. 
Lit: Bcrzins (1982). 

A. stichea f. intrasiimala (Oloisson), 1917 
I.oc: Winmurra billabong, Magela Ck, NT.; 

Coll: R. D. Tail. 

L subitlis Hairing & Myers. 1926 
l.oe: Winmurra billabong, Magela Ck, NT.; 

1 5. i v.l 980. 
Coll: R. D. Tait. 

Lecane tasmaniensis sp. nov. 
FIG. 3 a,b 

Maicrkit: II contracted females, sample No. 1120, 
in formalin. 

Hoiotype: loricate female, sample No. 1120. Coll: 
22.iv.l984. R. J. Shiel. SAM V3942. 

Ptrmrypes: SAM V3943, 

Type hcafify; Roadside pool, button grass plain 
near turnoff lo McAuliffes Weir on Lake Si Clair 
road, Tasmania (42 il'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 
plalc; ventral plate with distinct transverse ridge 
across first toe segment from which parallel lines 
run towards the anterior margin (Fig. 3a): posterior 
segment rounded eaudally; eoxal plates medium, 
rounded; first foot joint rectangular, distinct; 
second foot joint square; toes less than half body 

lie. 3. Lecane ia,sinunien\is sp. nov. a, ventral, b. dorsal, 

R. J. SH1LL c^ W. KOSTE 

length, parallel-sided ending in short claws with 
basal spicules. 

Measurements (/im): 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. 
eurvicomis (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. eurvicomis 
-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. tudico/a Harring & Myers, 1926 
Loc: Dam, Beaufort, Vic; 4.xi.l953. 
Lit: Berzins (1982). 

L. venusia Harring & Myers, 1926 
Loc: Coghill's Ck., Vic; 
Lit: Berzins (1982). 

Family Proalidae Bartos, 1959 

Bryceella voigti Rodewald, 1934 
Loc: Bombala, N.S.W., moss and tree stump mould; 

8. i. 1950. 
Lit: Berzins (1982). 

Proales do/iaris (Rousselet), 1895 
Loc: Riddel's Ck, near Sunbury, Vic; 6. Hi. 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 aL (1983). 

Family Notommatidae Remane, 1933 

Monommata maculata Harring & Myers, 1924 
Loc: Magela Ck, NT., 14.iv.1980; L. Grace- 
Norseman area, southwest W.A.; ix.1981; dam 
near Chillagoe, N. Old. 12.vii.1983. 
Lit: Koste (1981), Koste et at. (1983), Qld material 
coll. B. V. Timms. 

Dorystoma caudata (Bilfinger), 1894 
Loc: Yarnup Swamp, southwest W.A.; 8.x. 1981. 
Lit: Koste et ai (1983). 

Ultra viridis (Stenroos), 1898 
Loc: Reservoir, Trentham, Vic; 5.xii.l984. 
Coll: I. J. Powling. 

Eosphora anthadis Harring & Myers, 1922 
Loc: near L. Logue, Eneabba Rd, southwest W.A.; 

Lit: Koste et ai (1983). 

E. thou 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, NT.; 8.xii.l980. 
Lit: Koste & Shiel (1983). 

Reslicuia get Ida Harring & Myers, 1922 
Loc: Yarnup Swamp, southwest W.A.; 8.x. 1981. 
Lit: Koste et ai (1983). 

Notommatu cf. triangulata (Kirkman), 1906 
Loc: Magela Creek floodplain near Jabiluka, NT.; 

Lit: Koste (1981). 

Cephaiodella apocolea Myers, 1924 
Loc: Magela Ck floodplain near Jabiluka, NT.; 

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 
Loc: Clunes, Gippstand, Mt Buffalo, Vic; iv, v.1953, 

Lit: Berzins (1982). 

C. gracilis (Ehrenberg), 1832 
Loc: widespread, Vic: prob. perennial. 
Lit: Berzins (1982). 

C hoodi (Gosse), 1896 
Loc: Tarwin R., Vic; 1, 7.V.1953. 
Lit: Berzins (1982). Berzins also recorded C. remanei 

Wiszniewski frorrvCastlemaine, Vic. ( = C hoodi. 

For synonymy see Koste 1978b p. 351). 

C. intuta Myers, 1924 
Loc: Magela Ck floodplain near Jabiluka, NT.; 

Lit: Koste (1981). 

C. misgurnus Wulfert, 1937 
Loc: Magela Ck floodplain near Jabiluka, NT.: 

Lit: Koste (1981). 

Al'M K \l JA\ ttlTI II I RA 

C, mey,atocephala (Glascott). 1X93 
Loc: Wentwoilh Falls, N.S.W.; 22.\iXJ53 

Ik: Bcmns (1982), 

C. mvcrsi Wis/niewski, 1934 
I oe: Scrubby C k, WhiUlesea, Vic; 30.V.I955. 
I it; Ber/ins (19X2). 

C. nana Myers, 1924 
Luc: Creswiek Ck, Climes, Vic: 
Lit: Ber/ins (1982). 

C parasitica (Jennings), 1894 
Loc: Diggers c.'k. Ml Kosciusko, N.S.W.; Il.jv,l96l. 
Ill: Ber/ins (1982), 

C .WflOW (Gosse), 1887 
l.oc: Mage In ( k floodplain near Jabiluka. NT.; 

Lii: kosle (1981). 

C. tantiltoittes Hatter. 1935 
Loc: Bromficld Swamp, Qld; viii 1978. 
I ii: Green (19S1>. 

Family Irichoeercidae Remane, 1933 

Irichocerca hidens (Lileks). 1912 
I oe: Creswiek Ck, Chines, I1.iv.1953; L. Catani, Mr 

Buffalo, I0.xii.l962; Yarra R. Warbuilon, Vic; 

19.iii 1976. 
Lit: Itcr/jus (1982). 

/. ctulonta (Hauer), I93S 
loc: Mooiabool R. 4 Hallan, Vie.; 9,vii.|954. 
I ir Ber/ins (13*2). (7. myerst Hauer, 1939). 

'/: Jnxilonms Levandcr, 18 l H 
Log Magela Ck lloodpkun near Jabiluka, NX; 

Lit: koste {19X1). 

f. ?racili\ (lessin). 1890 
Luc: Solomon Dam, Old. Undated. 
Coll: \\ Hawkins. 

T. terms (Gosse), I8K7 
Loc: Yarra k., Warbtirton, Vic; 14, I9.iii.|976. 
I it: Bet/ins (1982). 

7' inennis (I inder), 1904 
Loc: Shcepwash Billahoiiu, Yea, Vic; 8.V.19S2. 
Coll: R. J. Shid. 

/. insukmu Hauci. 1937 
I oe: Magela Ck floodplain near Jabiluka, N.L; 

I il: kosle fISBli. 

T. mternwdiu (Slenroos), 1*98 
Loc: Bromficld Swamp, Old; viii. 1978. 
lit: Green (1981). 

/. ntaccra (Gosse), 1886 
loe: I. Caiain, Ml Buflalo, Vie.; 25.ii.l955: 

Gamboula Sin Dam, Cape Yorke, 12.vii.l983. 
Lii: Bei /ins (1982), Old malcrial eoll. R. V. Timms. 

T. moniana Hauer, 1956 
Loe: Winmurra Billabong, Magela Creek, NX; 

Coll: R, D. 'Tail. 

T. rnascahis (Ilauerh 1936 
Loc: Diggers Ck, Mt Koseiu.sko, N.S.W., I l.iv.1961: 

1 ake Giaee-Meeknlhaira area, southwest W.A.; 

iXi x.1981. 
I il: Ber/ins (1982), Kosle pj erf, (1983). 

T. rattus ehstata Halting, 1913 
Loe: Sunbury, WhiUlesea, Vic; 6.V.I955. 30.iii.I961. 
Lii: Bcmns (1982). 

J. ruttneri (Donner), 1953 
Loe: Dam near Chillagoe. Qld; I2.\h.l983. 
C oil: B. V. Timm.s. 

T. sulcata (Jennings), 1894 
Loe: Avoea R., Gowar Has!, Vic; 23. iv 1953. 
1 il: Bcmns (1982). 

T- ancinaia (Voigt), 19(12 
loe: Tabrabueea Ck, Bam'ngtou, N.S.W.; I2.xi.1953. 
Lit: Bcmns (1982). 

T. vemalis (Hauer), 1936 
Loe: Serpentine Ck, Bears Lagoon, Vic: 22.iv.1953. 
Lit: Ber/ins (1982). 

family Gastropodidae Remane, 1933 

Ascomorpha saltans imlica Wtilfen, 1966 
Loc: I i Barracoota, Vie., 20.1.1975. 
I il: Bcmns (1982). 

Family Synehaelidae Remane. 1933 

Synchaetu lackowif-jana Lucks, 1912 
Loc: Mooiabool R., Vic; 9.vti.l954; R. Murrav, 

Tailcm Bend, S.A.; I0.\.I9S3. 
Lii: Ber/ins (1982). S.A. material coll. R. L Shicl. 

Pvtyarlhra major Burckhaidt, 1900 
Loc: Coliban Res., kyneton, 06.iii.196l; L. Catini. 

Mt Buffalo, 25.ii.1955. 
I it: Ber/ins (1982). 

P. minor Voigi, 1904 
Loe: Magela Ck floodplain near Jabiluka, NX: 

tit: Koste (1981). 

Family Asplanchnidae Hairing & Myers, 1926 

\splunchna bright wclli asvmmctrka >pp. nov. 
FIG. 4a, b 
Material: 12 females, formalin preserved, sample 
#876; 9 females, sample #1005. 
fconotype: tropin, adult female. Lie. 4, sample //S76, 
coll. 4.U981 P. Hawkins. 
Paratvpes: SAM V3945. 

lype locality: Solomon Dam, Palm Island, near 
lownsviile, Old (18 "44 'S, 146 '35 I ■). 






Fig. 4. Asplanchna brightwetti asymmetrica ssp. nov, 

A, trophi apical, length 130 m, B, trophi compressed under 

Description: Typical saccate illoricate form; 
horseshoe-shaped vitellaria belonging to the A. 
bright we/fi-intennedia-siebo/di group (Koste 1978, 
Salt et ai 1978, Gilbert el ai 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^im (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 
(LI,): equal (equal); uncus (U 2 ): left only (2 
symmetrical); subterminal rami tooth: equal (left 

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 o^ the 

Fig. 5. Trophi of Asplanchna bri^htwelti (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| Hirst uncus; U^seeond 


ehaiaclerisiie asvmmefiical tropin in our opinion 
distinctly separate this [axon frprn 4. brightvcvf/i 
(C.osso). which ii moM resemble**. 

ispkH&hna herrhki De Gucrite, isss 
Lac: Albert Park, Vic 
i jn Evans [lMlj. 

Family Dicrnnophoiidae Kcmane 1933 

Albert* naidis Boitsfield, 1886 
Loc Maiida Ck floodplam near .kibikika. NT.; 

I5 r iv.I980i 
In: kosic (.1581). 

I V<7/</ "/"'/ (Hamn^). 1913 
LOCJ \arta K . Vic; I4an.l976. 
I ii- Bcrzins (1982). 

A. CtrcittUtar (Ciosse), 1886 
Loc. Plenty K. Morons s -. ViC;J 30.\.I954. 
I ii: Ber/ins (1982). 

A. th' Ber/ins, 1982 
LOCI Varra R, near Ivauhoe Vic: t4.lif.l97A. 
I it; Ikr/ins (1982). 

HaltUro rah'ns t tapaicde, [H67 
loc; Wimuuna Billabong, Ma-cla Ck, N ! ; 

I it: Kohc (1981). 

tXvnmaphbrm epifihark Hinting .v Myers, 1928 
Loci Magela Ck Floodplalh near Fahiluka, NT- 

1 if KoStC (1981). 

/>. hulhuciu iy»| 

i it. Wiimiiura Billabong, Magcla CL n;i,; 

I ii ki>Ste (1981) 

Q hcnulcs Wisniewski, 1932 
Loc* M;i'.vl:t Ck floodplam neui lahihik... N.I., 

I it: Kosu- (I9SD. 

IX fybHSlte Haiiinu S Mvers, 1928 
Loc: Noi-^ -KiiluoMiiic area* ^authwesi W.A.; 

I n: koste « fl/. CJS83). 

Lnccninon nmiion Ber/ins, 1982 
Loc: iiombaia, N.s.w. m moukl on a eucalyptus 

Mump. 2X.\ii 1949. 
In: Ber/ins (19X2). Doubtful SpctfcS Oil (he basis 
of poor figures. 

/:. fimsdettdrux Borzins, WK2 
Loc: liouibula, N.SW, In moukl on a eucalyptus 

slump, 2S.«ii.IM<t 
I ii; Bcrzins (W82). As above figures are inadequate 

I<m a new species' description. Doubtful record. 

Wlerzejskivtta vetox (Wis/mewski). 1932 

LoC; Yaimip Swamp. W.A., 8.X.1981. 
I il: Kme a al. (1983). 

family Jesiudmellidae Baitos, 1959 
Pompfto/vx uthulosit Ber/ins. 1982 

Loc L. Hurumbele, Vic: 29.vii.i953. 

lit: Ber/ins (1982). Poorly figured, '.'fornialin 
artefact, - P. complatuttu Gossc 

fcstudinellu xreeni koste, ) l >81 
loc: Leidlliardl Ril^bong. Mauela ( k, N [.: 

I it: Kosic (1981). 

lesttiilinvtlu husseyj sp no\. 
FIG 6:1a, h 

Mulerial: 13 formalin preset veil females, sample 


Hafatype; I adult female sample #998, coll. 

8*vit|9*3 B. V. I minis. SAM V 394ft. 

Pcnulvpcx SWI \ ■ 3**47. 

7V/V loiuiity: Dune lake near Cooklnwn. Cape 

Yorkc North Qkl (15 28 S, M5°15'E). 

Description: Sliapc of lorica oval; anterior margin 

with live tooth-shaped relatively long* keeled 

projections, twti on cither side of a median shorter 

tooth. In contrast, 7. /w//,/a fte/jra (Hermann), 

I7R3 (see Tig. 6:2) has only a soft rounded 

projection. The new sp. has at the anterior ventral 

louca margin a median, relatively broad noteh 

between two short keels, which is unknown in 

related species. 

Mc,/\ifrcincnf\ Lorica length 140-150 /^m; lorica 

width 112 fim; width of semicircular fbot-opening 

15 I8/inr. median noich in ventral plate II ^m deep 

Discussion; I he morphcrlogfcal group T. pufinu- 
nhlci tttuaonufa (after Koste I97N) have variable 
short tongUCS al the anterior doisal margin o\ ihr 
lorica. The new species mosj rcsembUsK / ohfeh but 
differs in the form Ql the dorsal and ventral ftnterioi 

lork.) mari'iii. T. ohfci Koste has only ihrce more- 
ronnded (loncuc-shaped) elongations of the dorsal 
anteritn margin [see Koste 1978a, b, ef. also /. 
patina tritahatu Anderson & Shcphard h\m\ I. kosfci 
Ofi Riddei (De Ridder 1983)]. 
Etymology: The new species is named alter Charles 
G Hussey, oi the Britfeh Museum (Natural 
History), in reco^ 01 Ins woik on the RotifCTS 
ol the IJ.k 

T- munda Bcrrina, I9S2 
LOC! C rcswick C k . Cluues, Vic; 1 1. iv. 1953. 
I. it: Ber/ins (1982). Not distinguishable Irom /; 
timplnnu llauei on the basis ol ihe figures and 

regarded here as synonymous. 



lig. 6:1 Testudinelia husseyi sp. nov. a, dorsal, b, ventral. 

6:2 Testudinelia patina (Hermann) from Tasmania, length of lorica 245 fim, width 216 ^m, ventral view. 

M si KM IAN KOI II |-K \ 



I \y ) Hr\„rfhm *>\\uns (Scrnov) Hi lateral, K ventral, c. vomral armltkc yppendapc, d, irophi apical, e. rami and 
fiilrunn, I, rami and fulcrum latctal, &, manubrium and uncus tceih. It, caudal projections oj //. QxyurtS from lasmama. 
i, caudal projections ul //. harramotiat Her/ins, I9N2 4-0 from lasmanian population, c-t tiom W,A, population! 


R. .1. SHIM. & W. KOSTE 

T. neboissi Berzins, 1982 
Loc: Creswick Ck, Chines, Vic; ll.iv.1953. 
Lit: Berzins (1982). Inadequately figured. Resembles 
T, emarginula (Stenroos) and must be considered 
a doubtful record. 

T. patina intermedia (Anderson), 1889 
Loc: L. Barracoota, Vic; 20.L1975. 
Lit: Berzins (1982). 

T. semiparva Hauer, 1938 
Loc: Magela Ck floodplain near Jabiluka, NT.; 

Lit: Koste (1981). 

T. vidzemensts Berzins, 1943 
Loc: Creswick Ck, Climes, Vic; ll.iv.1953. 
Lit: Berzins (1982). 

development and unci teeth number. The only 
difference between H. fennica and H. oxyurts is in 
the caudal armature. The caudal spine may be 
absent, as in H. fennica, or varying between 
individuals (both Tas, and W.A. populations) as in 
Fig. 7h (cf. Fig. 7i for a population described as 
//. harracootica Berzins, 1982). Most of our 
specimens had 7/7 unci 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, 
//. harracootica, which was described by Berzins 
(1982) as "very like H. o.xyuris . . . but trophi with 
only 6/6 teeth" apparently is a synonym of H. 
o.xyuris. Further, the ventral armlike appendages of 
both taxa have the same number of bristles (8) (Fig. 
7c, cf. Berzins 1982 Fig. 60a) 

Family Flosculariidae Hairing, 1913 

Octotrocha speciosa Thorpe, 1893 
Loc: Buffalo Billabong, Magela Ck, NT.; 8au.1980. 
Lit: Tait ef a!. (1984). 

Sinantherina ariprepes Edmondson, 1939 
Loc: Island billabong, Magela Ck., NT.; 15.iv.1980. 
Coll: R. D. Tait. 

Family Hcxarthridae Bartos, 1959 
Populations of a species of Hexarthra found in 
W.A. (#788) and Tasmania (#1146) were determined 
to be: 

H. oxvuris Sernov, 1903 
* FIG. 7a-L 
Syn: Hexarthra fennica var, oxyuris (Sernov), 1903 
Loc: 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. Shiei. 

Regarded as a variety of H. fennica (Levander), 1 892 
by Hauer (1941), this rotifer is characterized by 
inira-population variability in caudal spine 

Family Collothecidae Bartos, 1959 
Collotheca tenuilobata (Anderson), 1889 
Loc: L. Catani, Ml Buffalo, Vic; 25.ii.1955. 
Lit: Berzins (1982). 


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 1. J, 
Rowling (Melbourne), R. D. Tait (Sydney), Dr B. V. 
Timms (Avondale College), Dr P. A. Tyler 
(University of Tasmania). M. J. Tyler and an 
anonymous referee are thanked for critical 


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

aquatic communities in saline wetlands in Western 

Australia. Hydrobiologia 105. 77-84. 
Bi k/i\s, B. (1982) Contribution to the knowledge of the 

Rotatoria o\ Australia. (University of Lund: Lund.) 
Donnl-r, J. (1965) Ordnung Bdelloida. Besttinrrtungsb, 

Bodenfuuna Europas. 
DUMONT, H. J. (I9S3) Biogeography of rotifers. 

Hvdrohioiomi 104, 19-30. 
Evans, .1. (1951) The rotifer record of Victoria. PrOC. Soc. Vict. (195!), 4-7. 
FaDI EW, N. (1925) Materialcn /ut Kennmis der 

Rotatorien fauna Russlands. Trav. Soc. Ni Charkow 50, 


Gii in ki , J. .1. (1967) Asplanchna and posterolateral spine 
production in Brachionus catyciflorus. Arch. 
HydrobioL 64, 1-62. 

- & StbMBERGER, R. (1984). Aspfancluurhuhwcil 
polymorphism in the rotifer Keratetta slacki. I. initio!. 
Oceanogr. 29, 1309-1316. 

— , BlRKV, C. W. & WURDAK, E. S. (1979) 

Tuxonomie relationships of Asplanchna brightwetli, A. 

intermedia and A. siebotdi. Arch. HydrobioL 87. 

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

lakes. Ergeb. LimndL 8, 232-236. 
- (1981) Associations of rotifers in Australian crater 

lakes. ./ loot.. Loud. 193, 469-486. 



Hauek, X (1938) Die Roiatorien von Sumatra, Java und 
Bali nach Frgebnissen dcr Deutsehcn I.irnnologisehen 
Sunda-Hxpedition. I, Archiv. HydrobiaL SltppL 15, 

- (1941) Rotatorien aus dem "Zwischengebicr 
Wallacea". I. leil Das Genus Pec/alia Barren's. //;/. AVe 
i:cs. fhdrobtof. 41, 177-203. 

Kdsm, \V, (1977) liber drei rieuc For men des Genus 
HexanhraSctim&rda 1854: H.jenktnae H nukurv nX, 
//. hrandorfjin. sp. und //. polydonta soaplakeiemis 
n. ssp. Gcnaxs. Ahwass. 62/63, 7-16. 

- (1978a) Ubcr Tmudinetta ohtei Koste, 1972, ein 
Raderiler dei U-Ordnung Flosculariacca aus Uer 
Guuiea-Brasilianischen Region dcr Ncotrop'is. Arch 
/hdmhnd. 82. 359-36,1. 

(197Kb) Ofe Raderliere Mittcfeurofuts. 2. volv 
Borntraegtr, SiuttgafL 

(19X1) /ur Morphologic, Sysiemalik und Okologie 
\on neuen inonogonomen Ra'deriieren (Rotatoria) aus 
dem Uberschwemmungsgebict des Mage I a Creek in der 
Alligator- River-Region Ausiraliens, N.T. Tcil. I. 
Osnabruvker twturwiss, Mitt. 8. 97-126. 

- & Shii:i , R. .1. (19,80a) Preliminar\ remarks on 
the characteristics ol" the rotifer fauna of Australia 
(Noiogaeu). tiydrobiologia 73, 221-227. 

— &. - (1980b) New Rotifera from Australia. 

Trans, li. Sac S. Aust. 1115, 133-144. 

- & - (19X3) Rotifer communities ol billabones 
in northern and south-eastern Australia, Hvdtobiahxia 
104. 41-47. 

& (in press a) Plank ton ie and 

semiplanktonic Rotifers from Australian inland waters 
I. Introduction to rotifer morphology and ecology. Aust. 
./. Mar. hresh water Res. 

— - & - (in RrcSS b) Planktonic and 

semiplanktonic Rotifera from Australian inland waters. 
II. Svstcniatics and Bdclloidea. Ibid, 

• -& BttOfK, M. A. (1983) Rotifera from 

Western Australian wetlands wiih descriptions of two 
new species. Hydrohioio^ia 104, 9-17. 
Mi RftAY. J. (1911) Australian Rotifera collected bv the 
Shackleton Antarctic Hxpedition, 1909../ Roy Miaosc 
So<\ 31. 164-174. 

Rioolu. Mi Di (1983) Recberches eeologigues et 
biogcographiques Sur Ics Rotiferes de la Basse 
( 'asamance (Senegal). Rev. Nydrobiot. trap. 16.41-55. 

RL'smii, C. R. (1961) lire Rotatoria o( Queensland. 
Australia, trans, R. Soc. Y/. 1, 235-239. 

S\it, G. \V„ S\im\ni\i, G, I. and Commins, M. I. 
(1978) Tropin morphology relative to food habits in six 
species of rotifers (Asplanehnidae). Trans. Amer. 
Microsc. Soc\ 97. 469-485. 

Sfitrt. R. I (1983) The genus BradiionM (Rotifera: 
Biadiionidac) in Australia, with a description of a new 
species. Proc. R. Soi\ I'ict. 95, 33-37. 

& Komi. W. (1979) Rotifera recorded bom 
Australia. Trans. R. Soc. S. Aust- 103, 57-6S. 

IMi. R. IX, Stun , ft. J. & KOSTI . \V. (1984) Structure 
and dynamics of /ooplankton communities. Alligator 
Rivers Region, NX, Australia. 1 1 vdrohioiogia 113, 1 13 

WntitrRi, k. (I960) Die Radertierc saurer Gcwassct dcr 
Duhcncr Heide. I. Die Roiaiorien des /adlit/moors unci 
des VVildenhainer Bruchs. Arch. Ilvdrohiol, 56, 261-298. 

(1966) Rotatoria!) aus dem Slausee Ajwa und der 
Irinkwasser-Aubereitung dcr Stadt Baroda (Indien). 
i.tnwolouica (Hcr/i/u 4, 5 3-93. 



BY P. J. M. Greenslade 


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 epigaeic 
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. 


b_\ P. J. M. Qui-\ \si ,\Di * 


CiKii i mi , P L M, (iys5) Some cITccn, o) .vavc i ,-"■.■,.11.,.! L*pwt tin am\ iH>Miienopr 1 

1 mi idft hi, ih. M.. lolly Rdti'JL*-.. South VkttLUlIm ibtMS.K.Sftc.S \lt\i, I|W(U, I ,; |0 

Ami - '" ■ || -"' 1 li '" a yeai In EurulypWA huM-H-i- ^tiqua foresi at Bridgewiuei neai Adelaide 

1 |l1 ' 1 lw * ,l ' 111 "■■ ' "' pitfall [raps, g tc nn a well insulated hw 1 I 1 1 1 [ope, H n 1 oji a men 

: 1 om TIk- fauna u^s typical ol humid pa I , auh \u 1 ilia i 1 1 1 .. , > |he tol 11 

"' E '■'-.',■ in. h,.; id I uih LTvpn. .-mis (derive rrtamli wirbli 1 md liner! and :pi* > [acrivi 

an - 1 " 1 - 1 rtbui :lh' pHmitI '-■ 1 iieiiingciihcj-inftoil ffi irbd eally. One] -■;.■■,.;., | ,,,, ,, , -,. 1 

maximum in lito f>rm^ ind summer. GuctteruT cryptic specie ditti 1 h . 4ichi»arto ea I » irii ci 

' ! ' '"'■ M '" ' ! ■ | ' lL Btl '■!' MHil.'! 1 ; Oil li". ii-lil ' I I (Ope t I ,'IL'. I . ■ , i I n , I |j ,■,,; [.,1 

!' 'Hih jiswihiuKinM'xteu.iiM" i. ,,,,.,-, di icj 1 1 n Brideewaler predominated Eyi a ti ■, i 

!'''■ ■ n ' l| - -! iridvmvmiLX spt.vie i , ;ir,J ;.,- ,0i Lalcd LflXft v.cic Ir^n ■-., ' SOUrtlW I 

1 !■ I '.:■■! n Lti ti I ■■■ 1 ■ i, ! ; 1 1 .■ extendinjj into areas weird Ihui Urid i ■..- 1 ui. high i 

■ ||J 1 ■ ■"■" 1 [<n '"■"''" rob 1 '■ ■■■■'-'■■ i '"'i'" 1 ■ 1 .,; ■ !■ mi n\s iven lew, and iiass in 1 h ■■■ 1 

fui ■ :! i; " iiwnhtti lopi . a fri] fie in, icVi , e til erynuv iud nlhei ground Eivln) tin) ■ 

i;, ' ,L ■■' I " • ■ 1 equencj oj' irtdumww n . , , . 1 . ,,.,■■,,,, iv structure nn fhc 

«uutlwi ' 1 lope, I'm nt>! on the norttnvc 1 n , - tl h u ii m aiw L-ommi ■■ ill 11 I a I ■ 11 li< d ij 

; ■" ntt/ffw ohiiqiw \khkII uidinftuuhern Victoria a-, .. ,■■ h -i hisBntfi 1 am lisuiiy 1 - a 11 ■■■ iii mul 
"'■' 1 '■ ■ -"'"i' il (i idicnj til roii rail in .1 Vtl 1 oftj ftart^w ti, , 1 , , ,, . ,, , , , 

'''■'■ l ■''■ "'■' l ' ■ " t ■ ■ ' r 1 ■ I Utldol W, /»i7-(/i.'.'//'M;iiia triii*)t)ivm, K -\ nw[>i tl vi<- ■ -nMn: ■.-: Ii.;i n iy | ltd it- 

111 ll " i; mi 1 Hi ' ii' -li- ........nil,, ■ ,', i M ., rtel - |ani communilii I - no nper ilii 1 coo 

1,1 aipo 1 ■■■ ingl ■ ii.-i 11.' and unjciurdl w^ itKoi ui i|, I fri 1 "."■ \ .' h tc di uj 1 11 tn 1 

""■ rVfltion ol inwri bralt and theii us n en rim iual Li,\?n*ssmcnl 


The ;tm communiticso) South Australia's Ioiym-, 
and woodland have been described rectfuth in 
gi neral Lerrns (Grccnsladc, in press 1. Some ol tfo 
anis Di cine open lores! loeitlih 1 in the Ml I oily 
I'.ni , uv discussed here in greater detail. 
Although Genlrlli (1972) stressed ilie complexi!! ni 
theelinnuk iiiTluenees th;n ;iMcei southern South 
AuMrah.i, [Hj ■ [imflte "["die Mi Loftv Itai \ 
VleLuJe i> unequivocally mediterranean. Suinmcra 
ure hoi and dry. winter-, cool and WW, imptvn 
double mils- on biqra. Ginuui t be both 
moderated &rjd aecen tatcd b) topecL. The 
dis*nhiin\.n-. vi rtitr gucaiypius specie^ thai moke 
up ihe i.iivM-. and wdodteods ol" the Mi Lofy 
Rynyc^ ,ne deiem lined primarily hv soil t^pe and 
til Mddieni of increasing pteciptiaiion Ltun 
accompanies increasing elevation, but Speehi & 
Perry (J948) showed ihai aspeei is involved aii ^eil. 

1 ■ ed with "icnith-Facing ^iOTJct, noni, | 

arclikeli tnbcwaiiiterin winUi bul excessi !■■ 

hot jini '.h j mi 1 he ■-.Miinnvi. \>p.jLi aKu al !>'■■ b 
invc 11 brarc in this area, h ini tutrices ihe 
distribution of mounds 1 milt Vmutftermes 

■ I '1. 1 .IL, ■ ,.,,! |', |, ||| |:;;,u \, 

tl .1-1 i. - \\ ■ -'I.'. I 


r\ifioMtU\\\[\) rttbod ■ 1 K" I) and the strui 

1 ■■ lOpulatiCins and com n iii 1 ■:• ;i at ( 1 1 h 
1975, in press, P, I M Greexslatle & < 1 ■ ., 
\m\ ubi.) bflbciK nwj hi 1 omplc 1 he iiiLtTactinu 
ol'aspLei vegetation and irisulaiion determines ihe 
oil climate which, fn lurn, arflccts leading ni mb" 1 
oi ani communiiies with teeond'^n effect* CNi 
ui 'Ordinate speeies. 

in 1^83 M. ])r(. i ,.., ei used piitall naps at 
Bridgewaler rteai Vdelarde Ibi a v«ai to sampl« ■ 
poptrlaribn af 1 he iotrutlu d millipede 
QmwQtaiulus ww^/ev/'ttuca<)^nd rb rwmito 
poientiaJ ground liv&ig pieJauvs. i ins papc< 
reports ou the nnh taken fn t-heM irap 1 h ■'.,.. 
1 1. "Jed between a well n ■ ■' ted tturthwe$i ■■■ ii 

stopt and y more shacTed southwe^t-iacinc 

iheii ' ah \h i prewni anoi hi I >ppo ii j to 

invesj hi inn ,., :iion betw ■ 11 1 In eitcets of 

..1 and j'-.j.'.. 
1 1 1 iou svoi k invol\ ii bi 1 ihis site 

1 ! ■ cleall with iIk bioli - (X utmvleiii (Bake? 

19 -i ci and the Inputs dJ di id plan 1 ictuiI 1 

1 flooi and its di po i 1 b; h*rn fto I ] 
Butter 1477; Lee M" (mi el! P»~s ; i , 
in : oilier soil biota (I Ihimm, L v Witch lftS3>, Data 
and iiuiena! proceed b) iiakei and Unison have 

keen used to Jtow how diflen 

sampling soil lauiut can yield differinje ie- . i 



(Greenslade & Grcenslade 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 
c.IOOO 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 killing 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. eomm.). 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. 


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 
austral is* and the He tempo nera and 
Sphinctomyrmex species. The Amblyopone was 

For the few species for which specific names are available. 
authorities are given in Tabic 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 o\ 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 Iiidomyrmex sp.l 
which nests above the ground in hollow eucalypt 
branches and Iridomyrmex sp.4 may nest arboreally 
as well since I know it only, as in this case, from 
aiate 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. lb), 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. lc); on the shaded southwestern 
slope catches fell from December onwards (Fig. Id). 

(c) Northwest aspect 



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 



S;fiW^l? r *^ /v,s:? mtalrecot&m ttepes witknorthwwiundmuihwsiwects South 
Australian dfstribuiibn: W. occurring only in the wettest, mosi humid pans o» the State; O. occurring mainh in areas 
Uncnhanilu.Mnd.euaM-locah.N; M, at about the midpomi of their range on the raintall uradiem a. Hrid«cuaicr 

Vnl Species 


\fynneciu rfignscapd Rogei 
U. Jnrficutu (Fabricius) 

M. pyn/onnts I . SmitJl 

M. pitomta I-. Smith 

Amhlyopone australis L rich son 

Heteropun&v tntheflis (Emery) 

Rhylidaponeru 'nu j tadtca'\l. Smith) 

Spfiittctoniyrtne.X sp. 

ipttpostnitmt sp. 

Crewutoxasttr -,p.| 

C'tv/ntftoziistir sp.2 

i\!t'rutif>f)fits sp. 

Pheidole sp.] 

Phetdate sp.2 

CfwUmei sp, I 

Chi- Um or <-p - 

Chalarter sp.3 

Gen ct sp. indet.* 

tridomynmx purpureas (l- 

Indomyrmex sp.] 



\- sp. 

truiomyinu-y sp r 3 
IruloiityniK'x sp.4 
Ihdichoderus sp, 
\h'fo/>fu>/ii.s sp,l 
Meio[)hi)Kis sp,2 
Vci/p/icwj; sp. 

Cumponotas sitf/itsus t . Smitl 
C. \nnsohrinus' Frichson 
C, in nexus l orel 

Cawpunuttts sp.2 
GynppttottQ sp.3 


Record 1 

















as pee l 











Record 1 

as pec i 

*A damaged alate queci 










sonic species but can be tested only for those that 
were relatively frequent in both transects, i.e. 
Myrrneciii pdosula, Amblyopone uustmlis and 
Indomynncx sp.l. In neither transect however do 
the summed frequences of these species differ 
sigmficamly from the overall trend tor the whole 
fauna with both transects combined: northwest 
X- |n| = 4.60. p>(,).()5; southwest, x* ||rj == 3.95, 
p >0.0>, The differences between Figures leand Id 
therefore indicate thai the two sets of traps sampled 
different combinations ot species, i.e. different 

All but lour ot the species in lable I can be 
placed in one or other ot 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 are defined in the caption to Table 1 which 
compares transects. Treated as a contingency table 
this shows significant heterogeneity. Group \V 
predominates in catches on the southwest slope, 
group D on the northwest slope: the community 
here seems the best adapted to exploiting high 
Mimmer temperatures and withstanding drj 
conditions ( lable 2). 

The phenology of the cpigaeic members of 
groups O, M and \V is shown in Fig. 2 where catches 
from both transects are combined. Compared with 
group D, group W is aetive earlier in the spring and 
has an earlier maximum; catches decline through 



Tabi i 2 Catches in (he two transects with species grouped according to (heir distributions in relation to rainfall 
in South Australia: W, D, M — see legend to Table 1 lor explanation. 








Soul Invest 










P<< 0.001 



- 55.4 

the summer but extend further into the autumn. 
Group M is essentially a single speeies, Myrmec'ta 
piiosula (see Table 1), whose seasonal distribution 
in traps is closer ta group D than group W. 




Fia. 2. Seasonal occurrence of epigaeic ant species 
according to their South Australian distributions. See 
caption to Table I 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 
and status in communities. Here, distribution refers 

to the Australia-wide range of genera and species 
groups and not, as in Table 2, to the South 
Australian distribution of individual speeies. 'Role 
and Status', is explained in detail by Greenslade {in 

Taking the biogeographical affinities of these 
communities first, there is only a small element of 
speeies 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 
'mefallica' complex. As the frequency of cryptic and 
arboreal-nesting species increases in more humid 
climates, this /r/r/omv/'m^v-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 live 
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 Casuaritw- 
Leptospermum heath and adjacent Euealypitts 

Ml lol 1 \ RANGES' ANTS 


] ami i _v Structure oj itn/ cummunittes at &rid$ewwer Turn {genera and specie* groups) arc dossed according to 

llli ' lf / owaphical distribution* and role and status in communities. Role and status is described m detail b} 

ureensktde (tn puss). Dttfhei ( -) indicate ceils that are imomtpiuhlc ar apparently have no occupants in the Australian 
unt hmnu; cu. cell < "J ft unocatptahlc: if in not possible to have a 'donate specialist' that is widely distributed in 
respect i" cbmute. I fourth distributional category s Torresktn* fi.e. northern tropical/ is recognised but it is not 
represented by any species in these communities sv the fourth column has been on/it ted, / tactions Indicate numbei 

oj species number of records. 

Role and 

Biogeoeraphieal distribution 

A liyivan H Kalian C Wideh distributed 

Northwest Southwest NorthweM Southwest Northwest Southwest 
aspect aspect aspect aspect aspect aspcel 


fl, frido/nyrtnc.x 
b. Other genera 

2/5 II 



hoi .iniviiiae 


3 Climate 


3/3 (1 



4 c ryptk and/or 
predacious on 

? Generalised 

6 Opportunist 1 

7 I arge solitary 


5. N 

] 1 

I io i |5 


1/5 I I 



2 3 3/8 

/ popnstrutua 
( lielaner 

I 12 1/17 



2/10 4 14 


7 40 

1(1, 55 

2/J M 



3 22 3/4 


2 2 



4 11 4 26 

( rematogasier 

I 3 



10. 3M 

10 33 


Northwest Southwest 
aspect aspect 

5. IK 

-V 22 



4 11 

I. 3 

3 io 

22 K7 

V 17 

I I 

If A 

3 8 

V 19 

4 2fi 


4 14 


httxtcri woodland. Rainfall was similar to thai at 
f J .iid)icwalc-r, c.IQOO mm pci aniiinn, but c\enl\ 
distributed through lite year. Although the soil, a 
siliceous sand, also differs there are Similarities in 
ant I'attnas. Among Myrntecta for example. W. 
p\rilornit\, XL nlgtiscapa and M. piiostlla were all 
trapped at Wilson's Promontory (compare lable I). 

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 oceupiable cells in Table 3 provide 21 
paired observations (Table 4). At Wilson's 
Promontory, as at Bridgewater, there were no 

t-vwi 4, Comparison oj the structure of ant communities at bridgewater, South Australia, and Wilsons 
Promontory, l h torla (see tc\ti. Correlation coefficients frj for frequencies in pairs oj communities partitioned as 
m fable 3. Significance: . \ ■=■. p<(UrS, < u.OT. < 0.001. 

I oealitics 



\\ i I '-Oils' 

Bridgeware - ! 









o.;'o ; ■■ 






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. 


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 Iridomyrmex in dry 
areas, to weakened Tridomyrrrtex-dominzuce 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 purpureas 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 /. purpureas 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 
445 m with E. obliqua (one of the species at 
Bridgewater): M. forficata is present but M. 
pyriformis and /. par parens are not. At 350 m the 
Bridgewater ant fauna seems to be transitional; it 
contains both Myrmecia species as well as /. 
purpureas 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 /. purpureas and M. 
pyriformis, to M. forficata without /. purpureas 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. 


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. 


BAKER, G. H, (1978a) The distribution and dispersal Of 
the introduced millipede, Omtruuoiulus moretetii 
(Diplopoda: Iulidae), in Australia. J. Zool, Land. 185, 

(1978b) The post-embryonic development and life 

history of the millipede, Ommatondus moretetii 
(Diplopoda: Iulidae), introduced in south-eastern 
Australia. Ibid. 186, 209-228. 

— (1978c) The population dynamics of the millipede 
Ottunatoiuius moreletn (Diplopoda: Iulidae). Ibid. 180, 

Gl Mil 1 1, J. (1972) Australian Climate Patterns, ( Thomas 
Nelson (Australia): Melbourne). 

Gkefnsiade, P. J. M. (1975) Short-term change in a 
population of the meat ant Iridomyrmex purpureas 
(Hymenoptera: Formicidae). Aust. J. Zool. 23, 511-522. 


2 } 

(1976) Distribution of two farms of the meal aril 
tridowyrtmw purpureas (Mymenoptera: Lormicidae) on 
Kangaroo Island and Yorke Peninsula, South Australia. 
Ihul. 24, 55"- 564. 

— (1979) A Guide 10 Arils of 'South Australia. (South 
Australian Museum: Adelaide). 

(In press) Ants. In Wallace, H. R. (Pd.) "I he 
Natural History of South Australia's Forests and 
Woodlands". (Government Printer: Adelaide). 

& GkiiNstuH. P, (1983) Ecology of soil 
invertebrates. /// "Soils: an Australian viewpoint'' 
Division o\' Soils, CSIRO, pp. 645-669 (CSIRO. 
Melbourne/Academic Press: London). 

— & - (1 984) Invertebrates and environmental 
assessment Environment & Planning 3, 13-15 (L)ept. 
Of Environment &. Planning: South Aust.). 

Hitmin, U. R. & Viiicf), L. G, (1983) Mean annual 
population density of Collembola and Aeari in the soil 
and litter of three indigenous South Australian forests, 
Ausi, J. Fcol. 8, U3-126. 

Li f , K. E. & BuilLk, J. H A. (1977) Termites, soil 
orga.nk mailer decomposition and nutrient cycling. 
Eeol. Hull, (Stockholm) 25, 544 -548. 

& C'okkM 1 , R r L. (1978) Litterfall and its 
relationship to nutrient cydtnj in a South Australian 
dry selerophyll forest, Aust. J. Zool 3, 243-252. 

— & WOOD, I. G. (1$$S) Preliminary studies ot the 
role of Nasutitennes exitiosus (Mill) in the cycling o\' 
organic mallei in a yellow pod/olic soil under dry 
selerophyll forest in South Australia. Tains, int. Congr. 
Soil Set 9(2), 11-18. 

Mviir, .1. D. (197S) Studies on invertebrates in relation 
to bauxite mining activities in the Darling Range. A 
review of the first eighteen months of researeh. Lnv. 
Res. Bull. No. 3 (Aleoa: Australia). 

(19.S0) Report on a study of invertebrates in relation 
to the Kojonup fire management plan. Dept. Biol. Bull. 
No. 2 (W.A. Instiiute of Technology: Perth). 

O'D(Mi), I). .1. (In press) Seasonal patterns in the activity 
of ants in lielair Recreation Park, South Australia. //; 
Gieenslade, P. & MajcL J. D. (Eds) "Supplementary 
papers to I he Mediterranean and Related Ecosystem 
Source Hook". Dept, Biol. Bull. (W.A. Institute ol 
lechnology: Perth). 

SPECKT, R. I.. & Pi kkv. R. A. (1948) The plant ecology 
of part of the Mount Lofty Ranges. Trans. /?. Soe. S. 
Aust. 72. 91-132. 

Wood. T. Q, & I t l , K. L. (1971) Abundance of mounds 
and competition among colonies of some Australian 
termile species. Pedoluoloxhi 11, 341-366. 


by David M. Spratt 


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. 



by iv\Mi> m si \ 


'■' n vi U SI i IW$j S>; w« .".-..■■' "■■'.■'' ■■./'. ri i i m, ,-.. i ,, ,,,1 ■,, ,, ■-, ,,i ., i i ,,.,, ,,,,,11 

1 ■ 1 ■ 11 ul 1 [\| Li "■"■ ii'.i' Wun> ''■ - '■ I ■■■ IDWli, ' '■■' 1 .i ■■ i"- 

. ., I \h .v I ..vck.n. l%7) OuentiJi cV Krishiuisau... 1 1 : ., descrHwl frofl) il'ic 
Oesophageal ill u WW ol ih'j d&syurid marsupials Atitctiw m \tu rn/l M l< I RJ Phniftak' Inftrtmri (Tboiiia.s> 

lllld Swimh<>j>:;r, tpUfOpm (liT*) I I Mr. r. 1 1,,- I >j - , .1 , ! „ ■ . yp 1 1, ■ -polled linrn Au-.ri j I i.ia uair.c 

IIWi .ils, alilu>ii;'li |hc specu"> has been described Irom .1 mlcrOCXliroprprnri ( fh/>fu>:ou\ kurhhi-nsis DOl ■ 

in 1 'mm. 1 tml Ihrni lupaiid I nV/ww -/,■- (Dtaj'dk) nnvl lorisid (A(yi r/'i r/im couvan$ (Boddaerto prima* in 

\Mll 1\ ,,.1 

V ow/'i . ., . i miibt in morpholopv and m Mom to 5 1 vitiHtnw Mlepp, 1924) 

1 hi I wood, 193S f"ri>m pfatyrrhinc. 1 1 imaies and dlcWlphid lunrsupiali in South \nu rica. ( ephalic ytrycturci 
in thmi -si.i'-r jftrvafl 01 X tmruflwtfxtdemix from /l. shmrth are, in contrast w larvae of S. RUhtttPfjxii>\ 
identical to thaw to adult fbrmj, r>fTi rio :.. 1 n 1 ndciiee Ilia 5, uukutwib is the mwre Im-hlv pvoli 1 1 
nicnabci of thefcemj \llhoughil genua Spirttrn in probably of Gondwaiudand Origin, 11 IS unlikely thai 
V aumrtffubudenKu entered Australia via rhis rourd but rathet ftotfl the north with ( hiroptera cm with 
windhnrne hiMM.: |t$ vhmii H-c. !- mi Ausnatian uiui'.u[)iuls in interpreted as secondary transfci 

Kiv Spirurtt, Nunatoda, distrihuiion, Iioms, morphology, UasyuiJdaa 


I M t spiruroid nematodes of A I an 

ma wpials arc in need ot a thorough taxoncimu 

revision, A preliminary assessment pf the gr< 
based CM examination ol type jpe ton n.s held in 
Australian uisiit ut ions and oil material collected 

ino 1967 revealed thai one species exhibited 

morphological relationships more akin to iM \${ 

hi a microohiroptcran, primates and dldetplifrJ 

muisupials than (o those from otln t ,\ir ,,:, 
marsupials Details Of tin-; parasite, ffOm tl|C small 
dasyUrid marsupials Anh'i-htnus Mtwrfti MacU*a>; 

riunivaii' ingntmi (Thomas) and Stmnthopsib 
leucopm (dray) hnw the basil of tliis report. 

Materials arid Method* 

Native mammals wen- esammed Uuiiti^ iIil 
period March l%7- December |983 Irom the 
lollowni:' rri'.uins: Arnhem Land, Nortln-i m 
Territory; Quccndond; souTheastern New South 
\\;\\i\; uoiilua-hrn Victoria, the Ausiialtau t'apilal 
• ■ 1 1 itory; northern lasmama. 

Nematodes recovered by the author wer$ Used 
in hoi, l()".'(i neutral hnftn. <1 formalitl and eleaicd 
m hulophi'uol 

The Ibllowmg abhrevialnnr- fol morphological 
features oi adult worms and ihudsiujLie larvae are 
ii it in the text: 
I Length Of body, MW Maximum width of body, 

v Length >■ width oi pharynx, nk Length from 

Division oi Wjldiifeii Rangehnd.i Research, CSIRO, 

PO, Box S4. lyneham, AX T 26^2 

cephalic extremity \w ner\e riujz, EP Length llll,M 
iilic extremity to exetetoiy pore, D t ength Irom 
: ihalic cxiicmit) lo deirids, MO I ength of 
mu oesophagus, GO Length o( glkndulai 
oesopliagus, to Total length of otsophagu 
\<- B Length from ccphaJic extremity eo vemrftl 

cuiiculai boss (large thumb-like swelling o^ ventral 
■ " . i.i: | i b idv), V Length from eephaliL cxiremilv 
to vul\a, RS length of right spicule, IS Length 
of left spicule, T length Ironi caudal cxlrenuly lo 
anus or elt>aca, GP length from cephalic eMrennix 
i genital prlmurdmm of third-stage larva 

Measurements were made with the aid ol an 
ocular micTomeLer, drawing tube, and measuring 
wheel, and are presented in nucromctets uulev, 
otherwise slated. The range of measurements is 
followed by the mean, in pi renthesCS. Illustrations 
were made with the aid of a l.cit/ drawing device. 

Specimens have been deposited in the Australian 
Helminthological Collection (AHC), Queensland 
Museunr (QM), South Australian Museum (SAM), 
United States National Museum Helminthological 
Collection (CSNMHC) and Division oi Wildlife & 
kaneekmds Researeh, CSIRO (Wl HC). 


A i h iracteristic sptruioid nematode w;i', 
encountered rarely, attached to the oesophageal 
mucosa t>l smalt dasyund marsupials. Number of 
hosts inlccted/mmiber of hosts examined Irom each 
locality are shown in parentheses. The parasite was 
found in Anirthmus Mtwrttiiii Mt Nebo (1 5), Qld, 
and in the same host at Naduee Nature Reserve 



(section formerly part of Nadgee State Forest) 
(6/54), N.S.W., in the same host at Jervis Bay (2/28), 
A.C.T., in Sminthopsis leucopus at Timbillica State 
Forest (1/9), N.S.W., and in Planigale ingrami at the 
Smilhburne 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 & Lovckar, 1967) 
FIGS 1-11 

Material examined: from Antechinus stuartii, Nadgee 
Nature Reserve, near Eden, N.S.W., P. Havcock and L. 
Walter, It?, 1?, 3.xii.l981 in WL HC N1438; 1?, 5.iii.l982 
in WL HC N1519; Id" posterior, in WL HC N1630; 
Id 1 , 1<? anterior, 2? 2 posterior, 8.iii.l982 in AHC No. 13651; 
W, 15. and IV I0.iv.1984 in SAM Nos. V3557, 
V3558, V3559. 

From A. stuartii, Jervis Bay, coastal A.C.T., J. Kenagy 
and P. Haycock, ld\ 19, 12.ix.1983 in USNMHC No. 
78252; Vj, 2 L 3 , 7.xii.l983 in AHC No. 13652. 

From A. stuartii, Mt Nebo, Qld, D. M. Spratt, 1<?, I? 
anterior, 19.vii.i973 in WL HC N167. 

From Sminthopsis leucopus, Timbillica State Forest, 
near Eden, N.S.W., E. Walter and P. Havcock, Id", IV, 
I.xii.1983 in AHC No. 13653. 

From Planigale ingrami, Smithburne River, near 
Normanton, Qld, L. Owens, 1¥, 4.1.1979 in QM No. GL 


Short, robust nematodes, attenuated anteriorly 
and generally reflexed in anterior 14 or Vs 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. 


Male (7 complete specimens, 1 anterior end): L 
5.4-8.5 (6.5) mm; MW 180-390 (294); P 
50x30-80x50 (60x40); 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). Gubemaculum 
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, 1 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, V 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 34x25-40x30 (38x27), 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 leeth 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 

KfHkt A' I \l A' l \t, \/i\/)i \sts I KOM l>\S\ l iRICtAE 


Figs 1-1 1. Spiruru uu/iwxtjhutfcrtsis Irotn small dawurid marsupials. I, Cephalic end, adult, en face \ iew. 2. Cephalic 
end, adult, lateral view, h Cephalic end, tWrd-stage larva, lateral view. 4. Third-Mage larva, lateral view (arrow 
ventral cuticular boss), 5, Cauda] end male, lateral view, 6. OubernacuJgm, laiero-vemrul view. 7, Oesophago- 
inwsrinal junction, lateral view. 8. Caudal au\ female, lateral view. 9. Caudal end male, ventral view, |(). Cuticular 
bosses around vulval opening. II. Right and left spicules, ventral and latero-vential views respectively Scale lines' 
I lg$ I. K 2(1 M in: Pigs J. 6, y.n, so ^ ni; [:^ s 4 t s, 7 ( ioO „,„ 


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 
46x24, 45x20 (46x22);NR 120, 96 (108); 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). 


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 {Taphozous kachhensis Dobson) 
in India (Ali & Lovekar, 1967) and redescribed from 
tupaiid and lorisid primates (Tupaia g/is (Diard) and 
Nycticebus coucang (Boddaert)) in Malaysia 
(Quentin & Krishnasamy, 1975). 

Specimens particularly males, from Australian 
hosts are similar in size to those from /.' 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-cloaeal papillae and single median 
sublerminal papilla contra males with three pairs 
post-eloacal 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, 

Spirura aurangabadensis is distinguished from all 
species oi' Spirura except S. guianensis (Ortlepp, 
1924) Chitwood, 1938 (syn. S. lamarmi 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 o'i 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 oi' platyrrhine primates of the genus 
Sagiiinus in South America (Ortlepp, 1924; 
Cosgrove et at. 1963; Thatcher & Porter, 1968) and 
from the same site in the didelphid marsupials 
Caluromys philander (L.), Marmoset cinerea 
amarare Thomas and Philander (Metachiropsi 
opossum (L.) in Guiana (Quentin, 1973), are 

Nevertheless, S. aurangabadensis is readily 
distinguished from S. guianensis by the following 
suite of characters: (i) pharyngeal teeth terminating 
disially 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 larger than those o( outer 
circle, (iv) presence o\^ fine cuticular bosses 
surrounding vulva, (v) absence of longitudinal 
cuticular bosses on ventral surface of male tail, and 
(vi) much longer (2X) 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 
temurids 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 o( the 
cephalic cuticular collarette is rounded m 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 o^ 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 distinct 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 oi' S. 
aurangabadensis in .4. stuart'u exhibit cephalic 
features similar to those in adults and to those in 
third-stage larvae o( S. guianensis, supporting the 

SMKURA )t/< IV, [HAD/ vs/s I ko\i dasyl mini 


aigument that S, guianensis is the more highly 
evolved member of the second line (Quentin & 
kiishnasamy, 1975; Quentin, 1979). 

These authors argued that the occurrence of the 
most highly differentiated form, .S. guianetisis, in 
South America was due to its geographical isolation 
in ancient platyrrhine primates. Its presence in 
neotropical marsupials was attributed to secondary 
invasion at 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 
Chiroplera, or with windborne Insecta (e.g. Locusta 
sp.) which probably serve as natural intermediate 
host o\ the parasite. 


Thanks are due to l he following for collecting 
specimens or supplying small dasyurids for 
dissection: P. Haycock, .1. Kenagy, K. Nevvgrain, 
L. Owen and E. Walter. Drs I. Beveridge and J. 
Calaby offered valuable criticism o\ earlier drafts 
of the manuscript and Mrs. J. Rudd competently 
typed all drafts of the work. 


At I. S. M & I OVI KAR, C. D. (1967) On a new Spirurici 
Vaphozoia aurangabadensis n.g., n. sp. from a microbat 

TaphozoUS kacchensts. fnd. J. HvhtunthoL, Year !96fi, 

18, Suppl. pp. 68-73, 
Ojsi.kovi , C I-., Nu son, B. M. Hi Jones. A. W (1963) 

Spirant tanianni sp. n, (Nematoda: Spiruridac) from 

an Amazonian primate, Tamarintts niyricollis (Spix, 

1823). J. ParasitoL, 49, 1010-13. 
Otui tpp, R. J. (1924) On a collection of helminths from 

Dutch Guiana. J. Helminthot,, 2, 15-40. 
Ql imin, I. C. (1973) Presence de Spirura guianensis 

(Onlepp, 1924) che/ des Marsupfaux ncolropicaux. 

Cycle evolulif \nn. PantsitoL 48, 117-33. 

- (1979) Place zootogique dc Spirura rothsehitdi 
Seurat, 1915, Nematode parasite du Macroscelidc, dans 
revolution du genre S/nrunn Hull Mas, ttafn. Hist. rial. 
Paris 4 1 ' sen, A section A n /, 10^1-50. 

- &. KRISHNASAMY, M. (197^) Nematodes Spirant 
parasiiesdes '/itpaiact du Nscticebeen Malaisie. Ann, 
ParasitoL 50, 795-K12, 

I haii Hi k, v L-_ & Poster, J. A. (1968) Some helminth 
parasites of Panamanian Primates. Trans. Anter 
Microsc. Soc, 87, 1N6-96. 




VOL. 109, PART 2 





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. 




\\\K\i \ t, IV iV S< iiu\m k |, |), UVNsj i.rowih in Laptive hum ii : v> ttKlta (\"t>\ /»;% »/■*«> •-. r-< ■/*/> o i 
I htiwell i.i.hkJ: im ii.'M .nui laboratory tljjtfi*. /ro/w A' Sm X Iulvj HWfi. 5 i M 
7fi lima |v«5 

r.i|!hw Ihhm l%tj ttutta (\i>tahi\ on-r V17U7/M7/ from t h;i|i|K'U Maud But* Sirati rterc iVd 

\i\\ic,iff fihitmtt, GMWlh raft*-. (le-ldl lentil, .md weight) uric |\0|Ce thai of mainland ifrtftfl snaU---. and - lu 
iWTlKJi thai pfiltflUlffil populations mi other glands The puuernot increase, nt Mriggpi ^;,v e^ p^ncnliat ihmug 
the Qrsjl wi\ months mnd linear during the second six rliomhs of the .study. Orowth in length lui the hr-,i 
six mouth'. Wa* bent cvplamcd b> a linear model; hoih linear Bird loyaritliimic models explained RrflWlh 
jp length during the second six month* Regression ame>>as delected a significantly §K«uer increase mi leuuih 
relative to weigh! in the first six mom In Of growth, hut increase in weight iclanve u> lengih during Die 
second six months of L-rowih Aa estimate of gross growth efficiency indicated that metabolic rare wu* 
Maiiyely constant for these snakes under preset ibed conditions. Pattern* uffTgtfWth measured under ..oniioHcd 
conditions tna\ piOMdeiui insight into the evolution of life fusion i raits irt natural populations of liyn snake-- 
Kl\ WORMS Su.ike.,, gfllWlll canto e hushandiv, NottSCfliS VtV! vemww. 


( hapivil I BteAtfi Bfc$s Strait supports a popula- 
tion of Itger snakes iNutochh uter svrvenlyi) in 
which individuals reach adult body lengths 
apptouchiug 2 tn (Worrell 1958). The great si/c 
attained is almost twice Ihe average length of 
individuals on Ihe mainland (Shine 1978), and is 
rivalled only by a few other island populations of 
tiger snakes tn Bass St tail and ihe Nuyts 
Archipelago, South Australia. Although large adult 
\i/e in these snakes lias been attributed to their habit 
ol eating murtonbuds (Worrell 195S), no data exist 
to quantity their feeding habits or to determine 
growth patients bom birth to adulthood. The only 
pte\ ious studies of growth and maturation in tiger 
snakes involved dissection of museum specimens or 
estimates of these parameters from field sampling 
of the mainland species, N. SCUtatUS (Shine 1978). 

I lere we report a one year stud\ of feeding and 
growth th a brood of new-born, captive tiger snakes 
from I happell Island. The results are compared 
with previous studies of growth tares in tigei snakes, 
and with recent (unpublished) evidence for growth 
in a population of matked tiget snakes on the 
franklin Islands iu S. Aust. 

Materials am* Methods 

\ subset ol six snakes was taken from a brood 
ol 31 neonates born to a captive Chftppell Inland 
tiger snake of 1420 mm total length, on 3. in. S3. 

'■ ;>eiiM"[i Sireei, '\nlcci, Yu 5022. 
* StxJlh Australian Museum. Slltrih l^rraec, 
AOcUidc S- Ausi SQ(til 

Selection of the subset was biased by (he need to 
pair different colour morphs foi individual 
idem t float ion in cages, and to achieve CQUftl 
represeutatioti of sexes Mowevet, comparisons ol 
individual weights and total body lengths wilh those 
of the brood means lor these parameters satisfied 
(he null hypothesis that each individual was 
sampled from ihe same statistical population 
uf.O-S.df-.IOI ZfUl- is wtfghi 0.46 1,231; is 
length -0,49-0.68, Sokal & Rohll' 1981). 

The six neonates were maintained in pairs in three 
particle board cages rneasui ing 460 4 4ft0 '■■ 450 tnn^. 
The lid of each unit contained a gla^s witiciow 
measuring 420 ■ 250 nun. I ighl was provided unly 
from 'True-life' fluorescent tubes suspended al 
ihe cages, BflCh cage was fined with a blue, 4\t watt, 
incaudesceni light bulb controlled by a thcimosMat 
10 maintain a constant temperature range of 
JS-30 'C. Relative humidity was 70% throughout 
Ihe study. 

Snakes were ted entirely on house mice fMUh 
inusmlus) Neonates accepted day-old mice directly, 
or mice tubbed won faeces nl M.mud h/aub, (theif 
natural prey) Snakes were fed oldei. larger mice 
commensurate with increasing body sfcesand their 
ability to consume Jarger prey. During Ihe 12 month 
period, the -tiakcs were kd ml hhimiu to thceslcnt 
thai teiection of food indicated repletion Daily 
records were kept ot the quantity of (ooi\ accepted 

Snakes vsere weighed ano measured motvhly. 
• •nor ic. -A'cighiue snakes were denied food -o thai 
onuiihlv tVec£th!5 would be comparable among 
individuals. This achon reduced lite feeding time 
nl each sfUtkc by M-4fl rfays over the \2 ruonihs. 
C omplete data sets far iuonthl\ erowih m lencih. 



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 bt ), 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 2 ) was used as the criterion 
to determine which equation best fit a given set of 

Growth rates were determined using the 
instantaneous coefficient of growth, (G), calculated 
from the equation: 

where Y, = initial length or weight, Y 2 = final 

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. 


Growth in weight: Neonates increased from a 
mean weight of 8.7 g at birth to 1266.0 g at 12 

Tabil- I. Monthly rate a/growth in weight and total length of captive Noicchis ater ^crventyi. 














(111 OS) 

weight (g) 

SD (g) 

Range (g) 



length (mm) 

SD (mm) 

Range (mm) 






8.34- 8.98 

289 "' 


280- 298 







17.29- 21.91 





355- 388 







34.84- 67.48 





446- 549 





83. A 1 


43.11- 106.83 





491- 677 

IS. 5 






61.36- 154.18 





568- 799 







117.09- 233.03 





681- 914 







227.76- 430.93 












336.35- 550.48 












503.19- 734.49 












679.42- 968.74 






















1 266.00 










length or weight, t[=age in months at Y, and 
t : =age in months at Y 2 (Forsythe 1984). Relative 
growth rate was expressed as percentage increase in 
total length or weight per month, and was calcu- 
lated by multiplying G x 100. The velocity of 
growth, or actual increase in any length (L) or 
weight (W) was calculated as GxL, 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°- 7 *' 
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. r 

values are in parentheses. 






y = 47.2 ( - 78) ' 

y= -1.4 + 2.21nt 

y= -35.3 + 5U 

y = 435.6 a,wt, 
v=353.1 + 2.4lnt 
v = 320.5+ 101 .6t 

(.94)v = 7I9.3 |004)l 
(.75)y = 564.7 + 210.8lnt 
<.86)y = -578.6 + 156. 3t 

(.60)y=1076 Mm) ' 

<.76)y = 313.6 + 2.81nt 
(,99)y = 498.5 + 78.61 


(.67)y = 320.7 (ft,16 » 
(. 79)y = 230.6 +156t 
(.98)y- - 190.9+ !10.6t 

(.69)y-758.1 ,l(nH 
(,99)v = 224. 6 + 6. lint 
<.99)y = 308.7 + 99. 9t 



( M'llVI IK, I K SWM. ttkou 111 


AHei m\ months k;rowfli rates declined {lahle 1 1 
with a mean growth ratje of 26.()"'i* month from the 
Mh fofhe 12th inonlh.Growih for all snakes during 
this lattet period wax linear rim) besi desenbed by 
Ihe equation: 

VV(g) 578.6 l IS6I 
linear and (ogaiithinne models (Table 2) were 
similar (p-,09). 

Although rhc percent eaowih tares {% mo) 
generally deelincd after the first six months, actual 
mean weight per month (g< ■ mo) increased markedh 
a( this tunc (Table I). Monthly increases in actual 
weight averaged 37.6 g lor the- lirsi five mom hs; for 
the last seven months, ihisinCttSHSC Ltfti 207,0 g. This 
obseivaiion is important lor the discussion to follow 
because the increase in actual weight per mouth is 
correlated with a marked increase in feeding rate 
but not with a similar increase in monlliK erowlh 
in leneth (see below), 

Growth in lt j n,v(h; Neonates increase from a total 
lent-th of 2W) mm at birth to 1367 mm at 12 months 
( (able I).. Proportional increase, in length ("Vt/ino) 
were highest during the lirst mx monihs, with an 
overall mean ol 2$ ir M lot the fifM fwa months and 
15.6% lor the next four monihs. Thcrealiei, Ifi^Hi 
increased b\ mi average of only 7.4 u: m pel month 
liable I)- Unlike piopurtioiial weiyln gains, 
monthly increases in kuylh were poorly lilted by 
an exponential surve (Table 2). Variaiion in month!} 
length lor Ihe first ,m\.\ second sis months 
rospec lively, and for ihe emire study period was 
explained well however, by logarithmic and 
partiCUlArlyi linear ytowlh models ( lahle 2). 
C or relation coefficients lor all linear models 
differed significantly (p\<.05) from exponential 
models (Table 2), bin only the-|ineai equation tor 
the first sis months differed from Ihe lognn'thuni; 
model (p -..OS). 

It is well known thai length increases 
approximately as the cube root ot weight (Bmd\ 
1V45), and lhal length then must be irwres f] \ 
consistently in proportion to weight. However, i he 
average mm of increase It) length for these snakes 
appeared in lie at a much lower rale, proportionally, 
to ihe mean increase in wei.elu particularly diumv 
the second six monihs of the study. 

/ rtti'rh-wcixlti tvluhonshtps; The previous 
analysis su-.\uesled two different growth phases io\ 
weiuhl and length, o\K- dm me the first six months 
and anoihei (mm sis to 12 months. In particular. 
actual incicascs in weight (g/mo) weie highest 
during the last seven months, whilst proporuonal 
l'%/n)0) weight uanis deefmcU. A similar pattern 
u{ decline in proporlional growth in length during 
the last six monihs also was observed, bin actual 
increases in length (mm mo) did noi dii Fpi 

markedly tljioueuoui the study. 

Prom these observations it would appear lhal 
Height increases lus'.er relative to length during the 
lallei six months, This was tested using regression 
analysis of length veisus weigh I for the two growth 
phases, months J -6 and 7-12, respectively. Fiji. 1 
shows the plotted curves and their equations. ! he 
curve for ihe second six months of growth ts sleeper 
than the curve lor the first six months, and the 
ici'iesxion slopes (3 63 and 2.s>_\ respectively) arc 
highK Sign ficamly different <F[I,8| = I6.8 p< 005) 
This difference occurs at D total length ol 800 
900 mm. 


6 li mun*ft» 

HAW 9 ZB t.Vtflh 3 ' 55 ""-. % 

S monlhi. 

LFNOTM » 100 (mm) 

Fife I Lttti lojjplol ol length WeiLhi telaltoiiship )m lu.ih 

i i *\rh ph.iH^ wuh cqiiutiarta ibi line- given- Growifi 

phases basnl oil weight inCmaRi tj hm/i iviw<vn MOOllte 
5 I fable J). 

h»nl Inwke: Avcrayc weight of food consumed 
per month varied from 2fig during the first month 
ot the study Do 190 v _' during the last two months. 
I derail average weight of ingested prev for the first 
fi\._ rnonrhs was 94 g. However, for the Inner six 
months average consumplion o\" prey rose 
dramatical^ to 476 g, ffVC tunes thai tor Ihe 
previous months. I his siulden increase in average 
wciiihi of food consumplion coincides precisely 
kVlth tbe increase m weight observed ai the same 
I mie ( Tabic I; Tie. I ), and occurs at an average total 

Length of about soo-900 mm. 

Cru.w growth ifjtaencv: I he overall mcai* 001 
Ihl Lhappell Island tiger snakes led ud (thinm or. 
mi c was 4N.MSI) { >A). Willi the exception of the 
initial leeding intcival, CiCjF was neariv consraill 
ovci ihe entire study period I his value cumpures 
favour -ably with similat calculations for oiher 
.WLMii.stns 1 1 -Ms-. ihe 1984), and suggested thai 
jneiubolie rate was relatively eonMam ha these 
-aiakes under the prescribed conditions. 




Shine (1978) plotted monthly body lengths and 
estimated age classes and growth rates of mainland 
tiger snakes (Notechis scutatus) from N.S.W. 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 11th 
month of the study at body sizes of 1116 mm SVL 

(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 = 51). 
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 



11 - 


7 - 

5 - 





— i — 



1 1 






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. 

i \V< l\l i lt,l K S\AM i,Ki)\V in 

I ■ 

achieved in about 20 years (about 12 years in males) 
on the Franklin Islands. 

Several extrinsic factors obviously affect growth 
in the natural environment (Andrews 1982). Primary 
among rhese arc (I) the availability of food, (2) rhr 
effects ol temperature, and (.*) interspecific, and 
intruspecifie compeiirion and/or predation. 

Prey types vary markedly among tiger snake 
populations, particularly on islands, Diets are* 
composed almost entirely of fro^s in most mainland 
areas (Shine l*)7H), whilst Maud populations (except 
lot Kangaroo I. and laMiiama) are devoid ol' frogs. 
Tij'cr snakes on many islands, cac small mamma's, 
birds, and An li/ards (Schwaner, in press). Little is 
known of the effects of dilfeur-g quantity and 
quahiyol prey among natural populations of tiger 
snakes (Shine P//8). However, a high correlation 
between maximal body si/e and mean weight of 
available prey exists for several offshore island 
pupulations ot tiger snakes in South Australia 
(Schwaner. in press | 

Seasonal variations in temperatuie apparently 
afHvt tlie feeding habits of tfect snakes on the 
liankhn Islands (Schwa net, in prep.) Adults 
I > 1000 mm SVI ) do nor iced during winter 
moul lis apparently because they are unable to digest 
lan'c prey and apparently do not eat small prey 
L'ndci laboratorv conditions these snake-, 
a'gurejtated large prey ai body lenperatures lower 
than those measured when the prey was invested 
Survival ol island tiger snakes dining wititei may 
depend heavily on ihcir ability to store body ftjtd 
On the Iraoklm Mauds {and on Chappell Island) 
aduh lii'er snakes feed primarily on multonbirds 
tPuj [fums wnutrastth), a highly seasonal ptcv 
(Woiu-ll iy5S; Schwaner, in press). Adult female 
snakes are gtavid dining This season, but Slime 
tlfftft) and Schwaner (in press) indicated that iiravid 
UftOI snakes do not feed. Alter nativetv, the si/e al 
which l he laboratory reared snakes were observed 
to increase markedly in weight relative to length 
(I tg. I) is approximately the si/e at transition from 
juveniles to adults in both the Chappeh Kl.nul 
populanon and the tiger snakes on rhe franklin 
Islands Increased weight »o length latios suegestm* 
tat deposition jUjv therefore be iclaled to the ousel 
ol reproductive maturity in these snakes 

Most islands %uppoiimg populations oi liget 
snakes lack nalural predaiors Kangaroo Island has 
>M.unias f lorvfitfs rosenbetgt) and the urtdeseribed 
pvgmv copperhead snake (Au.sfwtufis >p.), both 
known ptedaiors of snakes. On Reevsby Island 
near Port I incoln. goannas have been introduced 
and may li.MW iwhiced directly numbers o\' lieer 
snakes (Mutschm \WZ). On St liters Island off the 
western coast ol byre IViiinsula near (edima, 

goannas may have reduced the numbers oH riper 
snakes indirectly bv I ceding on inuitonbnd eggs ( T. 
Merfeus, in //'//„ 3iii.K4), Mainland populations ol 
tiger snakes appear to reach smallet maximum bodv 
sizes than most island populations but no direct link 
with ptedators or compeiuois has vet been 
established as a correlated IncLor. 

inhere was an optimum nalural environment for 
feeding and growth in black tiger snakes, growth 
rales in those environments would he expected to 
exceed those ol the am Tier a I laboratory 
environment. Recognition that actual growth rates 
in natural populations vary due to particulai 
extrinsic factors (c;g. variable food availability) can 
be accomplished by comparative studies among 
different island populations both in the field 
(Schwaner, in press) and m t lie* laboratory. 

Whereas variable growth rales appear to be an 
adaptive strategy for ectothermic vertebrates (Poush 
ls»H0), and one which is highly affected bv extrinsic 
factors, a imporiani question concerns 
intrinsic inter populational (intraspeeific) differences 
in growth rates Andrews (1*)K2) has suggested that 
one method ot defining the relationship between 
growth a:ul Mines?; can be explored using 
experimental manipulation of food availability in 
laboratory sellings. C'eitain differences in growth 
rates may occur between sexes, as adaptations to 
particulai life styles (oi dilleieutial icsouice 
availahi lilies) and by wide variation amours 
individuals reared under similar condrtrons 
(Andreyvs 1982). Among island populations tiger 
snakes differ in maximal body size by as much as 
an order of magnitude (Schwaner, in press). Also, 
within and among populations, sexual dimotphisni 
in maximal body si/e is extreme for some 
populations, with males greater lhan females, whilst 
in oilier populations lemales equal males m body 
si/e, or are slightly larjjcr, This study has 
demonstrated that tiger Miukcs Wi\u<Utbi!i4>v ppcW 
at predictable rates Similar comparative studies of 
growth m captive snakes from other island 
populations with differenl maximal bodv sizes muv 
reveal whether these rates are adaptive or are simply 
environmentally determined. 

Ae It n owIim! k m rn ts 

Parts ol this study were s-u poor ted bv grants I tout 
the Souih Australian \\ ildlile Conservation Fund, 
and by the Australian Reseanh Ciianu Scheme. We 
thank D. Bradford and R, Seymour for the use al 
theit regiession analysis propamine and computer 
I da Schwaner typed the manuscript. 



Andrews, R. (1982) Patterns of growth in reptiles. In 
Gans, C. & Pough, E H. (eds.), "Biology of the Reptilia, 
Vol. 13!' (Academic Press: New York.) 

BRODY, 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. Zoo/. 202, 

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). 

Pough, 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. & Rohi.f, 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.) 



by Margaret Davies & Keith R. McDonald 


Topotypic Uperoleia rugosa were examined and their morphology, osteology and advertisement 
calls described. 


\\ Mak(.akli Dwus* & Kiim R. M<_ Don \t \y\ 


U.Wil-. VI., .V \l l>" \\ I K R ( I '.>S5 ) \ lui'itinilioh tf1 / /hn</ri»' f^Uffi <\ihU^, llM ) j \tnna: 
I rpn»J.ic;\ id.K-j Ww* A 1 far \' |m/_ UW£|- tt-42, IK lime. l*>K5.. 

lojuiivjik Upt$\>ft!ftt "<;;<■•../ suu ( £3 mined jiuI ilkii morphology flMPOlogy and .idwf lis* nu-nl eallr. 

Kl V Wo&l-W Moipliolm",, o^colorv, aUwl-i'UKiil cjIIs LfKtfkui fligftM 


I rogi ol die leptodactylid genus Ufwotcnt < iray 
are- small, slum limbed, iossonal animals exhibiting 
limited morphological diversity. A reconf revision 
and subset.)uciu papers by lylci, t.)av ies & Martm 
(l°<Ma,b,e) resulted in die reeoenilion oT IS species. 
I he revision was concerned principally wirh those 
speeies occult ine in dial patt ol' (he eonlineni 
investigated bv the auihois (die kitnbeilcv Division 
ol Wesiciu Ausitalia, and I he Noilhcrn Temlory); 
aueitiion to msa occurring in the eastern states was 
eoiiliued lo an examination ol' type material ol" 
speeies described from dun area. 

Beeause of tile limited inlcrspeeil it* 
morphological divergence , in L'/wro/riit. species 
concepts rely heavily upon biological data, 
particularly cell inl'otmation. 

Uperoli'iu ru^ (Audersson) is one ol' the 
eastern species recognised by l>ler cf at. (1981a). 
Since its description in 1916 it has been reported 
widely i'rom Uuecnslaud, New South Wales and 
Victoria (Barker & ftngg, tV77, Cogger el aL l c >8-U 
T I u- type locality of /_.•'. mgow is reported to be 
Colosseum Qld by Audersson (19I6>. Howevei, a 
label aLConipanvini: the type specimen states it w<J3 
collected at Mt C olosseiun. Colosseum is situated 
on the plains at the foul ol Ml Colosseum and in 
1912 was served by a now abandoned railway siding. 

Durilig a visit to i he area in December l t >84. one 
ol us (K.R.McD.) obtained a series of specimens and 
recowlcd the adverlisemeni call. The availability ol' 
this lopolypic material permits a study ot skeletal 
slruclwrc. Mere we describe the material. 

MiMcrials ami Methods 

Speeiiucns examined here arc deposited in Hie 
NaUuhistorisku Kiksmusccl. Stockholm tNRAM) 

pOlU Ol ?\>ftk}%y, I Iii ( vl'-iiI\ nt AJetaiJc, G P.O. Hn\ 
I'JS. A.lrliMib S \,,vi MM)I. 

( hi.i ii'.l.tna NiiltoiKil Park', <V Wildlife S^tvilc, 
i-.iii .iii'iida, Rmiwvillw, Olvl 4NM 

and the South Auvttaliun Museum. Adelaide 

Methods of measurement follow Tvler rf <//, 
(I9Ma). Measurements taken were: eve diameter (1-) 
eye to naris distance |HN|, internarial vpan (IN). 
snout to vent length (S-V) and libia Igflglti ( 1 1 ). 

Osieologieal data were obiained from cleared and 
Ah/ariu Red S stained specimens alter the method 
of Davis iSi Ciore (1947). Osieoloi>ical descriptions 
follow lrueh (1979). 

Calls were recorded with a LJher 4000 Report 
Monitor tape recorder and an AKO DI90 t ; S 
microphone. Wet bulb air temperatures were 
measured w'uh a Schuhheiss rapKl-readitiu 
ihetmonietcr wittiin 3 cm ol'ihc calling sue. 

Calls were analysed by means of a sound 
spectrograph (Kay Model 6061- B Sona-Graph) with 
the overall response curve maintained in the II -I 
position. Temporal and spectral ctiaraetcrislics oi' 
calls were determined I'rom wide-band (300 H/ 
bandpass) spectro^iams. Because Ol consistency 
between calls both within and between individuals, 
only «rintdc reprvsenlativc .calls of each male weie 
analysed dn<\ mean values calculated 

Ufuro/eni ni%osu (Audersson) 
FIGS 1-6 

Psiu-ttfttfttu^ nt' /'/f.i'mt/ AihJlts>ihi, l*>lh, K SVt?fl 
Vcten^kajisakaU. Hauttl. 52, 9. |> l_" 

Vp&'Ql&tfi'LlgtrW Palkel, 1M4U, NJmilrilL^ ^uol, 4^. p. 7(1 
IpaiU; Ivkr, l)a\ics ft Marlin I9KI, Aa>i. .1. /mil. 
Siif-pl, Ser. 7 { K i' 17, ( Lii'fci l*JS3, Rcpiilcs nnd 
AinplnlVuiH- til Au^lialia, 3iC) edlit, p. Hfr; ( ug^-i, 
( aou-roii iSi C'ojcyet I9fl3. ^noto.c.Kal Catalogue ol 
Au^r.ilij Vol, I Amphibia arwl Reptitia, p. ?4(pail.) 

MutciHil LXatHifiii/ 

NRAM 16.M), tinlnlvpc, uniwJ f , Mt { 'olm^nui 
\24 U. 151 3T') S.| Qlck MV12, I Miohe-i-; SAM 
RZ7U52-3, R2705 k-i IcIcMrod and -aainub. cullino .'-" . 
Ml -no ol former ColuaSffum railwav .-.taiiiMi t^l 211 
151 }~t '), '2Aii.|y,S4, K. R, McDonald. 



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


A small species ( <?<? 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. IB). 

Fingers moderately long, slender, unwebbed but 
slightly fringed with prominent palmar and 
subarticular tubercles (Fig. 2 A). 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 >L Metatarsal 
tubercles moderately large but prominent. 
Subarticular tubercles rounded, i Dorsal surface 
faintly tubercular. Parotoid and inguinal glands well 
developed; coccygeal glands poorly developed. 
Submandibular gland prominent, discrete (Fig. 1A). 
Cloaeal flap with tiny fimbriations. Ventrally, throat 
and thorax granular, belly smooth. Male with 
unilobular, submandibular vocal sac and glandulai 
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 


The addition material ranges 19.75-21.7 mm S-V. 
Relative lengths of tibia arc consistent (TL/S-V 
0.36-0.39). E-N/IN ranges 1.24-1.67. The specimens 
vary little morphologically from the described 

The snout is more rounded in two o\' the 
specimens, and the belly is faintly granular in 
others. Cream patches on the dorsal glands are 
slightly 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 hololype, 
although dorsal colouration and patterning can be 
observed. This feature conforms with Andersson's 
description (see Tyler et a/., 1981a). The 
pigmentation of the ventral surface was not reported 



Fig. 2. A palmar and H plantar views of the hand and fool o\ male Upcrok'ia rugma (SAM R27052), 

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

Osteology (based on SAM R27055) 

Skull well ossified, sloping anterovent rally. 
Sphenelhmoid not ossified dorsomedially or 
ventrotnedially; small portion ossified posteriorly 
to medial extremities of palatines, with ossified 
portion extending about 1/3 length of orbit in 
ventral view. Prootie not fused with exoccipital. 
kxoccipilal confluent posteromedially in dorsal 

view but not fused ventromedially. Crista parotic* 
short, stocky, not articulating with otic ramus of 
squamosal lying alongside lateral exi remit ies. 
Grooves of carotid canal present on frontoparietals, 
medial to cpiotic eminences. 

Frontoparietal elements extensive, crcnate 
medially, overlying spbenethmoid anteriorly, more 
widely separated anteromedially. Frontoparietal 
lontanelle barely exposed medially; anterior margin 
formed by spbenethmoid elements at level about 
1/5 posteriorly on length of orbit. Posterior margin 

-II : 


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

undefined because of lack of anteromedial 
ossification of exoccipital. 

Nasals large, closely applied medially for 4/5 
length, slightly separated postcromedially (Fig. 3A). 
Anteromedial extremities crescentic. Maxillary 
processes moderately acute, directed ventrally, not 
in bony contact with pars facialis of maxillary. 
Posteromedial^ 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 
posterolateral!^ 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. Quadrat ojugal 
robust, in firm contact with maxillary. Squamosal 
stock> 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, xiphisternum present. Sternum 

Fig. 4. A lateral view of pelvic girdle and B dorsal view 
of vertebral eolumn of Upero/eia rugosa SAM R27055. 

cartilaginous. Clavicles slender, curved, closely 
applied medially. Coraeoids well developed, robust, 
widely separated medially. Scapula bicapitate, 
slightly 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 > IV > sacrum > II >V = 
VI >V1I =VIII. Bicondylar sacrococcygeal 
articulation. Crest on urostyle extending approxi- 
mately 1/2 length. Ilium with tiny dorsal 
prominence and moderately large lateral protuber- 
ance (Fig. 4A). 

Kl 1)1 I IMI ion Ol UPERQUitA fti/GOS, 

I lumcrus wiih strongly developed anteroproximal 
Crest. Phalangeal formula of hand 2,2,3,3; small 
hony prcpollex, palmar sesamoid. Phalangeal 
Formula Of fool 2,2,3,4,3. Well developed bony 


\ mint ion 

A second lopolvpie specimen was examined 
(SAM R27056). The specimen varied in the nature 
Oi the extremities of the IVonloparietal elements; 
anterior evlremilics were truncate and medial edges 
weie straight, not civnate. The nasals were in 
tenuous conlacl with Lhe frontoparietals. The 
exoccipitals were noi confluent posieiomedially. In 
olhci features, lhe skeleton approximated the above 
description closely. 

Advertisement cull 

I he advertisement calls o( three individuals were 
recorded al Colosseum on I2.xn.1964. The cat! is 

6 1 


2 - 


~ 1 



I it*. 5. Scmagram of male .uhenisement cull o\ IfpetOkUt 
ru&ava SAM rUl7flJ3} T Temperature 22> <. , 

I \b\ i I. Advert tomtom call t'haravtm'$tic& etf'-tlpcrokSd lugasa tn Coiasseutn, Oiarns/und. 


No r 

61 pulses 


I'lil'a' rCpCtltlOJQ 

laie (pulses sec '] 


<H/) * 

Wet- bulb 

lemp, c 

SAM U27052 
SAM fc27053 
SAM R27(>55 





1 16-7 


3 f i.5 5 


22. 8 
22 2 


a short note of four pulses, pulsed at about 34 
pulses see ' and with a duration of about 
117 msec. The dominant frequency is about 
1*583 H/, A sonogram of the adveiiiscmctu call is 
ShOWn In lig. 5, and ehaiaclerisiics of the- calls ate 
listed in Table I. 

Ha hi! at 

liogs were located on lhe lower slopes and 
adjacent plains ol Mt Colosseum, the lower slopes 
arc disturbed forest of //. tcssellaris [-. Much., E 
tcreficorttt\ Suiiih, /.'. crehra, and Melaleuca 
quiliquenervJa (Cav.) S. I. Blake. Most of the plain 
is heavily disturbed and is mainly grassland with 
scattered large tress ol I., terotfcorni.s (Fig. 6). 

The frogs were found around temporary surumcr- 
filled pools and were calling in short grass on hanks 
1 6 m from water. C ailing was heard only following 
a thunderstorm of approv 25 mm rain. 

Oihci frogs heard and seen in the area were: 
Utoria atendca. /.. fallux, I. rubella, I . rothii, I. 
uier/ti/y L nttsuttu /.. ulbo^ut/nta, Nanidella el. 
deserucoUr .\detotu\ twvix fjttinedyriaste.s peroni, 

/_. icrtvetvutnae, I . omattts, I , tasmaniensis and 
bufo tt/ari/uts. 

Andersson (1916) described the habitat at 
Colosseum as open totest-eountrv. 


I [g. (y Hjhmil ill whi 

were found neiu I 

iilllilL! ni;il 

pcn>lct({ i 'v 

.timet tailwav Million. < olov.cuui. 




In revising Upero/eia, Tyler et ai (1981a) redefined 
the three named eastern Australian species, U. 
rugosa (Andersson), U. fimbriamis (Parker) and U, 
laevigata Keferstein, all from the type specimens. 
They resurrected U. fimbriamis from the synonymy 
of Li rugosa in view of their experience with 
restricted distributions amongst taxa elsewhere in 
Australia, and because oi its larger size. 

They also indicated the existence of a number o\' 
undescribed species in eastern Australia that await 
resolution. Cogger (1983) considered the eastern 
Australian species to be composite. 

Since the revision oi' Tyler et a/. (1981a), 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 Upero/eia 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 et ai (1983) gave the 
distribution of U. rugosa as N.E. coastal, S.E. 

coastal, Murray-Darling basin, Bulloo River basin, 
Qld and N.S.W. (A.C.T.). The A.C.T. records 
probably are based upon the work of Robertson 
(1982 1 , 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.). 


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 
oi^ 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. 


ANDERSSON, L. G. (1916) Results of Dr L. Mjobcrgs 
Swedish Scientific Expeditions to Australia 1910-1913. 
IX. Butraehians from Queensland. ft Svenska 
VetenskapsakacL Hancll. 52(9), 2-20. 

Barker, J. & CRtGQ, G. (1977) A field guide to 
Australian frogs (Rigby: Adelaide). 

COGGER, H. G. (1983) "Reptiles and Amphibians of 
Australia", 3rd edtn (Rcid: Sydney). 

, Cameron, E. E. & COGGER. H. M. (1983) 

"Zoological Catalogue of Australia Vol. 1, Amphibia 
and Reptilia" (Aust. Govt Publishing Service; 

Davis, D. D. & Gorl, V. R. (1947) Clearing and staining 
small vertebrates. Fieidiana: Techniques 4, 1-16. 

LlTTl [JOHN, M. J. (1967) Patterns of zoogeography and 
speeiation in southeastern Australian amphibia, 
pp 150-174. In A. H. Weatherly (Ed.) 'Inland Waters and 
their Fauna' (A.N.U. Press: Canberra). 

Robertson, J. G, M. (1984a) Acoustic spacing in 
breeding males of Uperoleia rugosa (Anura: 
Leplodaetylidae). 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. CapeUt 

1979(4), 714-733. 
Tyler, M. J. & D.wus, M. (1984) Uperoleia Gray 

(Anura: Leptodactylidae) in New Guinea. Trans. R. Soc. 

S. Aust. 108, 123-125. 

— , — & MARTIN, A. A. (1981a) Australian 

frogs of ihe leptodaetylid genus Uperoleia Gray, Aust. 

J. Zoot. SuppL Ser. 79 1-64. 
, - — ■ — & - — (1981b) New and rediscovered 

species of frogs from the Derby-Broome area of Western 

Australia. Rec. West. Aust. \1us. 9(2), 147-172. 
— , & (1981c) Frog fauna oi' the 

Northern Territory: new distributional records and 

description of a new species. Trans. R. Soc. S. Aust. 

105(3), 149-154. 

'Robfrtson, J. G. M. (1982) Territoriality and sexual 
selection in Uperoleia rugosa (Anura: Leptodactylidae). 
Ph.D. thesis. Dept of Zoology, A.N.U. (unpub!.). 



by S.J. Edmonds 


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. 


by s. J. Edmonds* 


LdmonUK, S. J. (I9S5) A new species of Phascolosoma (Sipunaila) from Soinh Ausiialia. Items, /< Soc. 
S, Ami H)<>(2), 43 44, 2K June, 1985, 

A new species of sipuneulan, Phascolosoma kapalum sp. nov, dredged from off the coasl of New 
South Wales, is described. It is distinguished from other species of Phascolosoma by the shape of its introvert 

Key Words: Sipuncula, Phascolosoma kapalum sp. nov., 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*, 151°56'E, 

off coast oi New South Wales; dredged at 710 in, 

during cruise of "Kapala" coll. P. Colman & D. 

Brown, 6.\ii.l977, 

Type specimen: Australian Museum Sydney: W 


Paratype: South Australian Museum: E 1694 


Trunk: Length 13-32 mm, maximum width near 

mid-region 4-7 mm. Subcylindrical to spindle-like 

and sometimes covered or partly so with coaling 

o\' very fine particles o\" 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 ^A-Ya 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 o\ 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, subclavatc 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, subclavatc forms 
0.3-0.4 mm in height. 

* South Australian Museum, North Terrace, Adelaide, 
S.Aust. 5000. 

Fig. t. Phascolosoma kapalum sp. nov.. entire specimen 
(scale bar = 5 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 pair 
more anteriorly from muscles 6-7, 6-9 or 7-9. 
Dorsal and ventral muscle on each side fused 




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 Va length and opening 
to exterior at about same level as anus or just 
anterior to it. 

Alimentary eanal: 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). 


Edmonds, 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. 



by Michael J. Tyler & Christopher A. Miller 


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. 


i>v Mich m i I. rvi bR & chrtstomii r a. mu i i u- : 


h i ic M I -V \i.) 1 1 •■, i \- NV&5) Sih taw iiKlHit\ini\' t»i iIil- Liois.d epidermis 111 tawratfuit I 

/a«k A'. w. \ 4w.\r, io^;:,i. * - *s, ::s June, I9B5. 

rwO aasiinct Umiic. yf surface archileepne ol ilii- epiJwriui mtiamOlft epithelium oeciu id \usiKiliaii 
hoi-- ui 1 1n Umilios I Ivlidac and l.eptoduciylidae: *<nc ennsisiiny <i! ail cUhnmie patient ot micioptie.i-.; 

i lie uita u nutoiilum lai'ii Fipftfjtt* exhibits cml) flic form bill *pisciwi \hn\ are cicely related phyjugcncticulb 

Eiiul ecologically may differ > n 'I 11 ' tialutt ol' the epithelial mm hue aKlniccMM.. 
K< V Wokom I lop, SIM, epidermis, ulltasinictitre. 


I Ik dorsal skin ul' frogs is an cMtvmcly complex 
tissue, containing mucous and grumilai glands 
whose contents are discharged to the external 
soi fecc via dUClS (Hkam IWEOi ThC mucous glands 
are imdet the conlrol ol sympathetic nerves; the 
niUCUS liberated assists tcmpeialutc control via 
cvapotalivc cooling and, hy maintaining ci moist 
mu lace, also enables the dorsal skin 10 function as 
a tespiraioiy surtacc (I illy while. I9?l>. In contrast 
the secretions of the granular glarufa ul many 
species are toxic, and at lord these animals 

protection agaifis4 some preduior^ 0^«ily A Myers, 


I he ventral skin is aelandular in the \asl majotily 
ol species and either smooth Ut distinctly granular. 
t iiih.ihiii.illv it is involved in waict Uptake, and 
&P&1& liable to c.\ticinc water loss adopt a stance 
in which granular skin is adpressed to the substrate 
during pciiodtol active restoration of body watei 
(Stillc, 1958; Johnson, 1%'M. 

I lie histoloev en tioe has been reviewed by 
I lias & Shapiio (1957) and I lk;m (19<58), Fhesc 
studies have documented uross struct 11 1 e and 
described the glands dispersed throughout the skin, 
In addition, the secretions of auurewal ions olptands 
10 hum parotoid Wands (as in $ufp species) or calf 
elands {in the Australian icptoduclylid species 
( i/i/!t(uh/nistc\ tiutitvtili) have hcen ihe Mibjccl of 
•.pc. .-iIk iiudtes (low, 1972} Croch & tyler, I9H1). 

AS is typical fill veitebrate animals, the outer 
epidei mis is composed of squamous epithelial cells 
attain'cd as a slratuiu cot ncum. ElCClrCfl 
iimioseopii sludies ol the outer siirlace ol 
squamous cptt helia fa various veilebraies has 
icvcalal till elaborate pallet n ot elevated 01 
deptcssed whoils and Mlawkes. [974j 
laluenbnch & Knutson, NTs; Andrews. 1976; 

Kaii-uiKuh. Harding & Siwin-. l^ftftj. 

Dfp.Miiiurii -it /on lO) I 'i i;\ ol Adelaide *' Ifl 

(.I'd.. \a,:i;na,. S Mist Wl 

llciv we have examined rhc ulirasirueture ol 
doisa! cpidennal cells in Australian frogs, I'Q 
establish the nature ot the diversity in surface 
urehilccluic and its |>ossible relationship 10 the 
systematic arrangement of those species It Includes 
more genera and species than any previous sm w-\ 
ol' skin architecture ol the Anura, ami is the fiist 
investigation ot Australian species. 

Materials und Methods 

Adult representatives ol the lollowiue 1 1 veueia 
and 50 species were examined: Arh'/otn\ hnns, I s\ ( / 
diHillfflOnh <- ritiiii ^eoiyjunu, Cyclnrutni uu^*iuh\. 

( . QUhriftiSt c. fofiglpe& C fflQttltosHS, < nwlttu 

(\ pttttycvptoaJubi iieieiopoms cuct, Lechrio(lm> 
fleieherl 1 unttodynostes 1 osvexfit/fi'Utu^ / • 

tfmttt'nit, I firm/ins, L sfieftcet% I . fasftttWiettst^ 
> UOftO ii'iihti'h'nsis, I allxy'uitaid, I hic<\lor I 
ct/L>tHle<{, /. ihtuns, /. K.'pLitidi, I <luhhi> I 
dent a lit, (- m f ingi t I. JitHuk, f. fr&y&itoeit, /. 
"judk-titti, /-. me/tuts. I pailicUh I pmmi I 
roitijonnfs, L mihfh I . ruhellu, I., nnmcu. 

Megislofotts IfenwitiSi tytxophjya f<tsci<>iam\ 

\eohnttuchu$ picM, NL sutt.n, NQMtdW hoincfft, 
V tnciatiosca/yliHs, P&eudopluynt* fiifmmh H 

i>uetttfwri, Kamtlcllu rt/inr lit, U. si.atnjit'u. 

Riwohaifiuhm tiluv, Urudai'iyttts twutirosirfot 1 

diwtins, ( 'ffvmlt'ia imtnthtfu and U. Wi'hydfitWil 

lite selection of species ict'lecis an aiu-mpl lo 
sample diverse genera, but was subjeci to the 
availabilitv 0\ utatcuvd. 

Por lions (5 X 5 mm) ol dorsal skin were removed 
fiom frogs that had been killed by contact with a 
_V'<> solution ol chloral hydrate, fixed \l\ W\\ 
lormahn whilst still In a relaxed suite and stored 
ul 65"" alcohol I he tissues were critical-point dried 
usmv the liquid COr amvl |iiocedtue in a Polaron 

L 3000 ci", drier, 01 else simple air-dried in « 

clesiccalot. Specimen^ were coaled with 15 rtttl 
carbon and 20 mil ,uo>U palladium |S0:20). and 
v icv\t_d in in Ml* i eaiinii 1 iii an microwopc 

ill 2(1 Kv. 



fig. I. SEM view of squamous epithelial cells of skin of Cyclorana main/ showing hexagonal shape. 

Fig. 2. SUM view ol dorsal skin of: A. Litoria freycineti showing tnicroridyes on ihe cell 
surface. B. Crtnia georgiarm showing a reticular cell surface. C and D. Sloughing of 
skin of Uperoleia irachvdenna. In D [he central portion is the aperture to a dermal 
gland duct surrounded by intercellular connections. 



In the species examined, the surface cells of the 
stratified squamous epithelium are polygonal 
(usually hexagonal) (tig. 1). Upon these cells two 
forms oi surface architecture occur. One is a 
complex pattern of curved elevated ridges 
corresponding to I he "microridges" of Havvkes 
(1974) and the "mieroplieae" o\' Andrews (1976) 
(Fig. 2A). The second form is in the arrangement 
of an extremely dense reticulum (Fig; 21i>, 1! 
appears similar to the type o\' surface that 
Kaltenbach et al (1980, p. 323) described as having 
"a spongy appearance". 

I he incidence of these two forms o\ surface 
architecture oi' the squamous epithelium in the 

species and genera examined is summarised in 
Table l. 

Tp establish the existence o( ini rasped fie 
variation, dorsal skin from six L'ttoriu cwin^t and 
six Ra/tidtdlu tiparia was examined. Within each 
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 variety 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 
cedysis. Examples arc shown in Figs 2C and 2L> 
The projections from the surface of the cells in Fig, 
2D are considered to represent intercellular 

1 "Mil i 1- Incidence of hvo forms <>l dorsal, dermal arch ffet (arc cfflfOrigsJ 17 gertew and 50 speck's of Australian frogs. 

< ilMIU 1 


No. o\' Spp. 

Surface architecture 
Microridees Reticulum 



I cchrioilus 
I imnodvnastcs 






f'scudo/dtt ync 

Rani del fa 
































cue ru leu 

















/t,u nanus 


seller i Hi 




bonnet ti 












d turn as 



H Hvlidae; L - I epiodaet\lidnc 



connections. None of the preparations examined 
demonstrated any evident intermediate stages, 
documenting the change between the presence o\~ 
the intercellular connections and the formation of 
the final surface architecture. 


The results demonstrate that two forms of surface 
architecture of the squamous epithelium oi' 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 et 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 o\' 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 (Cyclorarw 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. dah/ii were placed 
by Tyler & Davies (1978) in a species-group with the 
microridge surfaced /_.. 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. longipes 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. 


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. Kcmpster for assistance in the 
preparation of the figures, and to the referees o\ 
the manuscript for constructive comments. 


Andriws, P. M. (1976) Microplicae: characteristic ridt:e- 
like folds of the plasmalemma. J. Cell Biol. 68, 420-429. 

Crook, G. A. & Tvi i R, M. J. (1981) Structure and 
function o\' the tibial gland of the Australian frog 
Limnodvnastes dumenli Peters. Trans. R. Soc. S. Aust. 
105, 49-52. 

0\\\, J. W. & Myers, C. W. (1967) Toxicity of 
Panamanian poison frogs (Dendrobates): some 
biological and chemical aspects. Science 156(3777), 

Owns, M., Mariin, A. A. & Watson, G. F. (1983) 
Redefinition of the Litoria latopalmata species group 
(Anura: Hylidae). Trans. R. Soc. S. Ami, 107, 87-108. 

Ei i \s, H. & Shapiro, J, (1957) Histology of the skin of 
some toads and frogs. Artier, Mas. Novil. f!819), 1-27. 

Hi kan, E. (1968) Mucopolysaccharides in the an u ran 
defence against desiccation. J. Zool. 155, 19-53. 

I- ahrtnhm ii, W. H. & Knutson, D. D. (1975) Surface 
adaptations of the vertebrate epidermis to friction. J. 
Investigative Dermatol. 65, 39-44. 

Hawkl-s. J. W. (1974) The structure of fish skin. 1. 

General organization. Cell 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 ultrastructurc of the cornea and adjaeent 

epidermis during metamorphosis of Rana pipiens: a 

scanning electron microscopic studv. J. Morph. 166. 

Iji i vwhitf, H. B. (1971) Thermal modulation of 

cutaneous mucus discharge of evaporative water loss 

in the frog, Rana catesbetana. Z. vergt Ph\'siolo$>iel3. 

Low, B. S. (1972) Evidence from parotoid-gland 

secreiions. //;: W. E, Blair (Ed.) 'Evolution in the genus 

Bufo\ pp. 244-264. (University of Texas Press: Austin). 
Sin I i , W. T. (1958) The water absorption response of 

an anuran. Copeia (1958), 217-218. 
1'yi i k, M. .1. & Davies, M. (1978) Species groups in the 

Australopapuan hvlid frog genus Litoria Tschudi. Aust. 

J. Zool. Suppf Ser. (63), 1-47. 



by J. A. Long 


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. 



by J. a. Long* 


I i i* • I- \ i( (IW5) A tltfu dVLilfCOU-j chiiuuciid (Pisces UoKrtX'pllflll) Irani Souih \n ...hi. //<///, A'. 

> dupftodvn cvtvnsis Kp. ttov, in described from a su.ule leli mandibular looihplatc from the Aptun 
liiillJi)}! Shale, wovi ot Bopeeehce SiJinu,, northern South Ausltalia. E, eyren\t.\ is characterised hy aj) tfiyulftl 
oral margin mul by tbc shape and arrangement Of he lour triiors, ( OJllpftrfcOAb ftrf made wiih olhcr 
< 'iclaceous and Teniary chhuaerid dentitions. 

Ki s WORDS Pisces. Holoccphali. Cretaceous, South Australia, new species, tt/ttphn/t < rrtvr.wv. 


J he chiiiiaerids teached a peak of diversity 
during I he Mcs.voio, and though uumetous genera 
are known from the Jurassic and Cretaceous of 
North America, U.S.S.K. and t.uropc (Newton, 
JK7o. Woodward. IN<>2, P>I2; llussakofl, I'M:; 
Obruchcv, l%4) ihe only one*, desciibed from 
Australia aie Etfaphodon sweety E. mirahitis $j\& 
teckyaziUh mot font fton. the Tertiary (Chapman & 
Priiehaid, |907; Chapman & Cuduiore, 1924). Die 
ceiiir fuhi/i/iodttii ranges from harly Cretaceous to 
Pliocene, with most species being Laie Cretaceous 
in age. The specimen described here uas found on 
the floor of Lake Phibbs. just smith ol Lake Lyre, 
and Undoubtedly came from outcrops of Ihe nearby 
Apltau Bulldog Shale (I udbrook. 1%6). 11 is 
therefore not only Cite lirsi iccord ol a fossil 
chuuacrid Irom the Mcso/oic of Australia (and the 
Only chimuend fossil Irom South Australia) but also 
one o\ ihe earliest species ot EfJaphodVtt. 
Cretaceous ehimaetid tooihplates are also known 
lioiu New Zealand but these belong to 
Cttllorhymhtts hectoH and l.schyintus hrevirostris 
(Newton, 1876). 

Syslemaiic l\ilm , onloluy.\ 






Type species. Chimuvru numtelli Buckland I.S^.S. 
Cretaceous of England (Ward* 1973). 

Kemurks: Ihe specimen is referred to hdemhodon 
because of its broad syiuphysial lacei which 

Department ot ecology, l lie i\uttnaK*ri National 

I'nneiMiy, HO Hm 4, t ;.nhctru ( its. \.< .1., An Mi.iha 

expands in breadth posteriorly, the number and 
nirangement of its triiors and the absence ol a 
thickening oi\ iis outer lace (Hussakoff. 1912. 
p. 202). 

lulaphodon evrensis sp. nov. 
FIGS I, 2, 31, 

IMS. 1 A Ijiyr looihpkue ol ' LiLwIunlon tony, p. i\ 
|*)£4 htu/jfiodi'ti ->\\ I onj' A. himci. \\ Mo 

Etymology. Aflcr lake L v re, neai where the 
specimen was found. 

Diagnosis. An Edaphodon having a mandibulai 
loolhplale twice as long as btoad with four tutors: 
of which the two posterior and the single 
symphysial ones are large, and the lateral median 
one is small. Oral margin angular with anterior ends 
of outer and lateral median triiors forming right 

\fitfcriuL Only one specimen, the hololype SAM 
P24770, maximum length- 110 mm, 

Oixvrremv from the floor of the Lake Phibbs 
approximately 21 km west o( southwest from 
Uopcechee Siding, wesr oi Marree, South Australia. 
I owe. Cetaceous Bulldog; Shale (Apiiau). 

Lhscrifidofi. The bone is weathered, with the 
senndentine of (he irirors being chalky. Despite this 
lite overall shape of ihe toothplnte and outline ol 
the tritors are well preset vcd. 

The oral margin of the mandibulai toothplate 
tligs. I, 2, 3f) is quite angular as each of the two 
laical tritors fOUWf li'itoi, OT, lateral median tritor, 
I MT) form fight angles with Ihe bone in between 
them. Overall the toothplate is rhomboidal with its 
breadth exactly half the length. 1 he tsvo large iritors 
in ihe posierioi hall (outer irilor, OT; mesial tritor, 
M I ) are each about a quarter as broad as long and 
in contact lot close to half their lengths. The median 


.1. A. LONG 

r j 



;■ .- 







- .. - •' 

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. 

Nl \\ ( Kl I M. 1-dL'S t 1I1MAI KID 


lateral tritor (LMT) is situated vcnlral 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 Hatter in the posterior half. The 
symphysial facet (st) is broadest posteriorly, with 
the ventral margin being gently curved. 


Toothplates of ehimaerids show a wide range of 
shapes and tritorial arrangements which led early 
workers to confuse various genera and species 
(Huvsakoff, 1912). A scries of mandibular 
toothplates of Edaphudon mirijicus 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, 






Hg. :. A. hthsphiuliut eymtsfc sp. run., Cretaumus, Smith Australia, ffolowpe in oral view. \i. ti. .swceii Chapman 
<& Piiictmrd. Miocene, Victoria. Composite n?Moi*tion of left mandibular tpoinplai-e based mainly on NMV HI 6076*). 
I Ml. lateral median tritor; MT, Mil, M 1 2. mi-.ial iiiuu .: OM, oral margin? OT. outer tritor; sf, symphysial tacei; 
ST, symphysial triiot 

v i.onc; 

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. agassiz'r, D, E. tatigerus; E, E. 
back fundi; F, E. evrensis 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 E. 
mirabilis Chapman & Cudmore, 1924 both from the 
Miocene and Pliocene of Victoria. Fig. 2 shows a 
comparison between E. evrensis 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 

shape of the toothplateand arrangement of tritors 
can be restored (NMV P160769). E. sweeti differs 
from E. 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. E. bucklandi, E. agassizh 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 
Pachymytus (Woodward, 1892), a possible precursor 
to the line o^ Edaphodon. 


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 MeNarnara (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 

Ml \\ ( Rf i"\( [ "Ol.S CHIMAl RID 


Mary MacDougall for typing Ihe manuscript. This Rothmans Postdoctoral Fellowship in the Geology 
work was carried oui under the tenure of a Department of the Australian National University. 


Agassi?, I . .1. k. (184?) Recherchcs sur les poissons 

fOS$fles, Neuehatel and Soleive, 5 Vols., 1-188. 
Ben ki and, W. (1835) A notice on the fossil beaks of four 

extinct species pf fishes, referable to the genus 

Chiwaerut which occur in the Oolitic and Cretaceous 

formations of England. Proc. Geot $oc. Lond., 2, 


(1838) On Ihe discovery oi fossil fishes in the 

Baeshot sands at Goldworih Hill, 4 miles north of 

Guildford, Pud., 2, 687-688. 
CASH k, E. (1966) Fauna ichthyologique du London Clay. 

Brit. Mas. (Nat. Hist.), London. 496 pp. 
Cm \pm \n, I ., & CUDMGKC, F. A. (1924) Some Caino/ote 

fish remains with a revision of the group. Proc. R. Soc. 

Vict., 36, 107-162. 

— & Pkik hard, G. B. (1907) Fossil fish remains 

from Ihe ternaries of Australia. fhicL 20, 59-75. 
Copl , E. D. (1869) Descriptions of some extinct fishes 

previously unknown. Proc. Boston Soc. Nat. Hist., 12, 

310-317. ' 
HussAKoii, L. (1912) The Cretaceous ehimacroids of 

North America. Butt. Am. Mt4S. N#L Hist., 31, 195-228. 
It tin, J. (1856) Notice of remains of extinct vcrlcbraled 

animals of New Jersey, collected by Prof. Cook of the 

State Geological Survey under the direction of Dr. W. 

Kitchell. Proc. Acad .\ttt. Set, Philadelphia, 8. 220-22L 

Lost.. J. A. (1982) The history o\' fishes on the Australian 
continent. In P. V. Rich & E. Thompson (Eds), The 
Fossil Vertebrate Record of Australasia', pp. 51-85. 
(Monash University Off-set Printing Unit: Melbourne). 

- & TuRNhk, S. (1984) A checklist and bibliography 
of Australian fossil fishes. In \L Archer & G. Clayton 
(Eds), 'Vertebrate Zoogeography and Evolution in 
Australasia*, pp. 235-254. (Hesperian Press: Perth). 

LuD BROOK, N. (1966) Cretaceous biosiratigraphy of the 

Great Artesian Basin, South Australia. Geo/. Surv. S. 

Austu Bull. 40, 223 pp. 
Newton, E. T. (1876) On two chimaerid jaws from the 

Lower Greensand of New Zealand. Quart. ./. Geol, Soc, 

Land., 32, 326-33L 

— (l878)Chimaeroid fishes of the British Cretaceous 
rocks. Mems. Geol. Surv. U.K., Monogr. 4, 1-62, 

GbrucheV, D. V. (1964) Subclass Holocephali 
(Chimaeras). in Y. A. Orlov (Ed.), 'Fundamentals of 
Palaentology*. Vol. II; Agnaiha, Pisces, pp. 353-399. 

Ward, D. .1. (1*973) The English Palaeogene Chimaeroid 
fishes. Proc. Geol. Assoc. 84, 315-330. 

Woodward, A. S. (1892) On some teeth of new 
Chimaeroid fishes from the Oxford and Kimmeridge 
clays of England. Ann. Man, Nat, Hist., 10, 13-16. 

(1912) The fossil fishes of the English Chalk. 
Palueontogr. Soc. Mongr., 264 pp. 





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. 




Many recent new distributional records for reptiles in 
South Australia' - v reflect inadequate field surveys and 
poor documentation of the state's herpeiot'auna. Here we 




Ml Mi"i 





• i.'iLjKtitinaH^lcl , 




c3 ° \ 


. 400 



lip. I. Distribution of Lerista terdigttata. Area under the 
dashed line in I he inset denotes the previously recognized 
range of the species in Australia, Solid circles denote 
S.A. Museum specimens. 

report a significant range extension of the fossorial, scincid 
lizard, Lerista terdigttata (Parker, 1926), in South Australia. 

The range o( L. terdigiiata was "islands, coast and 
hinterland of the Great Australian Biglu ,u (Fig. 1). 
However, specimens recently collected from near Billa 
Kalina Hstd. 29 55 % 136*11 E (SAM R2099I-92, R2I022), 
and from the Ml Morris foothills, near Amata aerodrome, 
26 OS'S, 131°05'E (R25883). in the Musgravc Ranges in 
the far north-west of S, Ausl, extend the distribution oi' 
L. rerdigitata about 600 km N of its pre\ iously recognized 

Although the Mt Morris specimen (a male, SVI = 
51.5 mm, TL = 64.0 mm) has the normal scalarion of /.. 
terdigttata from more coastal localities 4 * the paltern is 
markedly different (l-ig. 2). Bold stripes replace the 
longitudinal series of spots on I he dorsum, and much 
larger spots (nol 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. terdigttata in the Musgryve Ranges 
suggests that the species also may be found in adjacent 
areas of the Northern Territory and Western Ausiralia. 
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 Mi 
Morris. Roman Ruehle provided Fig. 2, and Cila Sehwanei 
and Diana Massacci typed the manuscript. 

Miller, B. & Schwancr, I. D. (1982). Trans. R. Soe. S. 
Ausl. 106, 79-80. 

Setmaner, T. D. & Miller. B. (1984). ibid. 108, 215-216. 

Schwaner, T. I>. & Miller, B. (I9S4). Ibid. 108, 217-218. 

^Cogger, H. G. (1983). "Thc'Repliles and Amphibians of 

Ausiralia" A. H. & A. \V. Reed, Sydney. 

Slorr, (i. M.. Smith. L. A. & Johnstone, R. E. (1981). 

"Lizards of Western Australia, I. Skinks. University oi 

Western Australia Press & Western Australian Museum: 






. - rv : 

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 
MILLER, 10 Yarrow Crescent, Hope Valley, S. Aust. 5090. 


by J. A. Smith 


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. 



I he Onkaparinga estuary is Minted approximately 
10 km south pi Adelaide on Ileutieu Peninsula, and 
extends some 10.5 km Jiom lis mouth ai Pi Noarlunga 
on Gull M Vincent to the township of Old Noarlunga. 
In iceeiu times the estuary lias conic under renewed interest 
as a recreation site which has led to the dredging of 
sections in Pt Noarlunga. There is, however, some concern 
lor the environmental effects ot extending these works to 
Other regions. 

During the period May to July 1983 two pressure sensing 
tide gauges were deployed in the estuary (Pig. I), one 
5(H) m upstream liom the tnouih f A) and the other I km 
downstream from the head (B). This enabled a continuous 
record of levels to he obtained at each cw^i o\" the estuary 
over this period tFig. 2). Water heights were taken from 
each record at hourly intervals and used to derive tidal 
eonstaitls hv a least squares lit technique. The cstumv 
eonsianis were then used in a comparison with constants 
derived lor Pi Adelaide (Outer Harbor) using data from 
(lie same observation period. 

The constants lot the four major primary constituents 
< fable 1 ) show a general decrease in amplitude and increase 
in phase lag as the tidal wave progresses from the gulf 


){( *v c ■'--" " ; 

fJijAOLLjN L.A\ \\) '. -. 1-0 \ 1'"! 


/ nt.K A PARI U B4 \ \ 

n i ■: eh J i 

i . . , , l 
t i nLr - B )},, 



Pig. 1. Onkaparinga Estuary, South Australia; A, Site o\' 
tide gauge near rnoulh; B 4 Site of tide gauge near head; 
C, Shallow r-eeion. Mean river depth given in m. 


1 5 

JUNE 1983 




I ig. 2. Water levels lor period llth-27ih lime 19S3; (a) Old Noarlunga |B in Fig, 1); (b) Purl Noarlunga (A in fig, I); 
(cj Port Adelaide (Ouici Harbor). 

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. 

Tablf. 1. Tidal constituents for the Onkaparinga Estuary 
and Port Adelaide. 





A g 

A g 

A g 


17.3 19.1 

11.8 69.0 

9.8 94.3 


25.7 47.3 

17.8 82.5 

12.7 121.1 


51.7 106.0 

19.8 124.1 

11.1 176.1 


50.0 173.2 

19.0 190.4 

12.8 248.0 


0.7 158.6 

3.9 97.4 

4.2 243.4 


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 (cm) 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 ( VgH, where g is the acceleration of gravity 
and H the mean water depth), 1 and demonstrated the 
importance o\ friction in the estuary system. 

Tabi r 2. Wave travel times 









g hi -g x The phase difference between stations A and B 
( ), T.T. The wave travel time = S^'Su (hrs),<rThc wave 
speed ( 7msh). a 

This observation prompted a theoretical study using a 
finite difference numerical model to simulate the estuary 
flow. 2 

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 scagrass and other vegetation that presently exists in 
the estuary. 
1 Dyer, K. K. (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. 


by C. J. M. Glover 


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. 



Pour maiinc species (Prtstis sp., Ophisurus serpens, 
Jhporhamphus reguiaris reguiaris, Scorpis lineo(utus) are 
recorded from South Australia Tor (he first lime. A 
treshwatcr species {Amniatuha percoides) is recorded from 
the Stale Tor the first lime on the ba.sis of a known 

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 
water species {Pnstis sp.) is unexpected and it evidently 
constitutes & vagrant (see earlier reports 1 ). 

Amntataha penalties has been recorded widely in 
inland waters of northern Australia. It was listed for South 
Austialia in the W20's :, \ but (hat record was neithei 
confirmed bv any Known collection nor sustained by 
Subsequent authors. 

All specimens are deposited in the South Australian 
Museum (SAM) 

Fig. 2. Ophisurus serpens, SAM I 4566. TL BBS mm. 

Family Ophichthidae. Ophisurus serpens (Linnaeus, 1758). 

A specimen (Fig. 2) was collected from a rock pool EU 
Coobowie (approx. 35"03'S, I37' J 44'E), on 7.U970, by 
M. -I. Stanton 

serpens has been recorded previously in Australia 
from oft N.S.W., Victoria, Tasmania and W.A. S . It is 
recorded elsewhere in the Indo-Pacific and eastern Atlantic 
Oceans, and in the Mediterranean 4 . 

Family Hetniramphidae. Hyporfwrtiphus reguiaris 
reguiaris (Giinther, 1866) 

Ff& I. Snout of Pristis sp. (/? zlfam ■'). SAM F2064. 
TL (lota! length) 3^9 mm 

Family Pristidae. Pristis sp. (probably P. zijsron Uleeker, 

The lostral pioccss (Tig. I) ol a specimen caught 16 km 
offshore Irom Glenclg, (approx. 34' 59 'S, I3*T 20 'E), in 
1936, bs L. G. Kelly, is registered m ihe South Australian 
Museum. The intact specimen would have been about 
I 300 mm totaJ length. 

The rostral process has 29 paiis of teeth. This is 
consistent wilh that of P. zijsron, one of the three pnstid 
species recorded in Australian waters 4 . 

Pristids are found mainly in the world's tropical and 
sub- tropica I waters'. In Australian P zijsrun has been 
recorded in coastal, estuarinc and fresh inland waters of 
Queensland, New South Wales and western and northern 
Western Australia 4 *'; its range reportedly extends to the 
East Indies, India and Sri Lanka 7 . 

The present record is the first for (be family Pn*lidae 
from Australia's southern waters. 

Fifteen specimens (SAM P4803) were netted m Angas 
inlet, off the Port River (approx 34° 48 % J3K' J 32'L), 
on 30.\iii.iy84, by S. Aust. Dept of Fisheries officers. Two 
additional specimens (Fig. 3) were hooked at the same 
locality on 23.ix.l984 by I. L. Lloyd. 

H. reguiaris occurs only around the southern half of 
the Australian continent, where it has been recorded from 
southern Qld, N.S.W., Vic. and souih-we stern Wj\."\ 
Two distinct subspecies are recognised, one on the 
continent's south-east coast [H. reguiaris arc/elio), the 
other on its south-west coast (H. reguiaris regular -is) u) '. 

M. F. Goman advised me (pers. comm., 19K4) that the 
S. Aust specimens are I J, reguiaris reguiaris. This record 
therefore represents the most easterly documented 
occurrence of the western sub-species. 

Fig, 3. Hyporhumphus reguiaris reguiaris. SAM 14831 
TL 255 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. 

5. lineolatus has been recorded only in Australia, from 
Qld, N.S.W., Vic, Tas. and at Lord Howe Island. 8 . 

The present S. Aust. record represents the species' most 
westerly documented occurrence. 

Family Teraponidae Amntataba percoides (Clumber, 1864), 

Two specimens (Fig. 5) were netted at the Neales River 
crossing on the main road south-east of Oodnadatta 

Fig. 5. Amniataba percoides. SAM F4792. TL 54 mm 
(largest specimen). 

(approx. 27° 54 'S, 135° 46 'E), on 3.V.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. 11,12 . An early record from S. Aust. 2,3 
was not substantiated by any hitherto known collection. 

The present record is the species' most southerly 
documented occurrence 12,13 . 

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 

(1984). Trans. R. Soc. S. Aust. 108(2), 

'Glover, C J. M. 


2 Waite, E. R. (1921). Rec. S. Aust. Mus. 2(1), 1-208. 
3 Waite, E. R. (1923). "The Fishes of South Australia!' 

(Govt Printer: Adelaide). 

4 Munro, 1. S. R. (1956). Fisheries Newsl. 15(9), 15-18. 
5 Lindberg, G. U. (1974). Fishes of the World. (J. Wiley 

& Sons: New York). 
6 Whitley, G. P. (1948). Fisheries Bulletin No. 2, 1-35. 

(Western Australian Fisheries Department: Perth). 
7 Marshall, T. C (1964). Fishes of the Great Barrier Reef 

and coastal waters of Queensland. (Angus & Robertson: 


s Last, P. R., Scon, E. 0. G., & Talbol, E. H. (1983). 

Fishes of Tasmania. (Tasmanian Fisheries Development 

Authority: Hobart). 
9 Wheeler, A. (1979). Fishes of the World. (Ferndale: 


l0 Collette, B. B. (1974). Rec. Aust. Mus. 29(2), 11-105. 

"Lake, J. S. (1978). Australian Freshwater Fishes. 

(Nelson: Melbourne). 
12 Merrick, J. R. & Schmida, G. E. (1984). Australian 

Freshwater Fishes. (J. R. Merrick: North Ryde, N.S.W.). 
"Allen, G. R. (1982). Inland Fishes of Western Australia. 

(Western Australian Museum: Perth). 

C. .!. M. GLOVER, South Australian Museum, Norih Terrace, Adelaide, S. Aust. 5000 


byS. A. Shepherd, P. S. Clarkson & 7. A. Turner 


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. 



I Ins nuk- describe* the spawning cycle of Huhotis 
xvattim 1 cacti and Ihe settlement density ami growth ol 
juveniles a\ the species hi Wcsi Mand (35 37'00"S, 
MK 15 (K)'i:i, s-nith Australia. This spawning cycle differs 
marked t) from Mini previously described for the species 
□ I lipan.i Kocf'i SOUTH Australia and is therefore b) 
considerable interest. 

Samples oJ *> 12 serially mature female H. siufanv 
65 KO truti long were collected at Abatone Cove, Wesi 1 - 
at about nioiuhU mieivals from February 1983 July I9N-4. 
I In- entire viscera] tnaw was preserved in 10% formalin 
and sca-waicr mm\ later sectioned in the laboratory, t joss 
u-iiiiins ul the g0iia<j and digestive eland were traced on 
transparent plastic, the em out sections weighed, and the 
respective areas ol gonad and digestive gland calculated. 
A gonad index was then calculated with the tut inula 
CiOnild hide* UK) (Area ol gonad) (Total area ol 

section). I he annual reproductive cycle of this species at 
VVcm I, proved in Ik synchronous within the population 
so that measurement of oocyte-ova diameters to 
distinguish ilu- suites ol the reproductive cycle wqig 
uunccc'.saiy (sec Shepherd & laws' tor details ol the 

Sea surface temperature dala were obtained with a 
mcrcurv iheinioiueiei Bl about monthlv intervals a! We-KJ 

A hand lens, desirnc-d tot use underwater with about 
3 iiujnin .uioii (the opitcs arc described by Shepherd 
& TUmei in prep.) was used to search lor H, scahtris on 
oir-.tose COfflilLne algal substrate, ihe preferred substrate 
hw settlement 01 this specie* (Shepherd & Tinner in prep.F 
Searches were done lot b(J- 100 mmules at about monthly 
mietuits in the boulder habitat at Wesi 1. at 4 5 m depth 
and the lengths ol all individuals to about 25 mm long 
recorded to 1 mm. 

» . 


I ly I (lipfiri] mc:m innmhlv -.c.i -.nrlacL kmipeiuiuics 
(smooihcd curve); (mid) mouihK disiubuiiori ol 
gonad indices wait slandaul emu-., (lowct) density of 
-in. ill //. \a/h/rrs i.. S mm al iiub silc 

Changes in the mean monthly gonad indent ol // 
sculuris Irom February iyS3-May 1984, together with sea 
surface temperature dala arc given in Fig. 1. Ihe index 
Increases from wittier to mid summer and declines 
abruptly from late summer through the autumn. The 
increase indicates maturation and increase in size oi 
oocytes in the gonad, and the decline of the index indicates 
ihe onset o! synchronous spawning in the population. 
Spawning appears to have commenced latei in I^K? than 
in 1984 and to have been more complete. In \9$A ihe gonad 
never became completely spun but commenced to inciease 
in si/e again in July. 

Thus IL scalar \s has a late summer to autumn spawning 
season (February May) which is synchronous ihroug.houi 
the population, and appears to begin at aboul the time 
of maximum summer sea temperature. This spawning cycle 
contrasts strikingly with Ihe asynchronous cycle of //. 
scutum at I iparra Reel whem spawning potentially occurs 
iluoiighoitt the year. 


-■■1 a 

1 | 


1*. \U< 

13 \\> 

n 1 



'"■ | 

♦r- M»v i.i 

- 1 

.' . . I , 

- .. V 

I ig. 2. Length frcqtieno disuihuiions foi //, waldm oi 

WeM Islam! from Feb. nun 10*,! u , vi;iv [Oga, 

Of the various lauoi-. dial are known 
to influence the spawning cvclcs ol abalcnc eg. tem- 
perature/ photoperioj.' ami food abundance" only the 
laiiei shows a marked dillcrcncc between the two sites. 
Drill food algae are seasonally in short supply m I Iparra 
Reel but abundant iluoiigliout Die year a] Weil (., ' but 
this is unlikely Jo account for the differences in spawning 
beiweeu the sties. I miner studies of //. smhtns arc 
nn.cssaiv, especially in 01 her parts o\ its geographic uiugc, 
to elucidate the problem. //. rubra Leach also shows 
marked, hut unexplained, differences m spawning cycle 
between these iwo sues ' Ihe onlj other abalonc species 
thai wi; know of with a similarly variable spawning cycle 
between localities is Nations ntfesvvns Swain son " ' 


The density of small H. scalaris (measured in mean 
numbers of individuals s£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 

Y=2 99 ♦ 0043 X +0 000087 X 
R 2 = 99 

1983 1984 

Fig. 3. Plot of mean length (with standard errors) of 1983 
cohort of//, scalaris from March 1983 to May 1984. 
A polynomial regression of best fit to the means is 

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 

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. laevigata 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 1 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. & Laws, H. M. (1974). Aust. J. Mar. Shepherd, S. A. (1973). Aust. J. Mar. Freshvv. Res. 24, 

Freshw. Res. 25, 49-62. 217-257. 

^KiUuehi, S. & Uki, N. (1974). Bull Tohok. Reg. Fish. ^ £ *> * Dt ' Martini - J " ° < l977 »- Ca,if " ** ^ 

P I k ^ fiQ 78 GaiTie 63 ' 80 " 9 

KCS. L3D. 35, t>y-/». ^^ y y (1974) publ Mar uh Busan |7ish Co| j 7f 

'Kim, Y. & Cho, C (1976). Bull. Korean Fish. Soc. 9, 2 l-50. 

61-68. Shepherd, S. A. & Hearn, W. S. (1983). Aust. J. Mar. 

4 Giese, A. C. (1959). Ann. Rev. Physiol. 21, 547-576. 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 



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 Vperoleia (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 

McNamara, 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 177 

Shiel, R. J. & Koste, W. Records of rotifers epizoic on cladocerans from South Australia 179 






VOL. 109, PART 3 



by Wolfgang Zeidler 


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. 




/i 101 1 r, W. ((9R5) A new species o! crurtaceJUl, tSyiiearida: Auaspidaeca: ItOOBUDfldRt), tmm sinkhole 

and cavCS tn itW BOUth-Wl Ol South Australia. JYans. /<- She & .bo7. IWCV), 63 75, 29 NoVWflbfiB ' - 

A new species of syni fli Id I TOtai ean. Konnuuva vrenofMm sp. nov. is described from sinkholes and 
saves near Ml Gambicr, South \u-.traliu. The species oeeuis til IIS face V ! ClBttofl and to 8 depth »>1 4fl DL 
Superficially the new species resembles A., cunn/r hut females are at least twice as large, reaching Iciinihv 
exceeding 20 mm, (lie sternal ptO BfiS D| Ihfcl laic potASlDfl lacks 9 pOSfcriOJ pHnJcctiW *Pd the e\v ., 
abstrtt, A Uriel dlftCfiptiOfl Ol She dcvdOpOJCni Of tbfi tuali P£l ISIW and sensory organ on the antounule 
is given. 

Kiv Wokns SyiKJuda, Aiuspidaeivi. Kaormn&Q itv/iu/um <p. nov., South Australia, taxonomy 


In Australia, anaspidid synearid crustaceans arc 
known to occur throughout lusniama and Victoria 
and an interstitial species, Psammaspides wilfiawsi. 
has been found in the New England area ol Nov. 
South Wales (Sehuuukc lV>'/4). Hie l.r.manian 
fauna is relatively diverse and to date eight species 
have been described. Most of mainland Australia. 
however, is too arid to support anaspidid syncarids 
and besides ft wtlliumst the only Other described 
species are Knonunga cursor Sayee, 190K and 
StygOCartS gtselae Schminkc, 1980. Several 
undescribed species of Kootwnga arc known in 
occur in Victoria (Orummond L&59) and atso on 
King Island and if) north western 'tasmama 
(Williams, W. D. 197*1). Ii is likely that more specie* 
remain to be discovered as aquatic habitats are 
surveyed in more detail. K. cursor occurs 
sporadically throughout southern Victoria, usually 
in small permanent or semi-permanent swamps 
in pools in streams which flow only alter heavy rams 
(pflUnmond 195V) and i! has recent l> been recorded 
bom similar habitats in north-wcsietu Tasmania (De 
Deckkcr 1980). S. gtselae is an interstitial spt 
and is only known from the type locality. Battle 
Pointj l.mibo River, Victoria (Sehrninke 198(7). 

The species described here was found in sinkholes 
^\\k\ caves in the Mt Gambier area, South Australia 
(Fig, I). It was first recorded by Zeidlcr (1983) being 
the first record ol a syncarid from South Australia. 
It belongs to the family Kooiuinghtw n:. 
supei tiemlly tesemblcs A. cursor. 

Materials and Methods 

Specinwnti .wh i lei m ibej o\ 

sinkholes and caves in the south-east ot 

• Sotlth Australian Museum, NorlhTer tact fa I 

Ausl MM>a 

(Fig. 1) with a hand net from amongst algae and 
aquatic plants near the surface or by entrapment 
in a glass jar while .scuba diving. The "I " number 
given with each locality refers to the identities 
number used by the Cave Lixploration Group ol 
South Australia to distinguish each cave and 
sinkhole in t he lowei south easL A total ol 236 
specimens (60 o' and t76 ' wete collected and 

Specimen length is measured fa n ! • v 
rostrum to the tip of the telson. Specimen width 
is measured as the width of the thiid pleonite 
segment (usually die widesi segment). Meon length 
excludes the telson as the limit of the telson is 
ObSCUred by spines 

Material tepoucd hcie is deposited i" i 1 "" South 
Auslralian Museum, Adelaide (SAM), \usP 
Museum, Sydney (AM), Museum ol VictOI ia, Mel- 
bourne (NMV) and the lastiuunan Museum jrid 
Art Gallery, Hobai t ( I'M). All of the - ; 
preserved in 75% alcohol or 2"/i> h:Tti-aldL*i 

propylene glycol solution. Of the types onh 
holotypc, allotype and paratype r(5AM c ■ 

have been dissected (partially) and dissected 
appendages have been preserved wit 

The following abbreviations are u- -. | '«*\t 
Mxp. ■ maxilliped (f hotaaipoU l|, V I 7 - 
pereopods 1-7 (th<"-i- OPodS 2-8], t Jt 
pleopods 1-5, LWS - left hand side and KIl^ 
right hand side when viewed dorsal. 

Ko<Hfnni;.i emuvrw »p. no* 

surface at|gac and aqu amed 

•ankhoi; kno*T» as 'Tosol Cavr" U 81), Apr) 
5J km 3] anoola < aves i Pr nets 1 1 

S. Auar. bj v. Zddlet, I.lil.t982. 



tyjCBcr sam G3999 t^j> collected from same 

locality ny R I l-.-rnc. H.iii.1981. 

t>u>u!,:>c; AM CJ992 (>) collected P. Hnnic, 

S.ui.1981, guoe locality; SAM C399I tt), am 
061 i.i. P35W»wK NMV Jioxo^tvi. JI08IU 
i M ■ \IM% (vi, G2849 (ft collected by W. 
titer, ' "'.1982, from same locality 

Othrr mafertai examined Sa\ M C\1*W-1 (Hv'm. 

hum type locality, at 10 m depth by r. Hume, 2fM.I98J; 
sam C399MI5 fc*& from ivpc focaHry, l J Honw 
8 III lifcl; SAM C_W5 {70 Vs). J i urn type Locality, W. 
Z,-idl.:r. i.fiU?82; SAM C1996<*Us?V 4V*S). from rvne 
locality, W /vH.llrri.jMJ K I GMtflttt, l6.vii.l9K4. SAM 

C3W |J v>. Ihofc Cav* U230). jusl E ol type locality. 
coliecwd near surface by R Home, 6aU983; SAM C39W 

ll oi, Allendale Sinkhole (Lll), .entie of mum tCMUl 
AJIen*i 'U' t.'si approx. 20 km Sol Ml Gambicr, collected 
in 'nial darknc'-s at 27.5 m depth bv I* Home, i.ii 198J, 
sam r39W |4 a% Beaara Sinkhole iL32 33), neat 
••Ucndra". 6 km vvsw ol Mi Gunbiei, collected at 19 m 

dcpih. by K Home, livi I9N2. SAM C40K) (1 V), trom 
Mfno lot ahrv, ai 5 m tleplhj V Home, 27-KJ1.J9&4; sam 
* ' WOO (2 v\), CCibbyi Hole sinkhole (L46), approx. £L5 km 

w oi M Borleterr— tJ k«i sw <»i vit Gambler, collected 

.. lOmftptll »'> r Home 6ii.l983, SAM C'4001 (12 J\ 
2 .:.%). trom some lotatu*.. at J in depth t p. Hone, 
n ii l*82;tfAMC4<K)2<lrf| Devils IMnebtoawl sink) i 

I ■ ' ;;:■:>'; LfcfflSWof "BauiOolui"— jppro.v. 15. km 

sw ..1 \u Gstftttier; collected from wwtj ; : IJJm depth 
ty '- Hon r ! lu 1981, SAM C4003 (4Vsi, from same 
locality. Ai 40 m depUi, P. Home. i.iii.lV8l. SAM C40W 
[\ f, W*lnut One .L5M appro* 18 fctfi SW of Mi 
I will t. ,11-.] <1 km due \V of Ml Schank, collected at 3 m 
fjcpfh fcl) P Hmi.„, j:i ,. ivSt, SAM 04005 [2 r :\), '" lit I 
Koad cive <L6?) ( appro*. *i3 km NL or KapjpmfljB 
20 km S*V of Mi Qarotuer, cujlpcted near mruce by R 
Morn--. Kn I'>h: SAM r4<H)6 [2 6%^ Bcmlwto 
s.iiklio!. i; 64) in Caroline Iorm. apjinout 15 km SSE- 
Of Mr (tamrncr, collected ai II) m depMi rw H » j - i 
l^.'W.SAM.: 4l*j7<m H^vtnr<1-S|fr:nH. ,.„ (| 7||, 

appicu j'i v mi t: ol EwWJi PanUs--2J km SSL u 

m-.t. collected ne»i surface bv p Home, 22. m. a 
fc£AMC40OMf| v. t ,/k Mtuihole'ma.l.ui, (I <;) fema- 

• FdfKt, approv d km SI of lajiLanonta C.n^., 
tvtllecrcd at u m tiopm hv l^ Howe, ^ I i'»K ', s,\M r4lT9 

. I ./j. A.ii.vm. ( :rr sinkfuOv |LH/«2 
Mudtt"|*i collected SUTbcc ►«,- p H»>nn- Ju.mi.I98.?; 
5AM ■' lv!0 ii IvkKiw - inkhoU ti i:-i. S km 
W ol V.itlw lake, Mr C^ainhicr. collected at 3 m depth 

Home, il.xii.iw:. SAM < 40H i'. .'i, Mucftraoni 
k v,7 11.132), jppir-A 2 km l al Bwsnfi Rondv cotteowd 
Bl 3 in K F HQflW :-* I 16 3AMC4OI20 S), imnanicd 
Mnkrioic (Ll44k in M 1 GnmUa Rortth, amstne. 2 km N'w 

:.t Ml Cafltftltl AitpOft, c"l(uL'eJ nc:ir M.r'.*i;r h\ I* 

H< .-. I4.»j,19ft2i SAV1 ,:i " ),:i J, ' v:. ii- Shall |i 159), :*ppr«i\ 3 kin W oi 'Vfrndatc Dail, 
wollccteO at 5 rtj depth ^y P. Jlomc, M.ili I5"84, NAM C4014 
ti J|. from itac Niv. ai J ro depth, P. Morno 

OHO ?)l DW ftulliXk Hoi' Ml.k lull: 

it jfiiy, on **BfiiiHHiltir"pryip:;r!v -/ipp'tiv. M huSW ptl 
Mi Ciaintnci, collecteri at 27.5 rn deprh h) P Hum;, 
ft.ui.WK2; SAM C4027 44 ?\ I rf) t QlCtlCUe West i ive 

.. ,i. T t| ( t. v \ i :.v ftfl IP. H ,ih.. S9.HU |984;SAM 

I • i -i-m.i, .. ,n- ari 2i», Mi Qambien 

In 'm-II. 'h 4 I I985. 

mh: critfiem toWB a&fl heii«t siwhied in Morgan* 
Cave <L34> aj»d lh< Pine; Owe a <rl I, tonh 

Alleyns Cave; Hells Hole sinktiole (L40) just north 
of Bottlcbrush Sinkhole and Simpsons Hole 
sinkhole (L42) between Devils Punchbowl sinkhole 
and Gums Road Cave (Fig. I). 
Description of hohtype: Female measuring 
18.3 mm total length, 3.4 mm width General 
appearance illustrated in Fig. 2. First thoracic 
somite fused with head Percon wilh seven free 
somites and pleon with six free somites and a telson. 
All body somites wtlh one pair of appendages. 
Anus opens at posterior limit of pleomte 6. Pleon 
(excluding telson) longer (7.7 mm) than percon 
\h 2 mm) and broader than percon and head. Head 
Itucth equivalent to first 3itf pereonUes, broader 
than pcreonites 1-4, equal to pcreonite 5 and 
narrower than pereonites 6 and 7 Pleonitc 6 longest 
body segment, pleonites 1-3 broaden. Pleotnie 6 
with row ul six well spaced spines, near dorsal- 
posterior border interspersed with 1-2 fine setae. 
Body segment pn.gressively more robust from 
anterior to po&tetlor to about pleomiL* 4 

! rectangular (JJ ■ t8 mm) with short. 
pointed rostrum and distinct anterolateral incision 
atrovt attachment ol antenna, Pronounced shon 
mid-lateral transverse sulcus. Eyes absent but 
pigment (wore concentrated near an tero- lateral 


Antennuic (Fig. 4aj; poduncle of Ihree segmcnU* 
outet nagellum ol 56 (IMS) arid 48 (RHS> segments 
and inner flagclhim of 16 (LHS) and 15 fRHS) 
segments. Uuier llagellum about 3 5* length of 
inner. Peduncle and outer t1a>»ellum as tptkg as 3/4 
body len^lh- ftasal segment of peduncle broader, 
almost as long js following two combined- Basal 
segment of outer flagellutn with oblique dorsallv 
serrated inner margin Presence lk absence ol 
TUtoeyst m hasat segment of peduncle could not 
be determined wilh certainly. 
Antenna thig. 4b): sliyhrw more- than 2/3 length 
antennuic. peduncle ol 4 segments and single 
tla^eliuni (A 33 segments (I HS ft KHS). Basal 
%cc»i.eni short, segments 2-4 elongate, rectangular, 
Ntgrnerii 3 longest 

lippeT l.p tf.g. 3a): rows of short bristles on both 
sides ai extremity and small tentn*! depression on 
aboral .surface near extremis 
MfllldlblQS (PlgS 3c-U: 3 .segmented palp; middle 
.pi vtxaily enlarged, about twice lengrh brtsal 
demerit; tcimmal segment small, rounded; 
leathered setae ak<n= nin- i maigin tif segment 2, 
apically on segment V Base ol mandibles *tout. 
cnuiu^ in '.v r c(l developed molar arid inci'iOr pro. 
Molar process wilh small grinding surface 
sumuiuded by numerous \piniform setae. Ind&or 
proce* i" left nwodibk with seven denticies 
arranged in ^"shatH.-d iow. Oenricic nearenl molar 



Fig. I. Localities where Koorwnzu crenarum sp. nov. has been collected (•) or sighted (o) Scale bar - 5 km. 

reduced. Incisor process of right mandible with live 
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 ouier 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 

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 iobe 
with six feathered spinules of varying lengths 
terminally. Other lobes with more numerous (> 10) 
but similar terminal spinules which fan out laterally 
to oral and aboral surface. 

Maxillipcd (thoracopod 1) (Fig. 4c): 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 

V. /I 11)1 FR 

\ i : - 2i K0onun$a vrvnarwa m\ -v-*-. ItoIOl • :-?l>ar 

i distal margin Propodu* robust, slightly 

shorter than mcuis, with tWO oblique FQWS of strong 
spines on oral surface and r u ft of K I •«■ un 

abural, distal, outer cOi fttt. Daca ■ : u small, 
rounded, aimed terminally with one birgi and Ihree 

smaller si mil J claw like spines OJld FfiW BCtM I 

n i tnafgitt. 
TVrwipods (ihorar. i simil.u in 

structure to Map but with basis expanded proxi- 

h i- i as wide as coxa and becoming 

■i jsmvcK mote slender with all segments mora 

elongate Pi sliglutv s| 01 tCI thad MXP bul slightly 

, i*2 & P3 "t equal length P4 as long 

us Mxp and longe*! percapod. P3 shortest PC 

;fi it) ■ ,. .. nl Y shorter thai 

nilar in tenth to PI. All pcieopods 
Hexed between merits and carpus and in backward 

pusii' Pii which flexes inwards and P7 

n wards. Coxa wild setose tube Qi 

r, distal u"<nuT b-.M< .'.'■ ailsahle an 1*1 but 

pro?:* .^ccloptti to H un 1*6 bin 

.it on P7 P '•• with two unequal 

epfpn in Mxp. I'l 

segmented exopoclifc eonfciittng ol' large nasal 
hi reaching well past ischium and fiugeiium 

Of 14 |6 segments (table I) each segment be;n 
hvo long fealhered setae. P6, basts without rspoditc. 
P7 without cpipoditcs or exopodite. Dactyl Claw* 
similar to Mxp but P5-7 have one additional daw 
Spermatheca- two small, ovoidal plates, slightly 
raised from the sternum |usi lotwaid ol base of P7 
an last pertoaitc (Fig, 5a] 

Plcopods (Figs 51) -1): all of similar structure, 
lacking endopoditcs but with long muhi segmented 
cxppotfires '.(insisting of short-stout basal joint 
lollov | »ngei more slender one and flagellum 

ol 13 segments in PI 1-4 and 13 segments in PI 5, 
i bearing two long feathered setae. PI 1-3 equal 
in length and longest pleopods. PI 4 slightly shot ttt 
and K 5 -hones!, about 3/4 length PI 1-3. PI M 
also flattened laterally, PI 4 & 5 round in 

ross section. 

I impods (Fig. 5h): peduncle stout, rectangular, as 
long as lelson and almost 3/4 length plconitc 6 
[measured vcarratly), projecting to slightly more 

than L/3 telson length; few scattered short spines 
Dn dorsal surface and ueai ouiei margiu. Outer 
ramus almost I 1/4 peduncle Length* Slightly longer 
than inner ramus, with long lealheied setae along 
inner and outer margins and row of short strong 
upturned spines neai outer dorsal margin. Inner 
minus with long feathered setae along outer margin 
and partly terminally, icimuning terminal margin 
.villi three (LHS)or loin (RMS) long spines; dorsal 
Intra margin with row of Strong upward curved 
spines for about proximal J/J, steadily incrca-.v 
in size terminally followed bj comb <rf smaller 
spines, ceasing al first terminal spine. 
Telson (Fig Mi): triangular in shape with rounded 
apex ii' 1 Slight^ convex lateral margins; length 
(excluding spines) slightly less than width; margins 
wilh complex array of spines except for provimal 
1/4. consisting oi' close-set fringe of short, stout 
i 11 : . -iiir.dlv-. 20 evenly spaced, long spines 
directed posteriorly and about 14 slightly Shi 
spines directed dotsally interspaced with I \ 

■ i 
Colour: light tan with some specimens darker than 
others. Those captured in deeper watei and in total 
darkness do not vary in colour horn those caught 
near the surface. Preserved specimens only slightly 

Description oj otiotypet Male measuring 14. X mm 

in total length and 1.5 mm in width, essentially 
ami lei lo holoiype except lot the following 

diHeieik' i 

Anteimulc (fie. 4e). With characteristic 0V»l shaped 

ory organ ::t win from second segmerd of outer 
Ea Hum Basal stegmenl oi lonci flagelhinr saucer- 

iory organ Outer flagcllmn of 



Fig 3, Koonunau crenarum sp. nov. holotype V, mouihparis. 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 I; Scale bar - 0.2 mm. 



Fig. A. Koonunga crenamm sp. nov. holotypo r j. a. antennule; b. antenna; c. antennulc, allotype d"; d. antenna, allotype 
<$; e. Mxp: F-J - PI 7. Scale bar I mm. 



Fig. 5. Koomwva avrwrum sp nov. a- spermaiheci*, ? (SAM C39VI);b-f PI 1-5 holotype 9; g. tclson, allotype 
cT; h. ielson unci uiopott. holotvpc S; £ cursor; i. .tnienor ol 2 specimen (S.2 mm) from Railway Drain. Bayswater. 
Vic. (SAM 04016). 





lAHi.r l. MerfctiC lfot& fat koonunga crenarum xp. nov. ftoloiype (Hj, allotype (A) and paratypes /Pi. All 
mexsutvmettti (ram dorsal aspect. Unless mdkvted Otherwise /he number of segments refer to the ffagettum. 

"D" (lanutzeti or broken, 


H \ 

A .-f 

P ?s 




Length (nun) 




3.0 - 37 

2.0 - 2.4 




5.5 - 6.7 

3.4 - 5.0 




6.7 - 8.3 

4.4 5.8 

Tel son 


J. 2 

1.0 - 15 

0.7 - 1.1 

1 oi . :t 1 



16.5 -20.2 

10.5 -14.3 

Anicnnule LHS (RHS) 

13.5 (13.6) 

9.9D (13.!) 

in.9 (1L6)-14.9D H3.7D) 

8.4(8.21-12 (10.3D) 

Antenna LHS (RHS) P.*) 

7.5 (7. ID) 

7.5 (7.4) 10.5 (10.4) 

5.4(5,1) 7-5(7.8) 

RaiiO Picon Peieon 



1.17- 1.25 

1 16- 1.32 

Brcadllt (iviim) 




2.3 3.1 

1.6 2.1 

PcrfOttltc 7 

3 : 


2.5 - 3.0 

1.8 2 3 

Plconite 3 

3 4 


2.S - 3.2 

2.0 - 2.6 



1 « 

M - 2.0 

1.1 13 

Anicnnule (segments) 

Inner ramus LHS (RHS) 

16 (15) 

14 (14) 

15 (15) 17 (16) 

70 (11) 14(15) 

Outer ramus 1 MS (RHS) 

56 (48) 

39D (59) 

45 (50) 57 (57) 

43 (43)- 55 (43D) 

Antenna (segments! 


8 (35) 

27 (33D) 

34 (28>-40 (40) 

24 (24) -33 (37) 

I'euopods (*cgmems/e\opodite) 

1 I.HS(RHS) 

M (14) 


I3(I3)-15 (15) 

10 (I01-I2 (12) 

2 LHS (RHS) 

15 (15) 


15 (15)- 17 (18) 

11 (12) -14 (14) 

3 LHS (RHS) 


16 (16) 

15 I15)-t8 (D) 


4 LHS (RHS) 

15 05) 


15 (15) 17 (18) 

12 (11)14 (12) 

5 LHS (RHS) 

1? (15) 


14 <14)-17 (16) 

11 (11) 14 (14) 

Pleopods (-segments) 

1 LHS (RHS) 


15 (15) 

15 (15) 16 (16) 

II (11) 14(13) 

2 LHS (RHS) 



15 (15)- 16 (16) 

11 (1!)-I4 (14) 

\ I HS (RHS) 


15 (D) 

15 (15)-16(D) 

11 (ID-14 (14) 


15 (IS) 

15 i.D) 

15 (14)-I6 (D) 

11 (11) 13 (14) 

5 LHS (RHS) 


13 (13) 

13 (13)-M (Ml 

9 (D) -12 (12) 

Uropod spines 

Inner ramus — innct doc&fl] 

1 his (RHS) 

12 (12) 

11 (12) 


|| (11)12 (12) 

1 >nin t;mnifc— otilct dorsal 

! IIS (RHS) 


18 (17) 

1? (15H7 ,17) 

13 (13) 18(17) 

59 segments (RHS; LHS broken); inner flagelluiu 
of 14 segments (LHS & RHS). 
Antenna (Fig. 4d): left with flagcllum of 27 
segments, right broken bul still wiih 33 segments. 
Pereopods: similar to hololype bul all wiih foUf 
dactyl claws. PI 6 without coxal lobes. 
Plcopods: I and 2 with endopodite^ modified to 
form complex copulatorv styles (pctasma). PtetaSflM 
(Fig. 6f0 directed anteriorly against mid-ventral 
surface between, and posterior to, last pair of 

Fndopodite of PI I (Figs 6b, c) unsegmentcd, 
gfdoved for about middle half of external inner 
margin to provide sleeve for endopodite of Pi 2; 
cup-shaped terminally, directed inwards and towards 
body. Lndopodites joined about 2/3 from base by 

coupling spines on extension of internal, inner 

Endopodite of PI 2 (Figs fid, c) of 2 segments of 
about equal length, each as long as endopodite of 
PI ]. 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 
wit I) sternal process. Distal segment apically 
pointed, hollowed out on distal, inner margin to 
form concave depression directed towards body. 
Sternal process (Figs 6d t e): triangular in shape, 
slightly longer than wide, slightly less than 1/2 
length of basal segment of endopodite of PI 2> 
fitting inbetween basal 1/4 of endopoditcs ol PI 2. 
Telson (Fig. 5g): slightly more pointed apically than 

Rg. 6. tfoonungS CfMOrum sp. nov. a. pctasma, allotype J; b, endopodue Pi I ventral view; c. reverse of b; d. endopodite 

PI 2: ft reverse ol d; I i endopodites PI I (LHS) & PI 2 (RHS) ol' Specimens 7.9, 8.4, 9.3 & I ft J mm length 

, r tm-l>, A evrxor: J 6.6 mm (SAM C40I6); U enUcpodllfi PI I; k. reverse of j; I. cmlnpodiic PI 2; m. reverse of 1. 


in type, lateral margins straight or even slightly 

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 C3991. The inner ramus of the 
uropods usually has three large terminal spines but 
in SAM C399I 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 holotvpe. In the male 
SAM C3992, P2 & 7 are the longest and P4-6 are 
the shortest, equal in length and slightly shorter 

rhan the Mxp; the pleopods are like those of the 

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/pcreon 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 i0.5 mm (±95% c.I.). 
MaJes ranged in length from 7.8 to 16.2 mm with 
a mean length of 10.9 + 0.5 mm (±95'*7o c.I.) which 
is significantly smaller than that of the females 
(P<0.01). The female sample is larger (176/60 ?:<r). 
However, seasonal fluctuations in the proportion 
of males in the population may account for some 

'S 5 



• A '*€• «• • A 



/A «• • • 

Am jam • • 


▲ Atom • 

A ••» 

3 - 







■ ■ ' ' * i i ■ 


• • 

* • • ■ 

o • a o • •• 

• • r • • « • 

a • CO • 



Length (mm) 



Fig. 7. Koonunga crenarum sp. nov. Relationship between length and breadth (pleonite 3). Females closed circles, 
males closed triangles; open symbols refer to specimens from localities other than the type locality. 



of the observed Imbalance as 45/128 specimens were 
mules in July, 1984 compared to 0/11 in January. 
[Mb *W W Match, 1981 and 3/28 in March, 
1982— all from ihe type locality 

Development of the petasma and sensory organ 
oj mak' 

A relatively lane collection ol males from the 
lype locality in July, 19X4 {SAM C'3996) included 
a number of very small specimen? which exhibited 
various stages in the development of the petasma 
and also the sensory organ pf the iuiicnnula The 
development of these sexually dimorphic characters 
in syneui ids has noi been reported in the literature. 

The endopodires of PI I and PI 2 develop laterally 
from the base of the plcopods until they overlap 
and are fully developed; ihey arc 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 at a 
length of 7.9 mm (Fig. 61*). At this stage the 
endopodites are only recognisable as single, small, 
rounded segments and the sternal process is not 
visible, In the next stage at 8.4 mm (Fig. 6g) the 
endopodite of Pi 1 is beginning to differentiate but 
the sternal process has not developed further. At 
9J him (bijk'. 6b) the endopodite of Pi 1 is beginning 
to foim its characteristic shape; the endopodite of 
PI I has divided into two segments and (he sternal 
process is recognisable as a small ridpe. At 10.1 mm 
IF-ig. 6i) the endopodite of PI I is almost fjlly 
developed with coupling spines; the endopodite of 
PI 2 is 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 nun 
had a fully developed petasma and sternal process 
but rhe endopodites of PI 2 were still not linked by 
coupling spines. Other <peciinens at about 10 mm 
exhibited fully developed male sexual 

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. 8e). 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 t e). The earliest stage 
observed was at a length of 8.4 mm (Figs 8a, b). 
At this stage ihe 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 
flagellum is still unmodified. At 10.1 mm (Fig. 8d) 
the sensory organ is larger and more pedunculate 
and ihe basal segment of the inner flagellum is 
beginning to differentiate forming a small lobe on 
the dorsal surface. A specimen at 11.3 mm (Hg. 8e) 
had an almost fully developed sensory organ and 
the basal segment of (he inner flagellum had 
become broader and saucer-shaped with a fringe 
of long setae on the dorsal margin, partly shielding 
the sensory organ. 

Sayce (1908) in his description of K. cursor 
described and illustrated the sensory organ arising 
from the basal segment of the outer flagellum 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. iUrsor from Bayswaicr, Victoria (SAM C401n) 
demonstrate that these male sexual characters 
{Fig. 80 are similar ro those of K. crenarum sp. nov. 
and that Sayce (1908) was in error in his determina- 
tion of these features. 

In view of the above it is possible that specimens 
tes.v than S mm in length may be difficult to sex but 
as only 10 females were ibis small it would make 
link difference to the proportion of males/females 

Tamic- 2. Lh'sOnyuKshinp, characters of Koonunga cursor and K crenanim sp. nov 

Clin racier 

A. cursor 

K. ctenarum 

Specimen length 


1 \r; 

I' (<f| 

UiiCUl til 

rarefy cxeeed* 10 nun 
upprox. Us body length, when 
uiuvhett barely reaches pereonitc 3. 
evenly rounded anteriorly, antero 
lateral incision above antenna very 
^mall (Fig. Si) 
•^rnali but present 

sternal process with larac posterior 
projection (Pig?) t> '. ml Note attfd 
distinctive strm-iurr of endopo- 
|rc* of PI I & Z (RES 6j nil 
V 1-7 3 

exceeds 20 mm 

iippiux M body length * *»hen 
attached reaches past pereonitc 7. 
pointed anteriorly, antero- 
lateral incision above antenna 
distinct (Fig. 2) 

\tcrnal process without posterior 
projection (Figs 6d, ej Note also 
distinctive structure of endopod- 
iles of PI 1 & 2 (Figs 6fe-el 
P 1-3. - 4 V P 4-7 = 4-5 



Fig. 8. Sensory organ on antennule (LHS) Of—Koonttnga 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 & U.3 mm length respectively.— A', cursor 
(SAM C4016) I', dorsal view, from specimen 6.6 mm. Scale bar = 0.2 mm. 




Koonunga crenarum most closely resembles K 
cursor the only other described congener and can 
most readily be distinguished from it by the 
characters given in Tabic 2. In the lack ol eyes it 
also resembles Micraspidcs co/nu/ni NicholLs, 1931, 
the only other member of the family Koonungidae. 

A. crenarum amongst the Koonungidae 
(including known undescribed species) occupies the 
most subterranean habitat 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 roof collapsed 
e.g. Allendale and The Shaft. All previous records 
ol Australian synearids are Irom relatively shallow 
depths and even the Tasrnanian synearids which 
inhabit lakes lend to occur near the edge:- .md t\w 
deepest record is 8 m, recorded for Paranasphies 
tacimns Smith, 1909 (Williams, W. D. 1974). 

As sinkholes and caves are difficult and 
dangerous environments to explore it is hard to 
establish (he relative abundance of the species wnh 
depth although it would seem to be more abundant 
in the surface waters, especially near vegetation at 
I he edges ol the sinkhole or cave Specimens from 
the surface and deep water have been compared in 
detail and appear lo be conspecific. pigmentation 
is similar but deep waler specimens are slightly 
darker. A', crenarum does not appear to be a strong 
swimrnei so it is unlikely that regular migration* 
occur between the surface and the bottom of rhc 
sinkholes. Ii is unlikely thai the deep waler 
specimens represent individuals that have become 
"Iom" as many more specimens have been sighted 
in deep water than have been collected (R Home, 
per . comm.). K. crenarum therefore most probably 
occurs throughout the depth o\' sinkholes and cavtw 

The occurrence ol A. crenarum m several, some- 
times widely separated, sinkholes and caves (Fig. I) 
suggests that these environments must have been 
linked ai some lime in 'he past An Underground 
connection has been a popular theory but is not 
supported by hydrological evidence (Holmes & 
Waierhousc JVK3) and the animals arc too large for 

interstitial travel. It is most likely thai the habitats 
were once connected by flood waters although they 
arc not located near Hats which were subject to 
annual flooding before drainage (Williams, M. 
1974). It is possible thai they were isolated before 
Hutopeai' settlement Whatever the connection in 
the past sinkholes and caves are now isolated refuges 
for a once more widespread fauna, 


1 am extremely grateful to Mr P. Home who 
originally discovered the syncarid and collected 
additional material from a number Of sinkholes and 
caves. Prof, H. K. Schminke and an anonymous 
referee are thanked for critical comments. 


Ufc DhKklK, P <lsW)> New rtCOKteOf Kwrwngu cursor 
Sayee, 1908 (Svneamia, Anaspidaccat- Trans R Sac, 
S, Ausr 104(2), 21 -25. 

Pkummono, F. H £959] The syncarid crustacca. a living 
link with remote geological ages. Aust. Mus. Mag,, 13, 

Hoi mi s, I. W. &. WAifcitHOusL, J. L>. (I9H3) Hvdmlo&y. 
pp. 4949 In Tyler. M J. Twidule, C\ R., Ling, J. K. 
£ Holmes, J \V. (Lds) "Natural Hiuory ol the South 
East" (Royal Society of S. : Adelaide). 

Savc *_, (J A. (1908). On Koorwrtgu curs/tr, a remarkable 
new type of tnatacostratous crustacean. Trans. Linn 
Soc. London, II. l-fi, pis 1-2. 

St iiMir-JKi H K ii97<li Psummaspides willtamst gen. n., 
sp. n., cin Vcrlrcter einer ncuen Famiiie mcsopsammaler 
Annspidaeea (Crustacea. Svnearida). 7muL Sir. J, 

[19KT) 7.ur systemailk Uer Stvgocarididae 

(C rufilacea. Syncanda) und beschreibuuy sweier neuer 
artcn [SfygseattUu pJeoiehon K^n. n., sp. n. und 
v \ unruns giselae sp. n.). Bemifortia Jfl, 139-154. 

WitUAMS, M. <iy74| Draining Hie swamps, pp 178-326* 
fn Williim-.. M. 'M be making of the South Australian 
landscape" (Academic h're-.*.: London and New York). 

\V was, W. II (19741 hesriwarcrCruMueea. pp. 63-111. 

(n Williams, \V (hd ("Biography and ecology in 
Tasmania \ Monographtae htni 25 

Zii -iik. W. 1 19831 terrestrial and freshwater inverte- 
brates excluding insevU and arachnids, pp. 187-204 In 
lyler.M. J-.TwidafcC R.. Luig. I K A: Holmes. J W 
(Ed*| "Natural Histoiy ol lite South Mtt M (Royal 
Society of S. Aum.. Adelaide), 




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 



Sunn mar} 

Lodukook, N. H. 0985) Trigoninididae iMollusca; llivatvia) horn the Cretaceous of Lake Eyre North, 
South Australia Trans. R. Soc. & Ausi. 1(^(3), 77-82, 29 November, 1985. 

Pteil^u ev/emw gen. et sp. nov., a species of bivalve mullusc apparently belonging to the Trigonicrididue, 
an important family oJ non-manne Mollusoa in the Cretaceous of Lastcm Central and Somh-Lasi Asia, 
is described. The bivalves weather out from unconsolidated fine sand and silt mapped as Winlon ronuahon 
on the western side of Babba^e Peninsula, take Lyre North, in the southern part of the Great Artesian 
li.ism, No likely non-marine ancestral form* have been found among the Unionidac and ii is possible that 
ancestors nf Pk-ttyja ry/vasts mav be found in the Trigoniidae. 

kn Wouus: Mollusca, Uivalvia, t'rigoiMoi.lidac, Cietaeeous, Wmton Formation, Lake Lyre North, 
Great Artesian Basin. 


In 1982 a numbet of grey-coloured. thieV-shelled 
bivalves collected in July from an outcrop on the 
western side of Babbage Peninsula, Lake Eya* 
North, were shown to me by Mr Neville Pledge of 
the South Australian Museum. Similar material hud 
been collected in 1967 by Dr B, Daily of the 
University o\' Adelaide. Thought to be of possible 
Miocene age, those collected earlier were taken to 
the National Museum of Victoria lor identification. 
As their preservation and colour were similar to 
Cretaceous marine fossils from the Lake Eyre region 
and not to Tertiary molluscs of the area, I 
considered those shown to me to be non-marine 
Cretaceous bivalves which I had not previously seen 
m the Cretaceous of the Great Artesian Basin. The 
male-rial taken to Victor^ was then returned to 
augment thai in the South Australian Museum. 

Since the original material was collected, the 
LAKE EYRP I 250 000 map sheet has been 
published (Williams 1975), Outcrops of Winton 
Formation (Cenomaman) arc shown west of the 
fault on the western side ol Babbage Peninsula 
(fig. I). 

I he mollusc were found weathering out from 
sandy silt to unconsolidated fine sand mapped as 
Wmton Formation, towards the ba>,e ol a low bluff 
4 m high There is a possibility, not yet 
substanlialcd, that the sandy silt lias been reworked 
from the Winton Formation, Hie locality wa.s 
revisited in May 1983 bv O. W. Kiieg and P "■ 
Rogers of the Geological Survev ol South Australia 
who made further collections from the site and 
provided the composite section shown in Fig. 2. The 
Winton sediments are overlain by a layer of massive 
crystalline gypsum up to 1.5 ui thick in which 

*c/o Department o1 Mines and Energy, P£A Box 151, 
hastwond, S. Aust. 5063 

occasional bivalves were found reworked. This is 
followed by pale olive-grey gypsifernus sand and a 
compact gypsum crust. 

In all, 19 more-or less complete valves and ?.74 
fragments were collected. Some of the specimens 
were sent to the Geological Survey of Queensland 
and the Queensland Museum to see whether similar. 
molluscs had ever been found in ihe Winton 
Foi matron in Queensland, bul none were known 
to have been collected there. 

f hey seem to helong to the family 
Trigonioididae, an important family of non-marine 
molluscs iu the Cretaceous fauna of Eastern, 
Central and South -Bast Asia (Yang 1974), 
previously not known to occur in the Southern 

The family Irigonioididac and ils distribution 

The family name 'trigonioididae was introduced 
by Cos (1952) to accommodate ihe genera 
Trigonioides KobayasJii & Suzuki. IS36 and 
Hofjetrixontu Suzuki. 1940 in the Unionacea nil her 
than in the Trigomucea, Cos excluded from 
consideration or did not accept those genera IS 
belonging to the Trlgoundae where lltey had 
OJ iinally been placed. Later (lscSS'l Con withdrew 
the family TVigonioidid.-ie, considering Trigoniaides 
to be a normal representative of the Uniomdac. 
Kobayasfu (I956) restoied the Trigonioididac as a 
valid family which has been generally accepted since 
then and classified within the Unionacea. Kiehty- 
foui species have been describe*!, distributed anions 
ten genera and seven subgenera. A second family 
Feregrinoconchiidae with seven species was 
described by Qtl, Chen &. Ian (ui Lan I97M. while 
Martinson U994) erected the supci family 
Trigonicidea to include three lam- i 





Brown sand nifty and silt 

f.VpSMT- ClUnl 

'■ tffijfl 

ChannH MPt?fl '*"" | Wys _ 

■■ laywatj uli ric 



tos*n Meaiify 


Fig. I. Locality plan with geology of Babbage PcniiiNula (lake sediments blank) 

Trip,onioididae. Pseudohyriidae and Sainshandiidac 
(b Sainsehandiidae). Martinson's classification is 
at present available in abstract only. 

The present known distribution of iiic 
Trigonioididac in Asian non-marine basins is 
(Kobayashi 1958): Northern Asia— Lena Basin and 
eastern Siberia 

Eastern Asia— Amur River— Manchuria 
Japan and Korea 
Mongolia — northwest China. 

One species has been described from Colorado 
USA (MacNeil in Morris 1936). 

The Lake Kyra bivalves 

The bivalves from Lake Eyre resemble 
Trigonioididac from eastern Asia chiefly in the 
structure of the hinge; the sculpture differs in that 
I he radial or chevron palters of the coslae are al 
best vestigia! and the dominant sculpture is finely 
concentric. The interior of I he ventral margin is not 

According to Kobayashi (1958), the TYigun 
uudidae appeared 'sporadically and explosively hi 

the maritime basins of eastern Asia 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 
Piicatounio and Nipponaia (Ota 1959a v 1959ft 
1963). Kobayashi attributes this sudden appearance 
to some geological event which caused a change in 
environment, marine bivalves being locked in inland 
basms during the Sakawa cycle of orogeny. 
Nevcsskaya & Soloviev (1981) show the maximum 
development of the Trigonioididae to be from the 
Cenomanian to the Santoniau, with minor 
OC&urrence in the Campanian and Maastrichtian 
Martinson (1984) considers the Trigonioididac to 
be characteristic of the Early Cretaceous and the 
INeudohyriidae and Sainschandiidae of the Late 
Cretaceous. Reconciliation of these apparent 
discrepancies will presumably be found in the hill 
text of Martinson's paper when it becomes available 

The age of the Winton Formation at Lake Eyre 
\i considered to be Cenomanian (Ludbiook 1978. 
Moore 1982, Lorbcs 1982) or latest Albian to 
Cenomanian (Moore & Pitt 1982, Kricg 1982). 



Y^^Jv^jV^V_V_V^' Gyfuum cru«i 

Y -* •■' V--" V- • - \ p " arlv l,u " d M " v ,wl Hl medtum 5antl 'hoinughly impregnated 
"■''-'- l --|j, witfi ImrlV rrys'.alUntt tivpsimi 

- : ^ :. : . &. : :y :\ 

Y V . V- - -A 


- J, JgL irf ' \ 

,,-,..* ... '.",', , . '. \ ltl "' ,fm& ,,n 4cm ' ttvpstum cnniMBd quarti giaiwtM, lutily well wneft flnfw well 

"Y Y^i^ji* - V v """V^" V ^( to*'5fiv tfl RiflWuoi crystalling istriy unit* gypsum wild com in htm wrd *unm head 

j. y V- ^ V" V -V- ^V" 1 PMil, lotWNB (OSKtwn Ifflf SoOH parting parallel tu lidding wHli rftprrtiupctilwl 

j£ --z^--=---^_ ^^^ y ) C|r|t>UVTI as Imfe laminae porallol In parting airlatB!! Ai tiaxE «t iiypsum layer vniy Ihin 

•TT^y ^ u,V Y V ^ ( ^'^ °' < 0,| H n . Pa'" V*" nw r 'aV which Ihickniis alimg ilnkt) and iHtctts (ilsr* tit gypsum 

* V 2 V Y ^t—.y'^J '"V 11 ' vn ' v >'; !h,1 V i.ilr..rCljl!S 


-rTT'/r. ,'*. *^ T ~"~ :?r \ *""■ * Slitjhily bfBHRhed 

cc , 

'■.- - ,-,■---- -- - \X"- Wl 

" -. - ■ .■.•..' '^V' iw Male fltcv, r.layey and vcty finely sandy Ull passing down lo lias 

.'."..' '-• \ ' \ cbyoy, v(»ry line poorly sorted Mfltl, slightly nilEaaaniS and wtlh 

j '.'■'[■■ \ JB minur woudy liagmfciits Octianioital i outtdwd miutl/ granules in 

'■■■" -'*-*-.** -.-..----, . -* . •;;*-■ >-"\ \. J 

'■■'..■ ~ ' * -~ -~~ -~ ' .... \ V^?' 

- • ■'■■'.■;'.■■'.. .X WlNfilN FUHMAflON (Mmorhedl 

■<_ _ *• •• -• • •■ •- \ * «^. 

' ■ ^^ 

) _ 

\ ' jP 

' •" * -* " - . ~ „ BiV4KHt fragment* and occaaiDnal single 

,..'•-.-• .' *.','', '. ' ,'.,'*.."..■'..".."*..*'..■' '^v ™~^^ 

"',''!''* * ',' ' ' '"",''[,"''',"' '. "'. ', .•'-.■'•.'.• . - ■ v ( ^t^ 

.,".,',' ■-."..",.'..'■- .,..,'•*• _<>/•'-,' \„ • „ /\ -■ J -rrr- 

•*-.••.-•'..••..••■- ■ - . -.-•-, '"/N. f<S?c 4 

,,.,.,- . -. - - ^s^ x -:>v "-» 

. ■ .-.-.. ..... ii ■ . ...... ^^^ ;- - T 

.... . . . . .j*'. .•'..*'.•■_"_. .'.__'*. '...,* ,'.'',*,.'..*"..'', | N ^ -•■* 

' ' '" . "' . . * . . ..".'..■"..'..'"..'"."'. '■ ,'\~~-^r _ __ 


■_.' '■;' ' : ,ALj,'/I 

Fig. 2. Composilc section of fossil lite 6340 RS 4, Lake Eyre North. 

PalynoloKical evidence tor its Cenomanian age if 
given by Burger (1982). At this time, following the 
disruption of Gondwana and extensive marine 
Hooding during the L.arly 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 remnant (Ludbrook 1978). 

Systematic description 

Htimly TRIGONIOID1DAE Cox. 1952 
Genus PLEDG1A gen. nov. (fern.) 
Shell ovate in shape, elongate-ovate in the 
juvenile, very thick and heavy, sculpture ol 
concentric growth ridges and concentric lirae. Hinge 
with anterior teeth relatively short* transversely 
crenulaled, posterior laterals narrow, long, not 

t*lcdgi# eyrensis gen- et sp. nov. 
Materia/; A total of 19 more-or-less complete 
specimens 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, tragmenis P2399Vh-k, all 
RVs average length 33, average height 23 mm; 
paratypes P24000a-h. an ontogenetic series of 8 
almost complete LVs average length 30, average 
height 23 mm; P24639a-p, fragments all 
approximately length 30, height 20 mm; P24640, 
about 80 fragments. 

Geological Survey of South Australia (GSSA): 
paratypes 10184a, 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 
(est.) 32, aveiage of 3, length 33, height 22 m; 
10186-10192 104 fragmentary RVs and 61 
fragmentary LVs. 

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, 








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; I. 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; o. 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 



inner layers art composed of caleite, determined by 
X-ray dill Tactometer at the Australian Mineral 
Development Laboratories (AM DTI. Report 
G5S4-'K5). No aragonite was detected. Interior 
smooth, hinge arcuate with a dental formula of 

RV 5 1 PI Pill 

LV 2 Pll PIV 

5 is noi always present and is crenulated or grooved 
only in adult specimens, 3 is strong and high, ol 
moderate length and in adult specimens crenulated 
on both sides with a pit lor the reception of 2 in 
the left valve. PI and Pill 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 l>iii<tnioi(les 
(Uakinoa) Ota, ]%3 as illustrated by Yang (1983). 
Muscle scars deep, anterior adductor scar close to 
the anterior margin and to the anterior end o\ the 
hinge with a small deep pedal retractor pit above 
it past undei the hinge; posterior adductor scar 
larger, not so deep, more or less roundly quadrate, 
pallia! line entire, area within the pallial line clearly 
defined, ventral margin smooth, slightly bevelled 
but not crenulated 

Distribution Known only I mm the type locality 
£340 RS 4, Babba : j;c Peninsula, Lake Fyrc North; 
Winton f-onnulion, C'enoinuman. 

Nomenclature: The generic name is to honour 
Mr Neville Pledge of the South Auslrulian Museum, 
who collected many o( the specimens and first 
brought them to my notice. 

Discussion: The family Trigoniotdidae was based 
on the species TriRiwiotcies kodttirut now reported 
(mm many localities from die USSR to Japan (Yang 
1974), The motphojogy. ontogenetic variation and 
classification of this and related genera and 
subgenera have been extensively studied by 
Martinson (l%5) and by Yang (1974, 1976, 1978, 

1979, isfirjj, 

Pledgia eyrenst.s seems to have appeared suddenly 
in the t'enotnaniati in Australia, but, unlike ihe 
Trigoniotdidae of Eastern Asia as described by 
Kobayashi (1958), it is not known to have appeared 
"sporadically and explosively/' No likely non-manne 
ancestral forms have been found among ihe Uiuon- 
idac and rhere appears to be no close relationship 
between Plwixw eyn'nsts and the lYl&stfc noiv 
marine untonids (Ludbtook I%l) of Ihe Leigh 
Creek Coal Basm. Immature forms bear oulv w •■ 
slight resemblance in sculpruic to the Ncocomiau 
Protovtrxtts coatsi Ludbrouk 1%1, of which the 
hinge is not known hi shape, sculpture and thick- 
ness of shell the adult en a I form seems to be: most 
neatly telated to the undiagnosed "Trigotuid gfijL 
et sp/*ot Skwarko M963 pi. 6, fig. 5) from ihe Parly 
Cretaceous WaNumbilla Formation south-southwest 
of Roma, Queensland This trigoniid is known only 
from the single specimen figured by Skwaito, which 
is embedded in hard matrix so thai the mn-ricH tS 
completely obscured, It does wggc& however, ilwu 
ancestors of Pledgut eyten^y are most likclv to be 
found in the Trigoniidac 


I am most giaiclul u> Mi NcmIIc |46dge ijf Hie 
South Australian Museum lor showin? me the 
bivalve* and id Mr G \V Kricy and Mr P A Hogcrs 
of the Geological Sutvey ot South Ausiialur lea 
describing the section in which ihe\ occur and 
collecting further male-rial. I thank the Chief 
Government Cjcologist and Mr P i. G. Fleming oi 
the Geotagieal Survey of Queensland for arranging 
lor »|»e examination ol the specimen-, m Queensland 
and for the loan tfl specimen GSQ I 2559. The 
figures were drafted in tin l")ra (liny. Branch of the 
South Australian Department ol Muies and buet.e>v. 
and photographic priniiur, .tsMSiame w.i-- gfVCTI by 
the Biostrati^raphv .Section 6| ihe Occlo^-al 
Survey of Soulh Australia. 

Ref< *rem*es 

Bi-KolK. O 11382] Patynology t»t ihe Croraflnjjo IV, mm 
and its apptic;iiii>n\ In Moore, R S. A Mount, i I 
(compiler-.) "Erornangu Basin Symposium, summary 
papers" QtoJ <Soc. Aust. & Het Lxplor. Sac. Au>' , 
Adelaide. 174 \H2 

Cox, I . R, (1952) Note* on die Tl'igoriiJLlac, wild uvulmcs 
ol Hie tamilv, Proc, mutuc Sac, txmd- 25>(1&3), 45-70, 
pis M 

0'»"S) On rt« i»ITit)iiivs of the C rtnucrotr Umdli 
hr;tnvd ueneni TrigoniouJt"* and Hpffffrrfniwfa, iiotfi 

T-oroub. B. G. \\Q$2) Margin oi ihe Eramancn Ba$1n, 
South AUslralia: H fCVlMC in Mnorc, P. S. <V Mniint. 
T. .1. (compilers) "Eromanga Basin Symposi •. 
summary papers". Geol. Soc. Aust &. Met. bxnlor N.k 
Ansi , AuVUtule. (I0-12S. 

Ki .t:Av.-.-.H t , r 11956} On irV I n • /,.,.,.. 

-*i*ct i'* nit^hon 10 pd^Cyt^od - |i ■ > •''.'■ I 

(jy^H) I tv \ xov- ii'Hi-iiuiluc pclcwr'MKl-. 

hiiin ihe Natr. r>tnin^ \>,ur\ >.iio m i ,- i ..,, i.< 'aSdrW i>an 
• ihe Ivti'-rai PUtcnU, itijila'MJ, wuii u Dat£ OH |Jv 
rrigonioididac. Contnls.uion- w I hi .v^i'tw • i1 ■: 
piilUtoniotL-HS l*.I SoljHi^.4,1 ,s,,,j XJ.VIJ :.■• 
raiacom, SontJ^usi l,sw IV, 109-HK, f&j N\ \Mli 

& S"/'M. K Iiy36t Noi'-nijMiic Klvetlf; i>| tht 
NnMoni.'-\vitiMh.< Scries Aljcian i t&oK Otasr, \5, 
Kkiu., g W. t;iy;s:> Sir. } u^r.\\>u\ mil ivcauiun a \h< 

IXttt.o.jMC ,1'Htelfc, srm »pWt>| ' r'iHi,|iii;;i ''.;| fn 

Mooic P. S. at Mourn. I -l t (jcoitipikrs) "I -omu -■ q , 
il l io S) ii-o'-hiiii, ^noiiiM ItOol.Boc Au-.[ 

A i',i - -i i So Aksi \a:i,. tl K. 145 158. 



LAN Xiu (1976) Mesozoie lamellibranch fossils from 

Yunnan, China. Mem. Nanjing Ins/, Geo!. Palaeont. 

Acad. Sinica No. 7 (In Chinese). 
I udbkook, N. H. (1961) Mesozoic non-marine Mollusea 

(Pelecypoda) from the north of South Australia. Trans. 

R. Soc. S. Aust. 84, 139-147. 

(1978) Australia. In Moullade, M. & Nairn, A. E. 

(Eds). The Phanerozoic geology or the World' 1! The 

Mesozoic A., 209-249, 
Martinson, C. G. (1965) Cretaceous lamellibranchs of 

the family Trigonioididac and their classification, 

Paleont. Zhum. 4, 16-25 (In Russian). 

— (1984) The I ate Cretaceous molluscs of Mongolia. 

Matac, Rev. 17, 133 (abstract). 
MooRh, P. S, (1982) Mesozoic geology of the Simpson 

Desert Region, northern South Australia. In Moore, 

P. S. & Mount, T. J. (compilers) "Eromanga Basin 

Symposium, summary papers". Geol. Soc. Aust. & Pet. 

Esplor. Soc. Aust. Adelaide, 46-57. 
& Pitt, G. M. (1982) Cretaceous of the 

southwestern Eromanga Basin: stratigraphy, facies 

variation and petroleum potential, 127-144. In Moore, 

P. S. & Mount, T. J. (compilers) "Eromanga Basin 

Symposium, summary papers". (Geol. Soc. Aust. & Pet. 

Explor. Soc. Aust.: Adelaide) 
Morris, P. K. (1936) Central Asia in Cretaceous time. 

Hull geol. Soc. Amcr. 47, 1477-1534. 
Morion, J. E. (1967) 'Molluscs*. 4th (revised) edition. 

(Hutchinson & Co.: London). 
NEVESSKASA, Li A. & Soloviev, A. N. (Eds) (1981) 

Developments and change in molluscs at the Mesozoic- 

Cainozoie boundary. Academy of Science USSR. 

Paleontological Institute. Science (In Russian). 

Ota, Y. (1959a) Plicatounio of the Wakino Formation. 

Trans. Prot\ Put. Soc. Japan, n.s. 33, 15-18, pi. 3. 
(1959b) Trigonioides and its classification. Trans. 

Proc. Pal. Soc. Japan, n.s. 34. 97-104. 
(1963) Notes on the relationship of Trigonioides 

and Plicatounio, non-marine Mesozoic Bivalvia from 

Eastern Asia. Geol. Rep. Hiroshima Univ. 12, 503-512. 
Skwarko, S. K. (1963) Australian Mesozoic trigoniids. 

Bull. Bur. Miner. Resour. Geol. Geophys. Aust. 67. 
SUZUKI, K. (1940) Non-marine moliuscan faunule of the 

Siragi Series in South fyosen. Jap. J. Geol. Geogr. 17, 

215-231, pis 22-24. 
Williams, A. F. (Compiler) (1975) LAKE EYRE map 

sheel, Geological Atlas of South Australia 1:250 000 

series. (Geol. Surv. S. Aust.: Adelaide). 
Yang, S. Y. (1974) Note on the genus Trigonioides 

(Bivalvia). Trans. Proc. Palaeont. Soc. Japan N.S. 95, 

395-408, pis. 54, 55. 
(1976) On the non-marine moliuscan fauna from 

the upper Mesozoic Myogog Formation in Korea. Thais. 

Proc. Palaeont. Soc. Japan N.S. 102, 317-333. 

(1978) Ontogenetic variation of Trigonioides (s,s.) 

puucisulcat us (Cretaceous non-marine Bivalvia). Trans. 
Proc. Palaeont. Soc. Japan N.S. Ill, 333-347. 

(1979) Some new bivalve species from the Lower 

Gveongsang Group, Korea. Trans. Proc. Palaeont. Soc. 
Japan N.S. 116, 223-234. 

(1983) On the subgenus Wakinoa (Cretaceous non- 
marine Bivalvia) from the Gveongsang Group, Korea. 
Irons. Proc. Palaeont. Soc. Japan N.S. 131. 177-190, 
pis 38-40. 



by J. E. Jackson 


Final instar larvae and pupae of two species of Leptoceridae, Lectrides varians Mosely and 
Leptorussa darlingtoni (Banks) are described and figured. 


by J. E. Jackson* 


Jai ksu\, I E, (1985) Larvae and pupae of Let,tmie\ niacins Mosely and Leptoru&sa durlingioai (Hanks), 
(Triehoptera: Leptoceridae). IfttflS A*. -Soc £ .1w.s7. 109(3), ^-3-95. 29 November. I9K5. 

Final mstar larvae and pupae of two specie- of Leptoceridae, Led rides vuritms Mosely and Leptttni^ti 
diVimwoni (BanksK are described and figured. 

ki v Womos: TYichoplera, Leptoceridae, iarvae. pupae. 


The taxonomy of the immature* of Australtan 
Triehoptcra is in an early stage of development 
compared with that of the northern hemisphere 
fauna, for which keys of larvae lo genera and 
species have been developed (Hiekin 1967, Lepneva 
1964, Wiggins 1977). identification of Australian 
larvae to family is possible using the key ol 
Williams (1980) as revised by Drecktrah (1984)^ and 
a key to some free-living and net-spinning genera 
has been developed by Cartwright & Dean (1982). 
Few detailed desct ipiions of Australian larvae are 
available (examples are Riek (1968), Nehoiss 
(1977b, 1979), anil Drecktrah (1984)), and for many 
species the larvae are not known. No im matures 
of Australian Leptoceridae are adequately 
described, despite their dominance in the Australian 

faun* (Riek i97o>. 

The leptocerid genus Leptornssa Mosely is 
monotypic: L. russaki Mosely is considered by 
Neboiss (1977a) to be conspeeific with L. 
daritngfum. The species is recorded from 
Queensland. New South Wales, Victoria, South 
Australia and lasmama (Neboiss 1983); adulis of 
both sexes are described by Mosely &, Kirmmn* 
(1953) and Neboiss 1 1977a), Lecirtdes Mosely 
includes L variant and the recently discovered L, 
parilis Neboiss, the larva of which is unknown; L. 
vunun\ is recorded from South-east Queensland, 
Victoria and Tasmania (Neboiss 1983). Vaxonornic 
descriptions of the previously undescribed I trial 
mstar larvae and pupae of Lecrrides variant Mosely 
and Leptorussa dar/tngtunt (Banks) were 
undertaken prior to a comparative study oi the rwo 

* DepHiiment of /oolouv, UmverMh ot Adelaide, G-PQ 

Boa 498, Adelaide, S. Aust. 5001 
Present tiddresy 25 Austral Terrace Malvern, S- Ag^t., 

' Jackson, J, E (1984). Taxonomy, biology and case 
function of f-wtrif/i's vitruins Mosely and tenm mssu 
darltngtoni (Hanks) larvae ( Iriehoptera: Leptoceridae) 
Hoiionrs thesis, Department ol Zootomy, University of 
Adelaide. Unpuhl. 


Identification ol Lecmdes various and 
Lepturussa darlingtoni was confirmed by rearing 
larvae and pupae to adults. 

Descriptions of final instar iaivae are based on 
preserved specimens collected from a site near 
Heathfield. S. Ausl.. (139°39'E, 35°01'S); pupal 
descriptions are based on pupae reared ftom larvae 
in ihe laboratory, a few collected specimens, and 
pupal exuviae. Whole specimens were prepared for 
microscopic examination by mascerahori of soft 
parts in hot KOH (5 u /o) for 10-15 minutes* rinsing 
in glacial acetic acid, and clearing m clove oil. 
Specimens were Ihen dissected and prepared as 
temporary mounts in clove oil, or permanent 
preparations tn Depex. Drawings were made from 
slides with the aid of a drawing lube on a Wild M20 
compound microscope and a camera lucida on a 
Wild M3 stereomieroscope. Head capsule width of 
larvae was measured to the nearest 0.0 1 mm, at the 
level of the eyes, using an ocular micrometer. 
Morphological terminology ol larvae follows that 
of Wiggins (1977). 

Specimens are deposited in the Museum of 
Victoria, Melbourne. 

Lecirtdes xanans Muselv 

Material examined' 25 larvae, from near 
Heathfield, S. Aust., April 1984, 9 pupae from 
Heathfield, April 1984. and laboratory reared 

Final inttar larva 

Case (Figs I, 2). Constructed from roughly 
quadrate leaf fragments, overlapping in a dorsal 
and a ventral row; anteriorly, dorsal fragment 
projects further forward than ventral fragment, 
forming a hood; filaments ol plant material lill- 
between dorsal and ventral rows, and around ihc 
anterior opening. 

Larva (Figs 3-19). Body length 11-13 mm; head 
capsule width 0.78-0.81 mm, length I — 1*1 3 mm. 
Sclerotized parts golden brown, with distinct paler 




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 IX and anal legs, dorsal view; 6. thoracic pleurites. 
All scale lines in mm. 






Figs 7-13. Lectricies varians final instar larva: 7-9. head, dorsal, ventral and lateral views; 10-12, pro-, meso- and 
metanota, dorsal view; 13, thoracic sternites. AH scale lines in mm. 


U I\IKS< ■■- 

markings on head and thorax t darfc bands on legs. 
Head (Figs 7-9) with pale spots on parietals, margins and posterior half of genac; ovate 
in dorsal view, width about 2/3 length. AiHenuat 
about length of right mandible, short seta apicallv 
Frontoelypeus strongly constricted medially, 
expanded posteriorly to slightly greater width than 
at anterior margin; 2 shallow concavities pn 
posterolateral margins; the resulting irregularities 
emphasized by adjacent pale spots o\\ parietals. 
1 ves situated dorsolateral^, jusi anterior to 
frontodypeal constriction, surrounded by pale 

Labrum (Fig. !5)subovatc. anterior margin with 
shallow indentalion, dense tunge of short hair in 
indentation, ventral brushes of long, hair anlero 
larerally; median paired anicnoi setae short, pale 
in front of three pairs of setae in a transverse row 
Mandibles (Fig. 161 with rounded teeth distally on 
anting edges, m\ on left, five on right; (eft mandible 
slightly longer than right, inner surface deeply 
excavated, with two brushes; right mandible only 
shehtlv excavated, without brushes. MaAillo- 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 tobaie, with 
three Finger -like sensillae apically; submental 
scieritcs absent, two long submental setae present; 
labial palps conspicuous cylindrical, Ventral 
aporome (Fig 8) oblong, about L2 length ol 
parietals. width about 1/3 length, broad^si 
anteriorly, tapered to almost 1/2 widih posteriorly. 

Pronotum (Fig, 10) Anterolateral angles 
projected forward, acute, shghlly upturned, anterior 
margin concave, shallowly crenate with six pairs of 
marginal setae aiising in constrictions; tiansveise 
row of setae jusi anterior to middle of rtoturn; 
poMcnor nutiynt heavily sclerosed. Mesonotum 
(Fig. II) rounded posterolateral^. scMt sparse, 
three setae grouped in each anlerulaleial angle. 
Metanotum (Fig, 12) only paitially sclerosed; 2 
suhquadrate selerites anleiumedi'aUy 3nd smaller 
paired sclerilcs larerally, a single seta in each medial 
selente three in anterolateral sclernes, one long and 
two very shon setae posterior to each medial 
selente Prosternnm (Fig, 13) with dark brown 
sclcrotucd baron posterior margin, smaller bar on 
each side; pair pi mesostcrnal sclenles, utela* 
sternum with 3—10 setae in two curved transverse 
rows, most arising from small round selentes. 

Legs (Figs 17-19). Forelegs (Fig 17) short. 
segments decreasing in sue distally; COSH and lcimu 
subirianguJar, coxa with band of setae otlsei lioni 
posJerolateraJ margin. Forc-irochaniin horn 
sliaped. upturned apicallv. one seta on ventral 

margin. Trochanter with short spur on distal angle, 
disTalh a venrral brush of short pale setae. Femut 
length about 2 - width; pnM of short pale setae 
verurallv. Tibia cylindrical; anteriorly tfhee dark 
spurs, ventrally four; row of short pale setae 
veutrallv. Tarsus with three spurs venually. tarsal 
claw stout, cut ved, slightly shorter than tarsus, basal 
spin almost length of claw. MidlegtHg. 18) about 
I '■■ length ot foreleg, segments subcqual. 

Trochanter with ventral brush. Femur bears seven 
dark spins veui rally and comb tif minute spines 
basally on disial segment- Tibia with seven spurs 
on anterior face, five ventrally, Tarsus with four 
spurs, row of pale seiae distally; tarsal claw statu, 
slightly curved, basal spur short. Hindleg (big. 19) 
about 2x length ol midleg. Trochanter with tow 
of seven small dark spurs on distal 1/3, femur with 
seven dark spurs ventrally. two small spurs near 
division. Tibia almost 2x length of other segments* 
divided subequally, basal segment with live spurs, 
distal with eight. Tarsus with six dark spurs 
ventrally; tarsal claw robust, basal spur short. 

Abdomen (Fig- 3) dorsovent rally flattened; 
lateral Fringe dark, on segment VIII reduced to only 
a few hairs; segment VIII with lateral row o( 
spicules on anterior half, hairs on segments ll-IX, 
II, four dorsolateral; lll-VIL two ventrolateral; 
VIII, su dorsal two ventrolaieral, l\ t iwo dorso- 
lateral, Lateral humps (Fig. 4) each with c'avaie 
sciemc and three selae; dorsal hump conical, two 
pairs of Fine setae. Tensile IX (Fig. 5) rectangular; 
three pairs of marginal scrae posteriorly, each outel 
one shorter than others, iwo pairs of very short 
setae oMset from margin. Lateral ideates of anal 
legs uniformly brown Gills single, finger like, in 
lollowing positions: segmcnl I, posterior dorsal, 
segments H-V|, anterior dorsal, lateral and vemrnl; 
segment Vll, anterior dorsal and ventral (present 

PRpfr (tfgs 20-34). Lehwlh II.5-I2.5 unit. U$S 
without apical hooks ol claw, loir- and midlegs 
with dense hair h in.ecs. Antcnnal bases each with 
seven erect black setae. Labrum (Fig. 23 1 with laleial 
margins sttaighl, converging anteriorly, anterior 
margin founded: five pairs ot setae anierolaiemlly 
Mandibles (Lies 28-31) broad-based, slender 
apicallv; inner margin roughly senate, with or 
Without a well developed projection; paire-o ^ ( '' BKi 
on outer surlaee neai base lermin ilia ol abdomen 
(Figs 32-34) wilh Iwo small dorsal processes, each 
bearing four strung anterolateral^ directed setae; 
male (Fig. 33) with pair of fleshy clasper-shaped 
ventral plot uhcranecs with buibous process berweeu 
bases; female (Fig. 34) with ventral hump. Ana! 
processes rod-like, longer than 9th abdominal 
segment , apices curved dorsad and pointed. 2-3 
short setae arising Irom inner maigui subapicallv. 



Figs 14-19. Lect rides vartans final inslar larva; 14, mavjllo-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. 





a 4lh about 2/3 from basal end. Posterior angles 
of abdominal seyment I bear transverse oblong 
plates with thickened ridges (Fig. 24); segments 
HI— VI bear elongate anterior bookplates <Tig> 25, 
26) with 2-3 prominent posteriorly-directed hooks. 
segment V with rectfirattfar posterior bookplates 
(Fi>j 21} bearing three anteriorly-directed books. 
Seements 1 1— V 1 1 With paired T-shapcd selerites 
dorsally and vcnlralty, proximal bar o\' t reduced 
in ventral sclerites. 

Pupaf Case (Figs 20-22), Larval case modified 
bs ejoNute anteriorly oblique silk membrane, 
with which attached lo substrate; transverse vent 
in anterior dorsal p!atc T through which larval 
selerites ejected; posterior closure perpendicular leal 
na^ucnt, onentation of pupa reverse of larval 
oriental inn. 

Lepiorussa dortitiRtom (Banks) 
Muurml examined: 20 larvae, from near Hcaihtii Id 
S. Ausi., April, May l°84, six pupae laboratory 

/ tftul itistur furva 

Qjse<Fi&. 35). Length 8-12 mm, constructed ot 
sandgralns. cylindrical, tapering posteriorly, slightly 
curved; anterior margin usually straight or with 
small dorsal process; poMerioi membrane absent. 

Larva iRfeK 36— 55 >. Body length 6-8 mm; head 
capsule width 0.54 mm. length 0.73-0,77 mm. Herd 
arid thorax dark brown; distinct pale markings* on 
head, usually pale cross on fromoclypeus, thorax 
11101 tied. Legs golden brown with indistinct darkeT 

Head (Figs 43-45) ovate in dorsal view, width 
a hatl I Z/J length Antennae about length ot right 
mandible, bhW seta upically, Lrontodypeus sub- 
pauduiatc. lyes situated doi>olaiciaily, just ante-not 
lo Iron I o clypeal constriction, surrounded b> pale 

I .abr utn (Fig. S2) sunuvaic, anterior margin with 
h-ui setae in shallow indentation, one long seta 
ariMiig Irorn each anterolateral margin, medially 
three pairs ol setae in transverse- mw. Mandibles 
(\ ig.s 54. 5?) with prominent teeth distalty. st\ on 
tell. ITVb on riyrif ; left slighllv longer than right, 
inner surface deeply excavated and bearing two 
small brushes: ngli: onlv slightly excavated, without 
brushes Ma vi Mo- labium {Fig. 53): eardo blunt 
r>iv-.\!iri, ( lly. .ittenuyh'd lafern!ly, bears I media! and 
one distal seta, stipes with one lateral and one 

shorter medial seta on distal margin; maxillary 
palps three segmented, galea lobale, a single sensilla 
apically; submental selerites quadrate, two long 
submental setae; palpigers Z -shaped, vm narrow. 
Ventral apotome (Fig. 44) rectangular, width about 
2/3 length, slightly narrower anteriorly. 

Pronoium (Fig. 46) rectangular, anterior margin 
pale, with 6-7 pans of regularly spaced setae; lateral 
margins (Fig. 40) with row of about seven setae; 
posterior margin heavily sclerosed. Mcsonoturn 
(Fig 46) rounded posterolateral^, tour pairs ot 
scattered setae dorsally, about seven setae grouped 
in each anterolateral angle. Metanotuin (I ig. 46) 
mostly niernbiauous; medially two small irregular 
selerites, bearing one short sera on antcroproximal 
angle, laterally three long setae. Prostemal scienter 
absent, two small dark mesosternites {Pigs 47, 4N), 
motasternum with 4-5 pairs pf setae. 

l^g.s (Figs 49-51). Anterior face ol coxae much 
mote densely setose than posterior lace, other 
segments with tew >c(ae. Foreleg (Fig. 49) short and 
robust. Fore-iroehantin (Fig. 40) truncated distally, 
anterodorsal angle produced forward sliglulv, two 
setae on anterior margin Itochanter with two pale 
spurs near distal angle, brush of short pate setae 
distally. Femur width about 23 length, comb ol" 
small pale spurs ventrally. Tibia dilated disially, four 
spurs on ventral margin. Tarsal diameter about 1/2 
thai of tibia at joint, six spurs ventrally; tarsal claw 
as long as tarsus, basal spur about 1/2 length. 
Midleg (Fig. 50) almost 2x length of foreleg, less 
robust, segments (except for short trochanter) 
decreasing in SfZfc disially. Distal -segment of 
trochanter with ventral brush ot pale seiae. lemur 
bears comb of small pale spurs near base of distal 
segment. Tibia curved nasally, spur arising 2/3 From 
basal end. Tarsus with ventral row oi small pale 
spurs on disrai 2 3 Hind leg (Fig. 51) slcudet, about 
2x length of midleg, segments subcqual 
Trochanter with comb of small pate spuiN on diital 
1 1 2 of posterior face. Femur with similar comb at 
base ol disial segment. Tibia bears one sput near 
rjjsia] end. Tarsal spur arises medUlly: tarsal eUv 
lender, curvet) slightly to a tine point, length about 
J/3 lenglh of tarsus, ba.sal spur about 1/4 of length. 

Abdomen (Fig. *6) creamy white, slightly Jorsn- 
ventrally flattened; lateral fringe short. Fine; 
segment VHI with complete lateral row of spicules. 
lateral fringe absent; segments VI— V I M with dorsal 
hairs VI-2. VIM, Vlli-6. Lateral humps of 
segment I (Tig. _V7) each with a 1! tfltfctfd bar. 

Hys 20-14. tMfmtri varttm\ pupa 20-22. case, anterior end in ventral and dorsal views, poMenor doMiit. & tabrum 
Dflid aiiKvlviu .is. 4or$al; 24, tfcStU sclCfWWetf plaw 0f abdominal segment I; 25-27. right abdnmmal hBpkpla(v*< 

anterior bookplate dorsal arid lateral view, posterior liookplcac dorsal view; 28-51. mandibles, U?i\ wiihuui mnrt 
projection {IX 29) and wiih inner proiecliou tVt-M>. 32-J4, terminalia of abdomen, dorsal, male wnirul. and 
female lateral views. All .veaJe lines ir. mm. 




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 IX 
and anal leg.v, dorsal view; 40-42, pro-, raeso- and mctapleurites. All scale lines in mm. 




Figs 43-48. Uptorussa dar/ingfant final insiar larva: 43-45, head, dermal, ventral and Lateral views; 4<>, thoracic nota, 
dorsal view; 47-48, meso- and metasterna, enlarged mesosternite. All scale lines in mm. 



Figs 49-55. Leptorussa dar/ingtoni final instar larva: 49-51, fore-, mid- and hindlegs, left; 52, labmm, dorsal view; 
53, maxillo-labium, showing association wilh head capsule, ventral view; 53, labrum, dorsal view; 54-55, mandible, 
interior face. All scale lines in mm. 


■ i{ 

Taw l I. Arnwgetmmt of the gifts of L, dartingioni. 












p 3fa 


p Jbi 

a 3t?r 

a *br 

a 3hr 

a 3br 

a Ibr 

a 3bi 

a -Vni 


a 3br 

;i ?- il 


a |/3bf 

a Ibr 


a l/2br T present/absent 

a !.2hr 


1 Ibr, present /absent 

p-posrenor, a=anteno), br=branehed 

spinulose disially; a single medtat seta; dorsal hump 
(Fig. 38) flat, with medio! transverse scleroti/.ed 
band, roughened with minute spicules anteriorly. 
Tcrgtie IX frig. 39) width greater than 3x length, 
mostly unpigmentcd, with irregular dark spots; 
three selae laterally on posterior margin, medial 
setae much longer than others. Lateral seleriles of 
anal legs mostly unpigmentcd in anterior 1/2. lew 
irregular spots. Anal claw with one small dorsal 
accessory hook. Gills Filamentous, branched, 
arrangement as in Table I 

Pupa(Y\^s 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 limes around end ot 
abdomen, antcnnal hases without setae. Labrum 
(I ig. 61) semicircular, two short setae anteriorly, 
i'ivc long betac on each lateral margin. Mandibles 
(Tig. 61) pineet-shaped, distal 1/2 of inner margin 
serrate, two selae on outer surface near base. 
Terminalia of abdomen (1 igs 63-66) with 1 wo small 
dorsal processes, each bearing four strong antero- 
lateral^ 1 directed selae; male pupa (Fig. 63) with 
three bulbous ventral protuberances. Anal processes 
(big. 66) rod-like, longer lhan 9th abdominal 
.segment, apices curved dorsad and pointed; sparsely 

covered with shoit pale hairs, rwo long setae arising 
from inner margins stibapically, one shot! seta basal 
10 these and fourth about 1/5 from basal end. 
Abdominal segment I bears pair of transverse 
oblong plates with thickened ridges; segments 
lll-VI bear elongate anterior hookplates (lig. 5V) 
with 2-3 prominent posteriorly-directed hooks; 
segment V with rectangular posterior hookplates 
(Pig. 60) bearing 2-3 anteriorly-directed hooks. 
Segment 1 1 — V 1 J with paired T-shaped seleriles 
dorsally and ventrally, proximal bar of T reduced 
in ventral seleriles. 

Pupal Case (Figs 56-58). Larval ease modified 
by closure anteriorly with plate ol" sand grams 
bound with silk just in from rim (Fig. 57): 
posteriorly thick silk membrane, intact until larval 
ecdysis when trefoil shaped vent is formed (Fig. 58); 
pupa fn larval orientation, pharale adult escapes by 
opening anterior plate like trapdoor. Case attached 
to substrate with stalked disks tbig. 56) anteriorly, 
or anteriorly and posteriorly. 


I am grateful to my Honours Supervisor, Alice 
Wells, arid to Margaret Davies tor helpful guidance 
in the preparation of this manuscript 

References 0, I. & Di-an J. C (1982) A kev If) 
Victorian genera ot tree living and reireat-niafcinK 
cidtliblly larvae (Insecta; IriLhopltia) Mem. natnl 
Mus. \ -taoriu. 43. 1-13. 

l)Rt<*tK\n, (PiM) Description ot the miniature 
stages OX AHoei-elfa grhea Hanks fTrichoptera: Hclico- 
pbidac) and morphological charaet crimes used |d 
distinguish between larvae of Australian Calouclae. 
Conoesuudai* and Hclicophidae pp. 115-122. In .1. C 
MOfSC I Ed I 'PrfiC *tih hut-main. Synip. Trichoplcra. 
< JiMik The Mu±mjc). 

Hit kin, N. F. |1S*()7) "Laddie Larvae. Larvae at the 
British trichoptera'. (Hutchinson: London). 

I l t-M v\, S. G (1964) Fauna SSSR, RjUCheUilki, v6l. 2. 
no. I. Ljehinki i kukolki podoiryada kol'crvaiosh- 
chupikovykh. /oologiche.tkii Instiiut Akudemii Nauk 
SSSR, n.s. S8. (In Russian. Transtaied into L'ngh&h as: 
huunaof the U.H.S.R.; Inchopter^, vol. 2 V no. L Larvae 
and Pupae of Atttiuhputpio. Published by ihe Israel 
Program for Scientific Translations. W7nj., fvL £. & Kimmins, U L. (1953) "I he Irichoptera 
( Of Australia and New Zealand" 
pp 27<l-27fc_ (Biiiish Museum (Natural Hislotv); 

Nihui-v,, A, (1 977a) \ ra\ononuc and ^oogcottruphic 
study erTasmauian eaddisflies (Insccta: Thchopiera), 
Mem. tiainl. Mm, Victoria, 38, 1-208. 

(1977b) Anipk'Ltklidiie, a new eaddisfls family 

< Trichopterar AmpIccOdidac). pp. 67-73, In M. I. 
C "riehton (fid. f Proe. 2nd Internum. Synip- Tnchoplc.t ' 
(Junk: The I fugue). 

— (1979) A terrestrial caddisfly larva from lasmania 

(Caloeidae: Tlichoptera). Aust, F.nirtmof. Man. $, 90 -93. 
(1983) Checklist and bibliography of lhv Australian 

CaddivFlies (Tnehoplerai. Just. Sov. [ imnof. \pn\ 

F'uhL 5. 
Rn-k, E. F. (!96H) A new family ol caddis Hies Irom 

Australia (Trichoptct a: lasimodac). ./. Aost. t.m, Soc 

7 109-114. 




(1970) Trichoptera (Chapter 35): pp. 741-764. In Wiggins, 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. 


by A. Wells 


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). 


by A. Wells* 


WEILS, 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 Words: Taxonomy, Trichoptera, Hydroptilidae, Alligator Rivers, seasonality. 


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 
dicl 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 1 ) 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 NT. Water Division 
gauging station 821009), Georgetown Billabong and 
Corndoirl 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.56rf, Magela Creek, S. of Georgetown 
Billabong, NX, ll.vii.1983, A. J. Sharley. 
Paratypes: NTM 1.57-63 3d", 4? (including allotype 1.60) 
collected with holotype, NTM 1.64-89 16cf, 109, same 
locality, A. J. Sharley, l.vii.1983; NTM 1.90 \%, 
NTM 1.91-94 4?,, Georgetown Billabong, nr 
Jabiru, NX, A. J. Sharley; NTM 1.95 19, 30.iii.1983, NTM 
1.96 1?, 22.iii.1983, Corndorl Billabong, nr Jabiru, NX, 
A. J. Sharley. 

Other material examined; 7tf, 109,, 24, 2%, 4cf, 10?, 3.vii.l983, !<?, 2?, 5.vii.l983, 4d\ 99, 

1 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 




v Hf 




A V V:/l 



Figs 1-5. Hellyethira verulu sp. nov. I, 2.<f genitalia in dorsal and ventral views. 3. 9 ierminalia l ventral view. Tricho- 
leiochiton jabirella sp. nov. 4. 9 terminalia, ventral view. Oxyeihira warramungu sp. nov. 5. ? terminalia, ventral 
view. Abbreviations: ae., aedeagus,, dorsal plale;, inferior appendage; pr. ? paramere; VII, VIII. IX. 
abdominal segments VII, VUI T and IX. Scale bars = 0.1 mm. 



7 v.. 1983, U) &< SWI.M**, IcP, tt Mi 22, W, K, l 7 >« 1583. bfi I?, iy.vh.iMH3, Id 

2l.viilWV 4 ZVviilMSV Irf, lv\ 2S.Vtt.W83, I/, lv. 

2(».iii USA, .-, 23.lli.W5i 19, 2fi.iif.l983, 2?, 2B.iii 1983. 
Maittb Cted. S o| Ceor§ I > " | N.I , A. J. 


tkogttOStt Medium sized, with mottled lawn-cream 

Male Length ol anterior wing 1,5-1.7 mm 
Antenna .lO-segmeiwcd. Genitalia symmetrical, as 
in I i^s l. 2. (nierior appendages mulnlobed, 
ventral -most lobe elongate, slender, bifid distally; 
second lobe also slender and non^ate; upper lobe 
plntc-likc in ventral view, with apieotaicral due £S 
attenuated. Dorsal plate membranous, with p 
median set rale "tulT , \ Paraiuerc vender, curved, 
intersecting medially No subgenus! pUle evident. 
Jvmale. length of amertoi wing 1.6-2.1 rum 
Tbrrninalia (Fig, 3) short, uerutie VII .shjchtiy 
bilobed apiconicsallv. each lobe tipped ttt paired 

//. M£flfta shows close sitmlarirv to // eskensis 
(Mosely) and H, sentiw WelU (Well* J$79a> which 
have similar scissor-like paiBffltttS, TbK "" 
appendages more closely resemble thOiC pi //, 
eskenslSi aJ though the dorsal pUte of W. wuto is 
distinct from those Ol the oUiet two species. Hither- 
to, I considered H cskensis and /-/. seniiso to he 
close sister species; the precise nature '•' , ' 
ictaiiouship io H. veruta is unknown, but Ihc Ihrce 
species cleaily lorm a duiincl lineage witltin 

H ee'utoappears to begin i ■ .hsettr 

from late-dry season »■ .oiled ions, ran in iho.-.e ffOWl 
the wet season, ami uiosi abundant m ibe dry 
season samples. Most specimens were taken from 
Majicia Creek and only few from the two lajnmus. 
At the sampling siie Magela C 'reek has a white &&nd 
substiatc. and lacks the microphyte community 
iound in the lagoons; In the dry season the water 
recedes to form a small stagnant DOOl ' n r he si ream 
channel (Mahpahl per, OODAHl.), 

'tHchtiMadrhqu jxbinito sp, nov, 
FICiS 4, 6. 7 

Itdlotvftt'.' NTM L97,/. fooidoil UiII.-Uhhi*. in I;. I 

N.I., Saw 1983, a I Bbwlo 

fiarat^pCL NTM 1.98 let L99 (Allfltyj | ■ - Elected *lih 
Itoloiypc; NIM 1.100 IA U.wiJ9R3 ( NTM 1,101 Iji 
L&Bi,t9Sf, NTM I ItP-UU ;,/, Hmi,.14N1, Crnd.irl 

ibong, nr uhiru NT.; NTM l.un k', <ieoiy.emwn 

Bill.uvmy. nr AiNra, S.I , A. 1 S|Ujft«V, ? "..1983. 
Diagnosis: Medium i . •" rflOltltcl wings, lips 
ol wings uptioiie-J, R 2 and K, arise indcndemlv in 
anterior wing* 

Male UrnRth or .interim ping 2.2-2.5 nun 
Antennae- U^cgmcntwl. Qeniralia Jti irj FiftS 
Scgmem X short, sternttc with broad, deep, rnedwn 

HwnvaTian, lateral fobe#*£tertxlsad dbtally. Oorsal 

pt«^(c frilobed, membra nous, central lobe 
sul.K|uadiaie. Subv,eiuial plate membranous, broad, 
narrowly cleft apicomcsallv- Interior appendages 
stout, slightly extended apieomesalh to form 
rounded, sclerotised lobes. Aedeagus witli a btack 
spmc arising subapicaDf and projecting slightly 
beyond aprv 

female. UfttygXtl of anterior wine 2.5 mm. Antennae 
22-srjirnented. lerrninalia (Tig. 4) short, pale; 
segment VMI amplo, collat-like. 

T jabirclta is most closely allied to 7. fuiehs Wells 
< Wells 19X2) with which it shares the general form 

i \ male gentfalta and wing venation However, 
readily distinguished by the black spine on the 
aedeagus, broader Inferior apjiendages, and bilobed 
subgenital plate, Roth species were collected from 
the same localities in the emergence traps. T 
juhiwUa appears to be highly seasonal, being taken 
in the wet season (March), while 7. fidelis was 
UO)lect$d in ah seasons (Sharley Sl Mahpatil 
unpublished data). Although congeners were 
collected from lagoons only ut the Alligator Rivers 
region, T, fidelis has been taken from rivers and 
streams in NE Queensland. Bosh Georgetown and 
Ooi ndorl billabongs ha^c black clay substrates and 
luacrophyte communities and havie slow flow only 
in the wet season (Malipalil pets, comnt.). 

0%}Hhira iltefapfitrwhia) iwrnimunaa sp. nov. 
FIGS 5 f 8 t y 

HofotvfJ* NTM I UlScf, (Wvugefrwn Htliabong, nr Jabirti. 

k963, a. -i. Stearlcy 
Pamypes: NTM I.IU6*H» •■:, U. i including allotype 
MO?). 21.viit983. NTM 1410*112 3v. 27^1 1983, NTM 

■ i | tm L-r, 4v. 3*1.1983, NTM T.II7 lis M, 1?. 

! IW1 NTM 1 119 Iff, I9.v.liy83, NIM 1,120 I?. NTM 1.121 I^f. 27.Vll.W63, '^'T.emwn 
Uillabony. nr MlbWU, NT , A t Stwlcv; NIMI \21 Ifc 
Imi.1981, NIM I 123 iv. 29.vif.19S3, MaplaCrert, S. qi 
CicoT^eiown Billabong, NT., A. I Shmtey; NTM 1.124 Irf, 

■ Jorl hoIhN. u>i. nr tftbtru., NT, 233ri1 1983. 

Othrtmalerktitxmitned; i?. zvo.iyg), 2?, 5.viLl9K3, Ift, 
su'il.iwl. 2v. 23 vii.j9fl3. Georgetown Biltubmii? M Jaboru, 
N.T., A. J SSiarles NTM: IB, 26w.l$83, I?, J7-*U*Wi 
J8, > 1983, 4?, briLtSftf. 3t, 3*iiJ9S3, llf -vn 1983, 

3; 19^ M983 OT.I9W, MagdaCr^t.. 

S •' ciri'igetfAVTi B>ltabong, NT.. A. J. Sharley, NTM 
Uiusnosts; Minute caddtefltct with mottled 
vestUure: '.pur formula O.Z.4. 
Mate. Length of anterior wing 1.1-1.5 mm. 
Antennae 26-see.mented. Genitalia M m Tig^ 8, V. 
Segment VIII broad, rounded, sternttc sstJely and 
,[ . i lowly e^cued aptcomesally; tcrgite wtlh a broad, 
deep mesal concavity. Dorsal plate membranous, 
• Icct broad at base, uai rower dissalty Aedeajus 
stout, withOUl tiullator. litfcfftCM appendages discrete 
diMjIry. lused at base; pajred lobes d< terior 







i \ 

/ : 


:■ i 






a * 


Hi ' 


. ; .. .: 



j i ! ! 


■■ I ft || 

,<*") . 



> L; ! - 

8 n *. ■> f 

t ;■. " 

: ii -■ ■■ .- - 

*i : : ! 

ii hh I 

:. I 

; \ '■■ 

' ■■ 

Ii : ' 

3 ! 2 

:■' rf | I 



7 f\#lf%^— - 8ub -°- 

k-^ —, 




tins I0--I2. QrtHotrichia rurhinaia sp. nov. cf, 10. 7ih anicnnal segment, dorsal view; II, !2. genitalia, dorsal and 
ventral views. Abbreviations: ae., aedeagus;, dorsal plra.te; dorsal process oi' infciior appendages; infapp., 
interior appendages; pi , paramere; ids., right dorsal spine; sau., sensilla auricillica; sc, .-.ensilla coeloconiea; sp. t 
seiisiila plueodca; v.h., vesliturul hair; VIII, IN, abdominal segments VIM and IX. Scale bars as indicated. 

appendages may represent the subgenital plate. 
Female, length of anterior wing 1.4-2 mm. 
Antennae 20-segmented; stapes anout 2 * length of 
pedicel. Terminalia as in Fig. 5. Sternite VIII 
triangular, apex with a narrow selerotised band; 
tergite shallowly eoncave apically. 

O. warranwnga is clearly a member of the 
minima group in O. Dampfitrichia, to which two 
oilier north Australian species* O. artuvjliosus 
(Wells) and O, plttmosa (Wells), belong (Kelley 
IV84). 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 
stcrnitc VIM. The female terminalia of O. 
warramunga are distinguished only by the shallowly 
concave apex of tergite VHI, compared with the 
angular vertex of O. artuvilhsus, In features of male 
genitalia O. warramunga is probably closer ro O. 
plumosa than to O. artuvittosus. 

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. artuvitlosus were also collected from Magela 
Creek in the dry season of 1983 (Sharley & Malipatil 
unpublished data). 

Figs 6-V. Tnchalewvhtton jahMta sp nov. 6, 7. c? genitalia, dorsal and ventral views. O.wethiru wurramungu sp. 
nov. X, V. o genitalia, florae! and ventral views Abbreviations: ae., acdeaguv. dorsal plate; interior 
appendage; sub.g., subgenital plate; VJM, abdominal segment VIII. Scale bars I mm. 


A. Wlil.LS 

Orthotrichia eurhinata sp, nov. 
FIGS 10-12 
Hototvpe: NTM 1.125 d", Georgetown BilJabong, nr Jabiru,, A. J, Sharley. 

Diagnosis: Known only from a single medium-sized 
male with distinctive antennae: segments of 
proximal X A 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 
senstlla 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, 

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 inferior 
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). 


I wish to thank Dt Malipatil who kindly made 
the Alligator Rivers materia! 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. 


Ruin, R. W (1984) Phylogeny, morphology and 
classification of the micro-caddis fly genus Oxyethiro 
(Trichoptera: Hydroptilidae). Trans. Am. Enl. Sec, 110, 

Marsham, J, E. (1979) A review of the Hydroptilidae 
(Trichoptera). Butt. Br. Mus. (Nat- Hist } Entomoi. 39, 

Supervising Scientist For Alligator Rivers Region. 
(1984) "Alligator Rivers Region Research InMiuire, 
Research Report 19834984". (Aust. Govt. Publ. Sen. 
Canberra 1984). 

Weils, A. (1978)[1979] A review of the Australian species 
of Hydmptita Dalman (Trichoptera: Hydroptilidae) 
with descriptions of new species. Aust. J. Zool. 26, 

— — (1979a) A review of the Australian genera 

Xuiholrichia Mosely and Hellyctlura Neboiss (Trich- 
optera: Hydroptilidae), with descriptions of new species. 
Ausi. J. Zool. 27, 311-29. 

0979b) The Australian species of Orthotruhto 

Eaton (Trichoptera: Hvdropiilidae). Aust. ./. Zoot. 27, 

(1982) Trichohiochiton Kloct and Hineks and new 

genera in the Australian Hydroptilidae (Trichoptera). 
Aust. J. Zool. 30, 251-70, 

(1983) New species in the Australian Hydroptilidae 

(Trichoplera), with observations on relationships and 
distributions. Ausi. J. Zool. 31, 629-49. 

(I984) Comparative studies of anlennal features 

of Australian Hydroptilidae (Trichoptera). In J. Morse 
ed. 1( Proc 4th Internatn. Symp. Trichoptera" (Junk: 
The Hague). 


by Margaret Davies, M. Mahony & 7. Z). Roberts 


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 


i, y MA^OAfeBT DwIEfc*. M, MAiioNYt & J. D. Roberts: 


U^irs. M Mamomcv, M. & Roblris, J D. (1985) A new species of Uperohiu (Amite: Leptodactyliduc) 

from the Pilbara Region, Western Australia. Tram;. R. Soc. 5L Ausi. I00O1, 103408, 29 November, 1985 

Upivofoia xiandalasa fip, nov., is desurihed tYiifU the Pilhoia iciuon of Western Australia, The new 

Bptelos is nympfliriC with U. russeltl Call information and karyotype are included in the species Jcvripnnn. 

Key Words: Vperv/eia %tantlutosa sn. nov., ac'veitisemenl calk, Anura, Leptodactylidae, karyotype. 


lii revising tbfi genus Upervbda Cray, Tyler, 
P.ivies & Martin (f9Sla) restricted the distribution 
of the nominate species U. murmuratu Gray to the 
type locality "Western Australia", estimated from 
Gray's journals to be in the Prince Regent River 
area. Later they recorded the distribution of V. 
tntobaxi (Andersson) as confined to the Fit/roy 
River drainage (Tyler ei oi 1981b). Hence records 
of U. marmonnu and unsubstantiated records of 
V. mjobergi (as Glauerna ftijobergf) in the rilbara 
region ot Western Australia (Mam 1%5; Mam & 
Starr 1966) cannot be sustained 

A third species. U. russrih (1 ovcitdge) is recotded 
from the Pilhj , considered endemic to the 

region (Main 1965; Main & Storr 1966; Tyler el ai 
1981a; Tyler, Smith & Johnstone 1984). 

During field work following Tropical Cyclone 
Jane in January 1983, two of us (J.D.R, and MM.) 
collected two species of Uperoleia near Port 
Hedland. One of these species is U russelli and the 
other is conspecific with the specimen referred to 
by Mam (1965) ami Main & Ston (1966) as U 
mamtorata and which is undescribed, 

Here we describe the new species and include 
bi Id notes on its breeding biology, call and Karyo- 

Materials and Methods 

Material cited here is deposited in the Australian 
Museum. Sydney (AM), the SourJi Australian 
Museum, Adelaide (SAM I and the Wesicrn 
Australian Museum, Perth (WAMi 

Measurements of specimen* follow Tyler e7 it/, 
(1981&K Measurements taken (»n mm) were: eye 
diameter (E); eye to nans distance (E-N); internarxal 

span (IN); snout to vent length (S-V) and tibia 
length (TL). 

Osteological data were obtained from cleared and 
ah'rarin red S stained specimens prepared after the 
method of Davis & Gore (1947), Osteological 
descriptions follow Trueb (1979). 

Calls were recorded in the field on a Sony 
IC-510-2 reel to reel tape recorder, with Beyer MIOI 
01 A KG f>I 90 microphones at a tape speed of 19 cm 
s ' Sis call parameters were analysed in derail: (I) 
mean call period (seconds) estimated from the time 
m con to make 20 calls (time taken svith a stop watch, 
half-speed playback; (2) call duration; (3) pulse 
duration; (4) pulse period; (averaged over the whole 
call) (2-4, all to milliseconds and measured from 
oscillograms made on a San-ct 5M28 direct recor- 
ding oscillograph); (5) pulse number (counted 
oscillograms); and (6) dominant frequency, averaged 
over five calls by a Tektronix 5L4N spectrum 
analyser. For the tirsi five parameters, we analysed 
live calls from each recorded frog, Pulse durations 
were estimated from all pulses in the call Periods 
rather than repetition rates are reported as. these arc 
the basic anils measured; repetition rates represent 
a derived unit <Gaioni & Evans 1984). Rcpetirnm 
rates are a simple reciprocal function of periods. 
Data for each frog were averaged and the mean 
values were used to calculate an overall mean. Each 
individual thus contributes one datum point to the 
final figures. 

Mitotic chromosomes were examined from 
WAM R89489, using trie technique described by 
Mahony <& Robinson (1980). Silver staining was 
carried out following the procedure of Bloom & 
Goodpasture ^ 1 976). 

* Department of Zoolop.y, University of Adelaide. G.P.O. 
Box 498. Adelaide, S. Aust. 5001. 

i School of Biological Sciences! Macqu*tte Untv*f^ity* 

North Ryde. NSW. 2113. 
| Depart ment of Zoology, University of Western Ausiralia. 
Ncdlands, W.A. 6009. 

IJpewiei* je/anrfw/risa sp nov 

Upemleto murmwoiH Mam ls#» c R0ft£0l SdGtfl , - > - 

AnstMliH, p. 36 (pffl i Main * Slur', l%6. W AuM. 

Nat 10, p, 57; Cogger, 1983. p. M. 
Hototr„ t <; WAM KRS»4H^ an adult male collected 




Fig. 1. A lateral, B dorsal view of head of Uperoleia xtunduto:.a p. NOV. (SAM R27UHI). 

at Petermarer Creek, Port Hedland-Broome Rd, 
W.A. (21 c 236"S t LlS^S^rB), on 10.L1983 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 hototype: Maxillary teeth absent. 
Vomerine teeth absent. Snout short, truncated when 
viewed from above and in profile. Eye to naris 
distance greater than internarial span (E-N/IN 
1.36). Canthus rostralis prominent and straight 
(Fig. 1A). Loreal region sloping. Tympanum not 
visible externally (Fig. IB). Fingers moderately long, 
slightly fringed (Fig. 2 A) 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> I. Large oval inner and broad outer 
metatarsal tubercles. Subarticular tubercles conical. 

Dorsal surface with well defined parotoid, 
inguinal and coccygeal glands (Fig- 3). Dorsum 
faintly tubercular. Submandibular gland disrupted. 
Ventral surface finely granular. 

Male with unilocular, 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 Upemieia ghndulosa sp. nov, (SAM R2708I). 

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 scries 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. Vent rally whitish 
flecked with grey. I^wer surface of limbs flesh- 

This description conforms with our observations. 
Dimensions (in mm): Snout-vent length 20.2; tibia 
length 7.4; eye diameter 2.8; eye-nan's distance 1.9; 
intemarial span 1.4. 

Variation: The arc seven paratypes. si\ adult -■* * 
and one adult ? WAM R89490-2, AM R 114573, 



Fig. 3. Uperoteia glundulosu sp. nov. in life. 

collected with the holotype; SAM R27081, SAM 
R27082 (cleared and stained), 3.2 km NE 
Wittenoom turnoff on Port Hcdland-Broome Rd, 
lO.i.1983, M. Mahony, J. D. Roberts. WAM R22921, 
Mundabullangana (5 km E of Hstd), E9.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. IB). 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 

Table 1. Details of call structure of two Uperoleia glandulosa sp, nov. 

and range in brackets. 

Values are means with standard deviation 




Pulse Pulses 
Duration sec ' 







min ' 


(Air dry) 

(0.4, 2.0-2 


6.3 160 

(0.8, 5.7-6.8) 


(0.6, 13.4-14.6) 

(0.08, .69- 



(35, 2850-2900) 



water's edge ox in a similar position actually in (he 
water. Males with a similar call were also heard in 
the Petermarer Creek and around a Hooded elaypan 
about 3 km southwest of the Turner River on the 
Northwest Coastal Highway, in the Turner River but 


I ig. 4. Oscillographs (, l advertisement call 
fifatuiuhsa sp. nov. 

I Uperoteiu 

not on the Yule River. Males were calling on the 
third night after two days and two nights or heavy 
rains (114 mm at Port Hedland). 

Uperoleia russetii was calling at the same sites, 
Osteology: Skull poorly ossified, sloping 
anteroventrally. Small portion of sphencthmoid 
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 wilh cxoceipiial. 
Hxoccipital not ossified either dorso- or venlro 
medially. Crista parotica short, stocky, not overlain 
by otic ramus of squamosal. Carotid canal groove 
not developed on frontoparietals. Frontoparietal 
fontanelle extensively exposed, overlapped laterall> 
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 ramus of pterygoid. 
Posterior margin formed by articulating postero- 
medial eMremities of frontoparietals (Fig. 5A). 

Nasals moderately ossified, approximately 
triangular with maxillary processes directed more 
ventrally than laterally; widely separated medially. 
Maxillary processes not in bony contact with well 
developed prcorbital 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 (Fig, 5B). Alary 
processes expanded laterally, at right angles to 

Fig, v A dorsal, B ventral view of skull of Uperoleia xlandulosa sp nov ISAM R270X2) Scale hnr 5 mm. 



cullriform process extending almosl lt> level ol 
anterior extremities of media! rami ol pterygoids, 

Medial ramus of pterygoid moderately long, 
at 1 1 minute, not in bony contact with prootie region. 
Posterior ramus moderately long and broad. 
Anterior ramus in long contact with moderately 
developed pterygoid process ol palatal shell of 
maxillary. Cartilaginous quadrate present between 
base Of squamosal and quadratojugal. Quadrnio- 
jugal robust, in Firm contact with maxillary, 

Squamosal shaft robust; tiny /.ygomatie ramus, 
elongate uncxpanded otic ramus lying alongside 
lateral extremities of crista parotiea 

Maxillary and premaxillary edentate. Palatal shelf 
moderately deep with well developed palatine 
processes not abutting medially, and moderately 
developed pterygoid processes, Alary processes of 
premaxillaries moderately broad, bifurcate and 
perpendicular. Vomers absent. Columella bony. 

Pectoral girdle aieiferal and robust. Omosternum 
and xiphistcrnum present. Sternum cartilaginous, 
Clavicles slender, curved, poorly separated medially. 
Scapula bieapitate, about same length as clavicles. 
Sitprascapula about ! ■•? ossified. 

Tight non-imbncatc presacral vertebrae. Sacral 
diapophyscs poorly expanded Relative widihs of 
rransver>e processes I II > sacrum > IV > I J > V= 


fig. t). lateral view ot pclvie giuJle Ol t'l'trtt/riu 
ftlantlulusti 90 ROt ISAM R27082). 

Bicondylar, sacrococcygeal articulation, Urostylc 
wall crest extending : j its length. Ilia extend 
anteriorly to sacrum Ilial crest absent, large 
papillate, dorsal protuberance on small dorsal 
prominence (Fig. 6). Humerus with strongly deve- 
loped anleroproxunalcresl. Phalangeal formula of 
hand 2,2,3.3 Distal tips of Terminal phalanges 
knobbed. Small bony prepolles and palmar sesa- 
moid. Phalangeal formula o( foot 2,2,3,4.3. Well 
developed bony prehallux. 
Karyotype- The karyotype ol Uperoteia fifarululosa 
is shown in Fig. 7, the diploid number is 2n - 24. 
The chromosomes are arranged in two groups on 
the basis of si/e. Pairs 1 to 6 are large with relative 
lengths (R.L.) ranging from 1 5*Vo to 9%; pairs 7 to 
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 metacentric. 
A prominent secondary constriction which is silver 
stain positive, and thus considered lo be the 
nucleolar organiser region, occurs subtcnninally o\y 
the short arm of chromosome 4. A distinctive 
secondary constriction mid-way along the shorl arm 
of chromosome 8 was not strongly silver positive 
and is not considered to be a nucleolar OtJftnlSCl 

Etymology: The specific epithel is from the I aim 
glandule* in reference to the prominent parotoid, 
inguinal and coccygeal glands in this species. 
Comparison with other xpeaes. Uperoteia .atari- 
diilosa is distinguished from many congeners by the 
presence of a widely exposed frontoparietal 
lean .incite. This feature is shared by U. arerueola, 
(J. borealiSj U. inuntlatu, U. or ten (alts, U. russet ti 
and U. tatpa. All oi' these species, except for U 
ortentalts for which the data are not known, are 
"long call" species whereas U xtanthtlosa has a very 
short call. U. gtandulosa has a stronger back pattern 
and less extensive webbing than U ohentalis. 

From the three "short call" species in Western 
Australia, U. ti/honioda, U minima and U asfiera, L( 

IK n~u n n 

X 8 K <; X X 

X X 


Fit. 7 - Karyotype of Uperoteia planftulnsu sp. uov Arrows inotcaic rhe location ol ilic nucleolar organUer region 
of chromosome A; inset shows this rcpioii with silver Maifnny_ The local ion o\ a secondary CprntUftian on chromosome 
8 is not silver positive and is indicated hy an arrowhead 



gfandulasa ws distinguished by its greater toe 
webbing and greater frontoparietal fonranelle 


ll would appear that the new species that we 
describe here has been encountered previously at 
Mundabullaugana but mi&identificd 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 U. 
russeiii (as Glauertia rmseffi), considering that it 
might rcpicscnt Lf. mjobergt 

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 Pilbaia {U. Rlundulostt 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 striking conservatism of chromosome morpho- 
logy and, in particular, local ion of the nucleolai 
organiser region. While this conservation means 
lhal species of Uperoleia 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. 


Field work was supported by an Australian 
Biological Resources Study grant to M.M. and 
funds from the Zoology Department, Uuivcrsity 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. J. Tyler critically read the manuscript and is 
thanked for helpful discussion. 


Bijoom, S. fc. & OoooPAStxiKr, C. (1976) An improved 
technique for selective silver staining at nucleolar 
Organiser regions in human chromosomes. Hum. Genet 
34, 199-206 

Too tit r., Hi G, (1983) Reptiles and Amphibians of 
Australia. 3rd Edm. (Reed: Sydney). 

Davis, IX D. & Ooki , V. R. (1947) Clearing and staining 
skeletons nt 'small vertebrates, fidihuna Tech. 4, t-tft. 

< iAIQSr, S. I. & Evans, C. S. (1984) The use of ran- Of 
period to describe temporally patterned stimuli Arum, 
tiehtn: 32. 940-941. 

Mahony, M. J. & Robinson. b S, (19X0) Polyploidy m 
the Australian leptodactylid frog genus Neobatrachus. 
Chrom<M0M<* (Bert.) HI, 199 212. 

— &. (in piess) Nucleolar organiser region 

(NOR) location in karyotypes of Australian ground 
frofin (faroth Mrobatrachidae>. Genetic* 

Main, A. R, (1965) frog* ot Southern Western Australia. 

(Western Australian Naturalists Club; Perth), 
& StoRR, G. M. (1966) Range eviensions and 

notes on the biology of frogs from the Pifbata region, 

Western Australia. West. Aunt. Nat. 10, 53-61. 
Morfscaixhi, A. (1979) New developments in vertebrate 

cvtotaxonomy. I. Cytotaxonomy of the amphibians. 

G,w//n7 50, 179 193. 
TRtFti, L (1979) Leptodactylid frogs of the genu* 

Tefmotohius in Ecuador with the description of a new 

species. Copeia 1979(3), 714-731 
TYlbr, M. J., Davun, M. & Martin, A. A. (1981;*) 

Australian frogs of the leplodactvlid genus tfperoteto 

Gray. Auxt. J, ZooJ, Suppl. Sir. 79, 1-64. 
— , - — & — — (1981b) New and rediscovered 

specieS <>|" frogs from the Derby-Broome area, Western 

Australia. AV< . liiwt. AUSt. MUS. 9(2), 147-172. 

, Smith, I . A. &. Jomnstuni, R. E. (1984) "Frogs 

alWesr-m 'Xirsl^lj:*" f\V \ Muvmir P'-tltO 


by Ivor Lansbury 


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. 


by tVCik Lansbuky* 


Iansmrv, I. (I9K5) The Austialian Naueouaae (Insecta, Heimpieia-Heteioptera) with description of a 
new species. Trans. R. Soc. S. Aust. 109(3), 109 119, 29 November. 1985. 

The NauCorfUae ol Australia aic (cdcsctibed and keys to genera and species ate provided, A new species, 
t\twcttri\ \'uhaureu>; sp nov. is described from Western Australia. Leetotypes arc designated loi Sauvons 
itHsin/hcus Still and /V fubo/MKUS Moniundon. Distributional ami ecological noiex arc civrn for aJ] six species. 

Kl V Wokiv, N.iHtoiidae, \tuttons, Aphctutlwtrus, dislnhutiiHV 

ln (rod ir lion 

lliedesct ibed Austialian naucorid fauna of live 
specie^ (Naucoris ausfralicus Stal ( 1876), N. 
coayrcx Slal (1K76) and Aphclocheiras austraiicus 
Usinger (1936) from Queensland, N t sithopacui 
Montaudon (191.}) and N* rhizomatus Polhcmus 
(I9B4) Iron) the Northern Territory) is relatively 
depauperate compared witli that of New Guinea. 
I a Rivers (1971) lists 20 endemic species in eight 
endemic genera, ftom lhal area. 

Naucoris I abrieius is an "OKI World' genus with 
ca. nine species recorded from ihe Oi icnial Region. 
It ha> not been recorded from Now Guinea or New 

The inclusion Of Aphelocheirus Westwood in ihe 
Naucoridae is debatable. Clnua & Miller (1959) and 
Popov (1970) included ibe gL-ntis in (he Naucoridae 
subfamily Aphelocheirinae. However, Poisson 
(1957) considered Ihe genus merited family status 
and its omission by la Rivers (1971) from his world 
checklist of Naucoridae implies acceptance o\' this 
contention. A recent review by Hobetlandt &Siys 
<l*>79), retained Aphe/ochetrus in a subfamily of Ihe 

Polhemus (1984) briefly reviewed the Naueorinjc 
o\ Australia. In tin.-, paper, I redefine the Australian 
-P'Vies of the Naucoridae, including A, uustruitcus. 
and describe a new species oi* Noucoris from 
Western Australia, 

All measurements given are m mm. 

Key to Australian genera qf Nutuoruute 

Vnk\ L'Wtilv rounded between die- eyes (I ~(g£ l , 7 $ K>. 
Rwe Icinui inoassate (fig. J5f Rostrum shun, not 
icjK-liinji the lor.: eosae. Antennae, slion, thrce- 

ve^utcntcd and not visible lioni above 

, , . . .,.,,.,,.. Nuueons I -abriLiu* 1775 

Verks piod^ed helween the eyes (Fig, &2), fore lemur 
not iii'Jinxsale (,!•(£* 63 X h-tl. Roslmm IpMtt, reaching 

Hope 1 r1tornnlr-.1iL.4t < 'nlk-uioir-. I'mveivitv Museum, 
Oxford, t'k 

the m"id-eo\ac. Antennae long. four-segmented, oticn 
visible from above . Aphc/ochtirus Westwood >H33 

Naucoris I abrieius 
Body variably flattened dorso-vent rally. Tore 
femur broadly triangular and incrassatc, fore tibia 
folding into a shallow sulcus alone, femoral margin. 
Mid h\k\ hind leys "cursorial". Antennae three 
segmented lying beneath eyes on underside of head. 
Rostrum three-segmented, two segments visible, not 
reaching fore coxae lore legs separaied by a 
prominent pilose ridge produced eephalad. Between 
mid and hind coxae, smail plate like keel, not readily 
Visible when viewed from side. Head and pronoturn 
variously punctate light and dark brown, anterior 
margin of pronoturn more or less straight. C tavus 
and curium uniculorous or bieolored; membrane 
not always clearly differentiated from curium and 
embohar Fracture nol always entire I'ndersadr ot 
abdomen variably pubescent. Ventral latenvtergites 
shining, width variable, always becoming obsole- 
scent at apical margin of third sternite, posteriorly 
reaching seventh sternile. Sternilcs divided by fold 
or suture either side of mid-line (Fig. 2) Male 
genital capsule "boat -shaped", laierally heavily 
selerotised, apically membranous. Parametes 
asymmetrical. Male fifth stcrmie inedianly 
asymmetrical. Female operculum slightly rounded 
and distally emarginale. Wilhin Australia, Naucoris 
is of a fairly unifotrn appearance. 

Act H) ihe Auslraliun species of Naucoris 
I P >Menoi margin of pronoium not produced 

caudad ;n humeral angles (Figs I, 7, & 8">. Venter 
eii her appearing bare or with line short hairs .. 2 
Postenoi margin ot pronotum produced caiidad 
at huineidl angles (Hg. 41), AH abdominal 
veiitnies elothed in line golden pubescence 

V. stibuurva* sp. nov 

1 Mi S.uiellum and most of cinhcilium pale yellow 
(fig i) N aus matteus Swl 

SiJUtellum urnYolorou* daik iC-dtlish hrowu or 
black, embolturu basally vuriably pale yellow to 



FiftJi W>. h'aucoris uustralicus Stal: 1, dorsum and 2. venter of Leetotype 9 • 3, genital capsule and 4 t 5, left and right 
paramercs: 6, aedaeeus old", hid>vold, Qld; ms, median siernite: op. operculum; ps, parasternals; vtt, ventral 
latcro-tergites; hi ■ vii sternites. 

yellowish brown, apically always dark brown to 
black (Tig. 7) 3 

3 i.2) Posterior angles pf con next va 4-6 sharply 
produced (Fig, 10). Shining area ol 3rd ventral 
lalero-tergile hasally broader than anterior width 

of 4ih and brnudh mtuscaied (Fig. 39) 

_ _ /V, rhizomaTUS Polhemus 

PoMerioi angles of conneviva 4-6 not sharply 
produced (Fig, 9). Shining area of 3rd ventral 
latero-iergite nol basally broader than anterior 
width of 4th and not broadly iniuscatcd 
(Figs 35-3S) 4 

4. (.*) Dorsum of head and pronotum rugulose and 
rather dull. Shining area ol ventral latcro-tergitcs 
narrow (Figs 9 A 3S) /V. subupacus Montandon 
Dorsum ot bead and pronoium smooth and 
usually shining Sinning area ot ventral Intern 

tergitcs broad (Figs 35-33) . . a', cannrcx stal 

Naucoris australicus Stal 
FIGS 1-6, 26, 32-34. 51 

Ntiucoris tf usiruiictts Stat, 1 876, p. 145; Luudfolod, 1933, 

p. 62: La Rivers, 1971, p. 71; Polhemus, 19S4, 
pp. I S7-15S 

Lectoiype: Female, Austral boreal. Thorey; I rf and 

1 9 paralcctotypcs Australia Orient med. Thorey in 
the Riksmuseum. Stockholm, vtd. 
Distribution: Queensland. 

Male 8.0-9.6 long, width 5.3-5.7. Female 
(lectotype) 9.5 long, width 5.7. 
Colour: Head, pronotum, seutellum and most o( 
crnbolim pale yellow. Head shining with a triangular 
group o\^ shallow brown punctures. Pronotum 
shining, medianly faintly and irregularly 
transversely striated; anterior margin dark brown, 
groups of brown punctures converging from innei 
margin of eye towards the median line posteriorly. 
Between these groups of punctures, third group ol 
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 infuscatcd with irregular 
prominent brown spots. Underside and legs pale 




l-k"i 7 |0 Nuiii'oris spp.; 7, V. country Siul V dorsum. Moggitl Rarm, Qltl; 8 ,V. suhopacus Moiiumdon 9 dor-ami, 
I ngp Dam, N.I.; 9, A'. \ubopacus Montandon S abdomen, Adelaide River, N.7.; H», N. rfuromatas Pollienuis v 
.ilhlnmrn, Adelaide Kiver, NT. 

Structure: Anterior imerocular space almost the 
same as posterior. Pronolal humeral width 2.3-2.4 » 
median length, the latter between 1.4-1.8^ head 
length. Pronotal lateral margins slightly convex. 
Scuiellum between 1,5-1 K* broader than long, 
lateral margins simiale. Fabrum 1.5-1.7* broader 
than long. Mcsostemum conspicuous, .slightly 
produced eephalad. Posiero-laieral angles of the 
fourth and fifth eonneviva forming an approximate 
90' angle, sixlh and seventh slightly produeeU, the 
sixth mote prominent in the female (Tig. 1). 
Depression ol seventh parasternite conspicuous, 
almost reaching outer margin ol sternite. Male fifth 
sleruite (Fig. 26). Shining lateral margin ol Ihird 
ventral lateto-teigile narrow (Figs 32-34). female 
seventh sleruite about 2/3 median length of sixth 
(fig. 2). Operculum (Fig. 51) aboui I.Ssj broader 
than long. Male genitalia (Figs 3 f"0 
Material examined: Feetotype female, Austral 
boreal, Thorey (Stockholm) One d', Queensland, 
Fisvold; oik rf 'Queensland' (Hritish Museum. 
Natural History), 

FollteinuN ( 1 984 > gives data for a single male bom 
Qld, 14 miles NW ot Ayr. 2(1 m. 14.\i.!962, I S 
Ross, E Q. Oavagnaio. This specimen in the J. T 
rolticmus culleeiiou. 

Sial's 1876 account of Naueoris is confusing 
wncie it refers to Australian species. He 

distinguished /V. australicus from H tfttt&tx 
because the former had a paleseutcllmTi ("seutcllo 
pallido"). In the original description he refers to a 
male. The type series received from Stockholm has 
a female labelled f THUS, the other two specimens 
have dark brown sculella and are referable to N. 
enngrex The female labelled TYPUSj is hereby 
designated Feetotype and is labelled a< such as it 
is the only specimen which agrees with Stals 
description. The remaining male and female 
labelled "Australia Orient and Thorey" are hereby 
designated para leeto types ol V austral 'ictts 
although they are in fact N, con#rt\\. 

The slender data available suggest that N, 
austra/ui/s is confined to Qld- 

Nctueons australutn is similar in general 
appearance lo N. eonxtvx. Il is easily distinguished 
by the pule yellow scutelhnn and embolium. Across 
the widest part of the body il has three prominent 
pale spots, shared by no othet Australian naucond. 

Woodward et ui (1970) Fig. 26, 72c, figured N. 
uustralieus, nor N. eongrex as stated. 

Naucons cangrex Sial 
I K.S 7. 11-25, 27 28, 35-37 & 50 
Naucuris CODgr&x Sial, 1876, p. 145; Fundblad, 1933, p. 62; 
La Rivers, 1971, p. 71; Fothctnus, I9K4. pp. 157-159. 

Nolo type: Fe in a I e, M o re t o n Bay 
Riksmuscum, Stockholm, vicl. 

n the 



Figs 11-25. NaUcoris congrex Sfy\ male genitalia; II, 13, 16, 17, 20 & 21, Tasmania, Tomahawk River; 12, 14, 18 & 
22, Victoria, Yea River; 15, 1^, South Australia, Piccaninie Blue ponds; 23-25. "atlStntfiCUS" pwi of i\pe series 
congrex, (M-12) genital capsules; 13- 16 & 23, left para mere, 1ft, different aspect; 17-20 & 24, right parameres, 
20, different aspect; 21, 22 \ 25, aedeagus. 

Distribution: Tas., Vic, S. Aust., N.S.W. and Qld. 
Males 7.7-8.7 long, width 4.8-5.3. Females 8.5- \) 
long, width 5.4-6. 

Co/our: Occurs in two forms as fallows. 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 ot punctures coalescing into 
brown lines; lateral margins smooth. Sculcllum 
varying reddish-brown and black; inner lateral 
margins o( clavus pale. Emboli um basally pale 
yellow, apical ly verging towards corial colour. 
Membrane dark brown to black with many greyish 

punctures. Connexiva pale yellow, sliehtly 
infuscated posteriorly with dark brown punctures. 
Pro- meso and metasternum mostly black; lateral 
margins or mesosternal ndge yellow. Ventrites black 
with silvery hairs. Ventral parasternites dark bmw n 
with creseentie pale yellow indentations across 
ven! rites. Legs pale yellow. 

Pale form: Head and pronotum pale yellow back- 
ground, brown punctures on head reduced in 
density and coverage, wholly confined to basal half 
o\' head. Pronolal pattern reduced; brown stripes 
on postciior 1/3 almost obsolete. Clavus and 
corium brown, membrane black. Scutellum as in 
dark form. Presternum straw-coloured; coxal plates 
slightly pigmented Sternites dark yellowish-brown. 
Structure: An tenor interocular space slightly 



narrower than posterior space; inner lateral margin 
til eyes almost straight. Head width to length ratio: 
males 3.1-4.1 greatei than length, average 3.4\. 
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 1.5-1.7 x broader 
lhan long, lateral margins slightly sinuate. Labrum 
between 1, 3- 1. 8 k broader than long, average 1.5 ... 
Nhsosternum similar to N. australicus. Postero- 
lateial 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-iergite. Male fifth 
stemiic(Figs27&28), little variation between Tas. 
and O'd forms. Shining lateral margin o( third 
latcro-tergite broad (Figs 35-37). Female fifth and 
sixth sternites same length, seventh sterniie 2/3 
lenglh sixth stcrnite. Operculum basally irregularly 
cmarginate (Fig. 50), Male genitalia (Figs 11-25). 

Slight variation between Tas. and mainland forms. 

Material examined: Holotype female, Morcton Bay 
(Queensland) Rikstnuseum, Stockholm, las.: Georgetown, 
G. &. Cole; Launceston; Wedge Bay, C H. Hardy, South 
Australian Museum. PawJccna Lagoon, Coal River; Pond 
at karanja; Blackmails Lagoon; Tomahawk River; Cape 
Portland lagoon; Itena Lagoon; Moriarty; Ncwnham; 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 
(AN1C) Canberra. 

Vic.: Yallock Creek near Melbourne; farm Dam, 
Hammano road near Neerim (some adulls teneral also 
immature*); farm datn near Bunyip River, Labertouchc 
North (immature); Lake Purumbete: Eleucharis swamp 
aboul 8 km from Colac; farm dam about 2 km from 
Barwpn Downs (immature); billabongs along margins of 
Barvvon River at Inverleigh; "Sheepwash" lagoon between 
Caihkin and wesi of Molesworih, Yea Rd; backwater of 
Yea River at Yea (25-29.iii.1979, L Lansburv) (ANIC, 
Canberra); Noble Park, 6.K.I9IK, F. F. Wilson (SAM). 
S. Aust.: Adelaide, H. M. Hale; River Torrens, Murray 
Bridge, H.M.H.; Bridgewater, H M.H.; Lueindale, A. M. 





J2 I J5 

33 36 








Fibs J6-40. Ni/miffts SD|>- "iule 5th sternito of 26, A. amtratieus Sial, Qld; 27, A', cottgrex Stal, Tomahawk River, 
l.r-.,: ZB. V ctWgrVJ Stal, Moggill Farm, Qld; 29, A', suhaureus sp. nov. Millstream, Qld; 30, A. rhizotnatus Polhemus, 
i . uMiial i. ( r«ki NX; 31, 'V r yy(j0p</£u£Monlandon, Fogg Dam, N.T. 3rd ventral laterotergiteof 32, A', ausiralicus 
Stal, Lcilolypc Iciiulc; 33. V austmticus Stil, male, Fidsvold, Qld; 34, A' utistralictts Stal, male, Qld; 36-37, Naucarti 
congnx Stal; 35. Holotype female; 36, male, Tomahawk River, 1U\ j 37, male, Moggill Farm, Qld; 3K, N. suhtiputu* 
Mmiiaiidoii, male, Fogg Dam, INT.; 39, A. rht^oniulw- Polhemus female, Adelaide River, N.T.; 40, A. suhaureus 
'■.}■> no, lloloiype male, MtlUrrcanv, Qld- 'terminology p protonum; iii ps 3rd parasternite. 



lea: Roonka Sta., Blanehciown, in shallow water along 
edge ot Cumbunga Crock, \2\ .1973, R, Inns ISAM), Ml 
C.;unhjcr region, Piccaninie Blue ponds nearGlenelg River. 
20.iv.l979, I. Lansbury; "Ewcns" ponds near Mt Gambier, 
2l.iv.l979, 1.1 ..; Naraeoorte region, Hoik's Swamp neat 
Bool I RgOOTJi 2~\ iv 1979, II,.; same data, Bool Lagoon, 
23.iv.l979, I.L.; Kangaroo Island, river iust beyond 
Karfalla on road to Kingseotc. 9.iv.t979, U.. (A NIC, 
C anburTa). 

Qk\: OmnamulU H Hnidcjsile (SAM); Allietton Tabic 
land. S km from Matecba on Mollov-Marceba roftd, 
22.V.1979, I.L.; Brisbane, Moggill larm dam, b.iv.l'/N 
(some iiniTiiJiu.L-) I I , (ANIC. Canberra). 

A male from Tasmania, Tomahawk River, 
2.xi.l972, B. Knott has a pair of tunnel-like 
structures made with detritus and what seem to be 
short naueorid hairs lying along the 2nd/3rd 
cpisternal suture where the middle coxae articulate 
within the thoracic cavity. These structures are 
almost certainly phoretic in origin; no remains oi 
any organism were found. 

The broad shining lateral margin ol the thnd 
ventral latero-lergite in both sexes and the 
conspicuously asymmetrical fifth sternite of the 
male are diagnostic. 

Naiuoris wl>o/nuus Montandon 

FIGS S, 9, 31, 3&, 53. 7, 56-61 

Ntnworis suhofweus Montandon, 191 ), pp. 223-224; 
Lrodblad, r93A,pi$2; La Riven, I97i.p.7t, Polhemus, 

19K4, p_ 160, 

Lectofypt; Female and three 99 paralectotypes. 
Northern Territory, Adelaide River, in the British 
Museum (Natural History) \ id. TWo female 
paralectotypes, same locality SAM, vid. 
Distribution; W.A., NT, and Old. 

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 aigulose appearing dull, in 
artificial light rather more shining. I'roiuxum 
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 pitchy. 
reddish-brown to black with taint reddish-brown 
area medianly. Clavus and eorium reddish-brown; 
dark specimens from Fogg Daifl, NT. have obsolete 
yellowish-brown palches. Embolium apically yellow 
merging into corial colour. Membrane blotchy 
yellowish brown, membrane of left hemtelytion 
smokey-brown. Connexiva yellowish -brown. 
posteriorly lightly tnfuscated. Mesoslernum laterally 
yellow, otherwise daik brown to black. Meta 
sternum dark reddish-brown. Median sternites 
reddish-brown, ventral latcro-tergiles suffused with 

H ■ S 



igs 41-49. Naiwans \itbaureus yfp, nov. Holutype male, MilKtreani, QU\ 41. duisitiu: 42, genital capsule; 41-44. I. i< 
,' m imcrei 4i 46, righi pjramerr; 47, apex ol Fig- 45; 48-49. .t./'loirii'.. 



cicscentk yellow indvniaiions. Legs yellowish- 

Pah form: Head and prouotum paleydlowish-grey; 
brown spots on head nol scattered btjl in groups 
eubci side of niid-linc. Punctate area o\ pronotuui 
HOI m> w-tearlv differentiated from background 
CoJcjwr; pattern similar bill almost obsolete. 
SciUcllum paid Irregular yellowish marking'- on 
claviis and eorium obsolete. Conncxtva gtfWteh- 
wIImw wild rami iufuseation atony poslenoi 
inaigtus. Underside paler; mesostemum almost 
Completely pate yellow. metastermim ringed with 
reddish-brown along keel. Pigmental ion of dorsum 
dccidedlv fugitive, lading, rather quickly. Dark areas 
til head iwd pronotuni lade rapidly leaving 
.sculpturing and rugulosc areas. ConneMva lend t(3 
fade m unilorm^reyisli yellow and scuiellum loses 
as almost black colouring 

Strucnof Anterior inteaxiilar distance always lag 
than poslenoi mleioculai space- Inner margin ol 
eyes converging anteriorly |Fr"g. S>. Greatest width 
ol" head 1 4 ■ median length,; variation in pan dm 
it. t>osition oi head post-mortem; card-mounted 
specimens have head pushed forward in front oi' 
piouutum. fresh or "wet" specimens Lend to have 
head 'ktippinfi" below level of prouotum Pronota, 
hunteial width between ?.25 2S < median length. 
falter being vu. 2- median length Pronotum 
clcwucd oi "hump-backed", lateral margins diverg 
im\ sometimes siionglv rugulose with cunspicuou*. 
irfcjmlnr sculpt Urifltf over most oi disc. Seutelluni 
finely punctate; mule about I 7 4 hroader than Jang, 
female 1.6-1.9-.- Posteu> lateral ancles of conuexiva 
broadly exposed terming approximate 90 angle. 
I abriiiu 1-4 I 7, broader than lone Mesosternal 
ridge prominent, clearly produced ecphalad. Male 
fifth srenute slightly asymmetrical. Steiniies five 
and sis of male wiih long hairs; remainder of 
Mernites dark, tomeniose. Filth female median 
sternite slightly shorter than sixth, seventh not as 
lone as lifth (Ffg, 9). Operculum (big. 531 SWtlWg 
urea of third ventral Imeno-teteite narrow iPSft. 38). 
Mate rem m li-t (I i^s 5<S 61) 
Li't-lofYfjcih-yixruihotr l*he lypc scries ol NaucOfU 
m*/>< vwws comprises two species, V. subopttt iti tnd 
N. rht:.t»lunu*>- Momaudon did not designate any 
-.pecuneu as lype. A female labelled Noith 
AiiMr aha, Adelaide Kiver, Adelaide River SiaLion, 
Jul) 1-4 I St h. IW.M.J Walker k hereby designated 
and labelled U-ctotype- Three females with Identical 
data arc designated parateeiolvpes. All the 
loreefiui- aie ui rhc British Museum Natural 
Hlsrnrv, A liothrr It-mule paialecmiype in the 
Soulh \usimh<m Museum has been designated 
winch Was "M.rto.dh part of Moniandou ssyjiiypic 
-.cries. A -txtli tetnale from North Au-aralb, 

Adelaide Kiver. Adelaide River Station, K 13th 
August. 1890, J, t Walker is labelled and designated 
paralectoiype of N. subopacm although ii is in fact 
N, rhnamutus. All the preceding material was 
collected during the Voyage of H.M.S. Penguin 
lS t X)-lS93 and presented to the Btitish Museum by 
the Lords of the Admiralty, 1896. 
\htvnat examined! NT.: Koolpauyah. (I K Hill; 30 miles 
cum ol Darwloi G.r-.H-. sam. McMuuls lagoon near 
Dai win, 4-Kxv.W>. I, lansbury; F^Sfi &*** '^r Dapwin, 
4 »iSa P>7<>_ IX. ; Hltatjony near Nonrlauytt- Rock, KaLulu 
Nauotial Park, lU.v IM79, Id., Arntrun Highway, Mary 
Kivcr pools, I7.V.1979. I.I . ANIC Canhvtra. .lahiluka 
area, M*BGlfl C.rce^ OoOciplalUi Buffalo hillabony,, 
J2xl579 prnmaltire) K- Tair. vjme duia, ^flviiiy'V; 
JjdltKika Uilluhong, I7.7K1979 oimMittuio); -jiiw data. 
Ilv.1979 (iriimamre); Nankeen Billabonii, l-ViiiJs>79; 
Winmurra Billubon^, t4 lvdf)7V. K, laii. ANK Canbcoa- 
Qkl; Spin Rock, 14 km Mtitll of Uuta, 23-26Aa»W5, 
Li. B. Monteilh QM, Brisbane. 

W.A.; Uovcrlev Spin&s. I.iv .P)ft9, O. D (uiihani; WAM 
Naucufh suhafwitt \>, a '.mall robust species; iln- 

bioadly exposed connexiva and narrow shining ihird 
vcniral laiero-iergite disimguish N. UthopUCUS from 
the rc-M ol the genus within Australia. 

\0(,ttiw< rhizonwtus Polbemus 

HCjS 10, 30. 39 & 52 

Nattcarh ,nhofnuu\ Momandon, 191.^, pp. Z13-Z2A 

Xuui't>r^ rhizomaut'i Polhcmus. \$A4\ pp *— 1% 
ff/ilotvf.'t: Male. NT, near Darwin, ( oouiaiie 
t reek. II .sir. 1977, .1. I. Polhemus, in I he ANIC, 
t_ anbcriii. Paraiype male and female originally 
depoMlcd in 0\to,d (Polhemus, I9R4: 157) is now 
iu ANK , Canberra. 

Superficially this nancorid resembles A. 
stthopacus. The lauer has nanow shining latero- 
tcrgiles (Fig. 9); those o\ V rhtzomanis are broad, 
diaally miuscated. The lateral margin of third 
laiero tergite basally much btoader than apes of 
fouiih (Fig. 39K this leant re common to both sexes. 
Scutelluni rel oiwdy shorter ihuu iiuV suhopaats, 
always 2 -. broader than loug; latcial margins more 
rounded t\\\d apex far less acuminate Hian t|1 V 
subopacus. Me^oslernal vjdge muie produced 
ecphalad. Male fifth siernile <Hy. 30) Temale 
operculam (fie. 52t. 

MutirwUwHHiihi'ci:* : >ne S pani'.vMiiyp.;* a V \uho(>ui:ui, 
N, AnsO nli.i , VlcUtiOc RtV^r, -VdcluiJc Ki\cr Slaiuiit, 
g Imh \i^uO ( '890, I I WvitUr SAM. 

\.i.. JaftTuku i^itort. Magcla floodpbin, Buffalo 

liillabotiv, :2.i.lV^, R. Taiti I iramaHiu-\,iI 
murkmi!-- nuxnini-ni; same data, iO.vii.P/ 7 *', I ./tiiU-n svilli 

a* N Mi/llllHH-US. 

l\slhcmus (I9S4I describes Ihe habtiai whea' V. 
rhaonnilus was lound m Australia as a deep waiei 
crrck, rhe bugs be|t\fi found amongst the tangled 
roots of l\ttt(hmu\ Nonrom suhopocus tends 10 
be found in shallow weedx habitats The slender 



wvir. M a : " 

"57 59 

Figs 50-6). I'cmale operculum of 50, Nuau>ri\ C0tt£MX Stal, Holotvpe; 5l T /V. austraticus Stal, Lectotvpe; 
51, X. rhttomam Polhemus Adelaide River, NX; 53, N. subopacus XJomandon, Adelaide River, NX; 54, A 
subitun-us sp. nov. paraiype, MilJstream, QKJ S5, V subuureus sp. nov. foreleg, Holotvpe male. Midstream, Qtd; 
A. subopacus Momandon, male genitalia. 56, 57, 59 St 61 Fogg Dam, NX., SK & 60, 30 miles east of Darwin, 
NX 56, geniial capsule; 57, 58, right paramete; 59, 60, left paramere; Gl, aedcagus. 

evidence suggests that occasionally N. suhopacus 
and vV. fhizumutus occur in the same habitat. 

Naucttris suhaureus sp. nov. 
FIGS 29, 40-49, 54 & 55 
Hototypv: Male and paratype female, W.A., 
Millstream, 23,viU958. R R McMillan in WAM, 
Perth. TWo V V paratypes (one feneral without dorsal 
pigmentation), W.A., Drysdale River, 18-2J.vhi. 
1975, (14' 39S, 126 '57 T:), I. V. B. Common & M. S, 
Upton One S paratype, W.A., Drvsdale River, 
3-8.vii.1975, (15 C 02'S, 126 '55T), I.KB.C. and 
M.S.U. ANK., Canberra, 
Distribution; W.A. 

Male 7.0 long, width 4.6. Females 7.8 long, width 

Colour: Head yellowish-brown, median ly a 
longitudinal stripe of contiguous brown spots, 
eonstricted 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 lighily striated 
with inverted brown "V" shaped pattern between 
secondary groups of brown sculpturing; much of 
disc covered with shallow pits; middle of postenoi 

margin with short brown bar directed cephalad. 
Sculellum pale with broad brown band from base 
to apex. Emboliuni basally yellowish-brown apically 
dark brown outlined in pale yellow, Clavus and 
COfitun dark brown with confused pattern o\ 
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 latero-tergites basally 
dark brown. Legs pale yellow. 
Structure: Posterior interoeular space greater than 
anterior width. Inner lateral eye margins more ox 
less stiaight, converging anteriorly. Greatest width 
Of head 3 • median head length and just over half 
of pronotum. Pronotal humeral width about 2.5 x 
median length, lateral margins slightly convex; 
hUtncraJ angles directed caudad. Scutellum 
punctate, just under 2 >. broader than lone 
Hmboliar fracture anteriorly prominent, posteriorly 
obsolete. Clavus and eorium not differentiated; 
membrane continuous with eorium and coriaceous 
(Fig. 4)). Postcro-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 K broader 
than long. Mesosternal ridge elevated posteriorly. 

Al'STkAl l\N NAl C OklMAf 


crc.vl covered with shining yellow hairs. Mesoslernal 
carina vestigial Sternitcs co\ercd in thick shining 
golden pubescence Ventral I.jlct x»-icr^Ttcv narrow- 
(1 ie 40) female sixth \tL-mik 2 • longer than fif'h 
and 1.25 ■ longer than seventh, Op«CU|Ulil 
(Fig M>. Seventh paraslernhc with vest • cat 
depression. Male fifth steuiue (Fig 29). From leg 
(I ig. 55). Male genitalia (Pigs 42-49), capsule 
sclerosed, not membranous -iptc.illy, parameter 

Naiuons stihautvus is easily recognised by the 
thick golden pubescence covering the median 
\termic>. prO<IU«<JI pronorj arglcs and striking 
pattern on Iht clavus and eorium. 

I nmnwtns on distrtbuttrtn mri hialngp oj 

Australian Naueoris 
Within Australia I he re ate [vvo species pau> 
i\, uttsuvlicns/cotigrex and N, suhtt/waisTfuzo- 
tnatns. The firth species N. subaureus is anomalous. 

!\utu-(>rL\ iJH.siruiicu.s/congrcx: These arc supcr- 
Itciallv similar in size and general coloration and 
in having an "easterly" distribution. There are 
in'-ufl'n k*nr to comment any Further on N. 
oust rut tens. The most northerly iveord ol V. 
POttjf/W is Mareeba, Athcrton tableland. Qld. Its 
occnticnec so lai noiih may be as a rcNuh ol a 
"elntialic" anomaly caused by Ihe "high*" plateau 
with a mote temperate climate compared with ihc 
tropical climate of the surrounding area-.- V. 
eongnw is COMMON in las. and Jala surest l hat 
at Ihe soulhern end ol the range, U is norrnaih i 
voltiue. Occasionally a partial second generation 
may overwinter in the immature stages. Data b.< i( d 
on samples collected 1972^1973 from Tas, (small 
lake, f'awleena; Coal River 4 miles north of 
Richmond; small lake, uppet reaches So roll Knot 
and Ulackinaas Lagoon) lend lo support this h\po- 
thesis Mature adults arc present from 
Vlatvh i Mober, but Iyv then numbers appear to be 
|0W fairing flUd oviposjtion presumably take place 
in August-September, ty Nenernnei. 3td mstar 
nymphs are present Immature stages continue Lo 
be present unfit February, lencrai adults start 
appearing in January t>r earlier (no claia are 
available lor December) By March adults are 
niatnie and the immature are not usually 
ptesent. Immaimrs were collected m Mav front 
Blackmans Lagoon possibly representing an 
overwintering population of immature /V. ennxn \ 

In Tas \ cWffiWV has -< 'ow indes gift 
divetsilv. n»i example it ,V tonyrcx is COJujAlOH '' 
om COlUtWIC* OthCT wiitctbuus Ol soineM habitats 
sampled H fdtt$*& cvas found in ten. dfehl Ol 
which have live other species ol aquatic and semi 
agnatic Hercroptera Greatest diversiiy. was in 
Kla' kmans I aroon (nine species total) winch WB« 

sampled qtlUfi MMHlVtly On the niamla.Ki. v 
congrex is found mosL commonly in Vie. This 
probably ts due, fn part, ro lite greatci numhet ol 
freshwater habitats in Vic. compared with S, Aust , 
and also reflects lack of collecting in N.S.W, In Vic, 
V, con^tvx was found in nine out ol 37 babitals; 
greatest species diversity was 15, the lowest three, 
ol the latter, N. con^wx was rhc commonest species. 
Immature stages were collected from March-June 
in Vic and Qld. 

Notuons suhufuicus/rhizoinoms: These- I wo 
species are remarkably similar in size, configuration 
and coloration Roth have a "northcrly M distribution 
extending from NW, Australia, across the NT. lo 
Qld near Laura. Most o\ Ihe habitats sampled by 
the author in 1979 where V subQpttcus was found 
also had Difj/onvv/ius (Belostomaiidae) present. 
The bclostomaiid populations were always much 
gieatei numerically than those ol /V subopacus. 

fifaVCorlS nthuureu.y Does not belong lo either 
ol Che species pairs and is unlike other <\'uncoris 
species from SE Asia. The Millstieam Oasts south 
of the Great Sandy Desert if noted for its distinctive 
lautia. csr-eoalh udonata (AVatson 1981). Data on 
water-bugs are scanty. The Occurrence of V 
stthaurem in the Drysdak River area of NW 
Australia suggests ihat the Sandy Desert docs not 
impede the dispersal of water-bugs between the 
Vhllarcarn Oasis and the Kimbetley. 

AplwU'ihi-itus iiitstruhi-us Usinger 

FIGS 62-69 

Aplu-'hhfwirus attstratti'u.*. I srlngffr, 1^3 7, pp. ^41 ?42- 

\()hv'Un ht /n/w Woodward ci ol , 1970, \\ 4Sh (inciirioncil 


Useful dtagnostic leatures are: Head, pronotum, 
-i liielliim and most oi i. lau >s rHOFCOl less shining, 
Coralin and etultoliuni dull. I'ndcr^idc appcarint' 
.smooth and shinmg 

Body dorso-ventially compressed. Mead longer 
than hroad, antennae long, four-segmented, usually 
visible from above koMiwn reaching mid-coxae. 
Protloluiu transverse, latciai margins csplanaic; 
Mu-tTan length clearly shorler than rnedian head 
length, posterior margin roundly cmarginalc in 
from o\' scutcllum. Embolium basal ly broad, 
Upeiing uptcally along margin o\' coriunt. 
Membrane di.slincl from eorium C onne\ivum 
bn»adly e.v|n>sed. (Usjiiger's- iigme shows: lirsi visible 
eormeMva! segment mfuseated.) Specimens from N. 
Old have ihis segmeni pale yellow, not bioadlv 
itil'uscarcd, as ate remainder (Lip- 62). Femora u( 
all lees dorso-ventraKy flattened (figs 63, f»4) \or<: 
and mid lflg$ .'.like Male genitalia (f igs r»*> 6V); 
witfm. . apSUlt, pair of processes attached to lateral 
plates. oVuillv pioccsse* beavif, .cico.Uiscd >vit|i 
ititik brown spines apieally (fig, ftfc) These 



Figs 62 69. Aphelocheirus austrulicus Usinger. male. Cape York, Lockerbie, Old, 62, dorsum; 63, loxe ley; 64, raid 
leg; 65, geniia) capsule; 66, detail of spinose processes on lateral plaies; 67, aedeagus, 6S, 69, loft and riglu 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 Apehkv 
cheirines. The female from "Captain Billy Creek 11 
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. 
Mtiieriut examined: (all maeropterous), Qld, 142 45 F, 
1! 40 S. Dividing Range, 15 km west ot Captain Billy 
Creek, Cape York Peninsula, 5-12.n.l976, G. & Montcith; 
Upper Qld, Lockerbie area. Cape York, 11-27. iv.1973, 
G.B.M. QM, Brisbane. 

Aphelocheirus is normally thought to be 
restricted to well-oxygenated water; the bugs arc 
found crawling about beneath rocks and stones, 
Typical Aphelocheirus habitats arc 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. E. Woodward in 
lilt. 4.iii.l977). 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 oi 
water purity. The species ,4. aestivalis has an 
efficient plastron allowing gas exchange to take 
place without the bug having to surface, in contrast 
to Naucoris. The Cairns habitat lends 10 suggest 



that Australian Aphelovheirm has a similar plasi ron 
rcspiralion system, However, t he presence of the 
species in Cape York occurring in swamp sluggish 
creeks need no! be considered unusual as Thorpe 
& Crisp (l*>47) list a wide variety of habitats where 
Ajdie/ocheirtts aestivalis (K) has been found in 
Europe. Within the Palaearctic region, 
Aphelochelrus is dimorphic but it is not known if 
it is so in Australia. 


I wish to thank Dr P. S. Lake, Monash University 
and Dr A. Neboiss > National Museum of" Victoria 
for their unstinted assistance whilst I was in 
Victoria. OrO. Gross, South Australian Museum, 
Adelaide who enabled me to visit the Mt Gambler 
region; Prof. W. D. Williams ol Adelaide University 

who arranged the trip to Kangaroo Island. Mr 
Graham GnTlln ol C.S.I.R.O., Alice Springs and 
Or Gary Fit! (then in Darwin) lor their valuable 
assistance in the Northern Territory. MrC. Pcdersen 
Ol Norauda Mining Tor his extended hospitality at 
koongarra. Mr Wallord-Huggins for assistance in 
the Molloy region of Queensland. Dr T, L 
Woodward, Dr G. B. Momeith and Dr R. Kitehing 
for their great help in southern Queensland. Dr 
C N. Smiihcrs, Australian Museum, Sydney Tor his 
assistance. The work was commenced during the 
tenure of a grant from the Lcvcrhulmc Trust 
(London) and grants from AB.R.S. and C.S.I. R.O. v 
Canberra. Finally thanks to Dr W. Dolling, British 
Museum (Natural History) and Dr Lindskog, Riks- 
museum, Stockholm for the loan of critical types 
and Id Sandra Lawson of the Zoology Dept, Univer- 
sity of Adelaide for typing the final draft ol the 


Cmv\, Vv L. St Mil i i.K, N. C. L. (1950) Checklist and 

Keys 10 the families and subfamilies of Hemtplera- 

Hcteroptcta. Bull, Br. Mus. nut. Hist. L'ritotnofofiv 8, 

1-45. , 

Mow ki andi, I . & Stvs, K (1979) fumpocor'ts asiaticus 

Gert. and sp. n.-A new aphclochcitinc from Vietnam 

and further studies an Naucoiidae (Heteroptera). Six 

nur. \!u\. Prttte 33(B), 1-20. 
I \ Rjvhrs, f. (197II Studies of Naucoridae (Hemiptera), 

Biol. Sot: Nevada Memoir? II, I 120. 
1 i simi ad. O, (193$)- /'ir Keniiinis der HQUJIlletl uiid 

semi-atjuatileti Hcniiptcreu von Sumatra, .lava und Bali. 

Arch, tlvdrohiol. Suppl. 12 (Tropische Fiinncngewasser 

tW, 1-4X9. 
Mont"»M)onj, A, 1 , (1913) Hemipten-v. aujiaikiues. Nm»> 

el descriptions tie deux especes nouvelles. Bull. Sect. 

set. Acad, mum. 1, 219-224. 
Poisson, K. (1957) l-uitiic de FraACC f<IIW. 61 j Melenipicres 

Aquaticiues (Paul Lechevalter; Paris). 
Poi.141 vius, .1. I, (19S4) A review nfmc NaUCorinac nf 

Australia (Hetemptera: Naucoridae) J. Anst. ttif $0C: 

23, 157-lfiO. 
POKlV, Y. A. (1970) Notes on the classification of the 

Recent Naucoridae (I Icteroptera, Nepomorpha). Bull 

Acad, pot. Set, Ser. Set. Ot II, 16. 93-9H 
Stai . C. (1876) linumeralio Hcmiptcrorum. A', svensku 

VctenskAkad, llandi. 14(4), 1-162. 
Tiiorpi , W. H, & Crisp, 0. J. (1947). Studies on plastron 

respiration. 1, The biology ol Aphelochelrus (Hcmip- 

tera. Aphelocheiridae) and Ok* mechanism of plastron 

retention. J. c\p. Biol. 24, 227-269. 
LNiNf.hK. R. L. (1937) A new species of Aphelocheirus 

from Australia (Hemiptera, Naucoridae). Aust. Zool. 

8, 341-342. 
WvrsoN. J. A. I (19Kt) Odonata (dragontlies and 

damselflies). In A. keast (t.d.) Ecological Biogcographv 

of Australia; Vol. 2, 1141-1164. (W. lunk: The Hague). 
Wooiavaru, T. F , Fvans, J. W, & Fwn.p. V_ h (1970) 

Hemiptera. In CSIRO 'The Insects of Australia" 

(Melbourne University Press: Victoria). 




VOL. 109. PART 4 



BY J. H. Cann & V. A. Gostin 


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. 


by 1 H. Cj\^+ & V. A. GoSTiN) 


Cann. J- II. & Chimin. V. A. (W85) Coastal Hcdimeuuny iacies and forumitnfeial biofacies of the St Kikia 
Formation at Port Ciawtcr, South Australia. Trans- /?. Soc. S. 4ust, I0s)<4), 121142, 29 November, 1985. 

Thc Si kilda lonnaiion, WUIffti dull Si Vincent ami surcoundmu i oasial lands is redefined tu include 
.ill Holocene sediment- deposited under \Uc infiiiou-. 01 marine- processes. Upper surfaces ol the formation 
aic mil confined, hut rather ma> he uirkues ol uviue sediuieiiTaium. 

Port Ciawltr is an area wnere marginal marine ^edlme:!ll^ are accumulating under Ihc balHiuy, 'rapping 
and binding actions of scagrasscs, cyunobaslcrial maK niaiiimwes samphires and sullhusb. These plant 
communiries occur incssenlially discrcu Kpnc*, micccsmvcIv adjaceni and parallel io the tidal shore lines. 
Their innneiiee causes progiadanoii and ae^radaiion of rjiodostir catbonuie -quart/ sands and muds. Bivalves, 
gastropods and I'otarninilcru arc abundant and contribute ■unriinVanitv ro 'he carbonate component of 
sedimeni. Distribution of molluscs is closely related lo plain communities, sediment type and period of 
rjd&] tnundalion, I bus .i series of laterally .uIihocm sedmieiiiai v faeiesare recognised to be associated wilh 
the following .sub-environments: sublidal and irttcr'tdal sranass meadows; mangrove woodland; lidal 
disinbulaik's; eyanobaueuat mar^ o( the inner sMid Hat; high ride beach; supratidal lagoons or sabkhas, 
dunes and storm ridge*.; the estuary and associated backwaters ol rhc Gawter Rivci. 

loramini feral Diofacies, based on .selected sieved fractions of dense liquid flotation concentrates are 
established for llie various sedimentary environments. In (wo vibT»voo>, down-hole u^emhlages of 
forammileia are similar to those ftf modern Pbtl GawJcr envn'Onmeui,'.. PaUicocmhoiimcnlai interpietalion 
of Ihc cores, based on hnih luhology and assemblages ol tora minifeia , shows rliui the earliest scdimenis 
of (he Holocene Irans^rcssit wi wot essentially samphire muds and shelly SiOTOl debris. Later sedimemation, 
alter sea level stabilised, aacjL rfed through processes of pro>'ridation and aggradation, inextricably related 
to communities of seagru.^es, maitgrpvfif;, evanobactetial mats and samphires. 

Kiv Woros: Port t..i\\"e . ' mi'Si Vincent, Holocene, St Kilda Formation, loiaminilcra, molluscs, 
coastal sediments. 


The eastern coast of northern Gulf ft Vincent 
is normally subject to a low eiiery.v Wave ree. ftt 
This Tact, together with northward longshore cli ill. 
ensures thai ii is an area of aciive scdifuenianon. 
Low topographic relief of the coastal areas, and 
extensively developed sand and mud rials, result in 
broad tntertidal areas. Hieh Lidb associated with 
storm surges cause inundation ol not mall* 
supratidal environments. 

From seaward of I he low Liile slime lint Iw ih? 
supratidal areas there is a pronounced zonariutt o\ 
plain communities, These rangt from subrithd and 
interiidal sea grass meadows, through ifljoigjtsve 
woodlands and cyauobacieual mats, tosupiatidt I 
samphires and saltbush. In various wuv.s the plants 
baffle, trap and bind sedimeni causing gggra&ltjon 
and progradalion of ihc sand liars Molluscs and 
foraminilera areabuuOani and Iheu sfnih .uu' lo*-, 
contribute siynificanlly t(> the accumuUtin^ 
sedimem. Also, species ol these fauna ate 

* School of Pure and Applied Sciences. SA.t-AL. 

Smiihs Kd, Salisbury bast, S. Ausi. iiov. 
I Ueparlmeni ol tieoKu*y. I B1 Adelaide. UiW 

488 (iT'.O.. Adelaide. S. /\um. M)0l 

eksti naively distributed through the zoned plant 
communiiies and there are several consistent 
plant /annual associations. 

This papet documents relationships belween 
plant eomnumiiies and scdimetuaiy processes at 
Pfctl Gawlcr, northern Gulf St Vincent. A method 
of analysis, involving dense-liquid 
flumiion concentrates, and selected sieved fractions* 
is desctibed and used to establish loramim feral 
btoftaStf for the various sedimentary envirorunenu-- 
f hese dala arc used in pulaeoenvironmental analysis 
ot i wo tores o\ l-iolocene sediment- 
tor Pun ClawTer and surrounding coastal areas, 
it is proposed thai all Holocene sediments, 
deposiied under marine influence, riyjnly belong 
to the Si kilda formation. This usa^c includes those 
>cdimer,(s, Mthiidal, inleriulal and supratidal, 
forming al presenl. 

The Si Kinfj Formation 

In South Ausiralia considerable conlusion has 
prevailed in the interpretation oi' the St Kilda 
| ••>.!,. Uion since Firman <|966) first proposed this 
and other Holocene siratt.araphic units. 

On the one hand. Firman (l%6) defined ihc 
lormalion as 'Various lithologies tlehmilcd for 



mapping purpose bv ill* I. ... , r ract 

which eviends a mile til t . , , , |. , Kild& 

and elsewhere nlovm Ihe rrodcm coast." I hal is, hi 

included ihc modem wptfltidal ramphire sctttmeMb 

us a fades o\ the St KilCa hjritUJ rhtf 

interpretation was followed by ..m (1969) 

who referred "lighi-yrey shelly stranded beach tid\a 
deposits and shelly sills and sand overlain in places 
by modern intcrtidal and swamp deposits" io il.e 
Holocene St kilda rormaiian, aiul shOWCd ihcsc 
sediments io be distributed alone the eastern eoasi 

of Gulf St Vincent, attending inland abouJ S km. 

On Hie other hand Finnan's definiiiou iclaud 
sediments ot i In- St Kilda formation to ihe "raised 
sea-bed" of late (IK79. p. 69) and 10 the marine 
sands nnd clays Of the Osborne high sty lew -t 
(Aitehison, Spney & Cochrane 19541 Daily et »;r. 
(1976) elaborated on [his intCXpfClfltllHI, di 
relating the formatiou m u a high stand ol 
I -landrian sea when shelly sand . >• i ■ 
Their evidenee lot a supposed lilght i 1 1 ilot me sea 
level, about \,S metres ftboVe modern sea level 
consisted largely of stranded beaelnidgev, and ihey 
cite dates oi* J Kin) | 500 and I ur 
radiocarbon years 111'. icsjm_- I i Or Ihebawjind 
top ot Ihe Si Kilda lorniaitun. By implication, 
marginal marine sediment- n i - n... 
1 120 i 7S radiocarbon years B l\ wco. ihei-. 
excluded from the Si Kiltld Fbrnifllion This 
intcrpiclalion was reeenlly followed by Totelf (I'^.V 
in the Drafl Management Plan lot Use ftsrt Cittwlei 
Conservai ion Park \ figure illlftSI bfl '.••!•- 

stratigraphy, after Lindsay 11^65)' itfert to I 5 
metres ol modern sedimeni (black sandy day and 
mud with plant fibres) OVCttylUJ \}\ S- 

Kilda Formation (sands clay tdtti sh moil*). 

Thus ambiguity persists m Hit* use of the 
formation name. One inrerpretaunu ttrfudcQ the 
modern marginal mantle setKhiaiu together with 
Chose of earlier Holocvnraae The or Vi. based ftfl 
i tie premise of a hichei ifall! pi -: . 

Holocene sea, excludes those sediment* you 
than I 120 ± 73 radiocarbti IM> 

Bclperio et ul. (1981) rev.ewtd evidence relating 
to Holocene sea levels in Suulh Auvlraha and 
concluded thai there wa^ tittle evidence io nupporl 
the notion ot higher HcriucciK sea levels in the 
Adelaide region. I hey eiflpJta&hetl I ic Uiiporiu-i. : 
of processes ol coastal piogfddailiraAiu! Iiuattid i 
and supratidal deposition, and utuwidercd (hell 
ridges to rate poorly as evidence foi higher WW 
levels. In this papei WB follow Betpei o . I a/. (IWU 

l.indsav. r vt 1 1 '»)' : -i Stratigraphy i 

inierupalammoluiw tvl iHrOcdfiCfto I I Holt 

Gawk-i. S. aiisi IH-pi \l.m- K.| f ,..a. 53 -I 

i •■ reason io invoke any rceenl 

lowering Ol sea level and we are unable to 
,i ujjapfiicaR) differentiate the marine Holocene 
■-id' thin Cull St Vincent and suiiouiidiuti 

I fofC rccouruse (he Sl Kilda 
Porroatloi io include all Holocene sediments 
deposited under the iufluciicc of marine processes, 
Including csiuatine deposits, coastal dunes and 
aorm ridges, V- 5U0h« the St Kilda Formation is 
a rcadllj mappable unit, consisting internally of 
vanuus sedimentary fades, deposited under a 
-iv ol inarlUe processes. Sediments oi' the 
I ipsoi. RjrmniiOn and Semaphore Sand U irman 
IW.i ii,,v fhus be inteipieted as faeies ot the Si 
Kilda Foi illation 

Ihe base ol the Sr kilda Toimation is well 
del ill fid. Hie unit diseonformably overlies 
I', -ne sediments of the OiarlM (ilanville 
Formation (liunai, 1966; Cann I97K) and the 
alluvial Pooraka FtKUttatkid i%r-j EorJtest 
dcpuslUtUl u lllc Adelaide region kis been dated 
.oearlion yeaTs RP (Belperio et 
ul, IVX.t>. tn our prrscm usage, upper surfaces of 
Ihe Si Kilda Formation aie not confined, but rather 
may be surfaces ol active sedimentation. The 
neifi Bay founaiioit ol Spencer Gulf is 
equlvai£fil (hifU w at k984i 

The sludy area 

Ihc siud)' area ts siittaled about |2 km northwest 
the type location for the St Kilda 
Formation) I ie. U. 1 he aica is almost topographi- 
c-ally fltil with samphire swampland to the easi, 
[fussing throueh manf.rove woodland and open 
sAndflaO I ■ Cagr^SS meadows westwards 
Ininicdi.aeiy easovanJs arc sail concern r^t ion 
evoporafion ponds, the most Dortherl;. ul i ystetu 
lipa us soli extraction at Dry Greek, t iu- am 

inehi.K-s ihe estuary of theOawlei Rivet, site 01 ihc 

now dixu • • • u ■ hbon whai i Fie *) 

■- m bat lis SOUWt In the l*ara I-auli 
Bloc 1'm.i.cIv 4Q Km easi of Port Gawlci 

ti is an rtucrmittem stream, generally eooilned io 

n na indeifM|j chant I - iraverses the 

northern Adelaide Plain. Ai limes of heavy rainfall, 
fine ^tainvd clasitc sediments jtl transported inn. 
ihe estuary, from where they ao. redistributed 

ihrcnrt.,... iu ;i, a/ca by coastal marine 
pr.)L e , 

1 1 v region nl Stmih Aitsimlin which meiudes 
Pon pawfcr espenences a pronounced Mediter- 
laiican Ijpe U1 diffliHC Sununerx are mostly hoi and 
di v wiili iiiaviiiunn ..-inperaiurrs greater than 40"C 
UUtiDUimtfn. Winters are generally mild, m-in 
i. -iiiperaiurcs rarely falling let ft < Average annual 

I Jl -I 42 al Poil Gawter occurs mosilv 



• iv, i i iH.iiM.ii roap sivowbtg surface distribution uf Ihd 

Huloo-nt; Si Kikla and Pootaka Formations, u> used 
in I lii a- paper, and other simplified geolog) 

fix. 2. Rums tit Ufthon vvinni, Pom Gawky, rim, facility 

v.'.in irenoricNll; ptifiatu loj da- cvpun of grain arid 

■A.K.i in ih« juan r&3S u» imu. (Totefl <•/ al Hta-ty 

dutftlfl the cooler months, though Occasional 
summer thunder >rorms produce heavy falls of mm 
Ivaporation has been estimated at 1800 mm 
annually (Toiell L9B3). 

I lie are;).-, influenced by normal tidal change* ol 
sea level Bl I'on Gawlcr are extensive, The gradient 
M-awards from the samphire sail maisli U almost 
luipeieepiiblc and maximum astronomical lidal 
variations are about 2.5 m. Al maximum high tide. 

sea water covers the sand flats and floor o\^ Lhe 
mangrove woodland, and flows through udal 
distributaries to the samphire marshland beyond. 
Al minimum low tide, the sea retreats to lhe extreme 
seaward margin of the sand flat. 

Superimposed on norma! lidal fluctuations are 
the longer term effects oi surges with periods of 
1 to 20 days, and thought to relate to the passage 
D| weather systems (Provis Si Radok 1979). KelK 
(1984) has teported storm surge sea level 
nikUiaiions of 0.5 m along the southern Australian 
coast. When a storm surge coincides with a spring 
high lide, unusually high tides result. Abnormally 
Lure volumes of water move across lhe sand flats 
and through the mangrove tidal creeks, Hooding the 
Samphire marshland and adjacent (normally) 
supratidal areas. When these conditions are furl her 
accompanied by local winter storm activity, with 
strong south westerly winds, sea water is driven even 
more deeply Into the coastal environments. The 
combined eflects o\ tidal currents and storm waves 
in these circumstances can cause considerable 
erosion and redistribution of sediment. 

The wave regime ot Gulf St Vincent has been 
discussed briefly by Bye (197(0 and, more 
comprehensively and with partieulat reference to 
the Adelaide beaches, by Wynne tV al. (1984). I he 
configuration of the Gull is such thai, along the 
eastern coast, prevailing south westerly winds 
generate waves of maximum fetch south of 
Adelaide, Adelaide beaches are subiect to a tafWfil 
wave energy regime which generates significant 
alongshore transport of sand northwards towards 
lhe sludy area Execpi under storm conditions, wave 
energy from Port t.awlcr to flic top o\' the Gult is 
very low. 


The rote of vegetation in delerminiiHj the nature 
and distribution of sediments at Pon Gawlet is 
fundamental and is discussed m greatei detail below. 
In general the various eyanobacteriu, algae,, mangroves and other plants pet foi m tvvo 
significant functions, Firstly, they are the autotiopii- 
in food chains leading to organisms which secrete 
shells or tests of calcium carbonate, these Uiells 
and tests, either intact or comminuted, ultimately 
become part of the sediment. Secondly, in a variety 
of ways, ihey actually cause deposition of sediment 
trapping grains of calcium carbonate, quart/ and 
other mineral matter. 

rosidontu uttstrulis, the bioad leaf species, i-» a 
seagrass commonly known as "tape-weed". It is 
common in Gulf St Vincent, living from low tide 
level to depths of about 10 m. Pasuionta has tough, 
roughlv textured leaves that host a variety of 



epfphytes<\Vowrsky 1956). the flat ventral sill ; 

t%J (he m:;iiii lljlftl Nlibecuttitllf htiifi^.u ,,hliin! 

-! pDri OnwiCTi i, "«.n shows ^uiac like fcmpi bats of 

lit--. |i BVCS '.Va.vrsley A* Tlinn.r. | N76) 

u>iiMtieit*iI thai lew animals feed directly on tftC 

tatite ka'-cs. I he rhizomc/rooi system and 
shears of tlu> plain ate dense, fibrous and resistam 
'" -.'icanic decomposition. Pri\ifJomu UUMni'ts leaves 
an sui vivc unU lief periods of emergences! |gv» 
udes. Al Port Gawlcr it grows in i he outermost aro- 
i i the (and flat bi the lowei FmcrtJdal ;■■• h 
Tosieta mmeMeti ts another scajirass. of narrower 
leal ' '-nimoiilv referred W us "cet gtatf". Like 

Posukwita, (his plant has u iw o yxtem mud 
,i epiphytic algae and animals <Womcrsicy & 
I humus iy7f>» At Port Ciuwler, /■■tsferu, (oucther 
v.nli species flf Heterozosiem, leptfaeno and 
Ruppia (as described by Roberrsun WS4> t fotmv 
extensive, dense, scagrass meadowv seaward Of 1 
bare sand lint, and seaward ol. and adjac-iiii [g Lift 
i love woodland. Thesr seivgrasse*. can survive 
longer periods of emergence a' low tide. 

AvkwiWQ tfWWw/tS WW rcxhtifera is the Billy 
species of mangrove round in South Ahsi ialiu and 
oeeuis ac tern Crawlet in both marurc woodland and 
as younger colonising plants (fiuiler « oi i 
Harbison 1981; Burton I982tt! WWfc M»Wf I 
loteff 198?) Tidal distributaries form a dendritic 
pallet u through Oie mature woodland, St*.» wak-i 
fluvhe:; accumulated aalt from around tin: maiiurove 
POOtS fBllllCl W flf. W7) Mid disnibmes al 
sCHfir<iss and other organic llolsam. Much ••[ thk 
utean'L debits piobably provides nulrienl lor [hi 
niiintTjives ami \% il souiec ttf food for a vaticty of 
i ipodSi Mane, roves .nv ronlined in thcit 
rliMiibLiLion UftftMftttl* by the need loi then vertically 
protruding pneumaiophoi'.s lo be periodically 
CipOSCd in the air, add landwards by fll« need for 

tceuiat llmhlHg oi accumulated salt from the roots 
by sen water at high rtde (C hupman 1975), Juvenile 
mangroves axe colonising the />>sretu meadOM 
seawards of the kW* sand Hal ui Pon Claw let. 
Durlni the past decade we have observed itieti 
establishment pFQgrevuv^ southward- ovei sevcia! 
hundred metres from die older tftev. 

Mots of cyjinohacteru I b i en algae) 

colonise both ihc floor of the mangrove woodland 
andthemid ode bare sand lift] 1 "h/v atedull i fl 
in COlOUr ttfld slippery Mat*- are constructed at the 
sediment /water ntCtfaCC by one or more species of 

eyanobacu b > rrwry also contain a vtiriety of true 

bacteria and other algae. C'vunebaclci .al mat G 

,. mic the most productive Ol a<ju h - 

i/tiMlosyrulu-lH s : ^aerr*v (hVmld 19HIV \' 

.mm m oar-.-niiv feisty 

aropocK. Thdi cttvtrll I T hc open sand 

t,i «| ii d.jn«i.' L j by grazing gastrqpodG 
ieflwanfs, and b> the infrequeo^ and shoo duration 

61 hi.uli tidal inundation of the sediments rloscsi 
to the beach. 
liuiasuntu hafacn&MkFcL and Stirtvcortofa 

ifuiru/Ufffora are dominam plants ol the lamphlre 
•ill community. These arc low-growing, 
lleshy plants which have considerable salt lolctancv 
and occupy the /one inimediatcly landwards of the 

Srftinicnlalidn at Purl (iawler 

t. Posidonia-Pmna Facks/Zom 

At low tide it is possible to obset ve (lie seaward 
margin of the sand Hat. PtmrJania austrahs 
crows in patches 20-*0 tu : in area. Piflflfl 
bicoior\ P doluhrata and Suhifopirtna vtry_am\ 
IHuttei & Hiewster 1979), known locallv ., 
,4 ra/or fish" is conspicuously abundant, giowmp 
in . niton £ the scaj:iass. The sediment is coai\e. 
poorly sorted, shelly sand. Carbonate contem 
i-. ^ic-atei than 90%, much coming from linw- 
vecrenng organisms liosted by the Postrfont'a. 
Inraminitcra are pai tieularly plentiful amonj' 
the smaller grains; among the larger shelly 
ureai nuns, fi btCotw may grow to a leogtfl 
20 cm in a single year {Butler & Brewster 1479) 

Patches ol PostdanM on the outermost areas 
can be abfieived in various stages of burial by 

no^fle sand (I tg. JA) the banting action 

ol the fieagrass ar-parently 'raps the s<diment- 
Studics in the Bahamas by Scoflin (1970) and 
Neumann €1 aL (1970) show diat BCagr^fi^a can 
reduce water velocity ftorn ?0 cm sec ' 
(sullicicut to transport loose sand grtti ii 

the bare sea floor) down to /eio at the 
-seduuent/ water interface As ihe sedlmsnl 
^..ciiniulaf.cs, Pn.vfUmiti giows upwaids. but u 
constrained by its inabiliiy to survive low iilIl 
emerptjnee and it uldninn-U dies, laipcareas of 
the remains of receruly dead BWgr^Mt !._>^eihei 
with numerous emply fl&Wff Bhfllj, valves ^apiny 
and in life position, may be observed 

Mediately landwards of the present day lov 

wafer shore line (1 7 \g 3B). 

( arbonate sand is ihcrelbrc Hcuntulaoni: up 
to low tide level. c"Hi\iri? tUtt Oi'ici tfUTgifl | ll IJlC 
sand flat 10 plCgretde seawards. In some areas. 
after death of POsidOffiO, the uppermost 
sediment remains bate oi vegetattOfl aild 
consequently .abject to tidal and storm W&V4 
transpotT Such surtaxes become sLomed. 
chajtnellcd and rippled as the shellv sand \n 
redistrihuted according 10 coci i .lious. 

Iy ( however, tti.. si-.l-mein mass icmaiiLi 

i . ,, ccd by veagrww nhre, 




. ' . 

V ' ^>***£ 

\ ■ 

fig, 3A. Patch of partly buried, living Posidania, adjacent to remains of recently dead Posidonia; outermost sand rial. 
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 10 
fragment. Outermost sandflat. D. Dense, fibrous remains of dead seagrass (Zostera) after death and loss of foliage; 
section exposed in tidal channel; wrisl watch for scale is 22 cm long. E. Recently dead KatelysUi {\-A ), valves attached, 
convex upwards; bare surface adjacent to Zostera. V. Zostera in the seagrass meadow. 


]. II . i ANN & V A COM IN 

liv largi ill ti i d Wfl«d si'ciK, ai .' 

numerous ^:«iiu fiilv*-'.:'! Idled >IilII gf|| 

AIjmI 1'iov.iii cm the subsume helps to mini mist.' 
a .iii- i'v iiu;i!,ui!L''ji-- acid titana waves. Whs 
!■■■.. ikterabU prom Itfc " "■ -' ll * 

sand (Slllhlg Hi'- ; .:■■!-■ 

In the mid tidal area, the sand flat is colonized 
by n scagniss meadow, ponsistrng ttf several 
specie I rhhikri predominates 

(lu -I i |*bc dominant Nwlve mollis 

Ktttetysto so trip. 2E>. a:, scaktrina has been 

recorded from Port Gawtcr by Tutdf <I98?K but 
Uina species of Ratetym may be present 
|l udhtook 1WU). Di u'»J valves ol 

ftlc v r . have been widely redistributed 

ilit.MiL'hini Ihr study area 

L ike Posidot m. Zosferu battles and traps 
UMllfflC HI 1 U nucl i ida; area is less mfli:^ 

ill juJ tidal cm rents, m< gT£ 
is etrm>pomhrig!y MnJIer and [here is a 
irffcarri mud component Sediment is rich in 
uigauic luajtet, supporting bacterial reduction 
Of' BUlphatc "iiiv from sea water, below the 
setHmeiu/watet Interface It is Mack and has a 

• ronj sulphide odour (Tip,. 4A). Burrowing by 
small crabs Lends CO extend to the under Ivhie., 

grain supported sand ol the Positfonki-P Bioluibadou lends In, in pail, o\id.. 
Immoyemse ihe jUgUltt 

Aggradation of sediment in the /< 
ncadow continues to a level determined bj ihe 

period Of low tide emergence. I'Inmalely ihe 

plonfs on itu- landward side rftiic meadow are 
inadequately watered ar high tides and they die. 

On llieseawaid side, the meadow adVaW ■ - 
, |e old f'ttsiiloniu Pintw sulistta.r, and Ihe IlltCl 
sediment tacies thus pnoerades 

} MoTiwvf I tone 

\ Port Gawlcr wv have observed the 
...iIhhimdl' advance ot (uvea le mangrove*, 
tiuih vai. re\br< rtii 

•stem meadow (Fig jb>, 

t| ES evident thai the finer, muddier and 

i ntorich sediment crapped by tbb iea 
a ideal substrate foi Mangrove-cotonbatfoti. 
therefore represent a third major 

i lai i sediment uiceessidn 

Vbut!g mangrove tree* quickly establish a 

lateral root sysietn, earn WM bearing a series 
of ve- i ■ . ii umaiophorcs. bach 

Igti poi atlme 
both m^njnnvrvim .-.-■ exist butaslhc 

density Of pneumatophores' increases, so loo 
does the amount ol organic flotsam that ll 
tupped by, and adheres (o, these subacnal roots. 
I tie /astern thus dies under a blanket of 
accumulating marine compost. 

line sedimcnl continues to accumulate in this 
fades as the mangrove woodland grows to 
matuiity. Algal and seagrass debris bring 
epiphytic carbonate organisms which remain as 
grains in the sediment. Ihus aggradation 
proceeds to levels where Ihe floor ol the mature 
mangrove woodland (Fig. 4G) is inundated for 
only brief intervals during high tides. The 
sediment/water interface is then colonised and 
stabilised by a eyanobaelerial mat which 
Introduces new deposition^ processes (discussed 
belowX TJtC suh-tr.uc is intensively burrowed by 
the small mud crab, Hetice haswe/lianus, 
facilitating sea water permeability and aeration 
The mangrove sediment thus becomes oxidised 
and homogenised. 

Throughout tins episode of deposition a 
dendritic patlern of tidal distributaries develops 
and Is maintained. 

4. Tidal distnfmtary Jack's 

Tidal water movement is concentrated along, 
djstnbuiarv channels which aie best developed 
within the mangrove woodland. As a 
consequence ol Ihe dendnlic pattern Ol 
channels, current speeds are variable and 
sediment type varies accordingly. Major, shallow 
channels have coarse, shelly debris, rich in valves 
Ol kafelysiu and Anapellu (fig. 4F), 
disarticulated and convex upwards. Dcepci 
backwater channels have a high component of 
black sulphide mud. 

A major component of the shelly fauna within 
the mangrove Lidal distributaries is the small 
furrifonn gastropod licit Maria (/.eavumuntus} 
ihemi'ncnst.s (Ludhrook IV84). Where channels 
nicer the open sand Hat, shells of this gastropod 
are numerous and current aligned (Fig 4F), 
have been widely redistributed throughout 
the study area. 

5. CyatlObactertai Mat Satul hlat factes'/one 

Bauld (1981) has illustrated the ability of 
cvauobacierial mats to fix sediment in Speiicei 
Gulf. He describes how entangled trichomes of 
Mtcroco/eus sp., and their enveloping 
mucilaginous sheaths, trap and hind sediment 
in a thin, coherent layer. In addition, pholo 
synthesis ai the mat surface removes carbon 
dioxide from sea water, increasing pH and 
favouring precipitation of calcium carbonate. 



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 lo a tidal distributary. D. Tidal distribuiary with mature mangrove woodland, left, and 
inner sandt'Iai with cyanobacterial mat, right. E. Gastropod shells, mostly HatiUaria, 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 Katelysia and Anapelta are disarticulated and convex 


I II <_ WN 6c V A- OOSIIN 

( mats cover a large part of the 
otherwise unvegetatcd sand flaL the floor at the 
msiltSFOve woodland and some b£ the supruiidaJ 
areas. Ihese arc therefore sites .d vedimou 
aggradation. Sediments bound by eyanobaetcnal 
mats arc known to whhsiand currenl velocities 
up to five time* higher than ihose requited 10 
erode mat free sediment (Neumann et a/, 19701 

Sections cut through the active mat (Figs, SC) 
reveal a thin layer ol oxygenated sediment, beiov. 
which 15 black., sulphide-rich, laminated, 'silly 
sand (Fig. 5D>. Anaerobic bacteria, nourished 
by the organic mallei ol previous layers ol the 
mat. reduce sea water sulphate ions. Meuls, 
particularly iron, are fixed in this way. Black 
icdinienl turns to a pale rust colottr w 
piolongcd exposure to the ail. Calcium 
carbonate content may be as high av 70%. 

The cyanobaclerial mats suppotl a dense 
population of grazing gastropods, notably 
Safinafor sp. On the seaward edge of the bate 
sand Hat, persistent grazing appear-. io inhthii 
establishment of cyanobactena (Fig. 5IJ). In lllfc 
area, large numbers of the small bhmfvf 
■\napetla up., piobably A. cyiludca (Ludhtook 
1 0x4). live clustered together immediately below 
the sand sui taee (big. 5A). Disarticulated valves 
of Anapei/a are widely redistributed in sediments 

of the studv area 

1 he bare sand flat is sharply separated ffOtn 
a he Zosterv meadow to Hie wesl, ;tni\ the 
mangrove woodland to the south (Fig- 4D). by 
iriUHidal disrribumries. These (idal channels 
(WtVfi remained essentially unchanged for the past 

Hiyh ude bcaeh sediment at Pari Gawler has 
a calcium '..irbonate eontcm of about 80%. The 
texture is predominantly sandy, bui shells 
originating in all of rJic other lacics occur as 
constituents. Wide variation in grain si/e, from 
shell gravel to find sand, may be related io 
variable wave energy, which is a ft motion ol 
water depth and wind strength. Sections _ui 
through ihc Beach reveal horizontal io geiuU 
idled, well-sorted laminae, and, rarely, hi&hca 
auelc cross beds. 

I lotsam seagrass debris appears to stgr.i- 
n- . ."tijy a'-siM stabilization of beach vdiuienl 'o 
seaward, while sahbush and other plants SW 
established in, and fix, the upper beach sands 
(I jg 5 1£ V file high tide beach is therefore a mass 
of accumulating sediment, progradiuu seawards 
arrOM i he uiner sand flat. 

7. Dune Juaes 

Windblown carbonate sands occur some 
.50-100 m inland of the beach. This area has 
been considerably modified by human aciivily, 
but the tow dune lonns, stabilized l>> sahbush 
and oilier vegetation, are clearly identifiable The 
sediment is well sorted and consists ol line 
broken shell and entire small gastropod-, and 
iorammilera. Sections cut through the dune- 
reveal coarser, underlying sediments. 

H Storm ridge Jam's 

Storm ridges are formed at limes ol e.\trcmctv 
high tides and local storm activity "shelly 
material is down shorewatds under these 
conditions, and may pile up in a ridge nJ coarse 
poorly sorted shell debris, somewhat paralleling 
the shore line, bin seawards o) the beach, If the 
tidge is continuous, chat part of the sand flftl 
between the newly termed storm ridge and the 
prc-cxisimg beach is isulatcd from futibei wave 
action \ new beach facies is established on the 
ififlWard side of the storm ridge, and a new dune 
lactes to landwards. 

Al northern Port Gawler, extensive storm 
ridges cincc existed in ihe area between the tall 
evaporation pans and ihc present high tide beach 
and dune facies. Shell-grit mining has effective!: 
removed mosr of this sediment, the location of 
the ridges now being marked by a number of 
parallel, elongate., shallow depressions. 

immediately south of the Gawler River, a 
storm ndge some 1.5 km long and up to Z m 
high lies within the mangrove woodland (fig 6). 
H is situated several hundred metres landwards 
oi t and parallel to, the seaward margin of thl 
maneroves. The seaward side of the ticIbc is 
straight and steep, suggesting Ihc form o\' a 
stranded beach ( Tot elf 1983). On this suiface 
Harbison (pers. comm ) has observed nunicinus 
large, square-eut pieces of timber, >imila» \o 
ihose used in the construction iy( early South 
Australian jellies These ite-ms of drill wood were 
evidently elApbced at the time o(, oj ahortfe 
following, formation of the storm ridge. 

Alone the i nastii! plain, north of Port Gawler. 
numerous ninte oi less parallel storm nclgcs may 
he observed. 

L ' 'snprartdul lugoofl fades 

Areas of sand flats thai have been isolaied 
from op<-n marine influence may still receive %ea 
.I vi, i n,|,|| distributaries. Alternatively, water 
m .:, Iv supplied by seasonal hojvy run or rivnv: 
id water, Ofl the floors ol diese ,upr:ni.Jal 
lagoons, Dl,. '.yaii'-r-aclcriat maixedten 
flourish They maintain low diversity 



Fig. 5A. Dead Anapella bivalves ( x 1) in life position, immediately beneath surface of inner sand flat; 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 SC, Uppermost laminae are flushed with photo synthetic oxygen; dark lower 
laminae contain sulphide from bacterial reduction of seawater sulphate. E. Inner sandflat and vegetated high tide 
beach R Mature mangrove trees overhang the Gawler River estuary near the ruins of Lisbon Wharf. 



Fig. 6. Aerial photograph of the Gawler River estuary showing the locations of Lisbon Wharf ruins and the site 
of vibroeore 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 
foraminifcr Elphidium 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 

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 

10. Est uarine 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 vibroeore VC 138 and are discussed further 

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. 






_--** — ^*- v *.T* -• ' <v> ui,.*,.'. ••-■•-•-.- ■.'••i> > ^ 

^^ ^ ~* •* <* r : - v>>^rTJ 


.. ROAD 





♦ V 

v y £*, MANGROVE 

* v Y %>, FORE?T 

« ., '. '-..Ill -^ 

> •*",«* 


*/«ft3 f^f^rp 






i^Sj Zostero 

/'=?> ■ sg »^ 'l Posidonla 



Z7p Terrestrial sand and cloy 


i lg. 7. Schematic block diagram illustrating modern deposit ional environnn-in _, .mil ilIjioJ sedimentary faeies of 
i M Kilda formation at Port Oawler. Sites of vibrocores VC [3fi and VC 138 aic indicated 

TAHLb I. Summary Qf the set! i men ton Jot us uf Port Gawfoi 



Sediment type 

Posidoma — Pinna 


Tidal disttibutary 


sand fl;ii 


Stornq lidgf 

Supratidal lajioon 
or sabkha 

Mibtidal, outermost sand flat 

intcrtidal sand flat 

intcrtidal mangrove woodland 

throuphoui the tidal range 

high tide inner sand Pat 

high tide 

supratidal, back of beach 


supratidal. samphire 

Cawler River estuary and 
i"u k waters 

sarbonaic quartz shelly sand? 

muddy carbonate quartz sandy 

carbonate quartz silts, muds and sands with organic dbbril 

and cyanobacterial mats 

channel deposits with convex upwards disarticulated bivalves 

and current aligned gastropod shells 

line carbonale quart? sand bound by cyanobacterial mats 

shelly carbonate quartz sands and shell gravels 

well sorted carbonate sand, reworked from beach and othei 


coarse comminuted shell; lag deposits of this laeies undo he 

present day sand flat sediments 

gypsiferous carbonate quartz muds, silts and sands 

peat, orcanie mud and related s^aimem- 



In processing the laboratory wttnplav the high 
organic content made tl diiTieuh to determine the 
number of lorannnifcia with PCSpeCt 10 a •''• iw 
of sediment and this Imil- of invcsUga) is not 

pursued However, all sample* contain j suilicicnt 
numbers oi lotantmifera for relative p«ctent«gB 

After boiling m liesti watci for about an r I 
in break down the organic debris, samples 
sieved and sedinictn oJ sand ii " :"' 00 if- 1 mm) 
was retained. I orannuilcra were concentrated from 
these sand Tract inns by Flotation on 
tctrabiomoethanc. Residues wen: inspected 10 
ensure thai, most tests had been separated. The dried 
foraminilcral concentrates were further ,ie\id into 
phi grain size fractions lor microscopic 
examination. For radii wimple, pereentaee 
abundances of toraminifera specks were (tetcrmmod 
for the fraction. MXMfcSO mm and SO J5 nun 

Analysis ol these fractions only, rathei thai 
entire sample, has a number ol important 

Fllison (1951) has commented on the (i 
consuming nature ol quantitative foraminifcrnJ 
analysis in palaeoecolocical \ind .• I be wot'* 
repotted hoc was part ol n mi rtWC 

investigation involving several hundred surfai 
sediment and COW SOmfJcS It lYOS important I 

these sample* be processed effli d) Coarser 
fractions require lower powers of magnification, 
greatly facilitating identiRcaiipn and tin of 

individual specimens. 

In finer fractions, juvenile nidi. ida.i.s constitute 
a high percentage of I he loranimii-.ia. Sfli 
identification is more difficult Km 
particularly foi miliolid genera Stlmiiker 0967) 

observed lhat tests ol juvenile THlOCtfUtta i'i>n»ir,wi.>. 
asexually produced in laboratory cultures, diliercd 
significantly in fotm from thai of the parent. It is 
well known that some species of THtotlttlttG .1 
exhibit quiuqueloculine coiling as (uvcuilcs 
becoming triloculmc onlv as adults (c e., loe 
& t&ppan 1964), Such changes In form a** Fin 
com pbca i cd in microsphfirfc/mccalosphcric 


In natural sedimentary systems, lesb. ;-f -mallet 
foraminifcia arc easils winnowed I mm ilwir 
environments or origin to be deposited rfwwhcrc 

Msn, during iiUraenvituiuuciiial UaittpO I i lalkT, 
more Iragile spa u-sare mnr L prone l^aiUllfUfl Utd 
mechanical destruction than laracr, more rcihu«t 
forms. Finally, when cttfl&Ufc pics, pott 

diagencsis preservation will favQUl Iftftyei tomu, less 
prone to solution. In llusMudv, specie- •-•) Miuillcr 
genera sudj a- BvMniinakla { are consider.. 

■< arm «J ftlaia 

gtrwi . i-.v value aa environmental indicator* 

I larger genera ;,h as Peneropiis. 
In the Mudi. nstwenty-cipht species 

;-i kwaminifcra were observed The percentage 

numerical dtstnlMtii-.-ns of the more common 
species, with reference to the sediment a > 
euviionmenis ot Port Gawler. arc shown in fag. H. 
Micse Species are illustrated by scanning, electron 
photomicrographs in 9 and 10. 

Systematic notes on selected species 
of Fofaminifcni 

Nuhrcuiariu htfi/Ugti e cvIp.uicK vaiiablc in 
morphology, ics shape pftefl influenced by the 

t to w kidi it Adheres, Some form* are piano 

convex, tie Mat surface usually incomplete and 
revealing a planispinsl arrangement ol ehauibcis 
A jucat many individuals nave globular, twisted, 
tube-like UiSIS, with multiple apertures and were 
presumably unattached. Uolh attached and 
unattached forma STe recognised here as the same 

Peneropfh phtrrtitus forms rhirt, translucent. 
i n deeper water, but in warm, 

intenidal areas where salinity uses with high rates 
of evaporation, tCStfl are i link ami often aberrant. 
The n*nge ol' morphology lhat mav be I ibaOl ved in 
such populations Of Pnwmplis has been well 
illustrated by Sellier de Civrieux (1970) lot the 
Mediterranean and by H fi r larked KaIJ.(lW3) 

lor the IVisian thtlf. Mimiar Unnis OCCttl ill Port 
Gawler sediment and are considered here as a single 

Spiralomlitm spp. her:' meliales ,S. tJttttllurum add 
also form*) having ctl&fnbc i I Biorc quadrate 
scciion, tacking the numerous oblique costaeoi v 
annUurunL but besting distinct longitudinal rjdtgJ 
a? tnc edges . '.is, and often a thud, 

parallel, and I tt ol the Hat oenphcral 

ro rctei rarUe to S. tnCQSty 
1 5 I man & Todd, 1944 Some individuals have 

been observed to have features int bei ween 

those definitive ot thpse two spi 
Qiscorbto dlmfdiarm rs here used essenii<illy in 
..-use ut Medley w oi (!%"( i hey recognised 

ihai t'or-ris basing great I j OtOTphoto^y, such 

pcrtpdiei v. in: h mi I..-.V spires lari-c 

ur small win:.. . tly belong to a single 

specrev. All ul ibeu illustrated bums i:a.-. beCtl 

i:i-,; din I;;' I'mti i; i v. n - m itcfiaJ, and nbea 

spet ie.ii' are esGamlned ac one nine, it is 

apparent thai Vauittlidvi e. eoiilLnUuftlS. 

( \;?hiitiutv. tttarr i tl | EalJQB foi 

cans, di-so i ■■. i ■ i' -. 'ike lenin. 

iVttlt nunKU'us ip Hflatcd chambers. 

Siilui-.-. .-a. r&i&lfd 'nnbale. loiucd b- 





SfTr oloculina spp __ TZrr= 

.:" AWAR :■_. 




in < 
o lu 




so,- 1 







Fig. 8. Foramini feral biofacies: relative abundance ot selected species 
environments. Port Gawler, South Australia. 

through modern, adjacent sedimentary 


I 11. (ANN \ V A l-OI 

. slender relral processes. There is a promuieu, 
imperforate umbilical boss which is neither raised 
ru.r depressed The intwiotn^rginal aperture \% 
obscured i'v pustulfti carbonate f*w ,_ >ii thcapcrtuiai 

Ilie periphery bears a low narrow keel. Till 
sui face ia uniformly white and (t!os v 

i big sp< t -l".., which i*. COBUPOn m Smith 
Australian *u!f waters, has -only recently been 
described (MfcCutloctt 1981). h i> liU-ly th.< n. 
previous records of Australian Holoccnc 

FbJWiiniTem. \\ i as Efpfikiiiwi 

adwflutn Cushman I92i c u^hman n 
originally described E trtftift 
umbilical region, Etod lacking any significant boss 
or similaj material, However, in subsequent 
(1939) in.- rcfci red terms having a rhomboid section 

and il'i'.li umbilical plug (eg Urudv 1884, IM. C X. 
I'ijts. a, b) to the species. Australian workers (c? 
Chapman 1941; Parr 1945; Colli** 1*741 >"■ 

followed this later interpretation. 

Vella (1957) manna lied ;t d-lii.ii-.:-). between 
forms having a depressed iimbi h Email 

. pira! bossol clear shell maic!taL,atid those Iiimiis 
having the umbilicus covered b 
plu>;s no! pfOtrlldlng he ■■• I thi i 

for t he laiu-r iorri Veil* 1192 I i I 
LJpinttiwtoniun rhurlot; JW, I appears closely 

similar to the species I mured by Brady, cited jbove, 

McCultoch (is>«n also bellevi i simians 
i, in -r interpretation nf Fipfwlmm wtxtoum Included 
more than imu speuusand accordingly established 
Elphtdlum wcWif described above, Her 

descriptions a id figures agtoewUhsrp&iesoccu 

in the Holncei i ol Porl O&wfer 

Apiho'pA Eiphitftim fc (198tt, pL 26, 

" ■ 
and hci £ tfrfi mn [IWX pi. 26, flfr 10) may be a 
[tiven ll< "i Ik spec™ •.. 

ftirt Qawtee ttaravflna W IM ami \t 13ft 

v i i the rite? indicated *n 

Fifi, 7, VC 13* On (he open inuei titfid Hal ttfyc 

138 in the mangrove woodland, adjacent to ihe 
Claw lei River estuary. Both cores peneuau 
3 metres of sediment. 

In the laboratory) tbti cow wet *pti' lengthwise 
and litholoflical loss prepared The<e are pre.v. 
as Flgfi. 11 and 12. Prom one half, sapipk 
approximately 1 00 ml ol sediment were taken at 

m intervals nrul processed -in describe* $h\ 
Foi each wmpte, percentage abundance- 
k'u.niinifcra were determined for the stoe tractions 

1,0*3-0.50 mm and 50-0 25 mm. The percentage 
distributions it more common species wit: 

determined, these data arc shown down core for 
tbC productfw ! nsidcred as Si Ktlda 

I nation. In FlgSi i.* and 14. 

Each i'*"v represents a sequence ol near shoo. 
marine environments, recording both the initial 
i i.-cene transgression and some ol the taier 
episodes Of sedimentation that followed sea level 
lily Iln down core roramtnileral assemblages 
are, at least in part, indicative pf I bote 
environments, and may be compared with the 

loiauiimieial data obtained lor the various feocs 

ribed aboti 

I lie iciiability of the data was initially tested by 

comparing two samples from the cyanobatteria] 

laeies. ihe fust liom the surface transect, the 
StfCOnd from the top of VC 136. The sample 
localities were about 500 metres distant from each 
other on the mid-inner open sand flat 

Fig. 15 illustrates tins comparison m the form of 
histograms. The data fat the si/.c traction:-. 
;>.M)-0.25 mm arc remarkably similar. That for Ihe 
coarser fractfottS arc less CSOnviudPfc pcrhap> 
reflecting the Smaller numl -i •! specimens counted 
fel thai $Ut range. The number ol individuals 
lexxxvcicd from the fraction J. 00-0. 50 mm were SO 
and 123 for the transecr and top Of VC 136 
reNpec-iively; equivalcnl numbers t<u the finer 
fraction were 211 and 7J7. 

tor the seduneni si/e fraction O.sO-0.25 mm. 
where the number o( individuals composing lh 
haehtMi is approximately 200 or greater, the abundance of for;iminiIei;i species h 
considered to be a reliable environmental indicator 

Results ami inlerprcluiion ol* cores 

t r \ f<S 
The earliest sodimentsof the St Kikla Fortnation 
jed m wbio<-oie VC 136 represent the 
supraridfll faciei ol th< samphire /.on^t. rig.. 16 

'.ompares Hie pcteenlage abundance of specirs en 
. ainifera frOWl the modem samphire with that 

observc<i iit 5(1 cm in VC 136. The overlying coarse. 

shelly sand, sampled at 25 em, is interpreted a.s 

remruint -.loini driven shell debris. Although no 

foraminileral data were determined for an actual 
i n ridce* 'he percentage aburahmee Of species 

obtained tor the high tide beach, developed on the 
_.ul side of a low storm ridge, is e.ssentialiv 
.11 to thai at 25 cm in VC 136. The tup sample 

of VC 136 has been discussed above. 

lit: • ffata iM.iiiij.ini ll 661 wlimina ttuxto Cuithinan 1927 1 t, »'>; 

h. kM\ \.:,' ■ i-hJiii,' imtfi.yu Defntiici I&25 I j 40. SpiroJuokfiHa smliUmim lt>rbtfii) \HV9 li I. 5! 
ishmanA J.*'d.J 1944m iu*60. Trifavulim it\fhw (ft en; R2A o-q ■*>(». Mtltoflnella 
■ - * r r. .MO. Pcneroptis vfanu/Hs fflchicj A Moll) 1798 I 





_- \ ."^ 



Yellowish urey, mix«d quart; -r.aibonate, well aOrted, line sand 

Greyish yelluw, poorly sorted, coarse shell deons 
with some tine quartz sand. 

Dark greenish grey bioturbated mud with poorly sorted, 
muddy, shell debris and tine auaru sand 

Dark yellowish brown, indistinctly mottled, micaceous, sandy silt 
Silty, very tine, quartz sand 

Mean gram size Increases down care lo 

very coarse , angular, quartz sand with quartz pebbles 

Dark yellowish brown clay. 

Riiddish brown, stift, sandy mud 



Cyanobacterinl sandflat 
Storm deUna siiureld- e 

Alluvial flood pfBtn 

River channel 
Aituvirtl Hood plain 

Penetrated 294 cm 
Recover ttd 2 72 cm 

fig. II. Descriptive anU interpretive log of Vibmcore VI 1 30. 

VJbfOOOie VC 136 thus records the Holoeene 
transgression as initially marginal marine supralidaf 
sediments, overlain by storm debris, remnant of the 
episodes of storm ridge formation. The uppermost 
sediments are the result of aggradation on 
eyauobactcrial mats. 

VC 138 

The proximity of die site ol vibroeorc VC 138 to 
the (iawler River estuary suggests that sediments 
peneiraled by this core would have been deposited 
under eondilions of fluctuating salinity. However, 
ndal channels 2 km and 2.5 km south-east of Port 
Gawlcr probably indicate former sites of 
debouchment ol the Gawler River The present 
course of* the esluary has apparently developed only 
in later Holoccnc. 

Although only 51 individuals arc recorded at 
175 cm in VC 138, for the size liaction 
0. 30 -0,25 mm, the percentage dislribulion o\' 
loramimfcru species agrees closely with data 
established for supra tidal samphire .sediments 
(I ig Ifi) I In: distribution of species at 125 cm is 
very similar to thai obsei ved for die high tide beach 
and apparently indicates storm sediment. Thus the 
traflSglt&Slve sequences of VC 138 and VC I "U> are 
essentially the same 

From JO0 cm to 75 cm the increase in numbers 
of Spiroloailina spp,, MiliolmelUt luhtosa, Discorhis 
(iimkliatus and Llphidiunt ensptun, with deeieasiriu 
numbers of Penmptis pUittattts, is consistent with 
the development of scagiasscs (Pig. 17). i he 
increase in Ammcmh tmxarii, <\ noted poiyhaline 
species (e.g, Murray 1971), suggests increased 
influence of fresh water from the Gawlcr River. 

A dramatic change in the foramiiu feral 
assemblage occurs at 50 cm, as Trachammina 
iriflafu numbers increase trom almost zero to met 
70% and most other species disappear. 
Trochammintt inflaia is well established in the 
literature as an estuarinc dweller (e.g. Collins JS>74>. 
Aplhorpe (19K0) reported the speeies from low 
salinity waters of the Gippsland Lakes, Victoria. 
Cacti (1084) has found it to be a significant 
component of the fauna of the upper Onkapaiinga 
River estuary, south of Adelaide. South Australia 
The abundance of Trochummimi tnjlutu. reaching 
[OOto) o)' the fraction 0500 25 mm at 25 cm, 
mai ks the development of the Gawlcr River estuary 
at its present site. 

Finally, reappearance of most species in the 
uppermost sediment of the estuarine mangrove 
woodland suggests a return to conditions of 
tolerable salinity. If this is so, ihcrc may have been 

Mg 10- OtMoiht.'r ditiiiiJuji'us tParkvi & Jones) 1K(>2 a c, -AS; d, • 350, Ammonia heccarii (Untie) 1758 p-fc - 120. 
h, • i.uio. Bjptwfmm arttcufafum (d'Ortrigny) tS3 l > k i, .. 140, m, 1300. Bipkidhim crixpum (Linat) I75fl I I 

• 70. Klfhuimtn rtmtt'fiijormc MeCulioeh 19SI i\ o, > 60 lYkhohiutus trop'n US (Collins) 1958 p-r, <>5; s . 650. 

|Aiiiu>nfh I tropivu* abundance la lav> lltaa j*b It>rvu6h0tli both VC 136 and VC n*. h i, present In Mgnificatn 
numbers In the surface sediment ol tin* emwimc mangrove ( tjrta A* De P&CMcCr{19$l) reported 'his species 
hum ;i saline lake.| 



< 200 




n: 250 


Yellowlah brown, mottled, organic tlch 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 ahelts 

Live mangrove roots and ?sea grass roots 

Medium grey, poorly sorted, coarse shell and shell sand 
Disarticulated Katelysia. 

Medium grey, bloturbated (?crab burrows), tine, sandy, quartz sill. 
Live mangrove roots. 
Carbonate nodules. 

Light olive grey, micaceous, poorly sorted. 

very tine, silty, quartz sand. 

Carbonate cemented, tine quartz sand. 
Yellowish grey, moderately sorted, medium, quartz sand- 
Carbonate nodules and ?burrows. 

Brownish grey to reddish brown, 
stitf, sandy mud 


Estuarine mangrove woodland. 

Sea grass meadow sandllat. 

Storm debris shoretace. 

Alluvial flood plain 

Penetrated 308 cm 
Recovered 274 cm 

River channel 
Atluvial flood plain. 

. A ■ ■■_! |. ,: i .*>, 

Fig. 12. Descriptive and interpretive log ot Vibrocore VC 138. 

a real decrease in fresh water debouchment due lo 
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 
loraminifera 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 
/onation 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 oi' 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 trees coalesce with 
the existing mangrove woodland, and with 






\ :, 

D -, 

\ j 




























































































J , H . C . 

198 5 

Fig. 13. Percentage distribution of selected species of foraminifera down core #VC 136, open inner sand flat, Port 
Gawler, South Australia. 

Fig. 14. Percentage distribution of selected species of (oraminifera down core #VC 138, estuarine mangrove woodland, 
Port Gawler, South Australia. 




- 2 cm 

m £ q, 

B => E 

a °- fc 

~ a> o 

"V 'Z ~ 

•o E w 

* 5 E 


0-50 - 0.25 mi 

1.00 - 0.50 mm 

Fig. 15. Comparative percentage distributions of species 
of foraminifera for two samples from the inner open 
sandfiat. N refers to the number of individuals 
constituting the size fraction. 











































































Ui , 


N = 





N = 522 


50 - 52cm 

continued sedimentation, the northern area will 
come to resemble the rest of the tide channelled 
samphire and mangrove coastal zone. N = 289 

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 5. 
with various plant communities. 

4. The sediments of most facies are rich in calcium 
carbonate, up to 90% for the shelly sands of the 6. 
Posidonia seagrass facies. 



0.50 - 0.25 mm 

J.H.C. 1985 

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 sandfiat (Fig. 
13) reflects the landward sites of deposition and 
similarity of sedimentary processes. 

Distinctive, though sometimes subtle, changes 
in the percentage abundance of foramini feral 
species occur from one facies to another. 
Reliable comparisons of foraminiferal biofacies 
can be achieved using the size fraction 



N - 675 

76 -77 cm 

N: 1B0fl 



- D £ a e 








H 1 

-5 E N=I3CJ1 


50 - Q.2$ mm 

flff. I ■'. I Viventage distribution ot species r>| forttUtulfctfl 
(in siulihr vmiplrs from sea j«rasy meadows With a 
sample from vjppucone VC 138. < >nly the size fraction 

0.50 0.25 mm is considered. N refers to the nurnhei ot 

Individuals consuiuthuj ihe size fraction 

0.50-0.25 mm of floated foramiuileial 


Down core documental ton of relative 

abundances of foraminifcrai species, in 

conjunction with lithotogical studies, can he 

used to determine palacoenvironmeiils and thus 

l he sedimentary history. 


I he Marine Sciences and Technologies Granls 
Scheme provided financial support for the Coring 
programme The Council of the South Australian 
College of Advanced Education granted 
professional leave to John Cann for the preparation 
of the paper. Sharon Pro feres drafted figures I. 7 
and 8; Francis Gorostiaga assisted technically id 
recovering the vibrocores; Brenton Bowman 
processed samples fol lorarniniferul analysis; 
Richard Barrett prepared final photographic plates; 
Chris Moore typed the final manuscript. Some of 
the research reported here was completed by John 
Cann as part of a Ph.D. programme at the 
University ol Adelaide under the supervision of Dr 
B. McCtownm, We thank Dr B. MeGowran, Dr 
A. P. Belperio, Mr M. Lindsay and Professor 
H. B. S. Wouierslev for enlically reading the 


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by David M. Spratt 


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. 


by David M, Spratt* 


SPRAJT, l> M. (r985) RcdcsCripMom oFl*0 IfiCbWid nematode parasites of vertebrates iti Australia and 
Ftyua New OUJoca trans >■ Sm S I//./. Mivdi 143-150, ,29 .Ntfvembei tmms 

Anderson &. Bain, 1982 placed PartllrtchoSQm / fclhford & Muller, 1978 as a synonym of Cu^tlutiu 
fyemw /#//<) Zcdei. 1800 The reasons fat this synonymy arc discussed herein and C\t;»tUtrm cfQCOdikK 
I Asltibrd & Muller, 1978? comb. nov. from tunnels n 1 1 1 iu CpW«3 ntlS rjl Cto ffl^v'itt riovotyuinttte rwutesumrw 
Schmidt m PjpuA New Guinea is redescribed from co-type specimens. 

l>icM08omotifa& rwtolto ttfocva A. Auri?!- l%I i* described and illustrated liom tin cavity erf 
kaitus fusci(H", tWatct house) in southeastern Ncv, SoUtVl WaJes I niv represents the first nxord ot the pnrnsiie 
in Australia and /?. ,fitsdfi0$ ks I lie only mun.l "iom known <o dam. 

K.t-v Wokos. C'amtrkhosotnu. Capilbriu, th, hn\otrtoit1r\ . Nenuioda. distribution, hovts, morpHology. 


The Trichutidae ts a cosmopolitan family of 
parasitic nematodes which arc poorly understood* 
boih morphologically and biologically. During 
sine!) ot the uichurids o\' Australasian vertebrates 
the need arose to re-examine two unusual species, 
one described originally as Purainchosonta 
crocodtlus , AsMord <& Muller, 1978 from Crocodvtus 
novaegu'meae ftovaeguineac Schmidt at Moitaka 
crocodile farm, Poit Moresby, and Trie hoso mo ides 

nasalis Biooca 8 AurWi t%i i »rsi reported from 

ttuttus nQWegicttS (BcrkcnhonO in Rome (AtiHxi, 
1 95 H ) and not known to occur in rats in Australia. 
Rcdescriptiou ol' these species and consideration oi 
their taxonomic placement form the basis of this 

Materials and Methods 

Nematodes weie fixed in hot, 10% neutral 
buffered formalin and cleared in laciopbenoL 

Co-types of T. nasalis were kindly loaned to lire 
by Professor E BiOcca of the Istituto di 
Parassitologia dell* Univcrsita di Roma, loan of 
male and female pftatypes ot Parairfchosomu 
cracodilus was kindly arranged by Mrs E. A. frfatTll 
of the British Museum. 

Measurements were made with the aid of an 
oeulat inicromctci, drawing rube, and measuring 
wheel, and ate piesentcd in micrometers unless 
otherwise stated. Where possible, the range of 
measurements is followed by the mean. In paren- 
theses. Illustrations were made with the aid of a 
drawing tube. 

type specimens have been returned to [heir 
respective institutions. Representative specimens of 

• Division of WikJlilc& Kangeiari.h ReettkrCn < $I*0( 
P.a Box »4. Lyiicham. A.C'.I. 2602 

7. nasalis from R. Jusctpes in Australia have been 
deposited in the British Museum (Natural History) 
ifl.M.(NH) No. 1981/3537-3540), the istituto di 
r'urassitologia dell' Universita di Roma, the 
Museum national d'Hisloiic natutelle. Paris (MN 
47ft HB)» the South Australian Museum (V3244), 
the Australian Helnunthological Collection (13855), 
the United States National Museum Helmintho- 
logicsl Collection No. 77454 and the helminth 
collection of the Division o! Wildlife and 
Rangdands Research, CSIRO (N498, 717, 882). 

Capdtaria crocodiius (Ash ford & Muller, 1978) 
comb. nov. 
FIGS 1-7 

Material examined: from Crocody/us nwaeguimw 
nuvutxtdnetie Moitaka crocodile lann. Port Moieshy, 1c? . 
IV paratypes, BM(NH) coll. No. 197K/s»l 7-920. 


Long thin nematodes with attenuated anterior 
and blunt posterior extremities. Cuticle 
exceptionally thick, cuticular striae not obseived. 
Two exceptionally broad lateral and one narrow 
ventral bacillary hands. Hypoderma! gland cells ol 
bands papilla-like in appearance due to necessity 
for neck of cell to traverse thick cuticle to external 
pore opening. Lateral alae absent. Cephalic 
extremity rainuic, with minute dome possibly 
consisting of two lips; stylet and buccal capsule noi 
Observed oesophagus commencing as tiarrow 
muscular tube, broadening posterior to nerve ting, 
narrowing betore reaching stichosomc, passing 
through stichosome on dorsal or lateral surface but 
exiling from it on ventral surface as narrow 
muscular duct. Stichosomc not extending as far as 
intestine. Narrow muscular oesophagus forming 
unusual opaque junction with intestine, consisting 
Ol i wo rather than three- 1 issue %cemcn(s and not 





appearing as tmc valve Pto rrtesc ichymal cells not 
present at nesophago-inttslinal jiificrton 

Stichosome composed nf appMisur.aiely 36 slfcbo- 
cyies, much longer than wnlr, L-ave nuclei.- not 
observed ta Bach stfehocyte intestine large and 
conspicuous, lad broad and blunt, without papillae 
in female and wiihoui alae in uiaie. Anus and lonca 
[uinihM! fa female and male respectively. Male with 
short ejaeulatiuv dnct, brtfi cloaca lined ■ 
tuti-.iu, long roburt non-scleronsed spicule *fld 
spicule sheath (-ehm. setisu Anderson & Barn, 
1982) with robust blunt spines Or Mitt- Itetflk *1th 
vulva slightly posterior to ot-vophaey ndcMinal 
junction, vagina vera lined a ith cubrulRT series or 
scales, BJKg8 moderately thick shelled, unlanneil, 
shell without ornamentation, eggs ijj distal atcius 
not embryonaicd. 

Length 22.7 mm. Wid.t . nerve ring 10, at 
oesophago intestinal junction 57; maxifl 
width 66. Length of mu&culnr or- JOO; total 

length of oesophagus 13.6 ram. Length of 
Kticfroionw n.2 mm; >1 les 33 in iuup 

length of sUeftoeyteS 260-330, Nerve ring 50 | 
cephalic exttenm v. EbtaJ leiiyih ot cloaca 930; length 
of spiny spicule sheath isd; length of eiaculatory 
duel 100, spicule pouch entering cloaca 213 fttwfl 
caudal extremity. Spicule present, not SClCfOti 
450 in length, 8 mi maximum width. capknVaitl 

characteristic "bwl ; v ; ip d* htpad 

proxirnally, with broad hut tajvi iftg point distalb 
Junction of intestine and ejaculatoiy duct simple, 
both ducis same width at junction, Spicule siieatlh 
wnh robust but general!} bU>IM spit es t« scale-like 
ornarnenialion <not everted in 'his specimen). 

Spicule retractor rrttfwJe Lftsetttng 150 anterior to 

sphincter muscle delineating modification of distal 
vas deferens into eiacuhnory duct. 

Lateral and ventral haollaiv nueiici i\ 

in region of nerve rins; lateral bands 2 ? cell* v 
cells regular in distillation; ventral band off 
wide. CCllfi iW Btthtt in diMrihuium. I alerat bands 
bruademm*. i^ptdl v. |0- I ! • cM> >" width 
approximately S mm posterior to cephalic end and 
practically encircling worm; band*, narn 

Jpllj near tail ftp. 7 cdfc i T » width ai level of 
entry ol spicule- pouch lo cloaca.! hacillary 

bond with single column ol cells spaced irregularly 
oody length. 


1 engttl '■". c MiMii. Width at nerve ring 70, at vulva 
II. 1 , tnaXimUin width 135. length of muscular 
t.esupha^u.s 270; total length of oesophagus 
p.'.; T.n. length ol itJchosorae 18.7 mm; sticho 
yi in number; length of stichocyles 300-580. 
Nerve ring 60 fn>m cephalic evtremity. Vulva 
20.2 mm from anterior extremity. Vagina vera 109 
long, fined with thick utricular scales similar in 
etiology (0 those on male cirrus Keclum 201 
long, lined with thick cniide. Anus terminal. 
Lateral and ventral hacillary bands commencing 
gion ot nerve i fog. Lateral bands 4-o cells wide, 
cells regulaj in disposition; ventral band one cell 
cells irregular in disposition. Lateral bands 
broadening rapidly, 10 n cells in widih approxi- 
mately 5 nun posterior to cephalic extremity, 
prjLiieally encircling worm; bands narrowing 
abruptly approximately 0.5 mm from tail lip. 
pal hacillary band with single column of cells 
spaced irregularly along body length. 


My otntrv4(utits ddtei markedly from those of 
AsJifordA Muller 11978). This tre-hund nematode 
specie" bout epidermal tunnels in the abdominal 
skin of crocodiles possesses (i) conspicuous 
biuiltary bands comprised of papilla-like hypo- 
denuaJ gland cells due to the neck ol the gland cell 
having |d traverse the thick body cuticle to the 
fial pore openinR, (ii) a stichosome comprised 
til 3M7 elongate llldiocytes arranged in a single 
column, (fii) n Jon'- broad non-sclerotised spicule, 
.i spicule sheath with robust blunt spines and 
Male like wnajnentation, (v) a long muscular cloaca 
with chicle cnhcuhir lining and (vi) unembryonatcd 
egj5& in the distal uterus of the female, none of 
Which were icported by Ashford & Muller (1978). 
The caption to their Fia 5 contrndkts their plate 
menl On p. 216. "Ewti unembr>\>niMexi when laid" 
and should read, Tgg from skin, containing a 

I he abo\ , cmcntioned features arc characte | 
of the tridwii id nematode subfamily Capiilariinae 

I i' 1-7, ( uptllitftQcriWitnw [Adlfcnd & Muilrr, iy >i DOmb. rev,, j. Male posterior end showing junction ot intestine 
.lm'.I - lacutuu-rv duel with Jist-. MaK pOMCriof cr,t * showing capitulum ot spii nlr. Bplcule retractor and 

prormaoi rnusdexantlcloaCa. 3. Mai a ll Wfcdiltowinj .ir.i.i. spiny ^iculc sheath and distal end of spicule. 
4 Blum spmc mid wale Iffcc W Ba u arnlaUiiKi On spiCO*eShC»tu or male-, t, femnle anterior end showing oesophago- 
Int-ewinnl lunctii - &nd s-an'ma wilti Mralc in new. '■ Female caudal end. vemral 

virv. 7 ■ ' ' ,,; cwwb rwck ot BlamJ iIn^|bickeAilidetoe»ieriO!l poreopeninss 

vule li.iL-s: t ifv.t : ii> U t..,tu-, 2, 20wn;l 4 ^i I S.^fljun Abbi ■ il »hcwh i-ckrus sens* 

Anderaon& halt 1— " '■ w>'- i i pmttactor muscle 

• phade r i retiitm ".uic ■( -.i> 'av»u-. va— v^ginn. vg vulw. 



{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 tatu) Zeder, 1800. The reasons for this 
synonymy are presented above and the crocodile 
parasite is formally recognised as Capillaria 
crocodiius (Ashford & Muller, 1978) comb. nov. 

Despite the foregoing, C. crocodiius 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 terminal ing 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 o\' 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 

Ashford & Muller (1978) reported that two 
crocodiles, G novaeguineae novaeguineae and C. 
porosus Schneider were susceptible to infection with 
Capillaria crocodiius, that the parasite was rare or 
absent in crocodiles from areas of saline water and 
that transmission of the parasite does not occur 
under some rearing conditions on crocodile farms. 
Undulating nematode worm trails, probably 
attributable to C crocodiius, have been reported 
from C. acutus Cuvier (Garrick in Webb & Manolis, 
1983), C inter medius Graves (King & Brazaitis, 
1971), G johnstoni (King & Brazaitis, 1971; Webb 
& Manolis, 1983) C. m o relet HDumerti, Bibron and 

Dumeril (King & Brazaitis, 1971), G 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 latuj 
(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. 
J have recovered adults of species of Capillaria from 
these sites in P. nasuta and Perameles gunnii Gray, 
and from epithelial tunnels in the tongue of P 
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 
Rattusfuscipes 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 (5c=5). 

Material examined: from Rattus (=Epimys) norvegicus, 
Rome, Italy, T. Crapulli, co-type S containing 2 co-type 
c?c? in utero, fragments 2 co-type $$, 1958, in Istituto di 
Parassitologia dell' Universita di Roma. 

Figs 8-21. Trichosomoides nasalis Biocca & Aurizi, 1961 from Rattusfuscipes. 8. Anterior end, female, dorsal view. 
9. Oesophago-intestinal junction and vulva of female, lateral view. 10. Caudal end of gravid female, lateral view. 
II. 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 hypoderrnal 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 hypoderrnal gland cells of bacillary band in posterior half of female, lateral view. 20. 
Lateral bacillary band, mid-body region of female. 21. Lateral bacillarv band 0.5 mm from caudal end of female. 
Scale lines: Fig. 12, 10 M m; Figs 13, 14, 20 jtm; Figs 8, 10, 11, 13, 21, 50 urn; Fig. 9, 100 ^m. 




. I ■ , i - M i i . i mral l ri T 



©Mg ,, - (3) 




From R. fuscipes, Lee's Creek, A.C.T., P. Havcock and 
D. M. Spratt, 15??, 66dV, 22.i.l979; Bundarra Creek, 
Snowy Plains, N.S.W., D. M. Spratt, 34$$, 91d'c? , 
22.ii.1978; Emu Creek, Bondo State Forest, near Bondo 
N.S.W., P. Haycock and D. M. Spratt, 699. 21^ , 
24.xi.1978; Nadgee State Forest, near Eden N.S.W., P. 
Haycock and D. M. Spratt, 7$?, 23d"<? , 12.xii.1978; E. 
Walter and P. Haycock, 5??, 28rfrf , l.xii.1982. 


Female relatively long and narrow, with slightly 
attenuated anterior and blunt posterior extremities. 
Male minute, occurring in uterus and vagina of 

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. 


Members of the genus TYichosomoides are 
parasites of the mucosal surfaces of murid rodents. 
Only two species are known; T. crassieauda 



(Bellingham, 1865) Railliet, 1895 from the urinary 
tract, generally the bladder, of wild and laboratory 
rodents throughout the world, and T. 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 et 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. gerbil/is from 
the stomach of Gerbiltus pyramidum hirtipes 
Lataste in Tunisia but this was later transferred to 
Anatrichosoma (Pence & Little, 1972). Males of the 
genus Trichosomoides 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 
I he 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 T. 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 T. 
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 /im 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 

The records of T. 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 et 
al., 1970; Cross & Santana, 1975). 


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. 


Anderson, R. C. & Bain, O. (1982) CIH Keys to the 
Nematode Parasites of Vertebrates No. 9. Keys to genera 
of the super families Rhabditoidea, Dioctophymatoidea, 
Trichinelloidea and Muspiceoidea. Commonwealth 
Agricultural Bureaux, Farnham Royal, England, 26 pp. 

Ashford, R. W. & Muller, R. (1978) Paratrichosoma 
erocodilus n. gen. n. sp. (Nematoda: Trichosomoididae) 
from the skin of the New Guinea crocodile. J. 
HetminthoL, 52, 215-20. 

Auk i /i, A. (1958) Su un nematode del genere 
Trichosomoides parassita delle cavita nasali dei ratti di 
fogna (Epimys norvegicus) di Roma. Nuovi. Ann. Ig. 
Mien 9, 264-266. 

BERNARD, J. (1964) Trichosomoides gerbil/is n. sp. 
parasite stomacal d'une gerbille d'Afrique du Nord. 
Arch. Insi. 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, Hetmin'hot., R. T. Lei per 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. 

, Kundin, W. D., Jones, G. S., Gaba, A. M., 


Trichosomoides in the nasal cavity of Java rats. J. 
Parasitoi 56, 566. 
Gallego, J. & Mas coma, S. (1975) Capittaria garfiai 
n. sp. (Nematoda: Trichuridae), parasito de la mucosa 
lingual del jabali, Sus scrufa Linnaeus, 1758 
(Mammalia: Artiodactyla). Vie Milieu 25, 237-48. 

King, F. W. & Brazaitis, P. (1971) Species identification 
of commercial crocodilian skins. ZootogicaSb, 15-70. 

Lyne, A. G. & Sommerville, R. I. (1956) A nematode 
parasite, Capitlaria sp., in the skin of the marsupial 
bandicoot, Perameles nasuta. Ausi. J. Sci. 28, 205-6. 

Moravec, F. & Cosgrove. G. E. (1982) Pseudo- 
capitlaroides xenopi gen. et sp. nov. from the skin of 
the South African clawed frog, Xenopus taevis Daud. 
(Nematoda; Capillariidae). Rev. Zooi. afr. 96. 129-37. 



Pence, D. B. & Little, M. D. (1972) Anatrichosoma 
buccaiis sp. n. (Nematoda: Trichosomoididae) from the 
buccal mucosa of the common opossum, Didelphis 
marsupialis L. J. Parasitoi 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. Parasitoi 57, 63-71. 

Wade, S. E. (1982) Capillaria xenopodis sp. n. 

(Nematoda: Trichuroidea) from the epidermis of the 

South African clawed frog {Xenopus laevis Daudin). 

Proa Helminthoi Soc. Wash. 49, 86-92. 
Webb, 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, NT. V. Abnormalities and 

injuries. Ibid., 10, 407-20. 


byS. Skinner 


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. 



by S. Skinner* 


Skinnik, s (!«7X5) Australian and New Zealand species of Mavhtstu .md Fiafofftrix (lilachistaceae, 
Pbaconhyta). TfoHW. R. Sue. S AltSt, I09<4l 151 160. 2'v November, 1985. 

Three specif? at Fiachista Duby, E . tiaviomae sp. nov., / . australis .1. Ag. (Syn. Hottphtlhpia uustmlts 
(I. Ag.) Silval anil /_' nrhtmluns (Ohtai Skinner and btlC specter ol Hitfofbrt\ Reinke. H r(>hrf>i,r.ih\ ty 
tun arc desenhed trout southern Australia, t. autlrafis and E. findauert Chapman are described faun 
New Zealand, mid a key is given in the Australia and New Zealand ux.t ol tlachtsta. I he systematic* ot 

the r> 'His ftuilHMu ate discussed. 

Ki v Wouns: Uuchista; L ctaytonitu* Spi nov ; £, uit<>ntih\ f- nthnulmis, }■ tuahium; Hutt>thn\. 
H. tpkemenffk sp. nov.. Portphillipia; Llaehistaecac; Uiordariales; Phaeophyta; southern Australia. 


The family Hachistaccac includes 'hree genera 
and about 30 species, and is usually placed Iff the 
oider Cliordariales. The absence ol" any hair-like 
stiuetures and l he presence o I determinate cortical 
asMmilalory li laments r'paraphyses*') in 
combination with long, indeterminate 
filaments clearly separate this family Irorn other 
families in the order. The pulvinate thallus is 
composed of a filamentous medullary system which 
radiates from a basal layer and supports a Cpl BR 
v\ both determinate assirnilalory filaments and 
reproductive stiuetures, beyond which project the 
long, assimilatoi y filaments. The cells of 
assimilatory filaments of all three genera contain 
a large central nucleus, numerous discoid 
pbaeoplasts with many species having prominent 
pyieuoids praje&UIg from the phaeoplasts, and 
some physodes. All are epiphytes. 

The genera included in the family are jlafothri.x 

Reinkc, epiphytic on seagiasses, tdtchista Duby 
(including Syntphoricoccus Reinkc and 
frttlphilltpiu Silvai. epiphyiic on laieer algae, and 
Iferpo&tscus South (1974)| epiphytic on Durvifiaea 
Lluchisia species are found as epiphytes on various 
Phaeophyta and Rhodophytu. wirh one specie 
ZQSterae Noda in Noda &. Kuami 1V71 1 on a ruanuc 
angiospcrtn. PJurilocular sporangia, winch arc 
produced by most species, arc unisciiate, filiform 
and are borne on sporangiophorcN among the 
cuihcal assunilators. 

Hiihrhnx differs from Elachi&ta by having 
UHCteulaiv pirn llocular sporangia which develop, in 
p Minis, in cells wjthin the long assimilatory 
filaments It also shows relatively fewer phaeoplasts 
in the photosynthetic cells, and, at least iii H. 

*Depaitinem ol "Botanv, Umvi-i -tiv ill AdHaid'--. i .(' M. 
Box 49H ( Adelaide s. Ausi >aoi 

tutnhricalis (Ka'itz.) Rein be, a greater development 
of rhfcoidal filaments buttressing the medulla— 
both ol these dillereuces me ones of degree rather 
than contrast wilh fctachisia. The hosts o\' Hulothrix 
are usually marine angtospcrms y/ostera, 
Heierozostera and Phyllospodix). 

Etaehista {- Portpftiilipja) australis J. Agardh 
has been recorded from Victoria and Tasmania 
iVVomcrsley 1%7) and I . of hit ttluhs (Onta) Skinner 
from Western Australia and South Australia 
(Skmnet 1*>S3). I lie piesent study shows that there 
is also one further species of EJachisfa and at least 
one oi Haloihtixiw southern Australian waters. In 
New Zealand, two species of tlachisto have been 
recorded, E t austtvhs in the south, and the endemic 
E. lindaiteri Chapman, 

Asubamarctie species olEtoehbto, £*. atitarcticj 
Skottsbeig 1953, is also Included in the key because 
.if \\$ recent discovery on Micquarie Island (Rickcr 
pcrs. comm.) 

Key to the Gcucra 

I Ptant pul\ inatfl to spreodllVgi epiphvtie un liirger algae, 

♦viis of long ;issinHi.(ii)i > fiiamenu miji ounop&tous 

packed phaeoplusts, phtnlocular sporangia, when 
present, uniseriate and mimed on apOtatlgiDpUorffii 

IV "ising I'roni iTicdi''tai v I'datTicni • nnd within the 

Exnte& : . i — — — . - - 1 , . . I'lm hisiu 

*HanU pulvinate* eptphyltc -m tfrltrnz,<>sh-rii, celln ol' 
long ussimilmorv ftlaments with niinierous hin 
scattered phacniilasis. pint tlondai spoianuKj 
urn-Hillary ftl '-on on lent- tttfitnil&laft l'l;nncn(v 

. . Hahthrh 

Genus LLACHISIA Huby I83(h 972. umu. cons. 
I. Agardh 1848: 7; INK! 9 

c tuptuau 196; ux Kaniei wy> \\i k&tvq IW6j ^ 

Kielhnan 1807: J20, Kuekuek I929i 21 5fift ltW4 
t indaucr. t-hapmau iV Aiken 196k 214. Roscnvinc' 
1935: i9. SauvaKcau l9Jfi: 139. Skocuboq 1901 I 
1051 537 Sv'.*delii.s IMti 162 'QkaMUUui 1431a H3 

>amada 1928: I). 


& NMNM-k 

Svmpbom-octiss Krinkt I**'; 17. Kuekutk 1929: 32. 

KJcttman 1997: 219 
PoriphUtlpia Siva IS7D: 944 

Thallus epiphytic pulvin&tc lb glohose, 
hemisphe lea) oi iprtadidg, ttsunlfe ?-2Ummhiuh 

and aggregating in considerable numbers on (he 


Basal layer o! small isocliamelrie cells, forming 
a plate on the ho-st surface, often with short pegs 
tS extensions of the lower cell wall. Medulla a 
Filamentous cushion of usually closely packed cdli 
supporting and completely sheathed by the cortex. 
Medullary filaments of non- pigmented, eylmdtical 
to elongate-ovoid or subgiobosc cells with 
branching from most cells. Rhixoidnl filaments 
develop fiom cells of the lower medulla in many 
species and »wo species i£. QUstNtlh and £ 
lindauen) form interweaving, transverse supporting 
tilamenls IKLinimernreane (Kuckuck 192'). p. 19)] 
clampitons formed laterally from cells id lower 

CoUex ol short assimilatory ftJainetftftl and 
reproductive organs terminal to the medullary 
filanients. c onical assimilatory filaments 
determinate (associated with sporany.tuphorts), 
filiform to clavale, slraight to arcuate, with 5-20 
cylindrical to inflated cells, containing fewer 
phaeoplasrs than cells oi' long assimilatois. Long 
assimilatory filaments (sometimes reteired lo .t 
assimilatoiv bans) indeterminate (except in EL 
sit'iluris Arcscnoutf and ConocJla marchuntuc 
Setchcll & Gardner), protruding well beyond the rest 
of the tludlus with a meristem of short eylindiical 
cells, and above with usually elongate, cylindrical 
pigmented cells with thick walls. 

Uniloculai spCtatigfa in the curti ,hon 

pedicels. Oil terminal medullary* cells or from the 
h.ise o\ assimilatory filaments, ovoid ot pyitfonn, 
with a terminal poo-. 

Plurilocular Sporangia uiusetiate. filiform, with 
8-48 locuHi eithci in speeial./ed. coiymtose lateral 
hranches from the outer medulla or laterally from 
long assimilatory filaments. 

Type specks ElaChiste scutulata fSmilh) Areseh 

A genus of at least twenty species, found on a 
wide vauety <d hosts and known from temperate 

and colder seas. Thiee species occur in Australia, 
and two in Nov Zealand. Sexualit •' has nan bee (1 
dearly demonstrated in the life cycle, which, at least 
IJ3 some species, appears to be direct . 

Key to the species o/Elachjsta in Australia and NW 

I, Medullary filaments of one kind nnlv, uuiinculai 
spiifunpia present; plurilocular vpm.myui, when 
present, with 8-lh I ... 1 

Mcdidlaiy filaments of two kinds, unhranclu-d 
filaments which near the long asjimttaior) ntameuts, 

andsubdich.a ...s f ilnments which heat die cortex; 

1 I ulat .poiauri... jltM-.K, plunloeular Sporangia 
with fh-?4(-J8) locuti . . Ffaiijtsta i>rht,'uti/t>-, 

2. Medulla i ■nmpii,'i, lil^njc t as branching from mo*r 
yells, no transverse supporting filaments pn}3CfH 
cells ut tdtlfi assimilatory filaments at least as wide 
EU medullary cells 3 

Medulla loose, with sparsely branched tiLimcuu 
producing transverse supporting likmn-no; beltffl 
eelK 01 lOOfi assimilalory filaments narrowei the 
medullary eelK. 

3 Both unilocular ami plurihn n.,.i 5pOfA0fela hihv be 
present on the one plain, pi uiiJocu I at ipDtangra u.oialtv 
more comTtior.; cortic il Dsstiitllnrpry ftlaments 
cylindrical and pu ntfful tlvvhisiQ vlayionw 

Unilocular sporangia only piev-u I. COfUcti atintUl&torj 
ftlaments cUvate bill vpant thchistu antarctica ] 

i i elK ol medullary filarm'tu:-, 7 1(1 ,, affl 
in diameter; cbrriool n**tanlfe«D< y rilmucws filiform, 
Wflvy, Cttll muumw, .vhndncal and uniform 

Gtochtsm outittoiis 

Cells o( medullar*. idameniv OV -cylindrical, 

20-30 urn m dtmeici cmiteal a f -simii-m>rv 
il.-itiiemselavaieand arniare. upper cells larcull-. 

iliil.Ot i\ 

Efathisca hmiaucri 

] EfaCftf$la an tare ticct Skottsberg 195.? has recently been 
eollecled from Macquarie Island (Kicker, pen. 0OR1M t 
and may occur on otilfti subamareti-. elands south ol 

\ii\ir;iiiM and New Zealand. Skottsberg u*W3t considered 
ilui i Jus IAS0U did not produce conical awimiiarorv 
' i'UhhI MHsirnilators are present but uncommon on 
material froin Mueonarie island (Rlcfut MA-2M, 

J -i 1978; \OU A530S2). 

Pig. I. A-f. Hlochisto eiqjNonfm --v nov (ADD, .\508O9. Molotypci- A- Hui-ii, on bond of Sarjtassum %p. B. Upper 
medulla and COrieq showing sialkeu spherical cells amnn» pluritOCUjai spofBni^a < Unilocular and plurilocular 
sporimgifi D ' -liameni showing long asshsiikior and vjctuid confeal branches. L. Upper medulla supporting cone* 
and lone assimilatory AlarnenL I. Lower medurt&and Nbc.G-1. Eiachista australta i. Ak- (AIM K A34R09i G HaWt, 
on h'tpHaphara. H. Rhtmerttafaflwtog narrcAv transvern 'd«mcnu from lower nodes, medal lau) and long asilmlliuoej 
filament and conical branches with cortical a^snntlaiar& I. Coriest with uniloculai sporangia among cortical 
assimitatot.. t m Hato/hnx tyh&mtrttfis sp nov tAlui, M2A64« TVPE) I HatoU on HeiertK&stertr tantrtanita. 
K I.VJ f iLiPimi ihowinK uppei moduUa, conical t-i*.n lu-s with unilocular sporangia, meriMt m ami celts ot lonji 
HftjirrtflatOr* IBamcin with sorusol filunh^otar spdiro.i'.iN I Individual cortical asMtnitator. M. Mcsdulta :in»i \ww 
Of long Ts\inularor\ filament. 





Elachista claytoniae sp. nov. 

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. IF) of irregular, isodiametric 
cells 8-15 Mim 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 *m in diameter, L/B 2.5-4. Cortical 
assimilatory filaments (Fig. IB) 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. ID) 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.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 ADUA50809, but zooid 
masses were not observed in such organs (Fig. IB), 
and their function is unknown. 

Plurilocular sporangia (Fig. ID, 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 j*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 ^mdiametro, L/B 1.5-2.5; 
sporangia unilocularia ob pyriforma; sporangia 
plurilocularia uniseriata, filiforma in sporangio- 
phoribus corymbosis aut ex medulla superne aut 
ex fills longissimis. 

Holotype: ADU, A50809 {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.l971; MELU, 21207); Point Nepean, Vic. 
(Clayton, 4.V.1969; ADU, A50332). 

The specific epithet, "claytoniae", is chosen to 
honour the collector Dr Margaret Clayton, and her 
distinguished work on Phaeophyceae in southern 

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). PhitippieUa 
australis (J. Ag.) Silva 1959: 63. Lindauer, Chapman 
& Aiken 1961: 215, fig. 42. Womersley 1967: 229. (Non 
PhitippieUa Spegazzini 1 896: 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 titn 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 urn 
in diameter, L/B 5-7. 

Cortical assimilatory filaments (Fig. 1 H-I) 
flexible, sinuous, of 25-30 cells at maturity, 4-7 Mm 
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 urn in diameter, L/B 1-2. 

Unilocular sporangia (Fig. II) borne on a one 
celled pedicel or sessile on lower cells of cortex, 
ovoid to cylindrical with a terminal pore, 45-60 urn 
long and 20-50 Am 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). 



Distribution: Apollo Bay to Port Phillip Heads, 

Victoria, the north and cast coasts of Tasmania, and 

Stewart I , Now 7i?aland, 

Host ranee: Xiphophota chondrophyila and A 


Specimens examined; Port Phillip Heads, Vic. (Harvcw 

Alg. Aiist, l.xskc |Q1 5; AOU, *WS3l)j Apollo Bay, \ ic 

[fftmfittfctt 10.xii.lOt>V; ADti, A348Q9X Point Lom.a^k, 

Vic (Skinner, 4.1.1978; AlX', A4W>7 t . Uver-,tone. Lis. 

(Cordon, ou.iw*; AOU, A3O107); Belter ive. Oerwem B„ 

1;> [H'xhvuv, Del. 190ft AIMJ, A5HIS); gtO&t ^u.l 

N his (Hyi/^.-v/cv, irYx.1982: ADU, A5575U); 

'' ■ * * f *'i .! r.ivr, IM ni'. Eagtehwk Next-, |)u tparsomlk 
ilotnersify jn.v.m:. aOIJ, M39*i CHR, 399502); 
RlMiMMniti. Sicwurl t. ( Nw Zealand I ftbffwpfeft 4 i.1%6, 

Aim, A2&81Q) 

Llaehtsra oustrahs and £ lirtdouea Chapman art 
distuictbc specie. Both show i louse and eAtenstvc 
medulla with transverse supporting filaments and 
the development irf a subcortex by coriccnira'iop 
Of lateral branches in the upper medulla lo tttppui I 
Cortical asMimiatoiN. £ hndouen (see below) 
lias larger cells throughout; the medullary cclU are 
inflated rather than cylindrical ax ill E. australis, 
an! i In* i onical assimilators arc arcuate rather than 
sinuous, with the upper cells inflated laterally. The 
presence of the tran.sver M supporting filaments was 
used by Kuckuck (Kuckuck In Oltmanns 1922, 
Kuikuck 1929) to distinguish the genus PftWppiCi 
from L/ochista. There arc several olher species of 
J.luihtMa e-e. E tnoUis, }•_ \enoniant and E xlobosa 
Takamatsu 1938a, winch show a concentration of 
branching in (he upper medulla, although none of 
these taxa produce transverse supporting filaments. 
Tht retention of the genus Ponphttbpta Silva (* 
Hulippta kuckuck non Ktotzsch) with the inclusion 
til / ausirali; and E Hndauen, does not seem 
warranted, and ('. OUStfVlft is here relumed to 

EtecdfJTjfl l/mJuuen ( hapinan 1961, 16, fig. 2. 

1 indaucr, Chapman and Aiken 1961: 215. 

r IO. 2 A-D 

llullitN yellow hrown, cjobosc to pulvnuce. 
epiphytic on Xipbophora ehutatrttphyUa 

Basal I ivt r contracted to & very small diik. 
Medullary filaments (Rgi 2A. ft/ very lonv and 
sparsely hranchedv cells inflated-cylindrical, 2A-30 
Mm in diametei, L/B 2-3; lower Uterjls fojgjijjg 
rhi/oid-like, hunched, Uaaisvcise fclpbgrllng 
nk.iocnts wirh often more than Ofie ftoni i he start 
node; i-_3 tpper branches iv^. 2A, C, D) Ifcm 
BOdc, forming a aubumcv supporting Ltic conical 
aswmilarOTS and unilocular spoianyia Corneal 
assimilatory filaments sliimlly elavatc, areualc, the 
upper cells lateralis' ispanded and deltoid. I OHg 
n nniiaroiy filaments (I tj*. 2A,i io.u illy Icntfnctl 
lo ihe main avi- ,.' .3 medullars Munich', -villi a 

meristem at or jusf above the level o\' the cortex; 
mature cells eylmdneal, 12-20 em m diameter. L. B 

Unilocular sporangia sessile ovoid to pynlorm, 
30-50 *»m in length and 12-18 ^m in diameter. L/B 
3-G, borne laterally on lower cortical cells. 
Plurrlocular sporangia unknown? 
type locality: Bay of Islands, New A^aland 
(Lindauer, 6.xi.l9V > 
TjtyfS AKU (Herb, lindauer No 29). 
Specimen examined: Isoiype (ADU t A4573) and 
I on j Beach, Russel, New Zealand, lower culhtorul 
pDO B W-ontersley, 28.\.1969; AIX, A^.S()69). 
Distribution; throughout New Zealand. 

Chapman's (1961) description makes no mention 
of the Iran -v-.isc supporting filaments, and is 
uncomplete m the description ol branching in the 
-medulla His description of "plurilocular 
gametaueia tonned by modification ot ends of 
a.ssjmilarors", without a figure, is un: vbaantiated 
In the ^pcciriKTiv examined. 

I'(uc)ns(u orbicularis (Olua) Skinner I9H3; 98, 

Figs 1-3. 

GOtiOCfiti ortm-uhtris Dhta 1973: 2J. Tig. II. Noda 1975: 

kill n& i7- 

No further dala have been obtained to add to the 
species description ol Skinner (1983) 
IV pe Locality: Tappi, \onion Pre!., Japan \Ohta, 
I. i. 1 970), 

Tvpe- \Obio 133) in Herbarium, NLigata University 
Distribution: Japan, and southern Australia (from 
RottneM Island. W Aust. lo Port Hacking, New 
South Wales). This .species is present on its host 
throughout the year. 

Hostx Vndaria species in Upan, t.cMonio nidtittfi 
{€. Ai?,ardh) J, Agordh in Australia. 

The two most closely related taxa to (F.lachista 
orbicularis ate fi nigra 1938a, and 
Gonodia fu.\(fortru.\ Noda in Honda 1969 noni. 
FlUd i 5peciC6 o\' EiacfllSia) both endemic to Japan. 
While tin other species show the formation of some 
lateials from those medullary filaments ahich 
support the lon^ flxsimifdtaty filaments, £. 
ortnmlans develops sucJi laterals only very 
occasionally, and then from the lower most cells 

Genus HALOTHHIX Reinke l*K9: 49. 
rifrmvi 1935. 126. Kucfu.k 1929: 26, fig- 15-17. 
RoeeilVlntC J935: 37. Takurnauu ISOXb: 1MI VaniaUa 1928: 
S13 Yeudo 1909: 12.1 

Thallus pulvinaie. lieruispheriLal, loosely 
compacted, up to 2 em high, epiphytic on 
Zosteraeeae. Individual phmts may be densely 
clustered, giving the appearantv o| oik plaQl 

Basal layer a central disc of isodianietnc celts 
with, a1 the ruamius, a rhi/.oidal system from small 
isodUl Heine cells in the lowermost medulla. 



• --m and uAiomxrx in mjstaai *ma 


Medullary filnmeni 5 ot pyritorm or subglobose non- 
pigmented cells, with hunching confined mainly to 
upper cells. Conical assimitatoty filament* elavaie, 
arcuate or sttaigliL hi a lew ovoid or cylindrical cells, 
formed inuu.'di.aely hclow the iiiensremaue zone of 
the long assiinilatory filaments, on lateral medullary 
cells. Long assimilaiory filaments narrower hasally, 
extending in length many times beyond the iesl of 
the lhallus, with short cells |1 ft about I) with 
numerous scattered phaeoplaM.s 

Unilocular sporangia arising from upper 
medullary celts, ovoid-cylindrical or urceolatc, with 
a tCfiitmal pore and play. 

Pluritoculai sporangia in son on anticlinally 
divided cToups o! cells ft upper pails of long 
aSiiualatoiy I'iUmienlv 

/iV'f VM'ticx H. tumlinajlts (Rut/ ) Rciriko 

A genus of live described species, H. hmibncnlis 
from ihe north Atlantic and Baltic, and four specie- 
froni Japan, An additional new species is here 
described I mm southern Australia. The principal 
difference between this genus ami lilochisto is Mir 
specialized plurjloeular sporangia v.-u ihc suitaet of 
llic tOiVg asMnulaiory litamcrus in Halothris While 
) . kieltaris farms pturiloeular organs on long 
asMinilatutv filaments, ihcy do not involve the prior 
subdivision of tiu ifrfBok motliei -cell as in Huhnhm. 
IV-deiscu <|c)7M) lias shown that Halothrix lumbnotlis 
h;is a simple direct lite cycle. In a recent paper 
Kxlcrsen (I9K4) has suggested the inclusion of 
Halothrix in the Ciiraudiaccae on ihe basis of 
siniilanties in morphogenesis However, the 
concentration of branches in rhc medtilta 
immediaielv below the meristem in Halothrix is also 
seen in vaiious specie* rji Lfaclmta, notably in E 
mollis* l- claymniae, h gfofiasfl and £. rost&ue. 
More importantly the development ol the 
photONvmheiic pun ol the Gwnuiia lhallus. above 
the mefistem, involves an orderly sequence of 
penclinal divisions ol i he cells of each whorl or tier 
(Skinner ft Womersley, t9S4>, wluch im not seen in 
Ho/uift'Dc. Myrtoirichta species show possibly closer 
parallels to Qhutidm l as ihe axis of the lhallus in thai 
rcnus undergoes penclinal division ol" as.similau>r\ 
celts before the production of unilocular uml 
plutilocular sporangia tkuckucK, 1899), 

ffUfaffaA cphemetxh* S» IM 

lhallus hrown cpiphyri. on Urn ttg&Vteffil 
fasinuntva leaf blades (Pig, 1 1>, oiu-n so numerous 
as (0 covet tftQSI ol the blades, pnlvtnalt, loosely 
compacted, 5-1 s" mm high 

Basal plate of small isodiametne cells 1(1-15 i^rn 
in diameter, adnate to host surface. Medullary 
niatnoDQ (Fig. 1 KM) wiLh 3 or 4 lower tnpenng 
cells 20-25 Min in ditnncter, L/B 2 >~4 t inlrcqueivU 
branching, and 2 to 4 upper subglohosc cells 20-30 
**m in diameici, LB about I, hranchme 
subdiehotomously. Conical assimilafory tihiniem , 
straight, of 8-15 slightly inflated eelb. grading 
upwards from cylindrical to subglobose, 10-13 >un 
in diameter, L/B l-LJ (Fig. t k : , L). I oik- 
assimilaiory filaments (Fig K,. v: ., Ml many runes 
longer than the rest ol the lhallus, cells increasing 
in diameter upwards, short but broad, upper 
photosymhetie cells cyliudiieal, 30— rU) /on In 
Ctelttticr, LB 5-1 

Unilocular sporangia (Fig, IK : ) cylinilru;il. with 
a terminal domed plug and pore and lapeiing 
basally, 15-30 i«m m diameter, usually paired on 
Uttered medullary branches arising immediaielv 
below tht meristem ol a long assimilalory filament 
together with conical assimilatory filaments, 
maturing sequentially. 

PluriltKular *>porangkt (Kg. 1K,> ut iuteicalaiy 
son borne on 2-6 modified cells of |0ttg 
a ssi mi latory filaments; 1 1 ie soral mot her eel I 
undergoing two or Ihree anticlinal divisions, wiih 
individual sporangia (some with a basal cell) arising 
(torn periclinal divisions of these secondary cells; 
often present on plants belon; unilocular spoiangia 

Diagiiosts: Thalhis usque ad Halnthny ambivva, 
sod filis assiniilatiscotticatis stutti--. lililornUba^uft 
net arcnatis cum retinitis intlaiis. atque sporati^ns 
uniloculariis pyrifortuibu^, sporangus plurilocalanis 
mtcrcalaribus ah celtuhv fili longissimi. 
Typp locality; Aldinga red, S. Aust., m shallow 
pools and gutters especially rtea' ihe souMi-rn 
seaward margin ot Mu plaMorm. 
ffolntype; AOL', 32664 iUooiersley. 29 viii.1968). 
Diuribuoon: Only known from type locality 
(Aldrngareef, .S, Ausi) \Skinner 14j\.I977; ADD, 
A48249 and A48250; L5JU978; AOU, A49553). 
1 here arc two specimens, MELUI, 21072 and 21143, 
on Hneroz.mi&Q from Incalmes Id the Port PhilGp 
area of Victoria, which are labelled H. lr/tnhn'cali\\ 
but Ihe plants dr^ very small, WOJ very ienile and 
not idcnofiat/le heyond getlUG 
II oM; Helmaosteru tasnuinica, 
Specimens examottvi; /\\ above, 

The niactothalliat srage is present for a very short 
season, hence Ihc specific epithcl, It has been found 
only during August and September, for ;i period o\' 
less than six weeks. Thin species differs trom H. 

hig, i Hftihtsut Mifftfri I li-.n>nicin (ABU M50ffi) A to D all part ol one fiJaineniLnis qyglcTrL A tipper medullti 
vsiili union jfiO main )uiv\ias\irniiaii>r. U. liiwer medu&fl with IraOfiVprw ffVamcntfi «",0_ Mt^Jnll;iry:*ndrof|ic:ii side- 
brancli . ..-ion,.. 



ambigua Yamada (based on a specimen from 
Muroran, Hokkaido, Japan {Kurogi, 8.V.I978; ADU, 
A49376) by possessing straight, not curved, slightly 
medianiy inflated cortical filaments, and an almost 
cylindrical, not ovoid, unilocular sporangium. The 
differences between this species and H. lumbricalis 
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. ephemera/is 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. 


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 Maequarie 
Island. E. 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 Rottncst 
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 Ecklonia 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 E. 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 
E. intermedia Crouan & Crouan, E. 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. lubrica as modified long 
assimilators, not as hairs as Jaasund (1960) had in 
separating E. lubrica 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 1: 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 et at. 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-I6 loculi {E. scutulata may 
have 16-24 loculi). In E. lubrica Ruprecht, 



plunlocular sporangia may also form laterally to 
cells of tong iissuntlaiors (all hough the origin and 
function of these structures is interpreted otherwise 
by Pcdersen (197*))). Pedersen (1984) provides strong 
evidence of conspecility of L. lubrica and E. 
ftuirola. There are Uvo subsections. 

Subsection 2ft Medullary filaments close packed, 
branching from most cells, or with short inlcrnodes 
of 10 or fewer cells, with concentration of branching 
in upper medullary filaments (to form a subcortex) 
in some species, pluriloculai sporangia present in 
many species, laxa included: Gonodia murchaniav 
Setehcll & Gardner 1924; Eluchisiu coccophorae 
Takamatsu |938a. Nodd I9C9; E. antarctica 
Skottsberg I9$t} £ lubtua Ruprecht, Kuckuek 
1929; £ fuctcotu < Velley) Areschoue, Hauck l«H^; 
E mcridionabs Skolisbcrg 1907; B. scutulata 
(Smith) Arcschaug. KjHIman 1897, Hamel 1935. f: 
Jlavcida (Dillwyn) Aresehou^, Hamel 193:* 
Takamatsu 1938a; £ globosa Takamatsu K9fBa> 
Nuda (969; E. claytoniae Skinner sp. nov.; Gonodia 
johmtonii Setchcll & Gardner 1924; t. -o-terae 
Noda in Noda & Kitami 1971; L mollis Takamatsu 


Subsection 2b. Medullary filaments loose, 
branchine at intervals of (5-HO-I5 cells, with upper 
blanches forming a subcortex to support the cor 
assmiilators and unilocular sponmem, ;md lower 
branches of the medullary filaments modified to 
interweaving transverse supporting filaments; 
plutilocula/ sporangia unknown 

Taxa included: Etachista bndaueri Chapman I9rd; 
E. australis J. Agardh 1882, Kuckuck 1929. 

Section J: Plants epiphytic with discietc basal layer 
often with pcglike projections from basul celh and 
rhizoids from lower medullary filaments, two 
morphologically distinct kinds of medullary 
filamcnls, one with broadei cylindrical to ovoid 
cells, supporting the long assimilatory filamenK 
and the other of narrower elon^ate-ovoid Lelis. 
branching throughout supporting the cortical 
assimUaiorv filaments and icproductive organs 
unilocular sporangia presern jii some -.peuev. 
phtriloi ulaf sporangia 111 corymbose croups 

individual sporangia with i"Wi-)24-4-S IftCnfl. I Wit 
SCCOUd kind oi medullar filament may arise 
laterally from the first k«nd of medullars- filament 
or separately from ihc basal cells. 

Taxa included: Etacfustu mgto Takamatsu 19iKa; t 
intermedia Crouan & Crouan <= E. nettectc 
Kik kuck l929)Sauvugeau 1936; E. tenuis Yamada 
1926* Takamatsu 1938; (fed I paci/ica), Noda 
1969, 1974; Mvriuciula surpass* <Ycndo> Feldmanu 
J943. Noda !9M*j £. orbicularis (Ohtaj Skinner 
IWJ, Noda 1975, Ohia 1973; Gonodta fiisiformit 
Noda in Honda \%% nom. nud 

Taxa excluded from Eiactusta 

Etcihtt-ia ra/uosa Skotisberi; \Hffi = Gononema 
ramosa (Skonsbcrg I Skoi I sber^ 1921. /:, .taw™ .„■/<•-. 
Skotisberg 1921 = Myrtacntla roxariotde* 
(Skottsbcrg) Skottsberg 1953; £ pustltu Skonsbcrg 
1921 =■• Myricn tula pusilla iSkoihttGf^) Skottsbcrg 
1955. The following six taxa are described or figured 
gS i'dving true phacophycean half sand should thus 
b€ Included ill BentO if* t nc Lcatitestaceac: E. 
minuiissimalfyiCn 1928, 1960; £. 
1938a; E taeniaefonnh Yamada 1928; E MUasa 
TakamahiU 1938a; E. sadoerrsis Noda 196$; E. 
sargossicofa Noda I9f\9. 


Much of (he present wtk was undertaken as part 
of the author's doctoral research, supported by a 
University Research Gram at the University of 
Adelaide. The work was completed while Use author 
was a icscareh associate with Prof II. IIS. 
Woroersley with support from the Mamie Sciences 
and Technologies Grants Scheme. Much thanks arc 
due to 1>r M. Kurogr ol Hokkaido University 
Japan, Dr M N. Clayton oi Monash University, 
Victoria, and L>i R.J. Kmy •. ' n verity of New 

South Wales for the loan of specimens l'ai tlcular arc due to Prof. H. B. & NVonicrsley, and 
also Dr M J. Persons ot D.S.I.R., Chtistehurch, 
New Zealand, for icadinfi and discussion of tKe 

: 'YcikIo ||M20> described (pis IftWn ai Isovb^fi occauonul 
hair\. but Noda\ ilV64htrsi -riuiion rtnd UhiMraiimesare 
■\iiliout halts, ttiere may be two insni^-u \ . 


A(.MU)ii, J. G (IK48) "Species «ener« et ....wmos nt 1oni, 

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(1882) Till alfcrncs syMeinaiik.. Nya bidran Acta 

Urtiv. Lund 17, I M6. pK 1-3 ' " lv ' • - 1 - E. (IH30) 'BOiamCOtl tfatllCUfm 1 II U^"- | 

I. U (IMVSj "Sylloge Alparutn oinnnini 
i.m> ir>i|nr Lo^nirnrum.'" Vol. 3 tPaduaj 

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IhC PhmcophytCiW o( New Zealand. Bull. Res. 0>.^/ 
fsntrt, tfd n mil, i^-'.h, 

liniMANN, I 11943) L'ne nouvrttc rspe<.x: -IcWt/J' 

parHMir dii tjtoalarfa wrvtata I. Ap. iSutt. Sen- fif$f, 



Guiler, E. R. (1952) The marine algae of Tasmania. 
Checklist with localities. Pap. Proc. R. Soc. Tus. 86 

Hamel, G. (1935) "Pheophycees de France" II, 81-176. 

Harvey, W. H. (1846) "Phycologia Britannica" Vol. I. 

(Reeve: London.) 
Hauck, F. (1885) Die Meeresalgen. Vol. II. In L. 

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Jaasund, E. (1960) Elachista lubrica Ruprecht and 

Elachista fucicola (Velley) Areschoug. Bot. Mar. I, 


Kjellman, F. R. (1890) "Handbook i Skandinaviens 

hafsalgflora" Vol. I. Fucoideae. (O. L. Lamms: 

— — (1897) Phaeophyceae. In A. Engler & K. Prantl 

"Die Naturlichen Pflanzen familien" 1(2). (Englemann: 


Klotzsch, J. F. (1834) Ericearum eel. Adelberto de 
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Kuckuck, P. (1899) Beitrage zur Kenntnis der 
Meeresalgen, 5-9 Wiss. Meeresutersuch., Abt. 
Helgoland, N.E 3(1): 13-82. 

(1929) Fragmente einer Monographic 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, II: Phaeophyceae. 

Nova Hedw. 3, 129-350, pis 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 pis 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. 

Noda, M. (1964) Marine algae in the vicinity of the 
Shioyazaki Cape, Fukushima Prefecture. J. Eac. Sci. 
Niigata Univ. ser. 2, 4, 33-75. 

(1969) The species of Phaeophyta from Sado 

Island in the Japan Sea. Ibid. ser. D, 6, 1-64. 

(1974) Some marine algae collected on the coast 

of Kashiwazaki Province facing the Japan Sea. Ibid, 
ser. D, 11, 65-74. 

- (1975) Some marine algae further added to the 

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& Kitami, T. (1971) Some species of Marine 

Algae from Echigo Province facing the Japan Sea. Ibid, 
ser. D, 8, 35-52. 
Ohta, T. (1973) Some new and rare marine algae from 
Tsugara straits between Honshu and Hokkaido. Ibid. 
ser. D, 10, 11-28. 

Oltmanns, F. (1922) "Morphologie und Biologie der 
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(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. 
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Phaeophyceae. Mem. Acad. R. Sci. Lett. Dan. 

Copenhague, Sect. Sci. 9, 6(3), 40 pp. 
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Contributions. VII. Univ. Calif. Pub/. Bot. 13, 1-13. 
Suva, P. C. (1959) Remarks on algal nomenclature, II. 

Taxon 8, 60-64. 
(1970) Remarks on algal nomenclature, IV. 

Ibid.941 19, 941-945. 
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orbicularis (Ohta) comb. nov. (Elachistaceae, 

Phaeophyta) in southern Australia. Br. phycol. J. 18, 

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Ergebnisse der Schwedischen Sudpolar-expedition 

1901-1903. Vol. 4. 
— (1921) Botanische ergebnisse der Schwedischen 

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(1953) On two collections of Antarctic marine 

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Herpodiscus durvilleae (Lindauer) comb, nov., a 

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- (1938b) Halothrix aus dem nordostlichen Honshu, 

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34, 1-12. 



byK. 7. McNamara 


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. 



by K. J. McNamara* 


\1< N\m\k\. K. I <iyX5) I he spatangnid echinoid Limhw imiu flu Ian I ticenc or southern A.i-.n.a..i 
Trans, (i. Sot S. Attn. 100(4), 161-165, 29 November, I985. 

l-ivr I -| rumen,.. DUG from Hit' I ait EpCBIW Tniacliilla I iiacsUMu- \f\ SoUtH Australia, and font from 
[he Pallinup SiltfttOriC m !R>Utb*WC9lCrT1 AUfitWlHD lit described as anew species, Lnuhtu />///. Aw ..p ITOV 
I he absence n! j tateroaflul fascials is nm ihouyin t<> be "I JufTMeirl importance to warrant pLumc tliil 
Species in any jvnns oihn ihun .' Jur/f/tf t he concept uf lite gCJlV) Cvttftlt) W CfflCOdCd 10 IflClUde forms 
both with and without a fotent&ual F&CiOilfe 

Ki > W'i.kds: Spauin^oid CCttltOtd, <""/"■' BOOCHC, IWW KpCCWR. *mthwn Aumralia. 

Within Illc leitiaiv sequence "I SOUlhBTTI 
Australia the rich spalangoul echmoid lauua has 
yielded a number of forms which, at sometime or 
other, have been assigned to the genus Linthia. late 
(18S5) described Unthia aniiaustnilis from the Early 
Miocene of the Murray Rive! cliffs; II. L. Clark 
(1946) placed Mexulasrer compressu in / huhh; and 
Pritehard (1908) placed two ol McCoy's (U 
species. PerkosmUS ttetsotli and Pericosmus 
in I.ittthiu, alone with a new species* L. 
mooraboolensis. However, as discussed below, 
of these species belong in the genus Unthui 
Consequently, (his common, cosmopolitan, lettiaiv 
genus has not previously been described 

Il is the aim ot this paper to describe Wh 
only the largest known spatanyoid echinoid frOCQ 
the late Eocene Tortathilia I irtiestorte in South 
Australia and the Palllnup Siltsioue in Western 
Australia, but also to record the presence of / mthta 
in the Australian Tertiary. 

The material upon which this papei i, 
consists of a single specimen from the TbrlaCl 
Limestone in South Australia and four from the 
Pallinup Siltstoue in Western Australia. Although 
the Tortachilla Limestone specimen dots net l*AVC 
its adoral surface preserved, much of its aborat 
surface is in an excellent state of piesct vahoii. I hree 
of the specimens lioui the Pallinup Siltstonc arc 
preserved only as inteuial moulds, but with both 
aboral and adoral sut faces preserved. I he olhcr 
specimen is poorly preserved and Consist? Ol Ml 
external and internal mould. Sufticient details are 
preserved in the live specimens to be confident that 
they arc consprcific and represent a hitherto 
uudcNcnbed taxou. 

Measurements on the specimens WCTC made with 
a vernier calliper to an accuracy of 0.1 mm A 

We tern Air.h.iimi. Mus.-um, taancis Srrrrt, Penh, 
m Australia nkki 

number ul parameters ate expressed as percentages 
oi' [tUlXimUm M»l length ("'oil.). SpccunctLS arc 

touted m iht collections ol the Museum ofVjctoria 

(NMV.l; Western Australian Museum (WAM) and 
Geological Survey of Western Australia (dSWA). 


The Tortachilla Limestone is a yellow Ixown, 
preen and jeiey bioctastic limestone which in outcrop 
m Ik Wfllimga fcmbaymcnl of the St Vincent 
Basin, South Australia^ attains a maximum 
ikicXncss of 2 m in cliffs at the south end of Maslin 
Beach (Reynolds 1953, Cooper 1979). It la early t Ate 

mc in age <l udbmok 1963; McCiowrun I97X; 
I Intbay 19X5). It contains a rich marine invertebrate 
fauna, predominant amongst which are the 
e-chinoids. Most forms from this unit have been 
described, including the tollowmg: Sterrocidarts 
tutimoret Philip, 1964; .S: fasten Philip, 1964; $ 
Uiermla Philip, 1964; S. hispida Philip, 1964; 5 
intricate Philip, 1964; Sa/cnidia tertwna | Tate. IN I 
(sec Philip 1965); OrthotophUS tnitrwn Philip, 1969; 
Tatvchimis mtdtti Philip, 1969; Fibufaria Rre&utu 
Tate, ISM; Frhinufampas postWO&QSsa Gregory, 
1890 (see McNamara & Philip 1980a); Apafopygns 
VOtCtHtittUS I late, 1N91); Auxtrutanlhus tonyianus 
(Grc»or>; I8?90>; i^isoiampas cancinno Philip. 1963; 

•aster lerttttttUS (Ot^cory, 1890); G bettixute 

r & Philip, I97fc GUtechMus cudmaret Ml 
1964 (sec McNamara er at. in press). Hemtostet 
(Botbaslerj sp. nov. ( MeNam a r a , in press); 
Prrna\tvr aitfmgcnxiv Hall, 1907; Pwtewsfer 
prraustralis McNamara, 1985; Schizas(er (Paraster) 
rata McNamara & Philip. 1980b. 

Lhe Pallinup Siltstone> a member of the Wen ilup 
Formation within the Plantagenci < iruup, IS a 5*Tt- 
SKOtlT of spongolite which occurs discominuously 

. the vouthern coast of Western Australia (sec 
Darrajih & Kendrick 1980. Fig. IL Ibis richly 
rossinlriMic uriii is considered to have been 
deposited m a shallow she-it env i :Darraab 



& Kendrick 1980). Echinoids form a major part 
of the marine invertebrate fauna, but remain 
essentially undescribed. Gil/echinus eudmorei has 
been recorded from the unit (McNamara et al. 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). 


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 111 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 Brisasier; incomplete, as in Tripylus 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 & Thie'ry 
(1925). Bather noted how specimens from certain 
localities show the lateroanal fasciole (for example 
Bather 1904, PL 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 111 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 



i* * 




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 xl. 

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 



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 ot 
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 Hat; 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 

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 et 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 

A number of species from the Australian Tertiary, 
as noted above, have been placed in Linthia. 
However, none belong in the genus. Linthia 
antiausiralis (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 
moorahoolensis (Pritchard, 1908) to belong in 
Victoriaster. Consequently, the Late Eocene form 
described herein and placed in Linthia, is the only 
unequivocal record o\" the genus from the Australian 

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. pulchra 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 L pulchra, a deep ambulacrum 
III 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, P1.4, 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. subrhotnboidalis Herklots 
(Jeannet & Martin 1937, Fig. 61). This rather worn 
specimen differs from S. subrhomboidalis (see 
Gerth 1922, PI. 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. 


Prolcssui G. M. Philip and Mr J. NT. Lindsay are 
thanked for reading the paper and offering 
suggestion* Fur its improvement 



Kmolk 1 V (1904) eelmioicU from Sokoiu. 

COOL Mag Dec 5(1 1, 2V 2-304. 
t i Akk, II. i (1946) The oc dnodi m. lauiw "' \u*tt&Ua 

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1 «7 
1 OOKE-, C. W, {195(5) tcno/ok u'chinoiclv ot Eastern 

United Male*. Prof Pap. i ' $ Geol Skfl 321, 14(16 
CoOWA, K I. (1979) t : ot.cMc lo \1io,.r.h: ^r,u Mi-aphy uf 

Ihv W. Minima » mbavnicnt. Atep. Pep/ Mines Encix*' s 

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Haiir, H. $>& Philip, Q m (J878) Tertiary holastmiid. 

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I -i i 'in, it. 11922) i ahmoddea /« Wb fbsrittcn von Js 

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& Thifry, P P>C>9-I925 Lssai dc nomenclature 

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I IMI'AV, I M |I9MM A.-|H t «'. Q| 50lHh AiiMiaha • 

ternary form (feral biotirarigrflpltt *Uh cmplia*i& on 

iludiuol Mamihrm and Subboifatt. In I M i indxay 
(Ld.), Stratucrzptary, palaeontology, roatacDlog) i'.i|H q 
in honour of Dr Nell 1 udbrook . sjpc*: ftv/;/. ppm Af/wt ". 
Etmjp S. iusi -\ is' 
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1 1 Dma 'k. N. H. (1965) < orrelatlon t-t the Tertiary rocks 
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Tohokti (Ini. Set. Pc/'t. 2nd ken fGtaL), Spec. Vol. yva 

■ 141-261. 
McCov, I-. (1KK2» Piodiouius 61 'he palaeoniology 01 

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Volume: Lssays in honour ol hene Cicspin. Huff, Bur, 

Mit„r ReSOUr Geol G*O0hys *MS?> l'>2. B 
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living specie* oi Bteynki (Echlnoldea; Spatau^oidaj 

rrom Ausiralia. Rev. West. Aust. Mus. 10, 167-197. 
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— in press Taumoiiiy and evolution "f h-niary 

speetes of Hetnlustar (Spatangflhia: Hemiasteridae) 

from -\jj.siralia Ptiheo/itofogy 

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iV 11980b) AuMralianTcniarv (icbizasierid 

eehinoidv Alchennga 4, 47-6S. 

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% ffi KRUSEj P 1>, in press. Ternary brissid 

Cchinokls 01 iOUthern Australia. A/cherinxa. 
MottrCffSEN, T (19511 A monograph Of Hie I *'liui*m1ea 

5i-'), ^paiin^oidea II (Reitzel: CopenhaKen). 593 pp. 
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SttiaJ Qeol Milg d?e 5. vol I, 441-445. 
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CCh lAOidS, fttfueunfoluyy 6, 718*726, 

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Australia II. Cidaridac (2). Proc. R. Soc. t'u: 77 T 


(19t.S) i he Iciiiaiy t.vtniioio\ ot souih-easlci u 

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Camarodonta (I). Proc R. Soc. Vk. 7K. LSI 196 
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2^ 275. 
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Lifithia in Victoria, with description of a new tpeocfc* 

i'n.-c, R Soc. Vic, n.v 21. 302 400. 
RETIMQLDS, M. A. (1953) The Cainowi.- succession of 

Maidin and Aldinjia Bavs, Soui'i Au>fialia Trans R 

■S".- V AUSt 76, 1 14440. 

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Q. JL y.col. Sen ijon<U>H, 33, 256-258, 

(ihks> Ml$ccft$n«0U8 comtibuiious io ihe 

pataeoMiolo^v ■■! ihv older rock* of Australia. Stti. Sa, 
fa, N S. I, (-5 

llMVIi \ MUllography .m.l revised list of the 
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14. 270-2X2. 


by Robert T. Lange 



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. 


by Roblrt T. Lance* 


LanU:, R. T. (1985) Spuria] distributions ol stocking intensity produced hy sbeepflocks grazing Australian 
ehenopod shrublands. Trans. R. Soc. S. Aust, HW4), 167 174, 29 November, TW5. 

The distribution ol estimated sheep stocking, intensity (ha sheep ') is described from within each o! 
several paddocks (large wire-fenced subdivisions) of and /one station* (ranches) in Australia. The estimation 
method is based on demonstrated proportionality between time spent and egesta deposited flfl sheep tn 
the localities (hey visit, Lgesta deposition measurements of which the intensity estimates are transformations 
aie shown experimentally to correlate with immediate sheep effects upon the vegetation. 

The magnitudes of estimated sheep stocking intensity are shown in vary greatrv from place ID place 
wiiliitt the one paddock, In the studied region about one third of each paddock's area esperieneed intensities 
lusher than the paddock average, grading upward; to about su times higher, while about two thirds 
experienced lower intensities grading down to ?eio. Implications tor perennial vegetation in the paddock'- 
arc discussed. 

Krv Wmkiis: Chenopod vhiuhland, sheep grazing, stocking pressure variation. 


Sheep pastotaltsrn in the Australian arid /.one is 
notable lor its great wool production but also lor 
the damage which the sheep do to (he perennial 
vegetation. There is 3 general belief that there are 
sale slocking intensities which do not harm the 
vegetation and excessive intensities ("overstocking") 
which do. Many authors have used stnall-paddock 
experimental irials (of area one ha or so) to 
demonstrate the relevant relationships between 
stocking intensity and vegetation reactions 
(Cunningham & Walker 1973, Hamilton & Bath 
1970; Harrington 1979; Leigh A Mulham \96$; 
Letch $ Wilson 1969; Leigh, Wilson & Mulham 
1968. 1%°-; Trumbie & Woodmffe 1954; Tuppcr 
1978; Williams 1970] Wilson 1974, 1976; Wilson & 
Uiaet/ 1980, Wilson, Leigh & Mulham i%9; 
Wilson, Mulham & Leigh 1976). 

The application of (heir results to full scale 
pastoral paddocks is complicated because flocks do 
not spend Iheir linie evenly throughout the large 
ana;-, m which they wander (Waite 1896; Osborn, 
Wooil & Pahridgc 1932; Valentine 1947; McBnde, 
Arnold, Alexander <V lynch 19rV7; Barker 1979; 
Squirt* 1970, 1974; Whalky. Robinson & Taylor 
I97X), So although the paddock average stocking 
intensity may be known (PSJ = enclosure area 
divided by flocksize), the intensity on any actual 
part (SIP average stocking intensity of the part) 
remains unknown. Without some means ttf 
estimating SIP at given points within paddocks, 
there is no connection with (he experimental-trial 
literature. The difficulties of this situation were 

* Department of Botanv. University of Adelaide, Box 498, 
G.P.O. Adelaide S. Aust. 5001. 

pointed out by Rawe.s & Welch (1966, 1969), Ares 
& Leon (1972) and Anderson & Currier (1973) 

An approximate solution to these difficulties 
follows demonstrations by Rawes & Welch (I966 T 
I969|, Laoge (1969), Ares & Leon (19721, Squires 
(1974), Lange & Willcoeks (1978) and others, that 
quantitative relationships exist between Hock use 
of pasture localities and the amounts of egesta 
which fait in them. From these relationships SIP 
can be estimated. A particular basis demonstrated 
by Lance (1983) is used here to examine Ihe spaltal 
and temporal distribution of estimated SIP in arid 
/one paddocks of the Whynlla region, South 
Australia, during the period 1969-1982. 


Computations and Graphical Summuttes 
As in the study o\' Hilder (1964) the enclosure 

(paddock) is imagined to be divided into many 
equal parts from each ol which egesta accumula- 
tions can be measured. Over any observational 
timespan the exact average stocking intensity OU 
each part (Lange 198."*) then is. 

area of the part (ha) 

SIP (ha sheep ') 

enclosure v fraction of total 
flock size ftoektirne spent 
in the part ( = F> 

1 he use of ha sheep ' rather than its reciprocal 
follows Harrington (1979), Cunningham & Walker 
(1973), Rattray (1960), Goodall (1971) and Vcscy- 
Fit/gerald (1974) because in the arid io\\< tt mostl> 
has values >L Lxeept in small-scale calibration 
experiments (Lange & Willcoeks 1978; Lange 1983) 
thete was no practicable means of obtaining F 



directly because canopies of the western myall trees 
(Acacia papyrocar pa 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 

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 (r 2 = 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 2 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. 




i i i i i 

f1 27 sheep ha" 1 







w 4- 


Ol ° 

1- 1 



I '5 


-1 g« 

-0-5 » 


2-5sheepha 1 Lq.125 

Fig. I . 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 v 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 1 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-dimensionaI 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 triplex and Maireana with variable 
low woodland of Acacia, on undulating desert 
loams. Rainfall is very unpredictable and averages 
about 220 mm yr l (1940-1970). The whole region 
is fenced into paddocks approximately 1200-2000 
ha which are stocked continuously with merino 



r.\Kit I. List of studies undertaken in the Whyaikt region, South Australia, in determine the magnitudes and 
distributions of SIP (stacking intensity of the part of the paddock in ha sheep '} in each case. PS2 paftdotf 

average stacking intensity. 





A pea 


(ha sheep 1 ) 

Dates Number 
qT pans 









( >\ crland 






!'"'! djt 


6 5 

6 5 

Aug. 1:969 

Feb. 1970 

i nc . 

Ooi 1971- 

May 1972 


June 1972- 

Sept. 1973 


June 1978 1979 


Sept. 19K0 
Nov. 1980 


l;tu. W&i- 
Mareli 1981 


June 1982 
Sq.l 19«: 





• f 



sheep at PSI ot about 6 ha sheep ' (Lange, 
Nieolson & Nicolson 1984). 

Validation Experiments 

Validation that cgesta accumulation was roughly 
proportional to flocktime spent by sheep in parts 
ol I lie discussed paddocks was published by Lange 
& Willcocks (1978) and Lange (1983). 

In | he work ol Arcs & Leon (1972), validation 
pf cgesta accumulation as a useful variable to 
measure in pastoral research rested on the 
persuasiveness of its correlations with stock effects 
upon pasture plants. The same applied where 
distance from water was used as the stocking 
variable (Osborn, Wood & Halt ridge 1932; Valentine 
1947; Lange 1969; Barker & Lange 1969; Squires 
1974; Gracr* & Ludwig 1978; Barker 1979; Latchen 
& Lange 1979). To provide an equivalent soil of 
validation that egesta accumulations (and Ikik< SIP 
estimates) correlate with concomitant effects on 
vegetation in the Whyalla region, several 
experiments can be cited, 

Lange (1984) showed that the observed proba- 
bility of random outer shoots of saltbush {Atrtplex 
vesivaria Hcward ex Bcnth.) 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 

v = ll.SO i 0.17.x; r 3 = 0.78, p <0.01 

where v was percent loss of marked shoots over a 
6 week period and .v was concomitant sheep egesta 
accumulation in kg dry wl, on plots 60 .-; 40 m. 
Across the different parts of small experimental 
paddocks stocked heavily, the relationship on plots 
10 >- 10 m was much clearer, namely 

v = 0.416 3 0.004,v. r 1 - 0.91, p < 0.001. 

One further validation experiment* is reported here 
to extend demonstrations of the usefulness oi cgesta 
accumulation as an index o\ SIP, at least in some 
contexts. Stipa nitida Summerh. & C.L. Ilubbaid 
and Danthonia caespitosa Gaud, in Freye, art 
prominent tuft grasses occurring between hushes 
in mixed chenopod shrubland of 2-Mile paddock 
(map and description in Barker 1979). A tract ol 
this vegetation 30 « 200 m was fenced to include 
a drinking trough and was surveyed into Hi equal 

In each part all grass tufts >5 cm tall on a plot 
20 * 0.2 in were estimated for biomass (g. di y wt., 
technique of Andrew, Noble &. lange 1979) and 
measured For height (cm). The enclosure was then 
stocked with 10 merino Sheep for 6 days, the grass 
then remeasured and cgesta dropped by the sheep 
was collected from plots 28 - 2 m superimposed 
over the grass plots. The prediction was thai egesta 

* This experiment was par formed hy Chester J. Merrick, 
an Honours Mudeui in the authors programme. 


R T I \NU.l 

accumulations would index reductions in grass 
height and biomass via regression equations. 

Fuilscaie Studies of Paddock SIP 

Table I lists studies which were undertaken 
between I9tV>and 1982 (see Barker 1979, Noble 1979 
and Lange, Nieolson & Nieolson 1984 lor paddock 
maps). Wanga paddock lies east of Werugo and 
Overland lies north of 1-Mile. 


I iilkiuliun Hx per intent 

The experiment yielded the very highly significani 
multiple regressions: 


= 0.99, 

36.99 - 0.47.V 

/.. < 0.001 

final mean grass height (em), 

egesta dry wl (kg) 

initial mean grass heighi (cm) 

14.11 O.I5,v . 0.95c, r =0.99, 

p <0 001 

tiital jirass biomass (g), 

egesta dry wt tKe) 

initial grass biomass (gj. 

f-uli Scale Studies 

Data from the seven separate paddocks were all 
the same in principle, differing only in detail of 

Cumulative Paddock Area 

0-5 1 

1 I 1 I I I I I I I 













Not Visited w 

Pig T-, The cumulative dismhuiion ol 10JL -SIP for ilir 
50 puns ol < htrUind Paddock June I978.1anuary 1979 
inclusive (.Sludy 4). 

spatial pailein and particular history, A selection 
of data is presented here as a basis for discussion. 
None of the omitted cases tells a different story or 
contradicts the examples presented. 

Over/and Paddock (Study 4) 

Tig. 3 shows the cumulative distribution of log. 
SIP for t he 50 equal parts of Overland Paddock 
for the period June 1978-January 1979 inclusive. 
Pour parts accumulated no egesta during this period 
so SIP for them was do ha sheep '. Over the used 
parts or the paddock SIP values graded smoothly 
upwards from PSI, over about one third of the area, 
to a highest intensity about 2.5 cycles (6v ) above 
PSI. Over about rwo thirds of the used area they 
graded smoothly downwards from PSI to lowest 
measurable intensities about 4 cycles 06*) below 
PSI Only I of the 50 parts actually had SIP PSI. 




Fig. 4. Perspective view of a 3-dimensional graph 6hoMn£ 
the distribution of SIP values in Overland Paddock 
(StuJv 4). Swamp Ham i^ the southern warci poiui 

r-ig 4 shows the corresponding SIP surface of 
the paddock with log_, PSI as datum arid with 
class-intervals of I log, cycle. This spatial pattern 
had the following weather and flock management 
history. In June lv>7X drought had restricted 
drinking sources \or sheep in Overland paddock to 
Swamp Dam in the south part. The flack was 
observed to depend on Swamp Dam until it dried 
out. Then a temporary source of water was 
introduced (by pipeline) to a point at the west end 
of the north boundary of the paddock, to which 
the sheep were introduced. 

Two- Mile Paddock (Studies $ 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 these 
distributions compared with a theoretical normal 
cumulative probability distribution for the 52 pans 
o\ the paddock. The diagonals represent required 
lines for perfect lit and the dotted lines show 
observed fit. 

Rank order correlation ot spring and summer 
SIP stores across the 52 pans of 2-Mile paddock 
was not significant, implying I hut the flock shifted 



Cumulative Paddock Area 

Fig. 5. The cumulative disiribution of log. SIP lor ihc 
52 parts of 2-Mile Paddock in spring Trine line) and 
summer (heavy line) 19R0-8L 

its pattern o( paddock use from spring to summer, 
1980-KI. Hence the resemblance of spring and 
sumnicr 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 

HfartigO Paddock (Studies 2 and J). 

Fig. 8 shows the SIP surface of Wertigo paddock 
lor the peiiod October 1971-May 1972 inclusive 
with log_, PSI as datum and SIP in class-intervals 
ol I log, cycle. The flock was observed throughout 
this period to use only the southwestern water point 
and lo graye away from it in two directions, namely 
southeast wards into ihe 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 tfce 
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 
(/■ - 0.846, p < 0.001), indicating relative stability 

Theoretical Cumulative Probability 
t Normal Deviate ) 

Theoretical Cumulative Probability 

( Normal Deviate ) 

Fig. 6. The cumulative distributions ol log, SIP for 
spring and summer 1 9K0- 8 1 in 2-Mile "Paddock 
compared with theoretical normal cumulative 
probability distributions. The diagonals represent 
required lines tor perfect tit and (he dulled lines show 
observed lit. Spring above, summer be!ow_ 


Fig, 7. Perspective view ol a .Vdimensional graph showing 
the distribution of SIP values in 2-Mile Paddock for 
the combined period Spring and Summer 1 980- 8 1 

(Studies 5 and 6). 


R T I MV.,1- 


WP — 


, eiiveview t»fi hdiincwli nnl nwph showing 

il,r OLISCI iPUl Oi ^H* ^ulu:-. it. Wcrtigo rSttltt&Cfe It 

.i :■ I I Mav W2 fnc (Suicfy 2). 

! ig, 9, Perspective vn;v. <>t ;1 l-iiirn'-'tlMOnil Ciaph sli" 

.in h-ii iimhoh ut Ml' value ni Wi rti - Paddocli fal 
mm period rime \yii SW |,,M mc. (Study 3). 

,.| us.- pattern, kangaroo Ctfcsta R£OV< red Inuu I be 
■•. tl in. ,i i i .-,o ciinelalcd betVffCtl periodfi {r 

0.39 ,• Q.05) 


Tin- dcinonsiialions bj RaV^S A We!>.h i 196ft. 
t9$) and by Art's A I eun UV72) tl..u measurements 
ol egesia deposition in open nmllfocl 
index stocking intCfwiiiesol iheep, ate -Mui-ledby 
the present data I tiai extension U phi merel) iu 
the Australian arid /one context, but also toward • 
s itft>r* qtianrihilive R>otfni 

Data icportecl hen! ilio* *e*J r ,mi 
quantitative relanonshipv bdwecn sheep e| 
depositions and aimuHaneous effects on pl&ntt at 
1, valines thniughoul the range of the rrcHy m.-; 1 
stheep flocli 1 unherfHO>nj ihflKje relationships havp 
need demonstrated over time intervals u> brl 

to admit no scope fOI Interpretations 0«Hef than ui 

I. deposition a* an index o\~ sirouttasi 
stocking 01 anuinc presort The onli ^ I 
conclusion from the regiessfion equations ol 1 
validation experiment is th*t egetta accumulation 
(»ii plots indexed the sJnwftaneow Ions of gr&sa 
1 ;nui hiuiiKiss caused by the sheep, and from 
. .1 .in 1. m 11. 1 iinfiv (1984) thai it indexed 
tancous 1 1 tahhush sbi 

[b ,.. ; the SIP estimates reported lien! 

airtllrect Tiansformatkmspf the cgesta deposition 

data, (he SI P estimates also relate to effects of Btt )l k 
on vegetation. Ii is reasonable from the prexeii! 
IcttCfl to suppose, if plants are being BffeCW?d h 
stock in their paddock, iliai the plant* are hkclv 
be affected must where SIP estimates are highest 
and least when: ihey aie lowest. But the particular 
advance introduced here is Ihe capacity to place 
quantitative estimate* tha sheep ') upon the SIP 
at particular places within Hie ranges ol 1 locks, and 
thus to link those places with the extensive small- 
plot experimental literature mentioned in Ihe 
Introduction. ' he results of ejtantinirti ,luvsc ntJ ks 

will he discussed elsewhere; the aim here is to discuss 
inlrapaddock SIP loi its own sake- 
Much evidence indicates fundamental similarities 
hi the relationships of SIP in all paddocks examined 
from the Whyalla region and exicudinn, 50 '■" ftS 
data exfsi t 10 published CBSC8 from elsewhere In 
Australia, thus the logarithms o\ SIP values in e 1 
ol Ihe examined paddocks were appioximatcly 
normally distributed Something intrinsic in the 
behaviour of sheep flocks is suggested by that fact, 
constraining them to distribute their flocktims very 
UDeVenly, on average, throughout their range Lest 
that be attributed simply lo the vast range area 
available lo relatively small Hocks in Ihe Whyalla 
region, or to the piosphere ctlect (Barker 1979), il 
should be recalled that the same applied even to 
Hildcr's data concerning 20 sheep penned insni. ' 

That inflexibility has important practical 
implications The first is thai the more even spread 
of flocktime throughout each paddock, desired by 
station managers (I ange, Nicolson & Nicolson 
1984), seems less likely to be achieved simply bv 
farther fcneina Of paddocks into smaller areas, than 
was earlier thought. 

I lie second is In relation LO absolute flocksizc and 
paddock size, and PSI. Ir is traditional in the 
Australian sheep industry to make slocking 
comparison* just on the basis of PSI (in ha 
sheep ' or, in wetter mote productive place 1 ;, sheet) 
ha l ), Bill as shown in Fig. 5 ol Lanv.C, Nicolson 
& Nicolson (1984), any particular PSI (say, 6 I .1 
1. 1 p ') is set throughout the industry by widely 
different tombinalions of flocksizc and pauJo-A 
sue. Present results shoyv that even m ihe t olnt iv. Iv 
small paddocks of ihe Whyalla iccuni. S||> , | 
reflect the limiis to which (locks can Or will ral 
away fmm their walet -point. There is no evitK ) 
to suggest that those limits are different in big 
paddock s 

It ought to be evident from this, and bom flic 
inexorable tendency of the flock to even SIP val 
areata than f'si qvcj afcoui Ofictbit^ol the iiock\ 
. i, , rjnai i*si becomes owgressive^ mo« 

b. rni>dcadlQg as an index nl SIP. as paddocks 



and llncksizcs increase ai Ihe erne RSf, Thyt is unless 
the larger paddocks have- cot respond ingl> nunv 
watetpoiitls, winch in many Australian cases they 
tin not. 

Thcre arc olher applications o\ the present 
approach which arc independent ol considerations 
of wool production and pasture. Civet 60 species 
ol rare and threatened Australian endemic flora arc 
to be found scattered within the sheep paddocks 
that enmesh Ihcir remnant distributions in this 
region. Clearly I heir lale is closely related |o the SIP 

they experience, whether high 01 low. It is important 
to establish what then situation is, by means of the 
present technique, as part ol developing 
conservation measures for ihem. 


Ihe author mosi gratefully acknowledges the 
financial support of I Ins work piovided by Ihe 
Australian Research Grants Si heme, together with 
support from the Bioken Hill Proprietary Co. and 
Miisubishi Motors (Australia^ Ltd. 


AnniusuN, \ . W & D lU'.UR, NV. i. (197}> lA'duai'me 
/ones ot unli/atum. / ft$H&e Mtl*4)& 2ft, X7-9J, 

Asit.krw, VI, II . Nutdt, I. K. & I \K'.i, R.T. (1979) A 
noil -destructive method for estimjimg the weight Dl 
forage on shrubs. AusK kdttgd. ./. I, 225-231. 

Akis. J. Q A Ltffli H I. C (1972) An eeoio-Kal 
assessment ol ihe udhieuee ol gwz?o§ on pltim 
eummuntiy ^t i net li re ./. Uoi M), 125- 3.1(. 

Bakmk, S. (1979) Shrub population dynamics Jltdet 
gia/iny— within paddock studio*. In R Ciraet/ A. k. 
Howes teda) Studic* fll the Australian And /one IV. 
Chenupod Shmhlatids. C.S.I.R.O, (Ausi.) Div. of I ;md 

Kck Manag.. pp. 83-106. 

tSc I AM a , K. 11 (1969) L fleets ol modernie %heep 

stocking on planl populations of j biackoak«hluel»ir..ii 
a.ssociation. ,-l^s/. ./ jJW, 17, 5£7 53*3 

c:i'NM',<.i!\M f o, vi. ^ Wm Ktc, p, ., ll97|) CJit>wrt and 

survival ol itiulgu (Acttau arwutv 1 1 Ytuell. ex Benih.) 
in western New South Wales, Ttup Grossl. 7 69-77_ 

Iao Hew, 1. I. & tftwOE, R\ T. (1979) Piosphere pattern 
and dynamics in a ehonopod pa si ore graced hy cattle. 
?« K. timet/ & K. ho\vex(eds}Suidu M -ol the Australian 
\itil Ame IV c'henopod ShruMands. (_ S.I.K.O. t Ausi | 
Oiv, ol Land Res. Vlanati., pp. 160-169. 

(iOoom i , D W (1971) Rsiensive ^ra/ine systems, (_ h. 9L 
pp 171-187. In J. B. Dent & I. K. Anderson (litKj 
Systems analyse in agricultural mahngemem |Wi)cj 
New York). 

GJ?M t/. U U, & Luowio, J. A. (197$) A method fm rhc 
analyse, o( piospherv applicable io rnngc 
assesMnent Anil RtffffW! I. I, U6-M6, 

— . K Wiisun. \ l"). (1979) An a^wssmcin ot 
herbivore diets in iln: ihcnopud shrublands — hi K 
Criuel/.V k. Howes (edM-Nludte-- ol (he And 
Zone IV. Ctltfttopud ShruhUnds, C.S.I.K.O IAh-i j |>i\ 
Ol l-and Ko'-. M,in w. pn 144-159 

II \mh i<i^.. |) & U\ih, J. t.. (1970) IVrforni.iiKL'il -Ikvp 
■hhI cattle "i;t/._il ^'oaruk-tv and loueihcr. Ausi. ./ t ran 

Mi*™-, and An Hwh. in, i9-:ri ( 

llAUKiNMws 1 , (i N. (J979) I Itf* effect > of leraJ ftdflli and 

sheep on the irflrufa populations in a %enu arid WODcllai d. 

Aum RetnxM I \. 114 u.v 
H ii oti', U. .1. (1964) l"hc disinhuuon ol pi. mi n tl inem> 

bv sheep at paMuu*. Ptut\ Au.$i $pc \n f J rtni. 5, 

It- if, H W. &, M J. (1971). CeiKUOk 

sedirnenis, soils and climates ai Wliyalla. Si Ulll 

Australia, Ucoih-rma -Q, 7.75 30S 
I yva,R. I. (1969) The pmsphere sheep i rack and dung 

paticrus X Aft//'iv M(2riaff. 21, vj6-40i). 

— (I9K.U I'Miirialtoii ol >t>ecp stocking inivnsilv -U 
any locaoon tn and /o t ir p.<dt)ocU /r/y/rs tf S.., 
,1/m/ 107. M7 

(1984) teal nurkms' in um [i i.mtl g."nr/Lne itri|0iev 

Ihw; hi SW V 1rv//|08, tl-IZ, 

. Nitoisr^, A l> A Nuot,so<4. |> A i0.';s4i 
Y.'^'unon mana^eineitl ol cheiuipod ran^elainh. in 
South Ausiralia. Aust Hmgct J. ft, 46 -54, 

A VMi.iX'kKS, M_ C. (I97S) The relation between 
>heep-nmc spent and ej;esla aea4mulaied v\ ultin an and 
/one paddock, \n±r I. hxj-t. Auric. Aft Hush. 18. 

I tlGtii-J H. A Mm ham, W. }■. (I9WS) .Selection of Jici 
by sheep grazing semi arid pastures oil die Riveniu' 
Plain. I. A bladder .-.alihush (Atiiptt\ \feSfCnrj0\ co'roi. 
bush (Kochm up/hitd) aornmui\U\. A</s/. I f \p! \j>r\£ 
An. Hit^h. 6, 460-467 

k Wuso:,. A. 0. (1969) Ulili/auon ot .-ll///,'/r.v 
^ptcies _\v sheep. In R. Jones (ed-> The liiolofv til 
Aliiplex C S.I.U.O. (Aust.) Qlv, ol Plnm Ind. p^r. 97-KM 

, Si MniusM. VV I:. (I96M A ^uh\\ 1.1 

merino sheep yra/in^ a couon-busli {Kovhhi ttphviUn 
er.«sslan J (Stipa variabilis— Danihon'io CV&$0ltQ$tf) 
eommunuyon thcRivmoe f*Ii4in -\u<i 7 \ a He, Res. 
19. 947-^61 

, 61 (I'-'TV) A vluOv ot "sheep glit/Jtm 

a Hetah (Cusuuntm cn\Umo Kosewood (/Uii'ttK/cntlrutn 
oiciftttiuw) xtirtib woodland in Wesiein New South 
Wa)e\ Ausi. / -lent, AV>. 3H, \:Z} 1236, 

Nb tiKiuL, ("1., fosrjii*, o w., ai i\.\NLn .,. < i, \ 
1 vw H, J, J. (1967) Eeobspcal Aipccu ol Uie bt-haviom 
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N'.uii, I. K. t!979) Compuiei tinmlrdion models ol 
EtttensTvs chenopend shrubiand grozltiy Kisicm^ in k 

(naer/ and K. Howes ieds| Slndiys of the \UMrftltai1 
And /one. IV ilienopod Shruhlands- C .S.I.R,( ». (AUM.) 
Div. of I and few. Manne pp- t^5-«as 

D-ink' T'i. H.. W' Kid, I. u. .v P\i ooiM.i I Q.(f4&) 
| 11 ihe and rtLvelKti in LTu/my 01 ilu. p-:i t (ira;tl 
,sahhu-,ii lAiripla wttv/itA -an LeohjjjCrtl sOidy ol 
the bid 1. laetoi. Pun. I inn. sr>. . \SU,il>^ 57, 
I ' 402 

Kai'kss, I M (I960) The e.a-.s ,,-vct ol Alnea. RAjO 
^tflriC "siudi--.-; No, -i'K 

R wis, ^'l & Wi 1 • it, l>. 1 ("tin) T-iifihci Mudt., dtl , 

eia/Kiu. in the iVHihem l-'ennmc?. J. Br. Cropland iac 

2i, se 1 1 

\ (I969> LpUnd i-od.^iivtly o| ^.TC'aiion 

ipd Sheep at Moo. House NylKmal Naojie KcscfVCi 

Wfesimoriand, >vA isaLOrAo^ SuppkrmTaum It. ) 12 
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SMJ '•- < /V. B. (l9M)lUomaihfinai.L.-, ( ti.oh-,- ' hiflin 

i to. <lond.) 3rd Edit, ''U Pfc 



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byT. Sadler and N. S. Pledge 


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. 




Flic fossil clypettMertHd sea urchin, Arachnotdes inchia y 
was originally described by Tate' from ihree specimens 
collected from so-called Miocene beds at Ked Bluff, Lake 
Tyers, east luppsland. This unit is now recognised as the 
Lower Pliocene .Icmmys Point Formation. Fostei & 
Philip 1 revised the generic allocation 1 and indicated that 
(he species also occurs in the Pleistocene of the Parish 
of Wernkoo, Victoria. 

Recently, one of US iT.S.) discovered specimens at 
Willowhnnk, near Murray Bridge in the Murray Basin ot 
South Australia, some- 850 Kin from Ihc type locality in 
theGippsland Basin of Victoria. Subsequent investigation 
and collecting has produced numerous specimens. This 
note records the new occurrence and offers additional 
information on the species. 

Class Eclnnoidea 
Order t'iypcasteroidea A, Agassis 
Family Arachmmlidae Human 
Genus I'viloxlcr Durham 

ledustpr trrvisu {Tate 1893) 
Aruchnoides irnisa Tate, T893: 192, [)l. 13, fig. }j 
Fetta\termcixa(Vdte), Poster & Philip, 1980: 156, pi 20, 
fig. 4, 6-7., test fig. I. 

Amended diagnosis; Test very flat, rising slightly 
towards apical disc; almost circular, transverse diameter 
slightly greater than longitudinal diameter. Apical disc 
slightly anterior of centre. Ambitus sharp, incised at end 
of each ambulactal groove, between which il is undulose 
Ambulacra inflated with median depression and abruptly 
declivous at the sides; declivity decreases beyond end of 
petals. Petals reach almost two-thirds the way to ambitus, 
converging slightly at Ihcir ends on each ambulacrum- 
Ambulacra slightly more thar twice width of 
inicrambulacra. Ambulactal plates obliquely combed, 
lorming five chevron pattern pointing away from apical 
disc. Ornamentation minutely granular, most appaient at 
ambitus. Periproct supramarginal with shallow depression 
between it and ambitus no definite anal notch. Peristome 
centrally placed and round: Actinal area slightly concave 
with intorambulacrai bulges near peristome (Tip. I). 

Material; Seven complete specimens and several nearly 
complete measurable specimens have been collected 
(fable I), 'ogelher with numerous Iragments, some of 
which show the plate pattern dearly. 

Tmhc 1. Measurements. is, collection number. South 
Australian Museum; L, longitudinal diameter (from 
frontal lobe); T, transverse diameter; H, height; A, distance 
from centre of apical disc to posterior margin; P, distance 
from centre of periproct to posterior margin. All 
measurements in millimetres. 










M\ 3 






3 8 










3 1 












35 1 












34 6 


















6? : 




P2487 1 






Occurrence: Most specimens came from outcrops 
interpreted as Nor west Bend Formation, at Witlowhank. 
8 km north ol Murray Bridge, and others near Sunnyside 
Lookout, 3 km upstream, opposite Mypolouga. Specimens 
occur throughout the lower 10 metres of exposed Norwest 
Bend Formation at Willowbank (Pig. 2). fragments were 
traced as far as Ponde, 5 km souih Of Mannum. 

5^ QC -:^i v ; Jf CuMeni hfilddLl t nit 

Lt_ ^ > rS i \ * V T J Curtei't he-flded qtavel 
■- *-■ ■ N T 7, i ' 

5 R«0gY t «lt0r»i>W* »«0<J 

* r*"~ Mi«he%l r«Mealiit found 

ibssiwb cfltoaipoui !ni"rt'.iiM<* ncfi in •«!' 

J Cuirenl bi/ddarl gravel 

gwsM FeHaiti" ' cuntl. 
to grninoo mil ncu'Ut, t.nh.mi ftulih 
sanflstonf. with mm OevlSiiifc 
Worm tubes common «•»«< top. 

Mil »..»*« bull \u vMlttw ..»k».«,it|e 
Kiln numarous Itregufe' fsi daroond; 
&r>d m%r\ Hand-i HiehW flttyW'fJllUll 
win L jJtflitll lyr&ev- MgnviijSfU? 
ay^Ui'ts, t)»v«tv«&, Drjoioe** •""* 
otmowad tigniona 

Hsisiifl Doal (amo 

Fig. I. Vetlaster irtasa, ta) P24857, ad oral, (b) P24KSS, 

Fig. 2- Stratigraphic section at Willowbank. 


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, ajid 
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 

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 Fellaster 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. 

'Tate, R. (1893) J. Proc. R. Soc. N.S.W. 27, 167-197, pis 

^Foster, R. J. & Philip, G. M. (1980) Proc. R. Soc. Vict. 

91(2), 155-160, pis 19-20. 

Durham, J. W. (1966) Clypeasteroids. In Moore, R. C. 
(ed.) Treatise on Invertebrate Paleontology, 
Echinodermata. 3(2), Geol. Soc. Amer. (Lawrence: 

T. SADLER and N. S. PLEDGE, South Australian Museum, North Terrace, Adelaide, S. Aust. 5000. 


by S. A. Shepherd and Lisa 7. Hobbs 


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. 




1 h, !>lue-ihmnteil wrasse PseudoUtbriis (etriCH\ 
\ Rjiftardfton) is one of the man common Fwhea df inshore 
rock v reels on moderate k> high energy coasts of southern 
Australia. Vei except foi taxonomic and habitai notes 1 ", 
nothing has been published on its biology. This note 
describes (lie age dependent growth and length-weight 
it-lain«uships a\ t J tctnt tt\ 

This study is based on ihr analysis o| scales of 58 
individual* raptured by spearing at West island and 
adjacent areas hi Encounter Hay, South Australia, 
supplemented In lunoJI samples Of large individuals fiom 
<- ape Jcrvis. Yaukulilla and Uleasons Landing, Yorke 
Peninsula, The data lor another 17 individuals were 
uivcied because I he scale drcull wciv toe indistinct to read 
aenmtielv {tires scales were taken liom behind (he left 
pectoral Tin oi each fish, cleaned in sodium hydroxide 
solution and mounted between microscope slides. 
Standard techniques' were used for reading Ihe stales and 
for Hack-'.alail.iiHMt ol growth. Scales were examined for 
growl h checks oi closely spaced circuit by using a 
microfiche reader Photocopies of the projected scale 
unaues were made and mean distances from the focus 
aK>Pg '*0 diagonal posterior radii (where cireult are most 
clearly defined) to successive circuli and to the margin were 
taken fur each sample. The scale circuit have been found 
i<» be annular lor other icmperutc labrids'*'' and are 
iherclott assumed to be annuH lot this species. All lengths 
arc expressed as total length. 

I be relationship between scale iadius (R) in mm and 
total length [\ ) in cm was found to be linear, and the 
fallowing equatiou was filled to the data by Ihe leasl 
Kpurcs method dig. D- 

R oooas i MQ5 I (r 0.95) 
1 his eqnaiiori was then used to back calculate the length 

o\' llsh al the lime of lormaiion oi each arttuilus. 
Because sex inversion from female to male is size-related 

.iii.i minis at a length o\' ahout 35 cm (Shepherd & 

i • i 




rOUL l«»G7M \ 

t'larkson, in prep.) data for female* and males were 
combined for the purpose of the calculations. 

The number of annuli increased with fish size, and 
annul! maintained ihe same position on the scales of fish 
ol different ages. Subject to minor deviations discussed 
below mean back-calculated lengths agree with mean 
observed lengths at each age. Hence the back calculations 
validate ihe leehnique at least for the first five age classes, 
Older age classes would need validation by an independent 

The means of back-calculated lengths at age were 
plotted bs using the graphics facility of the Lotus 1-2-3 
microcomputer package and a von Berlalauffy growth 
curve was then fitted interactively by eye. The von 
BertalanlTy equalion for the length l. ( in cm al aye I in 
years is: 

L ( 46.0 (l-expt-0.22 II I O.M)] 

Table 1 presents the mean length at capture (those in 
the i * age group are aged between i and i - I years), the 
back-calculated length, their means and the filled von 
Uertalanfly points, the data and ihe Titled curve arc shown 
In Fig 2. 


? 30 



Tiy, I Plot n| scmIi- t -,(1,,,-, v t -r,n. rolal length tor P 

2 4 6 8 

AGE CLASS (years) 

Hg. 2. Plots of true loial length at age (a dash indicates 
the age class), lengths at age from ihe baek-caleulattons 
(crosses), and (he fitted ct>q Benalantfy cutve tot P. 
mrieus . 

The length weight relationship, determined by regression 
analysis was W 0.052 I*** (R : = 0.85: N 611 
where W is the weight in g and 1 the total length in cm, 

The mean back-calculated lengths up to age 4 for fish 
aged >4 years are greater than the true length (Table I). 
I his effect, if true and not an ariitaet of sample si/e, is 
the reverse of lee's phenomeu' , Possible causes are: 
l\\ non-iandom or biased sampling in which slower 
growing fish are under-represented; this may arise because 
of the gear used or Ihe habits of the fish; and (2) selective 
natural at fishing mortality 1 . P. retricus is not sought by 
commercial or recreational fishermen (although ii i* oiien 
a substantial pan of the catch taken ineidenialh by 
recreai tonaJ anglers"), and The sample was taken by 


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 

Some labrids show a point of inflexion on the growth 
curve upon the transition from female to male 10,11 . 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 

P. tetricus appears to have a similar longevity to its warm 
temperate congener of northern New Zealand P. celidotus 
(Bloch & Schneider) 4 . 

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 

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 from the fitted von Bertalanffy growt h curve. 

Mean total Von Bert. Annulus 

Age N length at calculated total 

capture (mm) length (mm) 12 3 4 5 6 7 8 

1 1 



2 + 



3 + 



4 + 



5 + 



6 + 



7 * 



8 • 















































Unweighted Mean 









Weighted Mean 









'Last, P. R., Scott, E. O. G. & Talbot, F. (1984) "Fishes 

of Tasmania." (Govt. Printer: Hobart). 
2 Scott, T. D M Glover, C J. M. & Southcott, R. V. (1974) 

'The marine and freshwater fishes of South Australia." 

(Govt. Printer: Adelaide). 
3 Tesch, F. W. (1971) in Ricker, W. E. (ed.). "Methods of 

Assessment of Fish Production in Fresh Water." IBP 

Handbook No. 3 (Blackwell: Oxford). 
4 Jones, G. P. (1980) Copeia (4), 660-675. 
5 Quignard, J. P. (1966). Naturalia monspel. ser. Zool. 5, 


^Beamish, R. J. & McFarlane, G. A. (1983) Trans. Am. 

Fish. Soc. 112, 735-742. 
7 Jones, R. (1958) Int Comm. N.W. Atlantic Fish. Spec. 

Publ. 1. 

8 Ricker, W. E. (1975) Bull. Fish Res. Bd Canada 191, 


9 Jones, G. K. (1983) SAFIC 7 (4), 9-18. 

l0 Warner, 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 




byRussellJ. Shieland WalterKoste 


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. 




In a review of the rotifer genus Brachionus (Rotifera: 
Monogononta), it was noted that some species typically 
were collected as epi/oiles on planktonic crustaceans 1 . 
For example, Brachionus rubens and less commonly B. 
variabilis 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. sessiiis was recorded as an 
epizoite of Diaphanosoma brachyurum, and the known 
hosts of B. rubens and B. variabilis were listed as Daphnia 
muiinu, D. pulex, D. longispina, Moina rectirostris, 
Ceriodaphnia sp., Polyphemus pediculus and several 
macroinvertebrates 3 . 

Although B. rubens, B, sessiiis and B. variabilis are 
known from Australian waters 4 , all were collected free- 
swimming from the open water of billabongs (Magela Ck, 
NX, and Murray-Darling River system, Vic, N.S.W.). 
Significantly, the host species listed by Kostc* do not 
occur here 5 . 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 x 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 dadocerans 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 bv sticky foot- 

gland secretions to the hosts' carapace; most of the rotifers 
were ovigerous, with 1-2 large amictic (parthenogenetic) 

Some animals with attached rotifers were preserved in 
4% formalin and prepared for light- and scanning electron 
microscopy (SEM) 6 . Representative SEM micrographs 
are shown in Figs 1 and 2. The rotifers have contracted 
into their loricas in response to the preservative, and some 
distortion of the lorica is evidem, 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 7 . 

B, novaezealandiae is not listed as epizoic by Koste\ 
however Morris' original description (= B. variabilis var. 
novaezealandiae) notes that his material came from a pond 
at Totara, N.Z. "where they were in numbers, parasitic on 
Daphnia thomsonf*. 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 
rnountant, 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 *m. 


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 9 . 

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 seen- 

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 

■Ahlstrom, E. H. (1940) Bull. Am. Mus. Nat. Hist. 77, 
Kutikova, L. A., Kosova, A. A. & Khodorevsky, O. A. 

(1976) Zool. Zh. 55, 763-767. [Russian]. 
3 Koste, W. (1978) Rotatoria. Die Radertiere Mitteleuropas 
(Uberordnung Monogononta). 2 vol. revision of M. Voigt 
(1956). (Gebr. Borntraeger: Stuttgart). 
4 Shiel, R. J. (1983) Proc. R. Soc. Vict. 95, 33-37. 

^Smirnov, N. N. & Timms, B. V. (1983) Rec. Aust. Mus. 
Suppl. 1, 1-132. 

6 Koste, W. & Shiel, R. J. (in press) Aust. J. Mar. 
Freshwater. Res. 

7 Pejler, B. (1977) Arch. Hydrobiol. Suppl. 53, 255-306. 
8 Morris, C B. (1913) Trans. Proc. N.Z. Inst. 45, 163-167. 
9 Koste, W. (1979) Aust. J. Mar. Freshwater Res. 30, 

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, Quakenbruck D-4570, Federal Republic of Germany. 



his excellency the governor of south australia 
Lieutenant-general sir donald dunstan, k.b.e., c.b. 


M. J. TYLER, M.Sc. 


T. D. SCOTT, M.Sc, Dip.Ed. 

C. J. M. GLOVER, J.P., M.Sc. 

R. H. FISHER, E.D., A.U.A. 


Assistant Editor: 


B.Ag. Sc, D.Ag.Sc. 



B.Sc, Dip.Lib. 

Programme Secretary: 
R. A. TEDMAN, B.Sc., Ph.D. 

Minute Secretary: 

Membership Secretary: 
P. M. THOMAS, M.Sc. 

Members of Councit 
J. B. FIRMAN, M.Sc. N. A. LOCKET, M.A., B.M., B.Ch., Ph.D., D.O. 

G. M. E. MAYO, B.Ag.Sc, Ph.D., D.Univ. C. B. WELLS, E. D., M.Ag.Sc. 

B. J. COOPER, B.Sc, Ph.D. 

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