VOL. 124, PART 1 
31 MAY, 2000 


Transactions of the 
Royal Society of South 
Australia 


Incorporated 


Contents 


Schumacher, R. K., Austin, A. D. & Floyd, R. B. Parasitoids of the autumn 
gum moth, Mnesampela privata (Guenée) (Lepidoptera: 
Geometridae) in south-eastern Australia, with coecieiny of 
two new larval parasitoids - -  - - 5 

Mackness, B. S. & Hutchinson, M. N. Fossil lizards from ie Barly Pliocene 
Bluff Downs Local Fauna -— - - 

Kolesik, P., McFadyen, R. E. C. & Wapshere, A. J. New gall rmidues (Diipters: 
Cecidomyiidae) infesting native and introduced Solanum 
spp. (Solanaceae) in Australia- - - 

Bradley, C., Beveridge, I., Chilton, N. B. & Johnson, P. M. Helminth 
parasites of the purple-necked rock wallaby, Petrogale 
lateralis purpureicollis, from Queensland - - - - = - 


Brief Communications: 


Parsons, R. F. & Browne, J. H. Causes of rarity in Abutilon oxycarpum and 
Hibiscus brachysiphonius (Malvaceae) on the River i ide 
floodplain, south-eastern Australia - - - 

Williams, C. R. & Kokkinn, M. J. Records of mosquitoes pices Culicidae) 
from the Cooper Basin in north-eastern South Australia -  - 

Dunbar S. G. & Coates, M. Range extension and new host records of the 
ectoparasite Pseudostegias setoensis Shiino, 1933 
(Crustacea: Isopoda: Bopyridae) - - - - - - - 


Obituaries: 


Alan Francis Bird, BSc (Hons), MSc, PhD, DSc - - - - - - - - 
Patricia Marietje Thomas, BSc, MSc, AO - - - - - - - = = = 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 124, PART 1 


TRANSACTIONS OF THE 
ROYAL SOCIETY OF SOUTH AUSTRALIA INC. 


CONTENTS, VOL. 124, 2000 


PART 1, 31 MAY, 2000 


Schumacher, R. K., Austin, A. D. & Floyd, R. B. Parasitoids of the autumn gum moth, 
Mnesampela privata (Guenée) (Lepidoptera: Geometridae) in south- 
eastern Australia, with description of two new larval parasitoids - 

Mackness, B. S. & Hutchinson, M. N. Fossil lizards from the Early Pliocene Bluff 
Downs Local Fauna - - - 

Kolesik, P., McFadyen, R. E. C. & Wapshere, A. J. ‘New gall midges Wiptera: 
Cecidomyiidae) infesting native and introduced Solanum SPP. 
(Solanaceae) in Australia - - 

Bradley, C., Beveridge, L, Chilton, N. B. & Johnson, Pp. M. Helminth péicwaiiak of the 
purple-necked rock wallaby, Petrogale lateralis purpureicollis, from 
Queensland - - - - - - ~ - - - 


Brief Communications: 


Parsons, R. F. & Browne, J. H. Causes of rarity in Abutilon oxycarpum and Hibiscus 
brachysiphonius (Malvaceae) on the River Bay Hoodplain, south- 
eastem Australia - 

Williams, C. R. & Kokkinn, M. J. Records of rridstibitdes (Diptera: Culicidae) from the 
Cooper Basin in north-eastern South Australia = - - 

Dunbar S. G. & Coates, M. Range extension and new host records of the ectoparasite 
Pseudostegias setoensis Shiino, 1933 (Crustacea: Isopoda: 
Bopyridae) = - - - - - - - 


Obituaries: 


Alan Francis Bird, BSc (Hons), MSc, PhD, DSc - - = = - - - 
Patricia Marietje Thomas, BSc, MSc, AO - = - - - - - - - = = 


I7 


37 


PART 2, 30 NOVEMBER, 2000 


Reed, E. H. & Bourne, S. J. Pleistocene fossil vertebrate sites of the South East region 
of South Australia — - - - - - - 

Brown, 8. P. & Wells, R. T. A Middle Pleistocene Herbotirate fossil assemblage from 
Cathedral Cave. Naracoorte, South Australia - - - 

O’Callaghan, M. G., Davies, M. & Andrews, R. H. Species of Raillietina Futivinanti, 
1920 (Cestoda: Hapenneigne) from the emu, Dromaius 
novachollandiae - - 

Cribb, T. H., Daintith, M. & Munday, B. A new blood- fluke, Cardicola farsiets) 
(Digenea: Sanguinicolidae) of southern blue-fin tuna (Thunnus 
maccoyii) in aquaculture - - - - - 

Kolesik, P. & Cunningham, S. A. A new gall midge species (Diptera: Cebidemyiidas) 
infesting fruit of punty bush, Senna artemisioides (Caesalpiniaceae) 
in Australia - - - - - - - - - 

Goldberg, S. R. & Bursey, C. R. Intestinal helminths of five species of scincid lizards 
(Sauria: Scincidae) from Western Australia - - - - 

Walker, S. J. & Goonan, P. M. Re-evaluation of the distribution of Geocrinia laevis 
(Anura: Leptodactylidae) in South Australia = - - - 

Davies, M. & Burton, T. C. Redefiniuon of the Australian frog Limnodynastes egress 
Tyler (Myobatrachidae: Limnodynastinae) - - 

Beveridge, I, Campbell, R. A. & Jones, M. K. New records of the eastode genus 
Pseudotobothrium (Try pangritynete Otobothriidae) from Australian 
fishes - - 

Nicholas, W. L. & Hodda, M. Beicyletvnas hasiyi sp. nov. (orslakgbaiae, Dorylaimida) 
a nematode collected from sediment in a freshwater rock-hole in the 
Northern Territory - - - 

Tyler, M. J. & Davies, M. Developmental biology and iatval bepholapy of the frog 
Limnodynastes depressus Tyler (Myobatrachidae: Limnodynastinae) 


Brief Communication: 


Clarke, R. H. First record of the Southern Right Whale Dolphin, Lissedelphis 
peronii (Lacépéde, 1804) (odenecet Tieenlieay) ), from waters off 
South Australia - -  - - - 


lnsert ta Transactions of the Royal Society of South Australia, Val. 124 parts 2 & 2, 30 November, 2000 


61 


91 


141 


177 


PARASITOIDS OF THE AUTUMN GUM MOTH, MNESAMPELA 
PRIVATA (GUENEE) (LEPIDOPTERA: GEOMETRIDAE) IN 
SOUTH-EASTERN AUSTRALIA, WITH DESCRIPTION OF 
TWO NEW LARVAL PARASITOIDS 


By R. K. SCHUMACHER*, A. D. AUSTIN? & R. B. FLOYDE 


Summary 


Schumacher, R. K., Austin, A. D. & Floyd, R. B. (2000) Parasitoids of the autumn 
gum moth, Mnesampela privata (Guenée) (Lepidoptera: Geometridae) in south- 
eastern Australia, with description of two new larval parasitoids. Trans. R. Soc. S. 
Aust. 124(1), 1-15, 31 May, 2000. 

Parasitoids were reared from cocoons present in larval shelters of Mnesampela privata 
(Guenée) collected at Altona and Shepparton, Victoria and Canberra (Lyneham 
Ridge), ACT. The most common primary parasitoid was a gregarious braconid wasp, 
described here as Cotesia geometricae Austin sp. nov. This species attacked host 
larvae at all locations and emerged from the fourth or fifth instar, pupating 
gregariously. 

Key Words: Autumn gum moth, Mnesampela privata, Geometridae, Glyptapanteles, 
Cotesia, Casinaria, Heteropelma, Isdromas, Mesochorus, Elasmus, Megadicylas, 
Braconidae, Ichneumonidae, Elasmidae, Pteromalidae. 


Transactions of tte Rayal Sectedy ofS, 


Avs (2000). 2401 TLS, 


PARASITOIDS OF THE AUTUMN GUM MOTH, MNESAMPELA PRIVATA 


(GUENEE) (LEPIDOPTERA: GEOMETRIDAE) IN SOUTH-EASTERN AUSTRALIA, 


WITH DESCRIPTION OF TWO NEW LARYAL PARASITOIDS 


by R, RK. SerumMacner’, A.D. Austin! & R.B. BLoypt 


Summary 


Scnumachiek, BOK. Austin ACD. & how RB. 2000) Pordsifoids ob Wie antl pun Meth, Maesetipreta 
privde (Coence) Wlepidoptera: Cienmetridae) ro south-eastern Austrailia. with deseripfien of wo new larval 
parasttoids, Trans, BR, See 8S. AteG E24C1), 1-15, 3.) Miyy 2000, 

Parasitoids were reared Tromt cocoons presenti hieval shelters of Miewenpele pei (Guence) eolleeted at 
Altona unl Shepparton. Vietoriv and Cynbern (Lyneham Ridgen ACT. The most gamnton primary parisitor 
Wilk U gregarious brigonid Wasp, described here as Cofresia veomenioa! Aust sp. nov. Chis species athioked 
Host turvae ub all loeniens and emerged Wom the (ou ih or APU ister popatigi grogarioushy A second bricciidl, 
deseribed here as Glyptupanieles ningsampeda AUSGn sp. noywas found ab Lyngham Ridge. This parasitord wits 
superliciilly simiku to. geome ieee, atieked varly-Tnstay Host lievae und alse emerved fromthe founhe filth 
stir Ta pupae wregariousty. One ether soliaey primary parasdaid, Casivaria aera Jevmuan & Cruld 
Uchncumonidie), emerged tron fourth or fifth instar arvile cine pupsited externally. At least five species al 
hyperparasitoids emerged fran the cocoons of Co geaaectereae collected if Vieturias two species of yedeenices 
Uehacumnonidae), Mursentea iis Sp, (I chnewmonidive ), Playas sp. (Elasmidve) wil Meeadicylas sp. 
(Preromialidaes, with one of (he Adromes speeies being most common. The hurval-pupal parasitoid /ereropehnt 
seuposun (Morley) (lehneumonidue) emerged from 10% of pupae reared from larvae eatleeted tn Whe ACT over 
the same peril as adults of M. private. Sex and viability significandy influenced the weight of pupae ol MW. 
provide but pupae parasitived by P/. xeupayun could not be sepurtited using Welln, Notes and ai HLUsttited key 


ave povided Lo faovitite te easy jdeng ication of te purustiloids. 


Key Woks: Aulumn stn moth, Waresanpebe privat, Geomenidoe, Ghypripanieles, 
Mevacioylan. 


Heteropelmi. idromes. Mesecharus. blaxnnis, 


Preromatidae. 


Introduction 


The aulunn gum moth, Muesumpela private 
(Grenee) (Geometridae) is endemie to Australia and 
vecurs throughout! the south-custern and south- 
Wester parts OF the continent as well as Tasmania 
(McQuilhan 1985; Abbott 1993), Larvae feed anu 
wile range oF eucalypt species (MeQuilld 1985) but 
prefer the juvenile Molle of trees iin the blue yam 
sroup (Bilion & Bashford L978), whieh includes the 
TMpoerkint phintation species Licealyatiay ofteny 
(Deune & Miuden) Minden and 2. e/obulis bLabill, 
(Abbotl 1993, Bushford 1993. Neuninn 1993; 
Phillips 1903). Maesumpela priypera ean oceur in 
hiph numbers in young. everawed stunds of planted 
cueulypts and cause severe deloliacon (Elio & 


DePUIINEROOD TEQGIOY SH cy ACERT ATE SHOE CATS 
Cohen ACO) (200) (Clipe dddhess® Depiinen ot Fanboy anil 
Tivtoeloey. Phe Dabversity ot Queenan Qhiday 2), 
Depetnent ot Applied & Motewutin Bosley. Wyite ¢ 
Universaly et Adelaide (len Ohi SA SUbe 
CSIKO) Entomotiiry, PO Bes E700, Canberra ACH JAH 

bh hirton, PR OW. PORTE OA study ah the sini site imap, 
Vieunelit pelo Chive. Chepidepient Geordie on 
Frreoypiiy fees Ue retrth Yoest DASH wkachted: Papost 
Tin iy Terria Repent: Pribaet pinnyiity$ 


Swaps The 


Crteniihy Cunineriit, 


Bruwomdac. tehneumonidue PE lasmidie. 


Bashlord [978 Roberts & Sawtell O81, Alliott eval, 
P9900; Abbot 1993: Neumann 1993: Phillips 1993; 
furrow ef al. 1994). Outbreaks cin be evonomically 
vostly as severe defoliation ean rediice the growth of 
ees (Floyd & Farrow 1994) ant miny eventually Kill 
them if defoliation occurs over several successive 
years (Came eral. LO74), 

Currently, management of outbreaks Of Ad. private 
coppists of browlsede sprayiig will don-specihe 
chentical insecticides (de Lillle POST! EO en af 
1990; Neumann L903: Phillips (O94: Neunan & 
Colleu 187). These chemicals have tndesinible 
effecis on The environment. miaty  exteerhitte 
outbreaks of ML privekt und other detoliaers hy 
eliminating mature! enemies and inay indice 
insecticide resistance (Huffaker LOO, Risch (987, 
Neumiunn 1992), Atternalive pest miinagement 
lechmiques such as biological control, larvet-specitic 
chemicals vind silvicultural methods have the 
potential To WUAMise enyvironirental dunked 
contribute 10 sustainable miunigement practices 
(Hutliker L980: Ohmart 19900 Floyd & Farrow 
}994), However, successfil iiplementition of any 
pesh munagement programme requires a thorough 
knowledge of Ihe eegloxy of Ihe pest and iS natural 
enemics (Qhmart 1990). Despite the importance 


1 ROK SCUUMACTIER. A. i, AUSTIN & R.B.PLOYD 


MZ. price as a pest there have beer few studies oF ins 
higlovy (ea. iio & Bushford 1978. de Little 
H98E Lukaes 19997) and information congerping its 
porusioids is fragmentary, 

The aim of this stuly was to identify parasitoids 
ind byperparasitoids associated with the hirvae of Az, 
peiveta iy the ACT ound Vietoria, conduet ain 
Tnveshition of parausiond Host age preference and 
examine the intlvence of sex. viability: and parasitism 
on the pupal weight of WA privad. A key to the 
panisitoid species reared. in this study is given und 
Iwo new braconid species, Colesia coomteniede 
Austin sp. nov. and Clyplepaateles smeyampela 
Aust Sp, fay. are desenibed, 


Matertals and Methods 


Std ey 

The study was undertuken ul two sites tn Victoria 
iy September (992 and one in the ACT during lite 
antirin aned winter [993 and 1994. The: sites: it 
Vietorii comprised a plantation of Ay glodielins 
wlobudun aw Aliana (37° 50° $, 144° 44" By. and a 
finn planting at Shepparton (36° 20° 8. 45° 13? By 
compresine Eo g. alohulis, 2. a. psendagtaliliis and 
Pow breostaiit. Boll locations tial heavy pifestations 
of preven (up to 100% defoliation by lirvae) Phe 
ACT site was ab Lyneham Ritge. Canberra (35° 14! 
S. (40° 6" 1), comprising mixed eucalypt speenes 
Which were only Trelthy uibested by Ad pri pete 
(apprasimtely FOG. defoliion by Larvae), 


Revirinie petresateanly 

Parsons were paared fron (ate-tstiin lirvae of 
A privade, (ne former hiv ing pupated 17 the leah 
in which tos! larvae sbeller during the day (EIHOLE & 
Busttord 1978: MeQuilhur 1885). Leiw-bugs 
conmining hile-insttr host larvae und parusitarl 
covoons were collecled from 44 £, 4, ulebuley al 
Altona. undipproximately LO Trees tearnprising &. 2 
wlubulia, by gs. pyendoglobuliy and Wg, Pteees teeter) at 
Shepparton. One leal-bar containing lele-instar 
hiewee ainel parasioid cocoons wits collected trom “4, 
0. Heese at Lynehuin Ridge i date August }994, 

Cel) parasitiid cocoon por group ab voeoons thal 
were apn together in the instances of iultiple 
parswisrad wes ified Fro he leatehog, placed in 
vw venlilited viel (2 en) dian, A 8 em tg) and 
incubalet al 2b 42°C. Approximitely a0) such 
sainples resulread from nnaterial collected Prom 
Altona (3 fruie Shepparton and one front Lynehwi 
Rideu. Vials were cheeked weekly andl parasitoids 
bevan to emerge and then every 1-3 dave for two 


‘hehe AT EMSU Terofagy te tuluni gan matty 
Viridian pre (Coen PID Tests. LivGretiy ab hawaii 
{api t 


weeks. A final check was made alter 18 days. Actull 
purasuvlils were removed as they emerged und 
placed in 70% ethanol. 


Hostage preference 

To investiga the host age preference ob hveval 
parasitoids, a range of availible instars was collected 
from EL. a Aicaytdla at Lyneham Ridge on eight 
vecasions in 1993, Collechons were mide 2-3 weeks 
upart. over the period thar hirvae were present (early 
May to late August), On ciel oceasion 2-12 trees 
Were Wspected dnd groups of |-19 (median of 5.5) 
jaryae collected from euch (ree, Larvae were reared 
in ventihited plastic containers (120 mnt dint s 95 
mm high), provided with a small braneh oF juvenile 
Foe bieostatd, Ihe stem of which wis placed throug 
a hole in the base of the contiuner and into water 
below, Folinve was chanved lwice weekly, Larvae 
were reared at 25 + 2°C under navural ligt 
conditions, When larvae or pupae of parasitoids were 
observed with a dead or dying M. prlvera larwi, the 
dale, number of parasitoids present and the instar ol 
the hosr larva were recorded. Larvae that died fur 
reasons other than purasitisation were not incldded i 
the analysis. Head capsule size was used to estimate 
the barval instar of MW, private CBMott & Bashtood 
L978). Parasiloid larvae or pupae were removed to 
mdividusl ventihwed vials (2 em diam. & 8 en highs 
und incubated ut 224+ 29°C until the adults emerged. 
Adults were-stored in 70% ethunol. 


Prpuel weight 

Mnesenipela privare larvae eolleered from 
Lyneham Ridge that pupated suecessfully wwere 
werfed Within tice days of pypation, Bach. pupit 
wus incubuled ina ventibkiedt vial (20 mim din, + 80 
mon high)-at a temperature of 4.0 + 0.5°C and 70% 
relative humidity until late Outober. In November. 
Papine Were placed outside inv nutiaral light un 
lemperadiire fegiies cand relative hunnidity wis 
mmuntiined ab 70%, Pupae were mapected imonthhs 
from mid-January lor the emergence of parasthonds 
Or udull M, privaia. Parasitoids emening (rom host 
pupae were storecl in 70% ethanol The ses of udull 
M. private was deteromoed ting the morphology of 
the Frenulumn (after Elhiou & Bashford O78) une that 
of noneviuble pupae by the position ol ihe aenieal 
sea (Lawwrenee ep al. 1991), Oneawany ANOVA was 
perlorined to test iP the ouleome of pupa ot Ad 
private (ive. theemergence oF either a male or lente 
HU AY. perveee an adult parasitoid er the death ol 
the pupae) iaflueneed weight at puparion (Salkal & 
Rony Ost wih significantly different groups 
separated by Schelté’s test of multiple contrasts (Zur 
JOS) A Chi-square gondhess-oPfit fest wis Used te 
compre The sea Tuties Of pulpit ane adults with an 
expected palio of Ty) CSokal & Rolf Tk)). 


PARASITOIDS OF THE AUTUMN GUM MOTH 


Peusson's chi-squire was perlorined to determine 4 
suecessful pupation to adult wis independent of the 
sex OF the pupa (Sokal & Rohlf F981). 


Results 


Parasioatd compen 

Cocoons Formed by twa species of priviary 
pundsitoids were present ih the leatbags collected at 
Allona and Shepparton (Table 1), The most common 
cocoon. made by Cates seemenmicae (Braconidae). 
wis white, ahout + mn in dength and Found in groups 
spun together with silk, The other type of cocoon, 
formed by Caswern anerd Jecman & Could 
(Ichneumonidae), was motled oringe-brown, about 


6 mm im length and solitary. Five speeies of 


hyperparasitoids emerged from the cocbons of C. 
geomenicde (Table {), the most Humerous being a 
species of fxdromasy (Ichneumonidae) (herealler 
referred to as Jvdromeas species A). A second species 
of dydremers (sdromas species BY. Megadicvlas sp. 
(Pteromalidae), Mevocharus sp. (lehneumonidae) 
wind Elasmis sp. (Bhistnidac) also emerged from the 
cocoons of C, geometrivac. Cotesia veometricac also 
emerged from cocoons present in the leat-bag 
collected wt Lyneham Ridge in 1994. However, © 
veametricae Was not reared from lurvae collected at 
Lyncham Ridge in 1993, Instead, another braconid, 
Giyptapaiteles mitesampeld. pupated in groups ef 
one or two white Covoons external to the late-instur 
hose (Table 2). Castiartea micro was also reared fron 
a larva collected at Lybeham Ridge i 1993 and the 
huval-pupal parasitoid, Mereropelmu scuposiue 
(Morley) Uchoeumonidae) emerged from the pupac 
OlM. privata reared over the suminer of 1993-94, 


tes 


Relalive frequency ef parasiigidy reared fron 
parasitoid cacoans present in leuf hays 

OF the 130 samples collected from Altona. 69 
resulted in the emergence of Co veamertece and/or 
its hy perpurasitoids (336 individuals) and LOowith © 
micra (Table 1) while 31 did nor yield any 
pirasttoids. The 13 simples from Shepparton 
resulted in four without emergence, four with © 
veamelricae and/or its hyperparasttaids (17 
individuals) and five with Co anerd. The single 
sample collected in 1994 at Lyneham Ridge yielded 
14 individuals Of C. gemeteicde. The cocoons in 
each group of C, veametrieue were nob counted or 
examined closely for prior emergence, so Twas nol 
possibly to determine the average number of covoons 
per group based on the number that emerged. or 10 
culeulaté avcurately the relative frequency of 
successful criergence. non-ynibiily and hyper- 
parasitisation. Beariay this in mind. about one-thire 
of the groups OFC. geemieiricue Covoons resulted in 
the exclusive emergence Of Co geonietricue, one 
third resulted ti ah ininal emergence of ©, 
geamenricue followed by the emergence ol fsdrenurs 
sp. A from the remainder of the cocoons (NL. six of 
these groups also yielded /yedrmas sp. Be 
Mevadievlas sp. and Mesechorus sp. alter an initial 
emergence of C, geonetricae), and one-third 
exclusively yielded hyperparasitoids. /se/ronnes sp. A 
tsiromas sp. B, Mesadicylus spoand Mesociiority sp. 
emerged from cocoons collected ut Altona, and 
Idromas sp A und Llasms sp. from Shepparton 
(Table 1), Thus, at the two sites in Vietora, C 
geomericde Was The most frequent primury 
parasitoid of farvac but about hall of its cocoons 
were hyperparasitised, primarily by /sclroniay sp A 


Vaset |. Parayiid species emerging from cocoons associated with the havee of Moesampela privat collectce in 
Vrona and Shepparton, Vietoria in 1992. *Totul number ol cocoons per aroup was not courted. 


Covoon type Biology Specics enicreed Altona Shepparton 
No. © oF total No, No, “eal total No, 
mdividiuials: todividuals individuals Tle idugals 
White, Primary Conese peomeniong 252 470 4 23.6 
SPOS UT LOLS parasitoid 
- Hyperparasiioid /seramteay sp. A 246 45,9 a) SN 
. Mesecharny sp. Is 34 () 
~ Iyclroanias sp. B \I 2, () - 
= Mexerdicyleay ap. 1) 7 a 
Plasmas sp. () . 3 V7 
Total No. individuals S36 17 . 
'Median No. 6 4 
of emergences/prouy 
Orange, Primary Cusiialiee Wer tt) 5 
solitury prursdsieane 


i 


R. kK. SCIIUMACHER, A, D, AUSTIN .& R. B. FLOYD 


TAnit 2. Parestisation of lurvee of Mnesampela privata collected at Lyuelan Ridge (Canberra, ACT) in P9097 Th 
/904 Cotesia geometricie wey reared from parasitoid cocoons present ica M. privata feef-biy. 


VL private collection 


Parusitoid emergence 


Date instar Dale Host instar No. emerged Parasttoid species 
2/05/93 2 22/06/93 4 | Glyprapantales 
pines pela 
W05/93 2 23/06/93 4 2 G nineseiipelit 
16/05/93 3 22/06/93 4 | Gimnexatnpelit 
TAT/YS 4 LL/O7/93 45 | Cuyinuria niche 


Cusinaria micra occurred less frequently and did net 
appear lo have any hyperparasitoids. Estimates of the 
relative frequency of Jarval parasitoids at Lyneham 
Ridee jn 1994 could hot be made as only a single 
sumple was collected and a comprehensive survey 
was not undertaken. 


Relative frequency ef parasitoids reared front 
collected larvae of M. privata 

The 426 larvae of AZ. privare collected at Lyneham 
Ridge in 1993 yielded only tour individuals of (, 
mnesampela and one of C. micra and neither species 
Was hyperparasitised (Table 2). The collection of M, 
private larvae prevented hyperparasitism of parasitoid 
cocoons und probably limited hyperparasitisn via the 
larval host, However, none of the larvae collected in 
their fourth or fifth instar (28%) contained secondary 
parasitoids, suggesting that the frequency of 
hyperparasiloids was. very low at this site. Fifty-six 
percent of pupae resulting from collected larvae died 
hetore mid January 1994, 34% pupated successfully 


to adult ML private and 10% resulted in the emergence 
of 7 scaposum. Vhus, the laryal-pupal parasitoid // 
scapasuam wis the most successful parasitoid 
allacking Lirvae at Lyneham Ridge jn 1903, 


Host age preference 
Three larvae collected in their second or third iistar 
yielded G. mesampela in the fourth instar (Table 2). 


‘This result shows that G. jiesumipela cab parasitise 


second instar hosts, ‘The possibility that first instar 
hosts can be parasitised was not confirmed and 
insufficient parasitoids were reared lo determine if 
later host instars are also vulnerable, The host age 
preference of C. wicra is unclear us only one host 
larva, Collected in its fourth instar, was parasilised. 
Heteropelna scaposum were observed attempting to 
parasitise both carly- and late-instar larvae of M, 
private in the field. However. percent parasitism: by 
MH. scapes did nol increase significantly wilh host 
uve suggesting that parasitisation of later instars was 
not as successful (Table 3). 


Tnte 3. Peron emergence of the larvel-ptpal parasited, Ueteropelma seaposum. ti refaiien to ue dustar at whted: its 
host, Mnesampela privata, wes collected Lychan Ridge (Canberra, ACT.) 


Instar of 
M_ private 
al Collection 


No, pupac 


HT sSeapesian 


( emergence 


M. private No emergence 


al 7 i| 
3 TB 1) 
4 SQ) te) 
5 29 te 


‘Wh dt A oe 
fmm 
wn a 
A mls we 


TAMA, Effect of the outcome of pipae an the weight of papae of Muesainpela privat diree das after pupation. 


Sp LOS) Scheffes test of multiple contrasts. 


Outcome 


Adult mile MW. peivafe &, adult female WL private. 
Male pupae that died v. female pupae that died 
Adult privaie ®, pupue that elted 

Adult MW private voadult (7, seaposun 

Adult (7 scaposan vy, pupae that cied 


Mean weight +. SE (my) F 
140+3,7 v. [2743.2 17.68" 
13043.) vy, 109+2.6 26.63" 
140#2.% vy, 1942.2 31.8% 
1442.8 y, 1425.8 O16 
1425/8 v, 119429 17.44= 


PARASTTOIDS OF THE ALTTTUMN GUM MOTEL 5 


Pupal weiaht 

The guteome of pupae (categorised by the 
emergence of either a male or female adult M- 
private. an adult AL seeposmm or the death of the 
pupac) significantly influenced the weight of pupae 
at pupation (P= 22.9; dl = 4.200: pp < 0.001), Pupae 
thatched were significantly lighter than both pupae 
thar successfully produced adult WZ. privata and 


in 
im 
we 

bill 


fal 


Nuoiber of individuals 


Mi 


mM 


4 


ct + 2 > = wy ™ 
a ri ri mH + bs $ a me 

r 1 c % > " te 
— el ¢ = at r ie 


Vays aller collection 


Bf geaieticae vacktuively 
Of peoetreay followed hy Feber sp A Trond Thee sete Gagarin ken ft 


LD tloney sp) tiner eine Tn aie acount sO weemedriene 


big 1. Temporal pattern of emergenee of the wresurious 
primary parasitonl Cefewa veametricn sp, dav. and ths 
liyperpurasitonl (vedramas sp. A. trom groups of eoeaons 
ussociited with the Janae of Maesauipele priya 
(Guence) collected at Aliana. Vietoria. 


[Ln 


Peh ite Mar |x 
Mus aque 


» LE 


Aaefore Tun) S dan 1h Feb. 15 


Patal ouniher 
NOP est 


Aller Ape 15 
Ve elo at thine 
Ch AN teint 
Nile prevvretee 
OO henythy at popu 
Big. 2. Temporal pattern oF emergence Of adult Mvreseinpe led 


privata (Guenée) and the huval-pupal parasitoid 
Heterapelma scaposunt (Morley). 


parasitised pupae resulting in adult AL seapasner 
(Table 4), Pupae resulting in adult fermale WW. private 
were significantly heavier than those resulting in 
miles. However. the weights of pupae resulting in 
adult //, Xeupasant were not significantly different 
from those of pupae resulting in adults of M7, privata 
(Table 4). The sex ratios oF pupae of M. privata (not 
including pupae that resulted in //, seaposuimn) and Mf. 
privata that emerged as adults were [21.04 (= 188) 
and 1:21.33 (n = 72) (male : female) respectively. 
Neither ratio was significantly different from bs x" 
= 0.085 und 1.389 respectively, dl = lL. p > 0.05), 
Successfal pupation was independent af thie sex ol 
the pupa (X27 = 1.61; df= 1; p > 0.05) 


Biology of parasitoids 
Family Braconidae 
Colesia geometicae Austin sp. ov. 
& 
Glyplapaiiteles ninesampela Austin sp. nov. 
(FIGS 5,9, 10) 


AIL known species of the microgasterine genera 
Cotesia and Ghyplapanteles are endoparusitoids ol 
macrolepidepterans (Austin & Dangerfield 1992) 
although there i fo previous record of either genus 
parasitising larvae of Mo pivvere. Vhe species reared 
during this study are unknown and described below 
as Two new species. Cofesie and Glyptupanieles can 
be easily recognised from other parasitoids 
assocrated with this host by the absence of venation 
inthe distal part of the fore wing, the absence of vein 
2mcu (Pig. 5), their small size and dark colour 
Superticially. they are similar to cach other und could 
be easily misidentified as a single species, However, the 
shape and sculpturing of the first and second metasomatl 
termes can be used 1 readily separate them (Figs 9, 
10) (see deseriptions below for further detail). 

Giyprapanteles mnesampela can parasitise second 
star larvae of M, private although tt is not known 1 
first or later instars are also vulnerable. Final-instar 
harvac of G. mesa@mpeld and CL geomemricue emerse 
from fhe penullimare or Final taste of AZ priveried. 
aggregate und pupate pear the host remains, Tits life 
histery is consistent with) other members. of 
Glyptupanieles and Cotesia except that not all species 
in these Wo Benera ure gregarious; some are Known ta 
be solitary (Austin & Dangertieh! 1992), Adult ¢~ 
geonerricae emerged 13-24 days alter the collection oF 
their cocoons at the field site in Altona, Victoria (Pig. 1) 


Family [chocumonmidac 
Caxinarta micra Jerman & Giuld 
(FIGS 3. 12) 


All known spectes of Cayinaria are solitary 
endopurasiioids of lepidopteran Jarvac. This study 


if) R. K. SCHUMACHER, A. D. AUSTIN & R. B. FLOYD 


provides the first record of C. pucra reared from M, 
privata. Specimens of a Casinarta were reared from 
M. private larvae collected by Elliott and Bashford 
(1978) but were not identified to species. In this 
study, C. micre killed late-instar larvae. This result 
differs from that of Jerman and Gauld (1988) who 
observed C. micra killing Mrnesampela (species not 
spevified) in an early instar. However, Allen (1990) 
found adults of C. micra emerging from mid- to hite- 
instar larvae of Uraba lugeny Walker (Noctuidae), 


The specimens reared from cocoons in Victoria were 
all solitary emergences which occurred 15-27 days 
affer collection. 


Heteropelma scaposum (Morley) 
(FIGS 6, 14) 


This species ts a common solitary laryal-pupal 
parasitoid with numerous host associations including 
Pararguda australasia (F.) (Wormerly Digelesia 


Figs 3-8. Wings. Fig. 3. Casinaria nucra Jerman & Gauld. Fig. 4. Mesochorus sp. Fig. 5. Glyptapanteles mnesanpela sp, 
nov, Fig. 6. Heterepelma scaposum (Morley). Fig, 7. Megadieylas sp. Pig. 8. Elasmus sp. Scale bars = 0.5 mm 3-5, 7: 1.0 
mm 6: 200 um 8, Abbreviations: a = areolet; pt = pterostigma; st = stigmal vein, 


PARASITOIDS OF THE AUTUMN GUM MOTH 


—_ 


al 


— 


a 


_— 


ee Lk 


4 


Figs 9, 10, Propodeum and metasomal tergites 1-3. Fig. 9. Colesia geometricae sp. nov. Fig. 10. Glyptapanteles mnesampela 


sp. nov, Scale bars = LOO yim. 


ausiralasia) (Lasiocampidae) and the agricultural 
pests Helicoverpa armigera (Hiibner) and 
Spodoptera litura EF. (Noctuidae) (Gauld 1984). In 
these associations H. scaposum parasitises its host in 
an early instar (Gauld 1984), It has previously been 
identified as a larval-pupal parasitoid of M. privata 
in Tasmania and Victoria (de Litthe 1981!; Lukacs 
1999), In this study, AM. scaposum appeared to be 
most successful in parasitising early-instar larvae 
(Table 3). This is consistent with early instars of M. 
privata not forming protective leaf bags and thus 
being more vulnerable to parasitoid attack and with 
the fact that larger late-stage larvae exhibit more 
effective defensive responses (rearing and 
regurgitating drops of Hucalyptus-scented fluid) 
(Elliott & Bashford 1978; Schumacher, pers. obs.). 
In addition, Lukacs (1999?) observed oviposition in 


first instar larvae of M. privafa but none in later 
instars. Larvae of H. scaposwm do not develop 
beyond the first instar until the host pupates (Gauld 
1984). The average weight of pupae that yielded 1. 
scaposum was not significantly different from that of 
the mean weight of viable pupae (Table 4), indicating 
that parasitism by H. scaposum does not influence 
the behaviour or growth of larval hosts. The temporal 
patterns of emergence of male and female M. privata 
and /7, scaposum: were similar (Fig, 2) with most 
emerging between mid-February and mid-March. 


Isdromas spp. 
(FIG. 13) 


Isdromas species are 
hyperparasitoids from 


commonly reared as 
small ichneumonid— or 


x KK. SCLUIUMACHER, A. D. AUSTIN & BR. B. PLOYED 


Figs 11. 12. Lateral metasoma. Hig, LE Meserheruy sp. Figo 12. Castmaria micra Jorman & Gauld. Figs 13, 14. Dorsal 
inelasoma (sculpturing not shown). Fig 13, Ivers sp A, Fig. 14. Heleropetma scaposin (Morley), Figs 15, b6. Hind 
les. Fig. 15. Elaymus sp. Fig. 16, Megadiovlas sp. Scale bars = 1S mm tl. 12: 0.5 mm 13, 15, 16; 1.0 mm 14, 


Abbreviations: cx = coxu, [= femur, 


braconid cocoons, particularly from microgastrine 
braconids. although they are also recorded as 
primary parasitoids of a range of lepidopteran hosts 
(Guuld 1984), There are about 30 species known 
from Ausiralia, all except three are undescribed 
including the two species reared during the present 
study. There is no doubt that they are 
hyperparasitoids in the cocoons of C. geometricae 
(Table 1), given that no other cocoons were present 
und the larval cackevers of M, privetd were not left in 
the rearing vials, The peak of the subsequent 
emergence of /xdremas sp. A occurred 11-16 days 
afler CL veamericde emergence (Fig. 1). 


Mesochorus sp. 
(FIGS 4, 11) 


Mesachoruy spp. are hyperparasitic on the 
endophagous larvae of Braconidae and Tachinidise 
(Gauld 1984). Within lepidopteran hosts. mese 
chorines will oflen attack gregarious endo- 


parasitoids, especially microgasterine hracontils 
(Gauld |984; Allen L990), In this study Mesechoriis 
sp. was reared from cocoons of CL geamerricde 
Species of Mesochorys also parasitise Cotesia 
uwahae Austin & Allen (Braconidae) and C. pilcra 
via U. /ugens (Austin & Allen 1989; Allen 1990), as 
well us the tachinid parasitoids of Piurupsix utomaria 
Olivier (Coleoptera: Chrysomelidie) (de Little |982) 
and Perga spp. (Hymenoptera: Pergidae) (Cure 
1969). 


Family Elasmidue 
Elasmus sp, 
(FIGS &. 15) 


Members of this genus are obligate 
hyperparasitoids of Lepidoptera, One species, /: 
australiensis Girault. has been reared from C_ aicra 
und two microgastrine braconids yia C/, dagers, by 
this study only three specimens of E/asniusy sp. were 
reared [rom C, veometricae via M. privata and they 


PARASITOIDS OF THE AUTUMN GUM MOTH “ 


ny be similarly parasitic on ©. 
USHOCHITON Was Hor confirmed. 


mifera althoueh this 


Family Pleromalicae 
Megadievlay sp 
(PIGS 7, 16) 


Mevadit las spp. 
lepidopteran larvae and puple in eocgors. er dre 
lryperparasitoids on them, They have previously 
been associates! with several lepidepteran timiles, 
as well as microvasirine braconid cocoons (Boutek 
IOXB). This is the first record of a species being 
redred fram ML privata. lt is most likely a 
hyperparasitowl of CC. veemelreue us no other 
vocoon types were moticed aod the cadavers of MW, 
private larvae were nor lei inthe pearing vials: Only 
one Species hus heen recorded from Australi 7, 
dibiins (Cina) but the association with MO prryvetd 
hus not been confirmed as this species. 


Other parasitoids insecrated with 
Mnesaimpela private 
(ehrieumenidac and Tachinidae) 


Apart from the species discussed above a number 
ol other parasitoids has been previausly reared trom 
M private bit these were not recorded in the present 
study They include the tehneumouids Eitherds sp. 
Meguvertie pagan (Morley), Pristiceroy spy. 
Cumpoplet sp. and WA raerty sp. uid an unidentilied 
tachinid fly (Elliott & Bashford 1978: de Little 
POXL > Crate TOKE, Lukaes 19997) (Table 5). OF 
these species, all have been reared from iW. prvvader in 
Tasman with the exception of ML. pagania which is 
known Only from Victor (Gudld l9s4y. However, 
Gaiild (1984) recorded an unknown species of 
Meeuceria trom MA private in Tasmania and i ts 


very likely to be this species. Based on the biplogy ot 


other species belonging to these ichneumonid 
genera, all of them ure probably solitary primary 
endopurisitoids (Craald L984. We have not been 
Hbke lo eXamine material of these species dnd so have 
nol Welded them in the key, although their 
distibution und biological charaetenshes are 
vompuredt with the species recorded in the present 
study in Table S 


Key tu the parasitoids of Mnesanipela privata at 
the three study sites 


ls Fore wing wilh relatively complete venation 
(Pigs 3-0) smi to Large SREP IE > LA amin in 
length... statin hendtreth wut nutnens 


bore wing ila pizmented venation pealuced la 
unlenor margin (Pts 7, 8): mimute tarsal 
species, <= 2.5 mong in length. breton sain 


are primary parasitoids of 


2 


fy, 


Pore wing with verution distal to plerasiyma 
wanting Cbig. Si vein 2mck absent 
(Braconidae) cs eleva naepfereres Al 
Fore wing with ‘listal veins S present sind well- 
Piemented, vein Jueu present (Figs 4. 4, 6) 
(ICHNGUMONIGUE), acjerer cere cestiee ten serenrseeey scenes eet 
Propadeum and metasomal tergiies { and 2 
virtually smooth: tergite L narrowing uploully; 
lergite 2 with subtriangular median field (Fig. 
LO) Gyprapertteles winesanpela sp. 10y. 


Propodeuin and Metasomal tergites | and 2 
with obvious dease punetation; fergite 1 
Y 


moderately broad, then ii yeEPE 

broad and rectangular (Pig. Oyu cescese 
waadeleridch doseaidenoshd ones Colesta geoinenivde Sp. hoy, 
Pore wing with an arvolet (Pigs 4, 4). 3 
Pore wing Withoutan areolet (p IPO) eeeseee ts) 
Seutumn and propodenmn densely panetate or 
rugulose: ovipositér very short, Hol protruding 
past posteror metusumal tereites. (Fig. 12) 
(body 7-8 mor in length, dark brown to black, 
legs reddish: @ genitalia without pair of lowe 
rods)... -Cusinarie aera Terman & Gaul 
Scutum and propodeun generally unseulp- 
tured (except for propodeal earinge and 
micropinetures assaciated with pilosteyh 
ovipositor about fi te ts length of metasenia 
(Pig, 14) (body 3-4 mm in length, yellow 
brown with darker markings; c genitalia with 
pair of long rods protruding posteriorly) csc. 
volMesoelaruy sp, 
Metasomal tergite 1 narrow apd very cfongate 
Fig, Jd): fore wig with radial cell elongate 
(Fin. 6902 boely length about 2 mim, 10-12 
mim, head and mesosoma bkick, legs und 
Metasama yellow to Orange brown). 
jeri ablererapedma seaposiar (Morley) 
Metasomial tergite | broadening apicsly Chis. 
13): fore wang wath radial cel short and broad 
(body feneth 2,3-3.2 mm not inclidine 9 


bagly db cieeds ies Lessa neta tee 


ovipositor: heac und mesosamnia  blick, 
incrasema either dark vor vei with dark 
markisgs) Usdrontas spp). tiipeagptetatteatih 
Seutin smooth: fore wands stigma of | S clear 


in posterior half, white anteriorly. ct ‘evenly 
Liinslacent lees yellow: metasoama cither dark 
and Sometimes with lighter transverse hinds at 
sutures or yellow with darker markings... 
_Asileonnas sp. A 
Seutuim ‘with dese nunetilte seulpruninns fore 
wing stigma of 2 and evenly darks fare and 
mid lees light brown, hind less dirk brown: 
melasome dirk brown (O bLMCK cso cen 

ai) sartastiesyancious wsdronias sp. B 
Hind « COX uy developed 4 us hirge flat dises hind 
tibia with Gistinet eriss-eross patlerth of sete 
(Fig. 15); lore wing with sigmil vein very 
short (Fig, Sy; body dark. tegula and lees 


R. K. SCHUMACHER, A. D. AUSTIN & R. B. FLOYD 


10 


- ‘ds weyMuposayy - - - . - - IRPIRULOIA 
ednd proysesed 
/ uMouyUN aseys 
‘ds “jsoy Uestaidopiday 
- SMUSP] - - - - - - Sepluuse[ ay BIA ploysesediadAy 
ednd proyseied 
/ umouyUN asejs 
‘sploqsesed pruoyarig jo 
“ds prouseiediadAy 
- SHAOYIOSI PW - - - - - - aepruowmauyay aes1qoO 
ednd prousesed 
/ umouyUN aseys 
‘splousesed pruoseiq 


SDUOLPS] JO pluowmneauyal 
- jo ‘dds 7 - - - - - - oppruowmnauysy jo plousesediadAy 
paral 
- jou andnd - - - ‘ds [x ‘ds [. aepruryory, 
‘ds “ds opundaAry 
- - SLIDUR - SIDUYG, SOLIS LA paynuaprup - 
wos parma wnsodns pupapd wunsodD Is ‘ds wasodpas ednd / easel {J Jo 
piujadosaja fy jou avdnd = putjadosaiayy VIIAIDSaW — Puljado.sajaH] DILAIDSI IA pupado.tola yy - aepluowmneauydy plousrsed AiPulig 
- - ~ - = - = ‘ds xajdodiup 
- - - - . ‘ds smoqueq - ‘ds snuoqgiq BAILY 
&T/ PAIR (77 JO 
- - - - - - - ds p1imuise) aeprluounauysy ployisesed Arewnig 
Pani Pag 
DIIDUISPD PLIpUISPD - - - - - - arpmuounauyy] 
pjadiupsauul 
sajajupdnjds}y - - - - - - - o-r1 
INILMIWUAIB aDIaUloas / teysut Apa jo 
pisaloyd vISAIOD - - - = - - aepruoonig plouseied Asewtig 
(ey) 
rola vSPOL AN (vy) (B]) 
» LOV SAS (BP) “StL § SOA ay [Pua 2 SBL WSEL MAN wSRLS 
pasiauta 
saisads prose Ayruurey / poyorne adels soy 


“peynuap! iou saisadg., “([sv Wl QOs <) apmiyye ysry = ey “(pse We OOS >) apN NE MOY = RY APMIS SIs “26GHGL SOBANT “FRGL PINEDOs "| 1 86L ALT @Py "SZ61 
PAOJYSLY 2 NOM[Ay “BVP JO sadunog “HIPusyAy U1 SUOUPIL] JUasaffip 1p eyeALd epaduresauy] YM Palp1ossy sployspavd [PdNd-]oAAV] PU [DAAD] fO UOSLIPAUO,D * ATAV, 


PARASITOIDS OF THE AUTUMN GUM MOTH I] 


2.7mm, ¢ 1.3-1.9 mm) (Elasmidae)......00.... 


F Elasnius sp. 
Hind coxa normal size; hind tibia without 
criss-cross pattern of setae (Fig. 16); fore wing 
suigmal vein elongate (Fig. 7); body vivid 
iridescent green or blue-green (2? body length 
3.2mm, ¢ 2.3 mm) (Pteromalidae)............. 
ronquapeinsidgdeguagange oecegagenghtertbseats Megadicylas sp. 


Description of new species 
Cotesia geometricae Austin sp. nov. 
(FIG, 9) 


Holotype: 2, Victoria, Altona, Dow Chemical Plant, 
10,1x. 1992, ex Mnesampela privata (ANIC). 
Paratypes: 25 292, 10 ¢d, same data as holotype 
but 10.1x.1992 and 8.vii.1994 (ANIC, WARI); 10 
22,244, Lyneham Ridge, Canberra, ACT (ANIC, 
WARI). 


Female 

Length: 2.9 mm. 

Colour: Head black; face, vertex and occiput with 
dull lustre; antennae and mesosoma black; coxae 
black, legs yellow-brown, apex of hind femur and 
tibia sometimes with darker patch, tarsi black: 
metasoma black except for laterotergites of segments 
1-3 which are yellow-brown; wings hyaline, stigma 
uniformly dark as are fore wing veins C+Se+R,. I- 
Ri, rand 3-Rs, these veins being darker than the rest. 

Head: Face, temples and lateral frons with fine, 
dense punctation associated with pilosity; medial 
frons and vertex between ocelli smooth and hairless; 
eyes densely covered with hairs, face slightly 
narrower than half width of head (3.1:6.5), inner 
margins of eyes adjacent to face evenly curved and 
slightly converging ventrally; ocelli large, forming 
slightly obtuse triangle, lateral ocelli separated by 
distance from lateral ocellus to eye margin; antennae 
moderately long, about as long as body. 

Mesosoma: Scutum with fine, dense punctation 
associated with dense pilosity, posterior half with 
smooth medial longitudinal line; notauli only very 
faintly indicated by shallow depressions: dorsal 
scutellum smooth except for a few. scattered 
punctures along lateral margins; lateral band of 
scutellum very broad and smooth; metanotum not 
filting against posterior scutellum so that phragma of 
scutellum exposed laterally; propodeum coarsely 
rugulose-punctate, with slight mid longitudinal 
depression and a few short striae radiating from 
ventral margin; lateral pronotum with deep dividing 
grooves which are very slightly punctate; meso- 
pleuron with fine punctation associated with pilosity 
in anterior half and ventrally below precoxal groove, 


the rest smooth and shining: precoxal groove finely 
punctate; hind coxa finely punctate in anterior */; and 
associated with dense pilosity. 

Wings: Fore wing stigma broad, 2.3 x as long as 
broad; hind wing broad, vannal lobe convex with 
long marginal fringe of hairs throughout. 

Metasoma: TI slightly longer than wide (3,3:2.9), 
virtually parallel-sided, smooth in anterior half, 
rugulose-punctate in posterior part but much finer 
compared with propodeum; T2 subrectagular, 2.3 x 
as wide as Jong, lateral margins curving inwards in 
anterior part, surface rugulose-punctate but slightly 
less coarse than on anterior part of T1, longitudinal 
mid-line slightly raised and smoother than lateral 
areas, posterior and lateral margins with single line 
of more distinct punctures; length of T2-T3 1.6:2.0: 
length of T2-T4 equal; T2-T6 smooth and shining, 
with long scattered hairs along posterior margin; 
hypopygium with a few scattered long hairs, 
posterior margin straight to very slightly concave; 
ovipositor sheaths very short and straight, with a few 
long apical hairs. 


Male 
As for female. 


Host 
Reared from Mnesampela privata (Geometridae). 


Comments 

The sculpturing on the propodeum and TI-T3, the 
shape of these sclerites, the form of the hypopygium 
and ovipositor, clearly place this species in the genus 
Cotesia Cameron. Cotesia has previously been 
referred to as the glomeratuy species-group of 
Apanteles s.l, (see Mason 1981; Austin & 
Dangerfield 1992) and it is the largest genus of 
Microgastrinae, comprising hundreds of species 
world-wide. In Australia the genus is both common 
and diverse but other than several species introduced 
from Europe and North America as biological 
control agents for certain lepidopteran pests (see 
Austin & Dangerfield 1992), the Australian fauna 
remains virtually unstudied. One other species, C. 
wabae Austin & Allen, is also associated with a 
eucalypt-leeding host viz. Uraba lugens. Cotesia 
geomerricae differs from this species in that it is 
gregarious, the first metasomal tergite is parallel- 
sided (not broadened posteriorly), the first and 
second tergites are more finely sculptured, the 
median field of the second tergite has rounded 
anterior Corners (not angled anteriorly) and the third 
tergite is smooth throughout (not sculptured 
anteriorly). This species is named after the family of 
its host. 


\ ROK. SCHIUMACTIER. A 1D) ALISTIN de RL BL PLOY 


Glyptapanteles mnesampela Austin sp. nov. 
(PIGS 5, 10) 


Haleawpe. V2 Australian Cayrital Territory. Lyneharn 
Ridge. 304.1993, ex Muiesampela private 
27 vi. [Y93 (ANIC), 


Parunpess | 2. 3 
(ANIC, WARI}, 


fd. same data as holotype 


Female 

Length: 2.5 mm, 

Colour: Head, antennae and mesosoma black pul 
propleuron yellow-brown; fore and mid legs yellow- 
brown, includin coxae, larsi darker; hind femur satel 
fibin yellow-brown, coxue black. tarsi darker (han 
tibia, distal tibia black; dorsal mietisoma yellow- 
brown in anterior half, black posteriorly, ventral 
mekisann yellow-browr inamtertor Lwo-(hirds, black 
posteriorly: wings byuline. venation nmod-erately 
dark, sGiginit uniformly dark, 

Head: Pace, temples and titeral Trons. smooth 
exvepe for scattered micropunetures associated with 
fairs; medial fromk. vertex between oeelli and 
vecipul smooth and hairless; eyes densely covered 
with hairs, diee slightly narrower than fall wide af 
head (2.6:5.8), Winer Margins of eyes adjacent to fee 
evenly curved: ocelli forming broad obtuse triangle. 
lateral ocelli separated by more than distuiee tron 
literal -ocellus lo eye mirging antennue long, much 
longer Thar body, 

Mesosorma: Scutum finely und diseretely punciale 
medially, becoming more densely punetale-reveulate 
towards lateral margin, densely covered with hairs: 
nou only very faintly indicated by shallow 
depressions; clorsal scutelliins smooth with ou few 
small sealtered punchiress lateral band oh scutelhin 
very hroad: ietavotum fiting closely against 
posterior scutelhion. phragm ob seutetun aaty 
exposed 1 Hiteral comers; propodduin smooth wath at 
few small radiating: strive postera-mnediolly; anternor 
hall al propodeanr with a few Tairs and associated 
INidroplineturess Mmesopleuron smooth, covered with 
Kies (anterior oneathird and ventrally below 
precoxal groove, precoadl groove unseulptureds Hind 
coxa finely punclite. covered with Tairs. 

Winws: Fore wing wilt stim broad, about 2.5% 
as long as broad: buse of 2-M piginented: hind wing 
browd, vannul lobe convex with long marginal fringe 
of hairs throughout, 

Menisonte Th mestly smooth with a few faint 
striae, lateral mnaegins evenly narrower, willl a few 
scattered long hairs: mediin field of T2 delimited 
hileralty cand posteriorly by cistinet suler. allvase ar 
vyuihuera! witigte in shape. length oF T2-Ts barby 
T? Te smooth dnd shining, T2 without hairs, T4-PA 


willl long scattered hairs; bypopysioni will a few 
seuttered long hairs: ovipositor sheaths very shor 
and straight, with o few long apical hates, 


Male 

As for female except as Follows: Antennae longer 
and more robust: head and scutum with denser white 
hairs: scutum with denser punehition: melasoma all 
black: hind legs with fermtu darker or ifuseate, 


Host 
Renred from Muexampela private (Geometridae }. 
the wulume eum moth, 


COMUMENTS: 

Giyplapanteley is i urge cosmopolitin: genus oF 
several hundred species. which comprises mosity (he 
viripennis and aelonediiiy speetes-2roups (sense 
Nixon 1965), a5 wells cbutnber of smaller groups, 
wll of which are extraclimilal except for rhe 
monuspecifie demeter group Tom New Zenlint 
(Nixon 1965, Mason 198) In Australia and 
Tasnninit there aire ai estiinated 100 plus species of 
Ciyprapanteles, only one of which ts deseribed., G. 
deliaya Austin & Dangerfield, The gents is. most 
diverse in the fropien! parts of the continent, sul 
uppears to be dispropartionately represented in {he 
microwasirine fain of the south eastem Pacilie, 
from where a number of aberrant species hive been 
deserihed, ca. G. denrerer (Wilkinson) Trant New 


Zealand, which is strongly dors ventrally flottened 


und G. ufimatiane (Pulliwiay) from Samoa, whet 
has weomplete medial propodest carina. vasiform 11 
aid stub of vein 3-Rs present in the fore wing 
(Austin & Dangerfield 1992), 

Glyptapunteles ineseanipeld can be distinguished 
from G, deflasd and most other undescribed 
Australian species by is colour shape of rictisen.il 
jergites | ane 2. and seulpturing of the sculuin and 
propodeuin, Although the degree of host speciticily 
of Chyprapaateles spp. is poorly Krown. ib is abso 
likely thal host associations provide a usetul initial 
buide to the identity oF many mierogastrine 
parasitoids and this iy probably the ease for (his 
species, G)\iprapanieley immesampela is named ale 
the host genus, 

Discussion 


This study has expanded the known parastbur 
complex of darvae of Ad peter in south-easiern 
Austedlia to inelude tbe primary parasitoids 
Cayinaria aero and bve new brieond speeies. OF. 
noiesampelea and Co eeameanricad und five species ol 
hyperparasitaid, m audition to several previotisly 
recorded species Ullioit & Bushlord 1978; de Little 
19O¥t!: Gaul 1984; Lukucs 19092) (Tible 3), The 


PARASTLOUSS OTHE ALLEL Ms GUM MOTH ia 


Parasol Conples OF gd host species is oflerr stable 
bebween geogeaphicnl locations wher (he host feeds 
on the same (or kexonomieally related) plam species 
(Askew & Siniw 1O86: Mills 1993). However, the 
puraotoids reared from WZ. privatia do not appear io 
follow (his vencrilisalton in thal the species reared al 
the two omainiand sites (Vietorit and the ACT) 
differed both front each other and trom those 
previously reported fron ‘Tasmania and Victoria 
(Mable 5), Th particular, Gr. vimesuimpela has only 
heen Lounel ii tie ACT bul no hyperparasitoids were 
recorded from this site wndiare also apparently absent 
i Risnumi Two speees of larval parisitoid 
recorded jn Tasmanti (Elliow & Bastitord 1978; 
Gaul (984) Tinve nol heen recorded) front the 
miintind and several larval-pupal parasiieids 
recorded ih Tasman and) Vietorat (Riot & 
Rashlord M78; de Lith: LOST Gauld 984: Lukdes 
19997) Were fot found in tity study. In addition, hwo 
diferent species of primary purasitoid were collected 
In consecutive yeursad Lyneham Ridge although i is 
possible that how spewies were present during both 
yours bit were not found duc to the low level uf 
PUMSTst ab ihe site. ba part differences: belween 
purnsionl compleses mity refleel cotlecting bias and 
an initil naive separation of different purasitoie 
species I the 1992 Victoriun collection (i.e, pupae 
were nal reared and GO. nitesemiprela may hive been 
comlised wih, seamedivae), However, the relative 
dburidinee af host cad syrniticanity influence the 
vomposiian und number of parasitoid species i can 
support (Mills 1900) MilIK a Kenis 1991), The 
frevler species Hchness of (he parasitoid complex 
Jou al Allona amt loo lesser exten) Shepparton. 
therelore. may alsa bedded the higher relative host 
ubundiinee ab Wiese loeanons ‘The paucity: of 
pardsitalels iy the de Little (MOST study |Table 5), 
whieh wis alse undertaken during a severe oulhreak- 
may fe beease only puasitouds seen oviposuing 
int hvac Were idenitmd, Dallerences tn Hast 
pheniloey may aso contribute to differences we (he 
make-up ok parastou! complexes (Askew & Shiny 
M88} ATIQOULR most pepaluuenis OF Ad, pay iais hitve 
a lonunmint aqulummoa winker populition: (MeQuillar 
IMSS) camel ain ilmest insignificant! sumnier 
popabiion (Lakees TYY8) (he furter study found 
Uiett populudions wt higher altitudes mt Tasemiia (> 
ADOH) asl) can aise lave a line stimmer—qutumnn 
POPULATION Ad dre poreataully: bivoltine, With Unis in 
Mond TH should be noted thal (he survey by de Little 
(29YRE') wus nde dering a sumer outbreak ala 
fineh altitude locahion in NW Tasmania whereas 
Pitiolt & Bashlord (1978) and this sruicdy examined 
Jower alittle populations present over cucu and 
wirer (NB. Lubes (59999) idk nop diferente 
benween Suntiner tind sutunit pOpURIiOns at best 
larvae or specifically reeonl the altinde el the 


collecting sites), Vurther jnvestizanan ol the 
influence of temporal and geographical variation of 
host abundance on the composition of the parasitoid 
complex dtilisiig farvae of MW, private will be 
required lo clarify these diserepuneies. 

This study found that Wt. private had an equeil sex 
eno at both pupation ind emergence. However, 
uithough the present results did not shaw a 
significant departure roma del sea ratio, the trend 
towards i greater proportion of female adules 
resembles that found by Elliott & Bushlord (1978) 
who obtuined 64% lemale adults tron reired larvae. 
The significantly lower weight oF pupae (hal died 
shevests thal hevae mast uchieve some critical 
Weight to survive fhe pupal pened. The degree of 
overhyy ol weights between wable male und fenmiale 
pupae of WL privat indicites (hit prediction of sex 
based on weight is nob lewsthle. 

The primary partsitoids of AQ, preven harvae 
recorded inthe uid otherstadies (Allion & Bashford 
JO7R: de Lithhe 1981! Gauld POR4: Lukues (9997) 
kill their host fn late tystars or ais papac. These 
punisdoids my reduce defoltation fo some extent as 
Juryue are most destructive in thea foarth and fh 
josnirs (tow & Bashford 1978). However in 
addition to their effeet during the Curren) seqson, 
larval parasitoids May reduce fle number ot MW 
private (hal emerve in the neal generon. The 
potential role of these parasitows in’ bighagieul 
control, therelore. is likely Jo be one al regulaion 
und prevention of onfbyeaks cather hai as a rethou 
OF CoOmLrOL when an Outbreak is Occurring, 

Although the causes of outbreaks et WL priveva are 
qouear i appears that large monveuliures af 
ocnelically sumtlir species aire especially Vulaerible 
(Neumann & Collett 1997), "The inefieetivenass af 
parasitisnr in such Situations (Ellow & Bostford 
IS7K: de Lathe IRE Neumiinn & Collen 1997) 
may be due to low otimbers al natunil enemies 
Irmited by the liek of allertative blood sources, 
coupled with rapid populition growth ol the insect 
herbivere iran ares of dense und abundant resources 
(Root }973, Alven & Letournedu LOS Alder eral. 
(DUS) Tn saldiion, the use oF nin-speeilie 
insecticides will Curial the momerual nespume al 
larval parusdoids and thus prevent their comipibutienr 
to Stub isiigs fast POpUlaliogs, 

Resource) inte tre sustiinuble manigenent of pest 
Inseets pS See USN imperiunt step to iniprove the 
CUrrenily poor exonomic reruns tron eueulype 
Jorests (Stone 1993). Curren teseureh in Australia 
has ineclided the evaluation of simins ob Bues/iy 
Nawtaseavin Berliner (llareourt a al | O06: 
Neumann & Collet 1997), the Use of maturedly 
resistunt species and pravenainees ob uc ayes 
(Parrow era. 199) ad Hiteneuous between the 
section ce parison, Zefequiats spo anil its host 


\4 Roi, SCLLUMACTIER, A.D. AUSTIN de RB PLOY 


M, private (Schumacher (997); Lukaes 1999%), The 
effecr of augmenting or encouraging natural 
populations of larval and egy. parasitoids of MW, 
private through practices sach us the ase of selective 
insectivides und ihe provision oF alternative Tood 
sources for adult parasitoids has not been 
investigated. allhough the benefits oF such practices 
have been shown in eucalypt plantations in South 
Americu (Brayunga er af, 1998) as well as other 
pest-parasitonl systems (Idris & Gratius 1995; Orr 
& Pleasants 1996), The necurate dentiication of the 
nittiral enemies of ML privat and an understanding, 
Of their ecology will be essential to the success of 
such researely in Australia. 


Acknowledgments 


We thank |. Naumann for initial identification of 
some of the parasitoids and WN. Stevens and P. 


Dungerfiekt for the line drawings. We ave grateful to 
R, Farrow for the collection of insect material i 
Victoria, M. Court for helpey coating and P. Hlart for 
vecess lo constant temperature eabinets (CSIRO 
Entomology). We thank SN. White for sfalishest 
advice and S$. Sehinnl and J. Seymour for 
constructive criticism of the manuscript. This study 
wits Supported im part by yrants From the University 
of Adelaide and the Australian Biologicul Resources 
Study to A.D.A, 


SUWPMACHERA R Ko C1MO7) A stialy OF Une piiristiiids af the cas 
SENG farsi UE (Ae AT TE ER resiegredin petite (len) 
Loepldepreny Geomenitac tan Fueddpni ilidaiin Labi subsp 
tucestata (Mote. Bhibely & SHIT a rkpth Oy Csinberra, 
Ansteatin Capital Terriory, GPSe hess, Austiahiiihy Stina 
Lmyersny Cuuput 


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FOSSIL LIZARDS FROM THE EARLY PLIOCENE 
BLUFF DOWNS LOCAL FAUNA 


By BRIAN S. MACKNESS* & MARK N. HUTCHINSON 


Summary 


Mackness, B. S. & Hutchinson, M. N. (2000) Fossil lizards from the Early Pliocene 
Bluff Downs Local Fauna. Trans. R. Soc. S. Aust. 124(1), 17-30, 31 May, 2000. 

The lizard fauna of the Early Pliocene Bluff Downs Local Fauna consists of members 
of the families Scincidae, Gekkonidae, Agamidae and Varanidae. At least three living 
scincid taxa, Egernia hosmeri, Tiliqua scincoides and a member of the Eulamprus 
quoyii complex, have been identified from dentary, maxillary and skull roof elements. 
Key Words: Pliocene, Bluff Downs Local Fauna, lizards, Egernia, Tiliqua, 
Eulamprus, Heteronotia, 7Megalania, varanids. 


Tresaerions of the Royal Spclery eS, Aust (2000), P2400), F740 


FOSSIL LIZARDS FROM THE EARLY PLIOCENE 
BLUFF DOWNS LOCAL FAUNA 


by BRIAN S. Mackness & MARK N. TutcHinson' 


Summary 


MackNrss, BOS. & PIPCIINSOB, MON. (2000) Mossi lizards from the Early Pliocene Blut! Downs Local Puan, 


Hans RoSec. & Ati. 1401), 17-30. 31 May, 2000, 


The tvand fauna of the Barly Phloeene Blull Downs Local Faun consists of members of the finhes 
Scincidite. Giekkonidie, Agamidie and Varanidve. At least three Irving scineid tina, Bgernia hosment, Titiqua 
weinemiles and a member of the Fi/ampruy queyil complex, have been identified fron dentary, taxllary une 
skull roof clements, The gekkonids are represented by a early complete dentary, whieh is lonkuively referred 
© the living genus /ereromaia. Agamids are represeated by unidentified dentay remains. A giant varamid, 
Mevafaiia sp. wae the valy extinel lizard found: other smiller varanicd remains were aso recovered The 
cubkonid is (he firstoublished Pliogene record of the Homily front Australia, The luna, consisting largely OF biaxat 
Will close phylerie hak to (he modern lizard fun eteastern Australia. as forther example of the evolutionary 
conservalisun of Australian herpetological assembhiges since (he Miocene, 


ivy Worns: Pliseene. Bla’ Downs Loeal Pauw. livers. Beene. Pilgue, ufanipus, Hereronona, 


Mervaliafua. varias, 


Introduction 


Squamates dre poorly represented in the Australian 
fossil record (Molnar 1882. J98ki.b, }Y8S, L991). The 
few reeards older than the K-T boundary 
(Bartholomar 1979) aie suspeet, as the groups ta 
Which they were assigned (Paliguanidae and 
Profactertiformes) are now thought ta be won- 
squiimite or at best, of uncertain ilfinities, Molnar 
(I985) reported (hat vertebrue of a small lepidosiaur 
(possibly a sphenodantid) had been found in the 
Cretaccous Toolebue Formation of Queenslaund. Some 
Iragimentary remains of lizards” ineluding a rib 
(Flannery & Rich 1981), a humerus (Molnar 1980) 
and an incomplete roaxillary (or dentary) (Molnar 
J985) have yet to he fully described. A. supposed 
lizard jaw from the early Cretaccous Dinosaur Cove 
has proven to be a fish «Molnar & Czeehura 1990). 

The eartiest Austrian Tertiary Heard mentioned mi 
the Hterature (bused on a submitted niuiuseripleited 
in Rich eral. (991) 1s front the Eocene Tingiuinurrit 
Local Fauna (Molnar 1991) Now published, this 
paper (Gedthelp es al 1992) reports no lizards. 
Unpublished date do-exist to indicate the presence ol 
squamiates al the site (MNEL pers, obs.) but the 
remains ure a) present, too Few and Tragmentary to 
reach conclusions regarding their rehitionships to the 


Scyeul of Wolugicat Sciences, b niverity of New South Wate- 
Kensington NSW 2052, Present deuress: PO Bos aot) Beerwoll 
Ol STO 7 nti: mew biniitercompaeerye.cuin 

Departing’ a Flerperology, Soulh Avstiatiin Musenit Serth 
Herder Adelaide SA HD, 


living fauna, Che first confitmed record of 
lacerulinns with modern day counterpuris comes 
from the Olige-Mrovene deposits of central Australia 
(Stirton ef al. 1967 (this record is actually ofa snake 
vertebra) (see Estes 1984) (Estes 1984: Pledge 
1984). The Miocene limestone deposits. of the 
Riversleigh area of north-western Queensland are 
Khown to contain Lossils of agamids, yviaranids. 
gekkonids and scinicids (Archer ev ad 1989, 199]; 
Covacevich ef af. \990; Hutchinson L992) Shea & 
Hutchinsen 1992). 

Pliocene reeords of lizards tnelude \eranis sp. 
indent. and Lilie sp. from the Curramulka Locsl 
Fauna (Pledge 1992), Tilignd sp. from Wellington 
Caves (Hind ef af, 1988), Megalania sp. trom the 
Chinchilla Local Pauna (Heeht 1975) and a Teaard 
from the Bow Locul Puuna (Skilbeck L980) Archer 
(1976) reported an avgamid ane Varad sp. from the 
Early Pliocene Bluff! Downs Local Fauna. A number 
Of NON-SQeaMatiin taxa bas been previously reported 
from this Kui (Archer 1976, (982: Bartholomai 
1978: Archer & Dawson 1982: Rich & via ‘Tes 
1982: Boles & Mackness 1994. Mackness [995 a.b: 
Willis & Miackness 1996; Thomson & Muvkness 
1900: Wroe & Mackness 1999), The present paper 
desenibes the lizards identified to date and discusses 
the implications for paliwoecology of the site and the 
pattern of Tizard evolugon mh Australia, 


Materials and Methods 


Fossil remains of repules were obtained through 
quarrying or through wWel-steving of sediments, 


Is U.S. MACKNESS & MON. HUTCHINSON 


Tani L. Mecsurement (mm) of dorsal ?Megalania vertebrae against Varanus giganieus vertebrae, Measurements 
as defined in Methods. Range (mean + standard deviation), Data fur V. sigameus are taken from Sinith (1976) 


Specimen No PrPo BW /PrPo CW/Pr-Po Pr-Pr/Pr-Po 
Vi giganteny 20 245-271 (25.74 14) (.54-0,.64 (QO. 58+,005) .55-0.63 (0.594006) QO 88-1,01(0,92+.007) 
Fossil 7 31.7-45.8 (36.44 3.3) O.48-O.457 (0.53404) 0.56-0.72 (0,63+,06) 1076 1.201, 144.06) 


TABLE 2. Measurements (nun) of individual ?Megalania vertebrae. Measurements as cefined in Methieds- 


(Meu + standart deviation). 


Specimen Pr-Po Pr-Pr Po-Pu uW CW Type 
POLAS 31,7 38.2 34.5 15.4 19,9 Dorsal 
£23234 40.0 46.5 13.5 22.) 25,0 Dorsal 
22235 33,7 - - - Dorsal 
F232356 32.1 = 20 16.8 - Dorsal 
F23237 32,7 35.8 157 19,9 Dorsal 
F2368) 380) ALS 32,3 21 Dorsal 
F23686 15.8 53.5 257 32.4 Dorsal 
F23084 38.3 - WA) - 143 Cervical 
(Mean/SD) (46.445,4311 (43,247.05) (46.3454) (1744.2) (24h 15.5) 


Specimens were examined using a Wild M3Z stereo- 
Microscope with eyepiece micrometer and drawing 
tube, X-ray microanalysis was carried oul using a 
JEOL JSM 35 scanning electron microscope and 
energy dispersive X-ray detector. Terminology for 
hones follows Romer (1956). 


Measurements 

Measurements made usiig vernier calipers 
wecurle to 0.05 min are summiurised below and 
largely follow Smith (1976). Statistical analysis of 
these measurements is provided in Tables 1 & 2, 

VERTEBRAL LENGTH (PR-PO) - measured as the 
greatest distance from the anterior cilge ol the 
prevygapophysis to the posterior cdge of the 
postzygapaphysis. 

PREZYGAPOPHYSIAL WibTH (PR-PR) - meisured ius 
the greatest distance between the edges of the 
prezyyapophyses, 

POSTAYGAPOMIYSIAL WibTH (PO-PO) - measured 
as the greatest distance between the edves of the 
postzygapoplyses. 

CENTRUM MINIMUM WIpTH (BW) - measured its the 
smallest distance across the centrum. 

ConpyLar Wipri (CW) - measured as the greatest 
distance between the condyle. 

Abbreviations for specimen numbers: QM) &, 
Queenslind Museum fossil nuinbers; SAMA, South 
Australian Museum, Adelaide: AR, University of 
New South Wales Research Collection, Site 
localities are listed in Archer & Wade (1976) ana 
Mackness (unpub.), 


Systematics 


Family Gekkonidae Gray, 1825 
Subfamily Gekkoninae Underwood. 1954 


cf, Heteronalia sp, (Gray. 1845) 
(PIG, LA) 


Material examined: A single nearly complete left 
dentary (QM F23655) from EVS Site. 


Characters 

Gekkonid dentition is characterised us isodont with 
aJarge number of cylindrical. pointed teeth confined 
to the marginal bones (Edmund 1969), Sumida & 
Murphy (1987) have amended this by reporting that 
nekkonoids typically have bicuspid tooth crowns, the 
cusps being separated by an apical groove. 
Allocation of this specimen to Heteronotia is 
justified below. 


Description 

The specimen is an almost complete lef dentary, 
complete anterior to the level of the posterior end of 
the tooth row and retaining mosi of the angular 
process, There ure 36 teeth or toath loci. The jaw is 
relatively slender and euryes mesially at about the 
level of the 18! tooth. Meckels's groove is 
completely obliterated by dentary overgrowth, The 
internal septum is not exposed by the splenial notch, 
which extends anteriorly to the level of the 29" tooth, 


FOSSIL LIZARDS FROM THE BLUFF DOW LOCAL 


Fig, | (A). cf Heteronetia sp. QM F23655 left dentary, lingual. Scale bar = 2 mm. (B). Agamid indent. QM E7812 dentary, 
lingual. le bar = | ram. (C-P), 2Wevalania sp. QM F23686, dorsal vertebra, (C), Lateral, (DO), Anterior, (E), Dorsal. 
(FP). Posterior, Scale bar = 20 mm, 


a9] B.S. MACKNESS & M,N. IUTCUINSON 


The teeth are delicate. sharply pointed and weakly 
bicuspicl with o fingual cisp ser oll hy a groove and 
lying lower than the apieal cusp. There is. little 
variation im lool size apart from eshght diminution 
posteriorly. The dental sulcus ts well demarcated by 
a Negual puraper. Labially, the jaw bears Four 
widely-spuced menuil foramina, the must posterior 
level with the 25" Looth: 


DUNC RSTOS 

Jaw depth (at the level of the posterior niust tout), 
2.6mm, length of tooth row, 12 mm: height of 14% 
(ooth. Q.4 mm, 


Remarks 


The dentures of vekkonids. are conservative, 
making idernfification difficult. Pygopods have 
Iimhly variable and specialised jaws (Hutchinson 
1997), none of which strongly resembles the 
speeunen uesedibed. Most of the larger 
carphodachylings have distinctive, lapering niult- 
toothed (> 40) dentaries quite untike this specimen. 
However most diplodaetylings and the Australian 
gvekkonmes have less specialised jaws which we 
supericially sinilar to each other, OF the genera 
examined (all Australian gekkonoids except 
Psendethecadaciviiys (=Rlacudactylus)). the most 
similar to QM F24655 1s Lereraitotia. AL present, 


(is attburion is based only on the canmbination of 


jaw proportions, tooth number and size; objective 
crileria for most vekkonid jaws have sull ta be 
developed, 

Hereranotia hinoes (Gray, W845) 1s a wide ranging 
complex of al least (Wo bnsextul species and 
numerous all-female parthenogenetic clones 
Members af the complex are found througheut 
muinkind Austrulia (Moritz 1983; Mority et af, 1989) 
Wiha varicly of habrats ranging (fon deserts to-closed 
lorests. Heterdnotid balver is noeturnal and feeds on 
arthropods (Bustin M968, 1970). The relationships 
Ol the war other species, A, ypefed (Kluge, 1963) and 
He plaicveps Storr, 1Y89 to the known chromosome 
races and jo each other are yer io be clarified. 
Pending the chirifiestoo of species boundaries, we 
cunimike no firmer allocation of the fossil speeimens 
Hun us a posstble Herermimeatio species. 

Fossil gckkonids have also heen recorded trom the 
Quatenpary of Quecustand (Archer 1978) 


Famuly Avumidive Gray, 1527 


LUindentiticd Material 
(iG. |B) 


Malena cevaerined, Two Trigmentiary  reght 
dentaries. the symphysial region of the jaw (OM 


F23056) BVS sie, the ether (QM F7S812). a purfial 
dentary beariny the niid-to-rear section of the tooth 
row from Main Site, 


Characters 

Agamid reptiles are distinguished by having a 
dentiuion comburiig one fo three anterior pleurodont 
teeth followed by aerodont tweth. Other agannl 
features are Summiurised by Estes (1984), 


Dexeriprien 

Both specimens are from relatively moderate-sized 
individuals, Phe partial right dentary (QM 7812) 
bears seven acrodont teeth. The lust implanted tooth 
is followed by an empty locus, The dentury is broken 
anteriorly at whut is esamated fo have heen about the 
mid-pointor the tooth raw. 

The second specimen. (QM F23056), ts the 
symiphysial repion of aw eight deotary, Ip bears Wwe 
pleurodont tooth loei (one tooth present, one absenty. 
followed by five partly damaged acrodont teeth 
Three elosely-spatced inental fortinina are preserved 
on the labial surlace of the specimen. the third being 
sited below the first aeradont tooth, 


De STONS 
Jaw length (OM P7812), 7.8 mm: (QM P2305), 


Sain, 
Remarks 


Archer (1976) suggested ihat the small right 
dentary (OM F7812) was similar to some species of 
Amphibalurus. Since that tune. the  generiv 
clissiication of Australian agamids has been 
considerably revised (e.. Storr 1982) and. further 
review of the phylogeny of the Australi diigons is 
likely (Greer 1989) Covacevich of ul. (1990) 
identified several problems in idenulying aganrid 
remains and we follow their cdudgen ino mit 
identifying these dentary fragments past family 
level, 

Dragon lizards ure divided Into three fireayes 
within Australia: the amphtboluroicds — (eevesi 
Hutemoson & Donnellan 1993), U7 ypartieris and 
Physignetiuy. The aniphibolurids inhabit nearly all 
coviraninents vaceph wet forests (Hutehinson & 
Doanellan 1993). Physienahis occurs along streams 
ina variety of habitats while Aypsitieas is restored 
to closed canopy forests (Witten 1993). 

Fossil ugamids hive been recorded trom the 
Quarernary of Queensland (Bennert 1876, Archer 
197K: Archer & Braysbaw 1978), New South Wales 
| Ryder 1974; Dodson ef a7. 199A: Balme 1995) and 
South Australia (dale & Tindale 1930; Hope et al 
1977; Smith J976, 1982: Sinitly PYK2; Pledge 
1990), 


FOSSIL LIZARDS PROM ‘THE BLUEF DOWNS LOCAL FALNA iI 


Family Varanidae Gray, 1827 


2Megulania sp. 
(PIGS 1C-T, 2A-C) 


Matevial evaiined: Two isokited cervicul vertebrae 
(QM P2364) JY Site, (QM £23233) Main Site, 
seven isolaled dorsal vertebrae (QM PY135, (IM 
23234. (QM 23245, QM 123236, QM 232357) 
Main Site; (OM F232686, OM F242687) DML. Site 
and condyle fragments (QM F23088 ). 


Characins 

Megelamia is characterised. im part, by having 
tise Porch and lombar vertebrae with weakly 
developed zygosphenes (absent in typical Varese) 
as well us small depressed neural canals. The adult 
jeeth Gl Mevalatio are large and slightly recurved 
distully. The anterior cutting edge is roanided and 
serrated distally, The posterior cutting edge is thin, 
blade-like and serrated along its entire length (Hecht 
1975). 


Description 

The two isolated fossil cervical vertebrae are 
sindki inaverall morphology to those frorn an extiart 
Varans varnis OWhite. 1790) (AR7641) bur are 
some 227% larver. Both fossil vertebrae lack neuril 
spines but bear hypapophyses with the grooved, 
knoh-lke extremities characterishe Ob lirge-sied 
varanids. The cotyle is more robust and less flatlened 
than that of the extant species and the condyle ts less 
ovale. Both these features may be allometre an 
nature, however. The dorsal vertebrae are much more 
massive than those of any extunt varanig, Tables 1 & 
2 summarise the measurements of the ?Mevaliantu 
vertebrac. The animal represented was clearly larger 
than modern varanic counterparts such as the 
perentie Vuraiiy givearitens with the largest dorsal 
vertebra being almost 70% larger than those 
measurements supplied Tor a perentic: by Srtih 
(1976), 


Remarks 


The assigament of the larger varanid vertebrie to 
Mevahuida is made on the basis of convention, 
Vargius and Mevelania are the two genera 
recognised from Austilia, bul the (vo are separated 
primarily on sive. Estes (1983) doubts whether the 
two ough! to be generically distinct. Megelania 
prisca Owen, LS60) is the only species currently 
recognised ang as significantly larger than all extine| 
andbextant varanids. Large varanid veriebrae are also 
known from rhe Chinchilla Local Fauna (Hecht 
1975) und these along with these from Blut Downs 
ure curtently being studied by one of the authors 


(B.M.). Hecht (1975) in his study af MW. prisce noted 
thilt there seemed to he fewer caudal and cervical 
vertebrae ussociued with renmigins of Mevalaila 
compared wilh the expectition based on Vera. 
This suggested to him that Megalunia may have had 
a proportionally shorter meck and tail thin extunt 
varanids. No caudals have heen recovered fron the 
large yarumd front Blull Downs, 


Verrudus sp, 
(FIG. 2D-1) 


Material evamined: Wolited dorsal vertebrae (YM 
£7774. QM F23238. OM 123659) Main Site: (QM 
F23683) EVS Site: isoluted cundal vertebrie (OM 
P7777. QM P9131) Main Site; (OM F23681, OM 
F23682) EVS Site; and tsolated teeth (QM F236a5). 


Characters 

Vuraids are recognised by huving vertebrae 
eharacterised by oblique — vondyle-cotyhir 
articulations, parlicuhirly tt the Horace und lumbar 
region with the vertebral centra consiricted walerior 
to the condyles. The bases of the teeth are expanded 
and sculptured with fe vertical Moting (hejervary 
1935; McDowell & Bogert 1954: Romer L956, 
Hoftstetier & Gruse 1969; Tlecht 1975), 


Dasevription 

The dorsal vertebrae show characteristic varanid 
morphology ‘and will a range of centr lengths 
(lable 3) which indicate a medium-sized goanna 
such as Vara gauldii (Gray, Laas). 


Tasie 3. Meeasirements (imncop centre af passil veaeantely 
and Varanus varius (AR F041) 


Dorsil Cervical 


OM F774 142 
OM P23643 18 
OM F23059 LON 


OM F431 Rae 
QM F23681 18.2 
OM F23682 15.7 


(OM F2323K Hos QM FI777 Lh 
AR 7641 (0) WG 
ARTO41(b) ThA 
ARTO4F T(E) ln4 
AR 764 1(d) 1h4 
AR7TO41(e) lat 
ARTOIIC) 15.7 
Remarks 


Given that there appears to be a marrow range ol 
measurement within the dorsal section of the 
vertebral column of individual goannas and that 
there are two distinct size chisses of fossil dorsal 
vertebrae, i is reasonable to assume thar either iwo 


23 B.S. MACKNESS & M,N. HUTCHINSON 


Fig. 2 (A-C), ?Megalania sp. QM F23686 cervical vertebra. (A). Lateral. (B). Posterior, (Cj, Dorsal. Scale bar = 40 mm 
(D-F). Varanus sp. QM P7774 dorsal vertebra, (D). Lateral. (2). Dorsal. (FP). Posterior. Seale bar = LOmm, (G-1), Verran 
sp. QM FYT31 caudal verrebra, (G). Lateral. (11). Anterior. (1). Dorsal. (J). Posterior. Seale har = 50 mm. (K-1.). Tilique 

6 OM 123247 Jeft dentary. ( 


.). Lingual. (L). Labial. Scale bar = 20 mm. 


POSSIL LIZARDS PROM THe BLO DOWNS LOCAL FALNA 


sympalric species or two sive morphs ure 
represented in the Taunme The caudal vertebrae show 
womnch wiler range of measurements which is 
typieal of this seeuon of the vertebral column. Once 
amain however, lwo size clisses can be inferred 
indiculing that cirber Iwo symipatrie species or twee 
sive morphs ure prescott. Only one of the caudal 
vertebrie (QM PYT4]) has wecomplete neural spine 

Mertens (1942) suppested a relationship between 
body lengify and jail length which Meee (1975) 
further extrapolated, sopwesting that the fom tls of 
some vardnids may be oa reason for the disparate 
appearugee OF cqudul vertebrae Over [hose trom other 
resions of the body in the fossil record. “An equal 
number of dorsal and caudal varanid vertebrie his 
heen recovered from Bui? Downs. Wilkinson (1095 | 
has recently listed a humMber ol Vertebral features hor 
different Vergnds species, hough potentially useful, 
hese characters have heen selected from single 
isolated speeimens und therefore cannobl take inte 
weCOONL Interspecific and intraspecific varkiion. in 
Vavanid vertebral characters. It 1s nok possible: tu 
identify the lossil vertebrag beyond Varuius sp, partly 
die tothe current lack of information and because the 
veriohrie recovered were not articulited and were 
possibly from several teividuals, The vertebrae alse 
came from two different siles, even though those sites 
we from comparable depositional sequences 
(Mackitess anpub.. The size of the animal cannot be 
estrapolited from the vertebrae given the problems 
identified whove is well as those identified by Hecht 
(1975), 

Varanies are found over a wide range of habstars 
eluding aquatic. terresndal and arboreal and fron 
tropical lorests to wil deserts (Cover & Houtwole 
1OSL). They ringe over a wide area and eat trost 
food dems tneluding inverlebriules, vertebries ane 
curnon (Ring & Green 1979, 1993 a,b: Losos & 
Greene JO8K: James eral 1992), 

Possil varanids are known trom the Quaterniiry ob 
Queensland (Archer 197s: Walters 1980; Tope 
(OST: Lorton 19K: Wilkinson 1995), New South 
Wiles (Tealird 1967; Aplin quoted in Flope 198 t) 
South Australia (Hale & Tindale (930; Mulvaney er 
al (9642 South (976. O82: Sith [Y82: Hope era, 
1977) Willams 1980) Pledge 1990) and Westen 
Avstrati (Apeher bO77 1. 


bunily Seinerdae Cray, (S25 
Sublamily Lygosominae Mirtleman, {952 


THlignta Gray. (825 
Tike servoides (White. 1740) 
(Fic: 2 Kel) 


Material exauined: Almost conrplete Telh dentiry 
(QM 123247) IVS Site. 


Io 
py 


Charravrers 

The closed Meckelin eroove dnd the presence wed 
form of several lange hemisphericalcoanieal cheek 
teeth uniquely characterise this as belomeine to the 
venus Tique (Shea & Watehinsen 1897). 


Deseviption 

Me speenmed is lel dentary minus angular 
process, a vertical broken edge runs just posteriorly 
to the fevel of the internal faeet lor the dentary 
process aT the coronoul bone, but (he complete, 
posteriorly projecting corenoid: process ol the 
dentiry is sill present, Sixteen lveth or alveoli ure 
presen. check teeth increasing im size posteriorly 
wath the largest being the Eh, 12" and fa" ater 
Which last three are abruptly sinallen Ao buttress 
supporting the miundibular symphysis rises abruptly 
below the denlary as a low keel al about the level of 
the 6" tooth, vistble as a Vold) running: posteriorly 
along the ventral face of the dentary to aboot the 
level of the anterior alveolar foranen. The anterior 
end ol (his foramen, that is, the apes of the splenial 
notehy is at the level of (he 10" tooth, 

The erowns of the enlarged cheek Teeth are 
markedly wider thad the tooth bases, «ult ales 
horizontid fatlened pedipheral ocelusal surfaces 
rising fo a central potol, Strive radiate over the 
occlusal surface from this ventral pone, 


Meashve nents 
foothe row length 22 mm, jaw depth at lewel of bitst 
tooth 95 min 


Remarks 


The Hflgad dentary was found lying on lop of the 
ground at EVS Site, in un area undisturbed) by 
quarrying. Possils are often exposed at ihe Blufl 
Downs site through min and other disturbances. The 
colour of thys particular dentary was different (toni 
olhers recovered From the site. raising doubt about its 
provenance. X-ray microanalyses of mineral content 
(Pius (4) owere undertaken-on phe Tiligna jaw trom 
EVS Sites un extunt Tilfqiue. as well as on fossil bade 
Irdgments oF turtle anda python, both front LVS Site. 

The results af these microanilyses showed that 
Siliea was a prommingatcorstituculof the lossil bones. 
whereas in the extint Vilquee, little sifiea wits 
presen. Since the calcium phosphate of the bone is 
afien changed by the wdditton or substitution of ether 
onaerdhs. such as silica, during fossilisatioa. these 
fesults indicate that the Wie jaw bream EVS Sile 
wis fossilised und probibly contemporaneous with 
the other reptiles sampled, 

The Blut Downs specimen is nob distinguishable 
from the living 7 sedueurdes and al least one other 
Pliocene THiywu. reported by Pledge (1992) fram 


na) KOS MACKNESS & MON. TUPCTINSON 


Cirmamulka, cin alsa be allocated to this species. 
Examination of these Species (partial dentary 
masta and fron) by MNEH shows them to be 
indistinguishable from Utose of the living species, 
However another mueb larger specooen referable to 
Tilia, recently discovered from the Pliocene 
Chinchilla Local Fauna (Autehiisen & Mackness 
unpub... is markedly different [rom any living or 
extinel species of ie genus. 

Tilique scincoides ant is sister species 7 gives 
(Schneider, S01) (Shea 1990) are the mast tropicul 
and forest-audipted members of this genus, Ulfque 
seruvaldes (an aidaprable species, found ip ae wide 
variely OF hubitat types and its presenee does oor 
have strong pala¢oceological pnplications. 


Evernia Gray, 1838 
FEoerina hosmert Kinghorn. 1955 
(PIG, 3A) 


Material examined: A partial cought anasid tary 
fragment (QM F23654) EVS Site, 


(Weare ters 

The maxillary is identified as a spinv-tiiled skink 
(hk. chaning’ (Gray, 1832) group Gsensen Harton 
(972)) on the basis of iis tooth morphology having 
compressed crowns wilh geclisal blades ariented 
stich thatthe teeth inthe jaws forma serrated ening 
cde. 


Deseripninr 

Fhe speermen tf the pesterior suborbilal portion af 
the right maxillary tooth row. The V-shaped notch for 
The jugal is almest complete as is the dorsal edge 
(orbital rim). Posterior ning teeth or tooth floor are 
preserved. The crowns of more inltet teeth are 
labiolingually compressed, with agulioy occlusil 
culiing edges. Crowns dre also somewhat fared in 
lingual view, producing an overall “aee-oFspades 
shipe. This tooth shape ts limited to membersoof the 
Bvernia cunminghame species group (orto 1972) 
which comprises Fo canninvehami, Lb, depressa 
(Giinther, [S75). 6. Aesmert and E, stokesin (Cieay. 
845). 

The four living members ofthis species group differ 
in the dekuls of their dentition, Myernia eiminiighame 
wid /. depressa Hive squared-off, somewhat chisel 
Shaped crowns. Egerntr stokes/Pand E. hosinert show 
ihe ereatest similarity fo each other and lo QM 
F23654, all three having teeth with linguo-labially 
flattened crowns whieh cise low medial po SAMA 
specimens of L. wrekesi? differ slightly from those al 
FE hasmert in being rather more flared in lingual view, 
his expansion being rue even of the most posterior 
teeth, In, hesmeri anibQOM F23654 the last few tecth 
are narrower and more acutely pointed thin those oF 


stokesii. The fossil is not distinguishable tron 7 
fiaymert and is theretore allucated to thar species. 


Measurenrents 
Length of specimen. 6.9 mm; depth al level ol 
Jugal suture, 2.) mn: heighr of largest tooth, 17mm 


Remarks 


Spiny-tutled skinks are all crevice dwellers, 
typically in rock outcrops but sometiines ilso 1 lows 
and stumps, The better studied spevios (2. 
cunningham’ aod Eo stokes) are almost entirely 
herbivorous in the wild (Brown 1991), 


Enlamprus Fitzinger, 1643 
Ludamprus quevti complex 
(FIG, Al F) 


Material esamined: A right demury (QM POLST) 
Main Site: A frontal (QM 623657) Main Site and 
several other fragments (QM F2365K) AB Site are 
possibly also referable fo this taxon. 


Characters 

Eulamipris comprises the larger, more generalised 
members of the Splhennnerphas Group (Greer 1979) 
io Auswahia. [is definition ts currently based mainly 
on scalution and reproductive characters but initial 
work (Hutchinson (992) shows Unit by asing loath 
crown Morpholowy us well as jw robsiness: al 
Jeast Lwo morphological groups are fecognisible 
osieolovically, the more gracile water skunks, 
bE. grey (Dummer! & Bibran, (834) and its relatives, 
und the miure robust tropical forest species such is 
E, murray! (Botlenger, (S87) 


Deseripuen 

The dentary is pearly complete. with a tooth row 
bearing 24 (ecth or tuoth deci. The denuded sulcus ts 
demarcated lingually by 7 pronounced parapet which 
diminishes and disdppears at dbout the level of the 
22" Wooth, Meckels groove is Widely open along the 
ventrolingual face of the dentary, The internal seplunt 
of the deniary is poorly developed and does nol show 
muuch posterior extension. Seven mental focarninw are 
present, the last at about the level of the 14% tooth, 
The teeth crowns are not fared or thickened, The 
lingual face of each (ooth crown is vertical, wath) an 
iInwardqprojecting bultress offset to the rear The 
junction of (he lingual face of the tooth crown with 
the occlusal surface is dernurcated by a groove which 
sepumiles lwo clasely appased low ridwed edees. 


Meaxnremints 
Length of tooth tow, 12 mm: depth of dentary at 
level of (he 20" tooth, 2.6.0m, 


SSIL LIZARDS FROM THE BLUFF DOWNS LOCAL FAUNA 


Fig. 3. (A). Egernia hosmeri QM F23654 right maxillary fragment. Scale bar = 2 mm. (B). Scincidiae indent. QM F23659. 
Scale bar = | mm. (C-D). Eulamprus quoyii complex QM F9137 right dentary. (C). Lingual. (D). Labial. Scale bar = 20 
mm. (E-F). E. quoyii cornplex. QM F23657 frontal. (E). Ventral. (F). Dorsal. Scale bar = 20 mm. 


26 B.S. MACKNESS & M. N. HUTCHINSON 


2047 


Counts 
ae] 


Fe 
A Vv 


2047 


Counts 


KeV 


10,230 


10.230 


2047 


Counts 


2047 


Counts 


0 
D 0.000 


Fig, 4. X-ray microanalysis of bone. (A). Recent Tiliqua scincvidey dentary. (B). Fossil 7. seénevides dentary. (C). Fossil 


turtle shell. (D). Fossil pyihon vertebra, 


Remarks 


This specimen is very similar to living water skinks 
and is clearly distinct from the more robust rainforest 
Eulamprus spp. such as EB. murrayi in that it lacks 
their deep jaws and somewhat large. durophagous 
cheek teeth. However, the fossil appears to differ 
from E. queyil, the living species in the area today. 
Specimens of E. gucyii of comparable jaw size have 
a longer, narrower dentary bearing up to 30 teeth, 
Thus, if this specimen were E, queyis, it would have 
to be regarded as having an anomalously low tooth 
count. The proportions and the number of teeth 
accord better with &. tympanum (Linnberg & 
Andersson, 1913) and £. fheatwolet (Wells & 
Wellington, 1984), both restricted today to south- 
eastern Australia, 

Water skinks are largely confined to permanent 
water with this habit enabling them to inhabil a wide 
range of babtats, They are diurnal and carnivorous, 
feeding on invertebrate and small vertebrate prey 
(Daniels 1987: Brown L991). 


Seincidae indent. 
(FIG, 3B) 


Material examined: Three fragments (QM F2365$) 
EVS Site: one small fragment (QM F23659) AB Site 
and an isolated vertebra (QM F23660) Main Site. 


Remarks 


Four tragments of lizard dentary and a vertebra are 
not sulficient to be assigned. The three pieces trom 
the larger skink compare well with members of 
Eulamprus and are tentatively assigned cf. 
Ewampruy. The other remaining fragment represents 
another type of skink but not enough remains for any 
generic assignment. The vertebra is identified as a 
scineid on the basis of characters outlined by Smith 
(1976). 

Fossil skinks are known from the Quaternary of 
Queensland (Trezise 1970; Bartholomat 1977; 
Archer & Brayshaw 1978; Molnar 1978), New South 

fales (Krefft 1867, 1870, 1871: Lampert 1971; 


10.230 


10.230 


HOSSIL LIZARDS PROM THE BLUFF DOWNS LOCAL | 


Thorne 19712 Marshall (973: Ryder 1974; Dodson es 
al. 1993; Baline 1995). Tasmania (Bowdler 1974), 
South Australia (Stirling. [S89 Hate & Tindale 1930; 
Tindale 1933; Mulvaney 1960; Mulvaney er al, 
1964; Smith, 1976, 1982; Hope er af 1977; Smith, 
J9S2. Pledge 1990, 19922) and Western Australia 
(Cook 1960, 1964; Bale ev uf, L878), 


Di: 


ussion 


The paucity oF information about lizards in the 
Pliocune Makes (the Blufl Downs omuterial 
particularly noteworthy, The fauna, in so fas i ean 
he identified. is canmposed of species or species 
groups that oeeur in the moder Charters Towers 
area, the only obvious exception being (he extinel 
Megalaitia, In this respect, the tizard fauna of the 
Blatt Downs Loctl Fauna is similar to the older 
Riversleizh deposits (Hulebinsen dapub.y in that it 
represents an early cstublishment af modern forms in 
the area. contrasting with the conteniporancous 
mammal fain Whiel meludes muny extinel laxa. 

Stow rules OF faunal turnover, when eompared wah 
mammals, seem to be the rule in Tertiary and 
Quaternary syuimites. This hus been addressed most 
recently by the detailed study of the Rameho La Brea 
snakes hy Lie Duke (1997). His explanation of low 
nies of extinction, evolution and lunal tumoever in 
reptiles (compared with mammals and birds) centres 
On Two eoncomilins OF reptile eetothermy: - low 
enerey requirements and small size enabling survival 


“AUNA 7 


of repuile populations in refugia too small for athe 
tapidly rnetubolising and generally larger endotheris 
(witness the reeent situation in Australia regarding 
nimi versus reptile extinetions). “Uhus. during 
periods of environimentul change, miuny endotherm 
populubions and species cun be driven to extinenon by 
habitat reduction, while the synlopic reptiles ane 
amphibians merely suffer range contraction or 
fragmentation, Restoration of former climatic reximes 
permits re-esttblishment by the former reptile 
populations bul may require evolutionary change or 
migration before a new minmmal fauna emeraes. 


Acknowledgments 


J. Mead and kK. Aplin provided helpful comments 
onthe mianuseript. D. Sewell und L. Durham, School 
of Earth Seternces, University of Melbourne carried 
out the X-ray dueroanalysis. The Smith family af 
Biull Downs Stalion continue vw provide help sand 
support for the ongoing research into the Blatt 
Downs Locil Pauna. The collection of the Blatt 
Downs material Was supported in part by an ARC 
Program Grant to M. Archer. a grant Tram the 
Department of Arts, Sport. the Environment. 
Tourism and Territories to M. Archer, §. Hand and Uh 
CGodthelp. a grant from the National Estate Progrun 
Grits Scheme toM, Archer and A, Bartholorai and 
avails in aid to the Riversleigh Researeh Projvet 
Trom Wang Australia, ICO Australia and ihe 
Austrian Geograuphie Society. 


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NEW GALL MIDGES (DIPTERA: CECIDOMYIIDAE) INFESTING 
NATIVE AND INTRODUCED SOLANUM SPP. (SOLANACEAE) 
IN AUSTRALIA 


By PETER KOLESIK*, RACHEL E. C. MCFADYEN? & ANTHONY J. WAPSHEREE 


Summary 


Kolesik, P., McFadyen, R. E. C. & Wapshere, A. J. (2000) New gall midges (Diptera: 
Cecidomyiidae) infesting native and introduced Solanum spp. (Solanaceae) in 
Australia. R. Soc. S. Aust. 124(1), 31-36, 31 May, 2000. 

Three new Asphondylia species are described from five Solanum species in New 
South Wales and Queensland, Australia. Asphondylia sturtiana Kolesik sp. nov. 
induces a stem swelling on Solanum sturtianum F. Muell., an Australian native plant 
with fruits toxic to sheep and cattle, Asphondylia paucidentata Kolesik sp. nov. causes 
fruit galls on the native Solanum ayviculare G. Forster and Solanum linearifolium 
Geras. Ex Symon, and Asphondylia obscura Kolesik sp. nov. causes fruit galls on 
Solanum chenopodioides Lam. and Solanum physalifolium Rusby var. nitidibaccatum 
(Bitter) Edmonds, native South American plants that have become weeds in Australia. 
The newly described gall midges limit reproduction of their host plants. 

Key Words: Diptera, Cecidomyiidae, Asphondylia, Solanum aviculare, Solanum 
chenopodioides, Solanum linearifolium, Solanum physalifolium var. nitidibaccatum, 
Solanum sturtianum, Australia. 


Fiscctions of the Royal Scien ap 8 Aust (2000), BACT 3-36, 


NEW GALL MIDGES (DIPTERA; CECIDOMYIIDAE) INFESTING NATIVE AND 
INTRODUCED SOLANUM SPP. (SOLANACEAE) IN AUSTRALIA 


by Pribk Konestk , RACHEL EOC. McPapyven! & ANTHONY I. WAPSHERE 


Summary 


Rortsik, P.McPariin, RG. OC. & Warsuinikh AOE (2000) New gall midges (Diptera: Cecidomytidae) infesting 
nitive vind intreduced Sofware spp. CSolunaecac) i) Sustrala Trey Ro Soe 8, Aust 1241). 31-36, 31 May. 


L000 


Phree new ehapliondidia species are deseribedt trom five Sadaiwn) species in) New South Wales and Queenshind, 
Australi Asphondvia supiiaia Kolesth sp. toy. iiduces a sien swelling on Sela sieetiqnue Py Muell aun 
Ansialan native plint with fruits toxic to sheep und cullle, Aselomdvlic puucidentaa Kolesik sp nov. causes 
Nuit galls on the native Seva cmetiure G. Forster and Solan Hnearifotinn Geras ex Symon, and 
Asphondvlit obscura Kolesik sp. nov. causes fruit galls an Sadan chenapedioides Lan. and Sedan 
Piysalifalium Rusby yar nitdikveccun (Biter) Gdimonds. native South American plitts that have becorne 
weeds in Anisiralit, Phe newly described gail mndges Tinie reproduction ef thei host plants. 


Kry WORDS: Diptera, Cecilomyliday, Ayphorndylia, Solana avicutare, Solana ehenuprodivicdes, selina 
linearifelivn, Selanue phiysalifoliin var anidibaccunn, Selene startin. Austral, 


Introduction 


Solanun — ehveagaifalinn Cay. silverleal 
nightshade, indigenous to central and south-western 
North America and temperate South America, is é 
weed OF cultivation and disturbed hand in New South 
Wales, Victoria and South Australia (Parsons & 
Cuthbertson 1992). Tt is a major weed in South 
Africa where it has beer a target of biological cantrol 
since the MY70s (Olekers d& Zimmermann (5), 
Biglogied! control has been considered in Austrslia 
but no agents have been introduced (Wapshere 
LOK), Consequently, there is a contiiuing imterest in 
Insects ullacking S. efaeegnifaliam and other similar 
Solanum species, Whether native or introduced. The 
coeidomyiid species described here were collected in 
the conse of Tivestigalions into insects as biological 
control ugents al sedan spp. in Austrahia. 

Solana saetianiua PO Muell. tharsemindab 
nightshade, is a Shrab weeurring in central-western 
Western) Australian, southern Northern Territory, 
South Austealio, south-western Queenstund and 
hovthewestert New Soutli Wales (Purdie e/ af, 1982), 
In Queensland, ik is more commonly found te the 
south-west bul uso occurs in the nerth (Henderson 
1Y97) afer winter run COR Jellreys. Queenshind 


Departed oP Horticutiiore ViRGHiine ine Oeretouy, Ware 
Connpus The Goiversty at Adehiide PMB} tile Osiiond SA 
SHO drial Voorn ohesi MO waite delle uli 

Quemnshined Bepwetient ul Naripil Besmninees, Ali Floto 


Keeewih Stn 27 Mauaving St Sherywaed Ql i7s. 
borat: ie haudlyerremedin qld ey ait 
CSTR) Category. TO Bas 1 /HO Cunmberra ACT teti, 


VTHUTE: Fay Wa pste crite coral 


Departnent of Natural Resources. Churters Towers, 
pers, comm, 1998), The ripe fruit is reported to be 
lowic 1o stressed sheep and cute (Cunningham en al, 
1981), 

Solan aviculare G. Forst. and §. la@arifoliin 
Gerus. ex Symon uve related species known by ihe 
common pimes of kangaroo apple and) mounkun 
kangaroo apple, respectively. Both species oceir 
indinty in New South Wales aed Vieroria with Sy 
aviculare occurring wdditionully in custern 
Queensland, South and Western. Australia, Papua 
New Guinea. New Zeuliund, Lord Howe Istana, 
Norfolk Island and New Caledonia (Purdie er al. 
1982), The ripe berries of S. evveulare were 
consumed by Australian Aborigines and jhe plant has 
been culliyaled as a source of steroidal alkiloids 
(Purdie eral. L982, Symon 199d Killipongpatina es 
al. 98). 

Solum chenapeatioidey Lam. known us whitetip 
Nightshade, and 8. plo yelifelier Rusby var 
Hitidrhacoorun (Biller) Exinonds are native to Sauth 
Amencau but are now established im localised 
populations in the eastern states of Australia with.s, 
Miysatifodin vite. nitdi bacon sporidic weed or 
agriculture (Purdie ef ef. 1982). In Australia 4 
physatifodinn var vitelibaecdnin his heen 
mistakenly referred (oS, seifucdioides Sent. ox 
Mart, (alsoa South American nitive) for many yeurs, 
Sefanum surracheides is now known lo -oeceur in 
Australi only on Montague Tshind (hepsehi 1906), 

The new gall midges belong ro the genes 
Ayphondviia. Together with A. enilecervidix Kolosik 
from fruit galls on Avitoeereis spp. Ukolesik et ul 
1997) Lepsehi etal (899), the pew species for a 


32 BP KOLESIR, BR. BC. MCPADYEN & A, J, WAPSHTERES 


nalural group ol Austtalian species associaled with 
Solanacene, 


Materials and Methods 


the new gall midges were reared on four 
necasions, In June 1985, adults (males only) of A. 
vbycure were reared (by AJW) from fruit galls ons. 
chenopodioides amd 8S. pliysedifetinnr ver: 
nitidihauccatim’ collected at Mt Tomah, New South 
Wales, Adulls and pupae of A. pancidenta were 
reared (by AJW) tron fruit galls on §, [imecdri/oliin 
collected aut Boyd Tower, New South Wales, in 
Novernber LORS und ond. avieulare collected in 
Bunya Mountiins, Queensland, in January 1986, 
Jarvae. pupae dnd uclults of A. scart were reared 
(by RECM) from stem swellings on $, sfurtianin 
vollected near Charters Towers, Queensland. jo 
September (998. The laxononmy in this paper bs the 
responsibility of PK, Canad balsany mounts of the 
specimens fae microscopic examination were 
prepared decording to the jechnique oullined by 
Kolesik (1995). The type series are deposited fn the 
Australian National iiscet Collection, Canberra 
(ANIC), 


Genus Asytiondyila Loew, }SS50 


Loew. (850; Dipterologische Beitriige, 1850: 21 and 
47 (as subsenus of Cecrdonvia Meigen, 1803) 


‘Vype species. Ceoddemvia sarethaaimi Loew, TASO: 
fo. 38 (des. Kursch, 1877). 


Asphondvlia ix one of the latgest genera of 
Cecidomyiidae ovcurring worldwide with about 260 
species known (Gagne 1094). Ircoatiios species that 
hive ou verntrodistul spur on the fost larsemere, the 
ovipositor with large basal lobes, the last four fenvale 
flagelomeres progressively shortened. ihe pouu- 
coxite bearing ( ventroupical lobe und a dorsally 
sited gonostylus Hat is aboutus wide as long with 
two basally merged teeth. 


Asphondylia sturtiana Kolesik sp, Nov 
(PIGS Lt) 


Holowpe |. Gregory Highway. 52 kny south of 
Charters Towers. Qucenshind (20° 25" S. 146" [2° 
Bb), venred from stent swelling on Sodan 
sli tian B Muell, gall collected ix. 1998, RTE C, 
McFadyen, 6176 (ANIC), 


Paruiwpes, &0 82. 3 pupal skims. 2 lieve Owith 
pupil skins inside). same das, 


Male (Vigs 1-5) 

Colour: antennae brown, eyes ditrk-brawn, pulpy 
grey, thorax dark-brown, abdomen with pon 
selerotised parts red and sclerotised parts vlirk 
brown, legs grey with dark-browl setae. weninili 
dark-brown, 

Head: Antenna: scape eylindrical, only slightly 
widened distally. lengthy 1.7 8 breadth at distal end, 
1.7- 2.0% length pedicel, pedivel slightly wider than 
long; first flagellomere 1.9 - 2.1 % length scape. 
Nagellomeres evenly cylindrical, circumfila dense. 
equally distributed along segments. Eye hieets close 
together, spheroid, eye bridge & - 11 lieets long. 
Frons with 16 - 20 setae pec side, Labella erescent 
shaped. laterally with 7 - 10 setae, setulose. 
Masallary palpus 3 segmented, segments suec- 
essively and progressively longer, 

Thoratx: Wing lenth 3.0 mim (ange 2.9- 3,0. 0 = 
2). width [2 mm (1.1 1.2), Ry Wterrupted 
proximally to areulus, with strongly sclerotised 
protrusion anterior (o arculus. Ventrodistal spur on 
fst tarsormere beac at midlength wt right angle, 
Clows ofall legs similar in size and shape, as long as 
einpodia. 

Abdomen: Genititia: ventroapieal lobes oh 
Lopocoxites short: teeth on gonostylus equal in size. 
large, symmetrical in posterior views pedeagies 
tapered distally; cerci large, hemispherical, sctose. 
selulose: hypoproct with several setae in distal halt, 
setulose, 


Female (Figs 6-9) 

Colour as i) male. Frons with 19 ~ 20 setae per 
side, Circumliliy sparser than in mile, Wing length 
4.4 om (3.3 - 3.6, 1 = 6), width |4imo (13 - 1d), 
Seventh abdominal sternite 14 4 (1.0 - 7.0) length 
sixth, Genitaliny ovipositor 1.9% (1.8 - 2.0) lent 
seventh sternite; busal lobes with small, distal 
processes in dorsoventral view, densely covered wall 
Jong setulae, Other characters as i uale, 


Papo Urivs 1 11) 

Colour antennal boros, frontal horns. abdominal 
spines durk-brawn, rest of body light-brown, Length 
3,0 min (2.8 - 3.3, 9 = 3). Antennal horns serrated 
along entire Wier edae, 278 pin (266. 289) long, 
Wilh small free space between theny basally, One 
upper und three lower frontal horns. Prothoracie 
spiracle slivhtly curved at midlength, basal third 
ubout 3x width term third, terminal third setose, 
tracheu reaching midlength. Abdominal dorsal spmes 
siiiple, straight, 2. 3 pairs on last segment curved 
laterally. 


Larve(Vigs 12, 13) 
Colour: orange-red. Length 2.2 mm (2-1 2.3, n— 
3). Head capsule with no posterolatentt extensions. 


NEW GALL MIDGES FROM SOLANUM SPP. 35 


a 
Ea e 
c 


f—_ 


Vigs 1-13. Asphoudylia sturtiaqna sp, nov. 1-5 male. 6-9 female, 10, 1) pupa, 12, 13 larva, Fig. Lb. Wing. Fig. 2. Gonostylus 
WW posterior view, Fig. 3. First tarsomere of middle leg. Fig, 4+. Head in frontal view, Fig. 5, Last three (lagellomeres. Fig. 
6. End of abdomen in lateral view. Fig. 7. Basal lobes on ovipositor in dorsal view (setae omitted), Fig, 8. End of 
ovipesitor in literal view, Mig. 9% Last five flagellomeres. Fig, 10. Anterior part in ventral view, Fig. 11. Prothoracic 
spiracle, Fig. 12. Sternal spatula with adjacent papillae. Fig. 13. Last two abdominal segments in dorsal yiew, Seale bars: 
a= Timm (Pig, 1): b= 50 um (Pigs 2, 11); ¢ = 100 pm (Figs 3.8); d = 100 pn (Pig. 4); e = 100 um (Pigs 5-7, 9. 12. 13): 
f= 100 um (Pig. 10). 


34 POKOLESIR, ROE. OL MCFADYEN & ALJ, WAPSHIERE 


Spatuhe with four unterior teeth, inner pair smaller 
than ouler, shafl long and nareow, broadened both at 
midlength and base, surrounded anteriorly and 


lulerally by extensive pigmented area. Gach side of 


spaluld With wo pairs of lateral papillite, all sctose. 
On the only specimen with dndamaged terminal part, 
Three setose terminal papillae. 


Call dnd hialogy 

This gall mdge induces 4 stem swelling on Sedan 
sturfianun, 3 - 20 mm Jong and 6 - % minh wide, not 
different in colour from normal stems. Inside the 
swelling are several chambers, each oeeupicd by one 
hirva. Pupation takes plive within the gall. 


Livinelosgy 
The name is derived from the specific name of the 
host plant, 


Asphondylia paucidentata Kolesik sp, Woy. 
(FIGS 14-23) 


Holvtype: &. Bunya Mtns, Queensland (26° 53’ 5S, 
151° 37° BE), reared from fruit galls on Selaiinl 
uvicnlure G, Morster, gall collected 24.7, 1086. ALT, 
Wapshere, 6177 (ANIC). 


Paraypes: 2d 2. same dain 3 ch, 4 pupal 
skims. Boyd Tower, New South Wales (34° 02'S, 
150° 03! Ey. reared trom fruit galls on Selanne 
lineavifolium Geras. ex Symon, gall collected 
2U.nrTO8S, ALJ. Wapshere, 


Male (Pigs 14 - 19) 

Wing length 3.3 mm (range 3.2 - 3.4 n= 6), width 
LA mim (1.2 - 1.3). Genitalin in dorsoventral view: 
gonostylus 14x (0 = 2) longer (leeth ineladed in 
measurement) than wide, distal edge slightly 
concave to stniight teeth on gonostylus asymmetric. 
Spur oe first larsomere bent gradually al 45 - 60". 
Other characters as in A, striae 


Female (Pigs 20, 21) 

Wing length 3.7 mm (i = 1), width Pim, Basal 
lobes on Oviposilor Wilh fo apparent distal processes 
in dorsoventral view, Otherwise as uA. sftardtade, 


Papa (Pigs 22, 23) 

Length 4.2 nim (3.5 - 4.7.05 4). Antennal horns 
474 win (360 © 385) long, with 3 - 4 teeth at the 
midlength of inner cdee, otherwise smooth, closely 
uflached to each other along entire length. 
Prothoracic spiraclhe strongly curved at midieogt, 
basal durd aboot 4 x width terminal third. Otherwise 
ibs A NeLedned, 


Larva Unknown, 


Gall and hivlogy 

This gall midge causes a deformation of Traits on 
Solanun aviculare and §, linearifelium. similiar to 
that cuused by Ayphondylia unthocercidéy Kolesik on 
Antheceruis litarea Labill. (Solanaceae) (Kolesik ef 
al, 1997) and A. aiiventiee Endl (Lepsehi er al 
1999), and Asphondviia ebscura sp. noy. ons. 
physulifolinnm van nitidibaccatan and 8. cheno 
podioides. Pupation takes phice within the gull. 


Erymalagy 

The name pencidentata is & compound Latin 
adjective from pues and dentiy, meaning “lew” 
and “tooth”, referring to the small number of leeth on 
the pupal antennal horns. 


Asphondylia obscura Kolesik sp. nov. 
(FICS 24-29) 


Holotype. &, Mt Tomah, New South Wales (34! 34° 
§, 150° 25! FE), reared (rom (ruit galls on Selena 
physalifalium Rusby var. aitidibaecainnn, gall 
collected 4.vi, 1985, A. J. Wapshere. 6178 (ANIC) 


Paratypes: 3h dy same dats 8 cod, sime data but 
from fruit galls on Selanun chenopodtoides Lun, 


Maly (Figs 24 - 29) 

Wine leneth 3.5 on (range 3,16 3.8.1 =9), width 
}4anm (1.2 - 1.5). Genitalie in dorsoventral views 
gonostylis L.7- 1.8% (n= 3) longer tia wade, distil 
edge strongly concave. Other ebaracters os in A 
paucidematd. 


Fenule, pupe, larve unknown, 


Gall and biology 

The wall midge causes a fruit gall on Sofia 
pliysulifoliun var. vittdibaceaii and S. chene- 
podioides similar to galls of A, poietdentara and A 
wuhocercidis, Pupalion lakes phice within (he gall 


Biyinalogy 

The name means “obscure” in Latin, referring, to 
the fact hat the gall midge was found on ton-native 
plants and therefore its primary host und original 
ecorraphical distiburion are ambiguous 


Remarks 


The three new species ire morphologically close to 
each other and ta Ayphordyvita anthocercidis, a 
species that cuuses fruit galls on Auihocerets linerea 
Labi, (kolesik ef a, L997) ond Antiaeeredsy 
anixanita Endl in Western Australia (Sokuiiecae) 
(Lepsehi ev af, 1999). Together, these Tour species 
form aw natural group assocnied with plants of the 


NEW GALL MIDGES FROM SOLANUM SPP. 35 


Le aye} 
27 29 


Figs 14-23. Asphondvlia paucidentate sp. nov, 14-19 male, 20, 21 female, 22. 23 pupa. Fig. 14. Genitalia in dorsal view, 
Mig. 15, Gonostylus in posterior view, Fig, 16, First tarsomere of middle leg. Fig. 17. Gonostylus in dorsal view. Fig. LS. 
Gonostylus in posterior view. Fig. 19, First tarsomere of middle leg, Pig. 20. Basal lohes on ovipositor in dorsal view 
{setae omitted). Fig. 21. End of ovipositor in lateral view. Fig. 22, Prothoracie spiracte. Fig. 23. Anterior part in ventral 
view. Specimens in 17 & 18 reared from Selayum Jineari/olium, remaining from Selunmi aviculare 

Figs 24-29. Male of Asphoudylia obscura sp. nov. Fig, 24, Gonostylus in dorsal view. Fig, 25, Gotostylus in posterior view. 
Fig. 26. First tarsomere of middle leg. Pig. 27. Gonostylus in dorsal view, Fig. 28. Gonostylus in posterior view. Fiz. 29- 
First tursomere of middle leg. Specimens in 24-26 reared from Solanum sarrachoides, 27-29 from Seleriun 
chenopodioides. Seale bars: a = 100 tim (Figs 14, 16, 19, 21, 26. 29); b=50 pm (Figs 15. 17, 18, 22-95, 27, 28): e = 100 


um (Pig. 20); d = 100 pra (Fig. 23). 


ite TM KOLESIBK, RL EO, MePADYEN & A WAPSTITOBT: 


family Sokiumecae that is morphologically distin- 
vuishable from other known Australian Asplondy {te 
spp. by the long. eytinedrical antenmil scape, three 
lower frontal pupal horns, a sctose pupal prothoracie 
spirucke died longeshatted: farval spatula. with) Pour 
anterior teeth, ophonedylre anthocercld/s ditlers 
{rom the itee new species in the gonostylus being 
narrow in dorsoventral view, in having a marrow and 
shallow posterior incision on the basal lobes on the 
ovipositor when viewed dorsoventrally and. the 
smooth antennal horns on the pupi Asphondylrd 
siitiane can be distinguished from A, panotdentala 
hy the ventroapical spur on the first uirsomere being 
bent af aright angle, basal lobes on the ovipositor 
ending in small processes, pupal antennal horns 
hein serrate alot the entire Hiner edge and only a 
shiehtlyebent prothoraeic spiracle in the pupa as 
Opposed to the ventroapical spur being bent at 45° — 
HO" basal lobes on the ovipositar with no obvious 
processes. pupal antennal horns with a small number 
of teeth in the middle of rhe toner edge and a 
Strongly-bent prothoracie spiracte in the pupit. 
respectively, Ayphandvlia paucidenraa differs from 
Ao alsenra sp. nov, in the ratio between the length 
and the width of the wonustylis in the dorsoventral 
view heing | as opposed to 1.7 - 1.4 ford, obscura 


Salanum chenepadioides and §. physalifotin var. 
nitidibaecatin, he host plants of A, ghyoura, are not 
mitive to Austidia, Although no Agplondvlia has 
heen known lo be associated with tMese plants i 
their waive South America (Gagne 1994), iL ts 
currently not possible to determine the primary bast 
and (he avea of distribution of this gall midge due to 
the limited Knowledge of gall midge fauna assouiited 
with Solanaceae in Australia and South Amenca, 

The new species restrict reproduction and growth 
of their respective plant hosts by turoing the fruit ioe 
a seedless wall and deforming the stem Further 
investigation is needed, though, to ghurify the role of 
fruil-gulling A. patordenteta andl, ebscura in the 
pollination of their Hosts, a phenomenon assumed in 
A. authocercidly (Kolesik etal, 1997). 


Acknowledgments 


We thank P. Cranston, ANIC Canberra, for ihe loan 
ol Aaphendylia pancidentata wid Asphenidyiia 
abscura specimens: and RR. d. Gagne. Systematic 
Pitomology Laboratory, USDA, Washington D.C, 
and Bod. Lepsehi, Australian National Herbarium, 
Canberra for their comments on an early death ol the 
rruinuscripl, 


References 


CinpinotiaM. GM Mubias, W.TL. Minton. Bob, & 
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GaAGsb RL Lh) The Gall Midges of the Neotropicul 
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noes and distribution” (Queenshand  blerbaun, 

Department of Environment, Indooroopilly, 

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KIC PONG bATAbAL NL Hock. B.S. & Portrk. JR O199%%) 
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Konusik, PB CLY8S) A pew species of Roelneticotiia Vel 
(Diptert: Cecidomytidiae) on Pacaipis fasemutise i 
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Woirrestanie, Re & Siach. Hh M. (1997) 
Asphonditia  andlecercidix, ou ew speeies Of 
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Trans, Ro See S Nast W240, TS7-164, 

Lresenth Bol (1996) Soke soprgefiortes Sendy, <a new 
when Aafia in Australia, J, Adelaide Bot Gaurd U7 
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Lony, HL CENS0) “Dipterologische Beitrdge, Vierter Theil 
Die Gallinticken, Zuo der dffentlighen Proline 
der Sehdler” (Das Kénigliehe Friediiel- Wilhelm 
Gyinnasioin zu Posen, Posen). 

Ovckees, Tk ZIMMERMANN, Fh Gi. C1995) Biologicu! 
control of “Sani’s bush” (Solana eddenenfoliinn a 
leap of faith pp. 425-429 / Delfosse. HS. Aeon, Ry 
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DSIRICSTIRO, Melbourne. 

Parsons, WoT. & Curimiagisos, E.G, (1992) “Noxious 
Weeds of Austiatia’ Cnkati Press. Melbourne}, 


Porpin, ROW. Symon, Dob. & Handi b. Choe 2y Pamily 
Solanaceae pp. 1-208 Fi Creare, ALS. (Rua Plant of 
Austratia” Vol, 29 (Australian Government Publishing 
Service. Canberra, 

Symons, DE. (194) ~Kiingaroo Apples 
Archaesolinuni (Apple Print, 
Australia} 

WaArsHenE, A. (M888) Prospeers forthe biological control 
of silverdeal nightshade. Solum elarenwfulin 0 
Australia Wise JAgie. Res, 39. S297, 


Soke Sechon 
Keswick, South 


HELMINTH PARASITES OF THE PURPLE-NECKED ROCK 
WALLABY, PETROGALE LATERALIS PURPUREICOLLIS, 
FROM QUEENSLAND 


By C. BRADLEY*, I. BEVERIDGE*, N. B. CHILTON* * P. M. JOHNSONT 


Summary 


Bradley, C., Beveridge, I., Chilton, N. B. & Johnson, P. M. (2000) Helminth parasites 
of the purple-necked rock wallaby, Petrogale lateralis purpureicollis, from 
Queensland. Trans. R. Soc. S. Aust. 124(1), 37-40, 31 May, 2000. 

Examination of 12 Petrogale lateralis purpureicollis from north-western Queensland 
for helminths yielded one species of cestode and 12 species of nematodes, five of 
which represent new host records. The diversity of the helminth community present 
was comparable with that found in other species of rock wallabies. The helminth 
community was divisible into three distinct groups, species known only from the 
various subspecies of P. lateralis, species found only in rock wallabies and species 
found commonly in the sympatric macropodid Macropus robustus, presumably 
acquired by host switching in shared habit. 

Key Words: Nematoda, Cestoda, Petrogale lateralis purpureicollis, Macropodidae, 
new records. 


Transactions of the Royal Society of S. Atist, (2000), 1241), 37-40. 


HELMINTH PARASITES OF THE PURPLE-NECKED ROCK WALLABY, 
PETROGALE LATERALIS PURPUREICOLLIS, 
FROM QUEENSLAND 


by C. BRADLEY’, |. BEVERIDGE”, N. B. CHILTON” & P. M. JOHNSON‘ 


Summary 


BrAbLty, C., Bivertibcr, 1, CHivron, N, B. & Jounson, P.M. (2000). Helminth parasites of the purple-necked 
rock wallaby, Petrogale lateralis purpureicollis, from Queensland. Trans. R. Soc, §. Aust. 124(1), 37-40. 


31 May, 2000. 


Examination of 12 Petrogale lateralis purpurcicolly from north-western Queensland for helminths yielded 
one species of cestode and 12 species of nematodes, five of which represent new host records. The diversity of 
the helminth community present was comparable with that found in other species of rock wallabies. The 
helminth community was divisible into three distinct groups, species known only from the various subspecies 
of P. lateralis, species found only in rock wallabies and species found commonly in the sympatric macropodid 
Macropus robustus, presumably acquired by host switching in shared habitat. 


Key Worps: Nematoda, Cestoda, Petrogale lateralis purpureicollis, Macropodidae, new records. 


Introduction 


The helminth parasite faunas of many species of 


wallabies and kangaroos are still relatively poorly 
known (Spratt ef al. 1991; Beveridge & Spratt 1996). 
Among rock-wallabies of the genus Petrogule Gray, 
1837, only the parasites of species occurring along 
the eastern coast of Queensland, members of the P. 
penicillata (Gray, 1825) complex (the brush-tailed 
rock wallabies) (.e. P. assimilis Ramsay, 1877, P. 
godmani Thomas, 1923, P. herberti Thomas, 1926, 
P. inornata Gould, 1842, P. mareehba Eldridge & 
Close, 1992, P. penicillata, P. sharmani Eldridge & 
Close, 1992) and P. persephone Maynes, 1982 (the 
Proserpine rock wallaby) have been studied in any 
detail (Beveridge ef a/. 1989; Begg ef al. 1995), By 
contrast, records of parasites from black-footed rock 
wallabies, members of the P. /ateralis Gould, 1842 
complex, the short-eared rock wallaby, P. brachyotis 
(Gould, 1841) and the yellow-footed rock wallaby, P. 
vanthopus Gray, 1855, are based on incidental 
collections from a very small number of hosts. No 
helminth parasites have been reported from the 
monjon, P. burbidgei Kitchener & Sanson, 1978, the 
Cape York rock wallaby, P. coenensis Eldridge & 
Close, 1992 or Rothschild's rock wallaby, P. roths- 
childst Thomas, 1904 (Spratt ef a/, 1991), 

The parasites of the purple-necked rock wallaby, P. 
lateralis purpureicollis Le Souet, 1924, from north- 
western Queensland are poorly known, with current 
records based on the examination of a small number 


Department of Veterinary Science, University of Melbourne. 
Parkville Vic, 3052 

" Queensland National Parks and Wildlife Service, Northern 
Regional Centre, Pallarenda Qld 48 10. 


of specimens from Dajarra, Queensland (Beveridge 
ef al, 1989). The parasites of this subspecies of rock 
wallaby are of particular interest, since they appear 
to differ significantly from those found in members 
of the P. penicillata complex from coastal 
Queensland (Beveridge ct u/. 1989). The different 
subspecies of P. lateralis occur in disjunct 
populations across the entire western half of the 
Australian continent (Briscoe ef al. 1982; Strahan 
1995) with P. /. purpureicollis being the most eastern 
subspecies of the complex. Petrogale lateralis 
purpureicollis is separated from the most western 
populations of P. assimilis, a member of the P. 
penicillata complex, by approximately 500 km and 
might therefore be expected to act as an indicator of 
the extent of differences between parasite faunas of 
the P, penicillata and P. lateralis complexes. 

It is not possible to conduct extensive samplings of 
rock wallaby populations which are uncommon or 
which occur in a restricted geographic range simply 
to investigate their helminth parasites. Therefore, 
animals which have died from other causes often 
provide valuable information about the prevalence 
and intensity of infection with internal parasites. This 
paper presents data on parasites of P. /. purpureicollis 
obtained from animals killed by motor vehicles in 
north-western Queensland and investigates the 
similarities of its parasite fauna with that found in 
members of the P. penicillata complex. 


Materials and Methods 


Rock wallabies were collected as fresh road kills in 
the Mt Isa (20° 44" S, 139° 29" E) and Cloncurry 
(20° 42’ 8, 140° 30’ E) regions of north-western 


ord C, BRADLEY) BEVYERIBGE. SN. B. CHILEON & 1M IOLNSON 


QOueenshind aid were stored at - 20° © prior to 
examination, Subsequently, carcasses wore thiewed, 
Ihe sex und approximate age of cach specimen was 
ned uml body mensurements were recorded, At 
wutupsy, beady cavifies were examined for flariond 
nematodes, the bile duets tor cestodes and the 
oesophagus for stronvloid nematodes, “The entire 
content of the stomichs wits preserved i 10% 
formaldehyde, The small and large intestines were 
pened and also examined for helminths. Any 
cestodes observed were washed in water and 
preseryed in APA (Priteliard & Krose 1982) while 
the remaining infestinal content was preserved |" 
10% lonnaldehyde, Cestodes were sluined with 
Celestine blue, dehydrated in cummol, cleared in 
methyl salicylate and mounted ip Canada balsam, 

The stomach content wits washed will) lap water fe 
remove the formaldehyde und the number of 
netaludes counted una S on 10% subsample. All 
nematodes in the subsample were cleared in 
Juclophenol and identified to determine the Lotal 
numbers of cuch species present, The content of the 
small und large intestines was examined micro- 
seopically for helminths: if oxyuroid nematodes 
were present i the colon, their numbers were 
determined by a dilution method. All specimens coll- 
weed have been deposited in ihe South Australian 
Museum, Adelaide (SAMA), 

Tie prevalence and intensity of infection of each 
species of helminth was ealeulated (Margolis en af. 
1982). The diversity of the conmmunily was assessed 
using the reciprocal of Simpson's Index (Greig- 
Smith l96d) und the prevalence classes of helminth 
species Were Used to separule “eore', “secondary” 
and “satellite species (lanski L982) Bush & 
Hulines 1986). 


Results 


Of the 12 Pf. parpureicollis examined, wine were 
males and three were females. Although ho parasites 
were found in the body cavities, bile duets on 
vesoplagus. one species of cestode, Triplotaenio 
mirahiliy Boas. (902 was found ia the small 
joresting, LO Species of nematodes Were found in the 
slortaich, all belonging (0 the subfimily Cloacininae 
Stossich, F890 ane consisting of one species of 
Rivophoryis Moennig, 1927 and nite species of 
Cloacim Linstow., 1698, while two species ol 
nenneode, the strongyloid Macrepostrong ylides 
fuvdiste (Wood JOST) aod the oxyuroid, 
Mrcrapovwuris sp. were Found in the large intestine 
(Tabhe |). 

The perechtages of helminth species in each LOG 
previlenve class were apptoxiimately tpimoadial in 
distribulion (Fig. tj; those helminttis ti the O-bO% 
prevalence cliss were classified as “satelite” 


Frequency |G) 


CHL P20 1 20-30 a Tesi ST ote TO Tee” RL OE 


Prevalence chiss (%) 


Fig.l, Vrequeney distribution of occurrence of Helmintil 
species po different prevalence classes. 


species, those i the 21-60% chiss as “secondary” 
species und those in the 7) 100% chase as “core " 
specics, following Bush & Holmes (1986), The 
classification of cach species on the basis ot 
prevalence is shown in Table 1. ‘The diversity: oF the 
helminth conpnunity assessed by the reciprocal of 
Simpson's lndex wis 9,83, 


Discussion 


The collections of purasites trom ff 
purpureicullis veported here significuntly increase 
the number of parasite species Known from this host 
Beveridge ef ai (l989) Tound seven species al 
helminths in the aniinals they examived (thre cestode 
Triplotaenia: fimbriata Beveridge, 1976 and the 
nematodes Cloecime enabelle Johnston & Mawson 
1938, 0) fivdrifarmix Johnston & Mawson, 1938. 0 
pearyon? Mawson, (Y7 1, Co eaeits Beveridge, 1998 
(=C, sp.l al Beveridge ef al. 1989), Pharva- 
qastrenuylis feat Mawson, 1965, Kagopliuaryis 
alpla (lolinsion & Mawson, 1938) (syn, Ay caairetey 
in purl). Subsequently, Spratt eral (L991) reported 
an luitidentified species of Papileastrangyias lobaston 
& Mawson, 1939 and Beveridge (1998) reported C- 
pelragale Joliston & Mawson, 1938. CL pari 
Johnston & Mawson, a8 and ©, freqiterts Johnston 
& Mawson, 1938 lrom this host, Phe current study 
adds (be cestode Tiplitiaena adrahilis Boas, 1902 
and the nematodes Co meacrupodts Johnston & 
Mawson, 1938. CL. fungefabluta Johiston  & 
Mawson, 1938, €  celidne Beveridge, 1998 ane 
Macroapastranestiides baylisi (Wood, V931) ba the 
parasites Krowi from BL papurereolliy, Uhre 
species of Mecrupoayurisy found in the alan of one 
rock walliby also represenis a new record bub is Hot 
considered Turley because of rhe unresolved status 
ola number af undescribed species within the genes 
(Beveridge ef al. 1992, 1998). 


PARASTTES OF ROCK WALLABIES 


ay 


Taner |. Hebait parasces af Petrogale lateralis purpureicollis from north-westeru Queensland: 


Species Prevalence Intensity Group? 
(Se) Runge (mean) 
STOMACH (Nematoda) 
Kugopharyax alphe 75 GU-10710 Cc 
Gohnston & Mawson, 1838) (1400) 
Cloacina coenis Beveridge, 199s HOO S59)-13240 Cc 
(3210) 
Clodetna echidie Beveridge, 1948 ah 32-030 2 
(206) 
Claedcinag chnabetla Ma Al) 1358 C 
Johnston de Mawson, 1938 (428) 
Choacing frequens a 20-65 2 
Jolnston & Mawson, 1938 (40) 
Cloacing lonwelabiata 42 40-97 2 
Johnston & Mawson, L9F8 (60) 
Cloacing macrapodis 50) 32-1060 q 
Joliiston & Mawson, [99K (203) 
Cloacina parva 58 60-206 3 
Jolinston & Mawson, (998 (100) 
Cloacina pearson? Mawson, 197] 75 36-040) Cc 
(304) 
Cloacing petrogale 33 20413 2 
Johnston & Miawson, 1935 (138) 
SMALL INTESTINE (Cestodit) 
Tripletaenia mirabilis Bous, 1902 25 - 2 
LARGE INTESTINE (Nematoda) 
Macropostrangvloides baviisi (Wood, (YA) ss) | s 
Macrepoxyuris sp. 9 725 S 


© classification based on prevalenee «C= core species. 2= secondary species, S= satellite species 


The diversity of the helminth community in P. /, 
purpureicolliy (reciprocal of Simpson's Index = 9.85) 
is comparable with that found in other rock wallabies 
such us P assinitty (10.87), Po godniani (13.89), P. 
herberti (10.04), Po inernata (11.24) and P, 
persephone (14.30) (Beveridge ef al. 1989; Begy et 
wi }99S), ws well as those of the small wallabies such 
us the northern oailtail wallaby, Onyehoealed 
wienifera (Gould, 1840) (10,9), and the spectacled 
hive wallaby, Lagorchestey conspicillatus Gould. 
[s42 (11.8) (Beveridge ef a/. 1992), This lower than 
values found inthe réd-legged pademelon, Tiylogeale 
stumaticad (219) (see Beveridge ef a/, 1992), the 
swamp wallaby, Wallabiea bicolar (17.5) and various 
species of Muacropus found in north and central 
Queensland (14.4-26.6) (see Beveridge er ul. 1998). 
The helonth community of Pf. purpureivalles is 
thus moderately diverse in comparison with most 
macropodids and its diversity is comparable with 
that found in other small wallabies and rack 
wilhibien. 

The examination of prevalence chisses suggested 


that the helminth community present in Pf, 
purpureicolliy was broadly divisible into three 
groups, "satellite". "secondary" and "core' species. 
utilising the terminology of Bush & Holines (1986). 
Beveridge et al, (1989) used the same terminology 
for helminths of rock wallabies of the penivillata 
complex, alihough the prevalence limits of the three 
classes differed. OF the satellite species present in F 
(, purpureicollis, M. haylist is a common parasite ol 
the wallaroo, Macropus rabustus Gould, 1841(see 
Beveridge ef al, 1993) which is abundant in the area 
in which the rock wallabies were collected. The 
secondary species encountered included the cestode, 
Triplotaenia mirabilis, Kaown only from the 
MacDonnell Ranges race of Po lateralis and P| 
purpureieollis, and the nematedes CC. petrogale, 
known only froin the rock wallabies 22 leterafix and 
Po brachyetis, and C. echidne, C. Jrequens, C. 
longelabiata, C. macrapodis and C, parva, which are 
primarily parasites of ML rofisriny: (Beveridge 1998; 
Beveridge ef al, 1998). The core species included &, 
ulpha and C. ernabella, whieh are restricted to the 


40) CC, BRADLEY, b HEV ERIDGD. SB. CHILTON & TM. LOTINSON 


MacDonnell Ranges race of Po lateralis and Fob, 
purpurcicoltis, us wellas Co pearsoni and © caenis, 
Which occur in most aspects uf Perragale from which 
helminths have been examined (Beveridge 1998). 
The helt comimunity ob Pf purpurercelles can 
therefore be calegorised as cormprising UWiree distinel 
wrotps. A small number of species, 7 mirabilis. C. 
erndhella and Ro alpha is known only from the 
MacDonnell Ranges race ot Po datrulis and PL 
purpureicvllixe additional species, Cy caenty, ©. 
petrygdle ined Co pedesant, oceur tn most rock 
Walhiby species examined to dite bul i oo other 
Macropodids, while a significant Suile OF parasites 
(MM. Pelion Coechidne, ©. frequens, ©, macropoddts, 
Co lwelubin, C2 parve) has apparently been 
ucquired Trond AZ. febaseds whieh is the most 
Hbindint macropodid host with whieh the rock 
wallibies aoe broadly svmpatrie, OF the additional 
parasile spevies recorded in the literature trom Pf, 
purpurcollixs, Pleo lamlelia is common ino MM. 
rabuvma (sec Beveridge er af, 98) and CC. 
hvdiitornis and Pap llosironglis spe are comune 
parasites af the red Kangaroo, MM, rufdy (Desimuest, 
1422) (see Beveridge JY8G, 1998). a species whieh is 
also abundiuit i the Mt Isa region, The community 
of helminths present im Pood. prirpareicadtiy tho 
consists Of aw siiall mimber ot species. including all 
of those identified ws core species, which ure speeitic 
to rock wallabies wath a number of additional 
species. (denrticd within the community is 
secondary or satellite species which have been 


decived from the Most camo synpattie 
mucropodid species M4, rubusins, anil a smiather 
number af species Tram ML cufiny, 

While two speees of Claaeina (Co caenty,. 0. 
pPearsea)) appear to be commen I hwst species of 
rock wallubies examined fo dite (Beyeriee 1994, 
C. rberisi Jahnston & Mawson. 1939 whieh ts 
abundint in members ol the F. penicdlare complex 
(Beveridge ef af, 1889) was absent trons oninuls 
examined in the present study, Sinmharly, 
Rusophoryay cela otmston & Mawson, 1939) 
Which dS a Common purisite of members of the # 
penicillata Complex was replaced hy Ralphacin Poh 
propureicalliy, Consequently, the helytoth parasite 
community oF Poof pinpureteolliy  siuseests 
similuriiies with that ofthe MacDonnell Ranges rice 
al FB faterefix rather than with those of wie 
peniciliata Complex from the Guster coust of 
Queenshind. Further definition of the relationships of 
the community of helminth purasites of Pf, 
pupeivelliy vequires more cetuled studies of the 
parasite COMMUNITIES Present i ofher subspecies al 
Whe P. kiteralis cooyplex in the Northern Territory and 
in South ane Western Australia, 


Acknowledgments 
‘Thos sity avas supported finunehily by he 


Austrian Resewreh Council Thanks are due to Mr 
C. balmomstone camel Mis Re Pheri Lor assistance. 


References 


Bho VL BV reat EO ChnaiOrs NG TOTES, PEMD dk 
MC aciian, M GC. (1905) Pursites of the Proserpioe 
rouk-wwallaby, Pelagia pervert (Marsupialise 
Maeropocidied Aust, Memnieal, 18. 48-34, 

Brevik. 1 (986) A review of the penutade tribe 
Macropesttonsy lines Liehtenfels, 1980 (Strangyloiden : 
Chaseinitiie) Tony) AUSTIN fiasupials with) the 
ercelin of u new tribe, Coronustrongy lined Ball Mis. 
meal TLS atat , Paviv, be vr & TT TINS, 

YOR Taonenic revision ob Te ovis 

Cheering von Linslow (Nematodins Strongyloides) tren 
Imcropodid marsupials, Javert, Taver, 12, 272508. 
; _, Chipros, SB. & Anbeiws. Bob (10a) 
Sibling spevies within Macrpesmenevivides Davtint 
(Neruda Straneylovdeay fram  maereyadid 
Wirsupials, Af Parasital, 23. 21-33, 

. _Jonssom. PML Setanis, Lo OR. 
Spry oR. & APRATT, DOOM. ClOUS) Thelimethy parasire 
CONT es OF aingirooa cn wallithtes (Via rapes spp 
und Wallabies dYeeforh from aorth aul central 
Queenslind, Aw 7 Apel, 4, 4732-195, 

_ See awn, Ra Jorpsson, MoM, & Seteary, 1. M 
(1002) Helle parasite conmminites OF tiaropodant 
niarsuphils al lie poner Gpsypryniieelet, Aap eyes, 
Tlivingale, Onvehowaled, Lugerchestes and Pendralagus 
frown Queenstand. Wiki Way, 18 389-376, 

& See) BOM Y96s Phe herman tiie of 
Australasian miesuphabs: urns sane oval torniey 
below, defi, Puresital, 37, 136-254, 


, Chose. KR. Lo BaRkuk &. Cook 
SHARMAN Cr. B. CLO89) Eelininth parasites af raiek 
wilhibies,  Petevale spp (Mbrosapiebind learn 
Quvenslund, Aust Wildl Rey [6, 273-287, 

Ideiscon, DOA, CALMIY. do Th. Ghosh, Ry dae Mayans, &, 
M  Mnersnn eS 1 & SiakmMan. CL Be (1087) lohan, 
Dilragression and Geneue varlhon in rock wallabies pp. 
TA-87 In Groves. ROVE & Ride Wo, LBs) "Species, 
at Risks Rosewwel in Australia CAustritian Acidery of 
Scienee. Canberra), 

BUSTACOL A TOMES IOC. EToSO) Hitestingl hells al 
lesser avaup duckes patterns of ussoctatan Com a Zeal 
4, | A dh, 

Cumic-Surin, FOCM964) “OQuanlihuive Phin Beatogey” 
(Giterworth & €r,, London). 

HANSEL Le C19R2) Dyngodes of reeimal distribotins tne 
core and &alelHite species hypothesix. Gihiy aR, 21)22| 

Marcos. LG. Gacr, GW. Tloumes. $C) Koni. A.M. 
& SCHAD. GAL (M82) The lise of eeologion! terints 1 
parusilolowy  Pardsitel a8, bal 43 

PRITOUAREL ML Ho & Kresge, G. OL Wo (TONd) The 
Collection and Preservation ob Anion Parasites’ 
(University of Nebraska Press. Nebraska, 

Sreaih, DOM. Bavnkingi, bh & Waptias Hob. (aia 
cululaguie bf AUStrolusiin Haiteirernies und jaysuprals 
dnd their rewarded helminth paeisites: Roe a Aye May 
Mono See bb v05. 

Sve Aw Re PL CIVOS) "The Mavmidls af Australia 
LRecd Books. Cliatswaenh 


CAUSES OF RARITY IN ABUTILON OXYCARPUM AND 
HIBISCUS BRACHYSIPHONIUS (MALVACEAE) ON THE RIVER 
MURRAY FLOODPLAIN, SOUTH-EASTERN AUSTRALIA 


BRIEF COMMUNICATION 


Summary 


A central problem in deciding conservation priorities for plant species in arid and 
semi-arid areas is that some species appear briefly following particular environmental 
triggers and then disappear for years until the trigger recurs. Such transient specics' 
are often listed as rare but are not necessarily threatened as long as the natural cycle 
continues without interference. So far, work on such species in semi-arid Australia 
has been concentrated on post-fire pioneers'’*. However, many transient species are 
triggered by infrequent rainfall events’. Because their ecology and conservation are 
poorly understood, here we use long-term data from 1983 till 1999 to explore the 
issues involved, including causes of rarity and the amount of rainfall needed to trigger 
germination and growth. 


Transactions of the Royal Society of S. Aust. (2000), 1241), 41-44. 


BRIEF COMMUNICATION 


CAUSES OF RARITY IN ABUTILON OXYCARPUM AND HIBISCUS BRACHYSIPHONIUS 
(MALVACEAE) ON THE RIVER MURRAY FLOODPLAIN, SOUTH-EASTERN AUSTRALIA 


A central problem in deciding conservation priorities for 
plant species in arid ancl semi-arid areas is that some 
species appear briefly folowing particular environmental 
triggers and then disappear for years until the trigger recurs. 
Such transient species! are often listed as rare but are not 
necessarily threatened as long as the natural cycle continues 
without interference. So fur, work on such species in semi- 
arid Australia has been concentrated on post-fire pioneers'”, 
However, many transient species are triggered by 
infrequent raimfall events’, Because their ecology and 
conservation are poorly understood, here we use long-term 
data from 1983 till 1999 to explore the issues involved, 
including causes of rarity and the amount of rainfall needed 
lo Wigger germination and growth. 

Abuttlon oxycarpum (BF. Muell.) F Muell. ex Benth, var. 
malvaefolium Benth. (Malvaceae) (Fig. 1) is a herb which 
may reach & em and which behaves as an annual in Victoria, 
Data are also presented for ///biscus brachysiphonius F 
Muell. (Malvaceae), a subshrub to 15S em high which 
resprouts from a perennial rootstock. Both species have 
their main growth period following rain in the warmer 
months of the year, in our area principally from November 
until April. Both taxa occur in all mainland states and al 
least as far north as 20° $" 

Abutilon oaxycarpum and Hy, brachysiphonius were 
unknown in Victoria until 1983 when they were recorded in 
close proximity at Bottke Bend near Red Cliffs, Vic. 
(Browne 1986; Fig. 2). This was in the first week of March 
following 32 mm of rain. For both taxa, the Victorian stands 
are widely disjunct from those elsewhere in Australia and 
represent their southern limit’. Neither taxon is threatened 
in Australia and frequencies greater than 50% have been 
recorded for them in some central Queensland comm- 
unilies. However, in Victoria, A. oxyearpum is rated 
vulnerable and H. brachysiphonius endangered’. They will 
be referred to as Abutilon and Hibiscus throughout this 
paper. 

The known Victorian range of Abatilon and Hibiscus isin 


an area less than three km across on grey cracking clays of 


the River Murray Noodplain within 0.6 kim of the river near 


Bottle Bend (Pig. 2). Abutilon and Hibiscus occur together 


a sites | and 2 and Abutilon occurs at sites 3 and 4, Site 4 
is ua depression dominated by scattered 
Muehlenbeckia floriulenta Meisn. (Tangled Lignum); all 
other sites contain Lucalyplus largiflorens F. Muell. (Black 
Box) woodland with a sparse understorey of occasional 
chenopods and Zygophyllum spp. 


The climate is semi-arid, with a mean annual rainfall of 


290 mm at Red Cliffs, the wettest months being from May 
to October, The rainfall is highly variable, especially in 
summer when rain usually occurs as heavy downpours 
during thunderstorms. Summers are hot and winters mild, 
with frosts occurring. 

Site 2 was inspected in January 1991 (12 plants) and site 
3 in January 1984 (4 plants). No further inspections were 
made due to track closures. Other sites were inspected at 


shrubs of 


least annually in summer-autumn between 1983 and 1995 
and again between 1997 and 1999 (Table 1), Areas of the 
same habitat in nearby Kings Billabong Wildlife Reserve, 
Lambert Island and elsewhere were searched intensively for 
the two taxa without success, Voucher specimens are 
lodged at MEL. 

The plant data recorded in Table | refer to Abutifon plants 
less than one year old and to resprouts from perennial 
rootstocks of Hibiscus. Rainfall data are trom Red Cliffs, 
less than LO km away. 

The first stands of both taxa seen (at site 1) were in an 
area where trees had been felled and soil disturbed by 


Fig. |. Abutilon oxycarpum 5 em across at Bottle Bend 
Reserve, showing flower and fruit. 


- 
i 
i 
AYR racwen 
i VICTORIA. 
. 
s 
= 
g 
= 
a Red Clitfs 
bay s 
G 
a i) 
2 a 
Go 
SS 
142-15) E {ae BoE <= 
L t i L 


Fig. 2. Location of Bottle Bend Reserve, showing the only 
four sites where Abutilon oxycarpum has been seen in 
Victoria. Sites | and 2 also carry stands of Hihisous 


brachysiphonius, 


42 


Taub b. Meanidily catifall totals (ain), sienificant catifall episodes and numbers of Abutilon aud Hibiscus et site 
wunibers band +4, Bottle Bend Reserve for November te April 1982-1999, 


Raintall (mm) 


Plant humbers 


Your Nav Dee Jain Feb) Mar Ape Major raintall Auton Abiutilon Uthiscus Noles on 
episodes Site | Site Site] Site 

[982-83 4 | ( | 48 10) 32 anm, 2-7 Mar 4() 40 x 

(OR3-84 3D Ah [4 2] 28 mm, 1-3 Jan 100 47 (3 

Lo84+-85 4 0) 7 0 3 ia) None {) () {) 

Y85-86 0 BA OAS OO lI QO 6 26mm, 6 Nav; () 3 5 
24mm, 4 Dee 

Dso-s7 200 4A 1 15 [4 9 21 mm, 8 Dee () {) {) 

9s7-88 40 LD 12 38 = No falls abave 16 mm () () () 
until Apr (23 mm) 

988-89 59 27) 120 2 74 66 37 mm, 14 Mur present present (i) 

939-00 34 25 11 9 & 35 26mm. 4-6 Nov: {) i) () 
26mm, 21-24 Apr 

1990-1 | 4 9 ss 0 Q 22 0 42am. 7 Jan: present i) 200+ Flood, then 
39 mm, 24 Jan dense Fleaoharis 

19Vt-92 3h 3 | 9 6 12 26mm. 26 Nov 0 0 () Dense Fleacherts 

9V293 56 94 TF TO th 0) Falls of more than (ry 0 present Flood, then 
30 mm in Novy. dense Fleacharts 
Dee & Jan 

993-04 23 2 SS 95 () 3 29 mm, 22 Dee: (y* 0 present Dense 
49 mm. 11 Feb: Eleocharis 
27 mm, 14 Feb 

99d-95 15 1h S420 O18 19mm. 6 Jan: 0 0 present Dense 
iQmm. 16 Jan: Fleaelaris 
16mm, 30 Jan 

195-96 No data 

1996-97 No data 

1997-US 3 9 37 26 O 47 No falls above 17 mm 0 0 present 

ORO 19 3 IF BBS 2 mm, 12 Feb present present present 


"The Abatilan part of Site | Hooded briefly in November 1992. resulting ma dense Ciiloriy stand in 1993-94. 


tractor to lay u telephone cable, suggesting pioneer-type 
behavieur us in some other Malvaceae’. However. all 
subsequent stands found have been in intact vegeuiion 
lacking soi) disturbance. 

When observialions started, the area was lightly grazed 
by Wesiern Grey Kangaroos and cattle: these animals 
readily eal both Abutilon aad Hrhixcrs and marked gracing 
effects were observed. 

The falls of rain tabulated usually resulted from single 
Warm-season storms, but sometimes significant rains over 
two or three consecutive days are given (Table 1). Over the 
15 yeurs studied. Abufilon appeared in 5 years al any one 
site und Mibiveus in 0 years (Table 1), At least one episode 
of rain of more than about 26 um between Novernber and 
March seems to be needed for plants of both species to 
uppear i the study area. In seasons like 1994-95 und 1997- 
98. a serics of smaller falls than this scattered through w 
month is enough lor Vifiseus but not Abyilon lo erow 
(Table 1). Single falls of 26 mm (e.g, 1991-92) or Jess 
usiully do not produce cither species (Table |). 

Following germination in Abufilen, development is 
rapid. After the significant rainfall of 2-7 Mare 1983. 


Aburilon had flowers and fruit in 5 weeks and mature seed 
in 7.5 weeks despile little extra rain. Plants only 4+ em high 
can produce seed, Most /7ihixcus plants were partly enized 
oft, but Hower buds were produced by 2 weeks. 

Of the original eight re-sprouting Hihivers plants seen i 
(983. all were still alive in L989 and four were still alive in 
1099. viving a minimum lifespan-of 16 years. In the stuiby 
area, cattle graying was discontinued in) IYER anil 
following 32mm of rain on 7 January 199), a further 200 
resprouting Hibiscus plants were found in an area 
adjoining the original stand and which had been searched 
previously for this species. We believe that this occurred 
through a combination of the raintall and the absence of 
cattle grazing. with the grazed-otf plants being 
undetectable in the presence of cattle, This more recent, 
larger population continues to upper given appropriate 
mintall, Since 19s8. it is clear thar some kangaroo sraziny 
of both taxa occurs, especially of Abiutilog at sile 4. 

In the second half of 1990 the lowest site, site 4, was 
Hooded by high river Jevels. Tt was still Hooded) and 
dominated by a dense stand of the sedye Mleveharis vente 
R. Br. in Noyember 1990. This almost certainly caused the 


dbsence of Advan ty TO00-9F despite sudable rail 
(Table 1), 

November 1997 had the highest river levels since LY 1! 
Mooding ste 4 again und the Velen and par ot the 
(bios shindar site 1 Hence dense Zlencharis continued 
fo dominate site watihat toast 994.95, accounting Lor the 
absence ol Abarilon tn the three years 1992-95, despite 
sumuttle pai. Noosmall plans occur within the dense stan 
and rhizome systens of Bleordaris, presiimably because 
they dire offteompered., 

Atihesie } Viadtaestind, higher than site +. the flooe 
Urdined away quickly, A dense stand of the grass C b/s 
(iid Vested, aceouoting: lor the absence of Abudlon 
following goed mins inthe twer years 193-95 (Table 1), 
By dhe lime of the 1967-99 yecords of déwelan and 
Tilusouy (Vable Uh Chloris amb Alea feares bind 
disappeared frat the: sites 


We camelude that the major ablotie determmianty al 


Alito mitlire plant gcourrence in the area ure warm 
scuson VAG ind tloodiiir the former necessary. foe 
recruimmien! and the liter temporarily exeluding Aburiten 
by promoting vlenke shins of competing species. 
Assuiiiiig that ain episode of rain of more than 26 gan 
berween November dod March is needed for germination. 
mescvernolthe US years sie} expevieneed rainfall suigible 
for Vhvaleeoccdrrenee, bul la iwooal these AAutilon wis 
eseluded hy post-fMlooding competition (or for three at 
them in the oase wF the lowerlying site dp A ddscay 
Brow th owas stimiated under drice conditions thin was 
Thidlon germination. possibly heewuse of ips perenuial 
Poabstowk 

In some desert speeies, Fills oF aims low as LO rene eia 
prodiice perinanon! While 26 crim was the mipinumn 
rammlall eeorded here which produced secdling emergence 
of Abuitlon. the actual quanriry of rain deeded will be 
Meaty mTuenced by fhe huh chi content of the study 
UPL WHICH Tas an unfavourable fluence on walersupply 
i THes Climate due tothe diverse texture effect). 

Phot species of arid and semiaurid creas ace often 
PrOUped THO “Summer Hore) “water Mone or those that 
oH keriinile nd esiiblish in both seasons’. The species 
studied here ure clear examples oF summer Cora Tee pure 
SHY are, Sade Wichapode (Malvavene) behaves very 
Hiteh like (Ay seds. TH Horth-westort Vietorid. probably alt 


‘Lang, J. & Krachenbueht, DON, (1990) pp, (7% 18S 
fy Noble do 0% Joss. POL ke Jones. CK, rEdsi “The 
Mallee Lands. A Consepyition Perspective (CSIRG, 
Mahourne), 

Parsons, K.P. (]OU0) pp. 25-28 fi Noble he C2. dose Pd 
& hoists, GEOR. (Eds) The Miillee Lanes, A Conservation 
Porspeerive”™ (CSIRO, Melhounie). 

‘Parsons, Ry BOX Browne. 2 HL 1982) Biol Conserv 24. 
[RA-|O2, 

‘Wheeler. Re (1992) yp. 200-244 de Wheeler J, RR. (Ed) 
“Won OF the Kunherey Reto (Western Australian 
Herbarium, Perch, 


43 
species OF AbWion and Sedo ake of the sainmen snowing 
rype’ The nia ol Abrrlon and (ifisens studied here aun 
occur hares with less than ZOO mn mean anual rainfall, 
much lower than the stady area", Further work is eeded ta 
explain why they da net extend farther soni, lower 
Incidence of suMTeH fun, Jower lemperdture regines 
Hrocludinu Frost effects) or both may be lnvalved. A fiither 
Taportank heron in Abilene aid Hibiveus oecurrence is 
priving wilh calle removal hiving beneficial effects at 
least on the /ibiverts population, 

Abunton iso Wansient species which can be absent (ror 
ose for at least six years before reuppearing. “Chis 
behaviour almost certainty relies on hard-eouud seeds 
forming a persistent soil seed bank as it does in Adiilon 
fheaphrasd’. Parsons & Browne give exaniples of longer 
Intervals before re-appearanve. 

Many Ininsient species regenenme profusely during 
Hivourable ciciimishiices heir vbsenue aver years during 
dry seqsons does. nol ridin thar they ave threatened as lone 
us seeds aire produced offen chough la ensure that 
2ermmabon oecurs during fivausible seasons. Many such 
Specics dre ok listed as either rare of threatened! 
Nevertheless, more recent work poms out that if the 
habit is not contiouusty Fie for aceupution because: at 
Irinsicnt climatic Conditions, this can increase extinction 
tisk and se dhs lacior is bial inta the tew piles for 
chissilyia such risk), 

While Aburilen avveatpin is fot threatened Australia 
wile, there is congern about the Vieloriin suinds, 
especially becuse they ure widely disjunct fram the 
species core ren sui are al its southern limit, The most 
serious (hreatis grazing: While parts af site | and all al site 
J were fenved gains) stock te PUM4. the Tenees aire Hol 
haniaroo-prook and: kansirao vrai prevented seed set 
by most AAiilen phinks i (YOO and sae PleWinua Veurs 
Vhe het thar the pasinun number ul Ahuelen plants 
recorded wits well below 250 marine pnedividials in any ene 
SeusON Meuns That he specias qualifies ae cadwagercd ip 
Vietorm using the basie IUCN Red List rules for risk 
ussessimen) The most impornin management recon 
Mendation Tollowiie Fron aun work is provision af 
kunuwitro-proal fencing, 

We tank Mo. Bartley: tin preparing Pig) 2 and he), 
Michand hh W. Viorgan For comments cimmdhe maniseript 


‘Cunningham, Gi. ML, Mulham, W. 1. Miltlorpe, PL, 
& Lelgh, ) WE (i981). 8 Phints of Western New Siuil 
Wiles” (NSW Government Priming Office. Sydney). 
Vessop. hb BOX Toetken, Th OR. (186) “Mor of South 
Australia’ (South Austhitian Goverinent  Printin 
Division, Adelaide), 

Hensham, Rep. (199) Ranuelwnd J, 2b, 82-10% 

Gudlan, PK, Cheat D.C.& Walshe. G. C1990) oRure 
or ‘Threatened Plants in Vietoria” (Departineny al 
Conscrvalian ahd Livitoniienn Mehbourmne: 

“Clark, T&S Parsons. Re 1 (1994) Proc Ro See, Viet 
10h (29-145. 


44 


“Baskin, C. C. & Baskin, J. M. 
(Academic Press, San Diego). 
"Parsons, R. F. (1994) pp. 291-319 /n Groves, R. H. (Ed.) 
“Australian Vegetation” 2™ Edn (Cambridge University 
Press, Cambridge). 

"Noble, I. R. & Crisp, M. D. (1979/80) Israel J. Bot. 28, 
195-210. 

“Browne, J. H. (1984) Vict. Nat. 101, 228-233. 


(1998) “Seeds” 


“Browne, J. H. (1986). /bid. 103, 150-157. 


'Brandle, R. (1998) “A Biological Survey of the Stony 
Deserts, South Australia” (Department for Environment, 
Heritage and Aboriginal Affairs, Adelaide). 


“Briggs, J. D. & Leigh, J. H. (1996) “Rare or Threatened 
Australian Plants” (CSIRO, Melbourne). 


"Keith, D. A. (1998) Conserv. Biol. 12, 1076-1090. 


R. F. PARSONS, Department of Botany, La Trobe University Bundoora Vic. 3083 and J. H. BROWNE, 52A 


Fitzroy Ave Red Cliffs Vic. 3496. 


RECORDS OF MOSQUITOES (DIPTERA: CULICIDAE) FROM 
THE COOPER BASIN IN NORTH-EASTERN SOUTH AUSTRALIA 


BRIEF COMMUNICATION 


Summary 


The mosquitoes (Diptera: Culicidae) of arid and central Australia are not well 
documented. Although Lee and Woodhill', O’Gower’? and Marks** have provided 
limited distribution records for some species, most data for mosquitoes in this region 
stem from the investigations of Kay*® and Kay et al® in Charleville, south-western 
Queensland. Given the abundance of industry and agriculture in the region and the 
presence of mosquito-borne viral illness’*” a greater understanding of the vectors in 
this region is required. There is also a suggestion that mosquito-borne encephalitis 
viruses periodically invade south-eastern Australia from more northerly foci’”’. No 
published information exists for mosquitoes in north-eastern South Australia (SA). 


Trousaetiens vt the Rayet Sacre of 8 an (20001 PCL, 4527, 


BRIE COMMUNICATION 


RECORDS OF MOSQUPLORS (DIPTERA: CULICIDAE) FROM THE, 
COOPER BASIN IN NORTH-EASTERN SOUTH AUSTRALIA 


The rrosqiitoes (Dipten: Culicidae) of uri and ventral 
AUSHOTIO are not well documented, Although tee and 
Woe) OFGower and Marks live provided: lied 
Jistribulion records for some speeies: iiest date for 


Hhosqiiloes in [is rexton stem from thes investivatiome of 


Kay ound Ray ef alo in Charleville, south-western 
Queenshonl Given the abumlanee of industry ane 
wrigudivre ine ie teehon and the presence oF imonquite- 
hore viral illwess: Ob greater afderstandinng of the vectors 
IW Uns femion is required. There iscakso a sugwestion. (iat 
Mosquilo-borne vrieephaliis viruses periodically inyacle 
SOUT -GuNter Austalia fron) mere northerly foci") Nu 
PUbTshed Lae ekists Ter MOSQUITOES in nrth: 
casteyp Sout Australie (A), 

Prom December 1YO% la December 1909, we conducted 
a study oF mosquitoes at (itr sites in north-eastern SA, in 
(he lowality of the Mooniba ane! Vierawarra oil and ans 
Highly. A History of seasonally ugh mosquitt, nanibers ia 
this drew’ und heavy rainfall i Septeniher (YOK ware 
Cunilysns Cee thins stuly, the aim al whieh wits to detenntine 
fhe species ol mosquitoes in the area 

The sites wens located wathin the Cooper lvndsenpe 
reo a! Barlow!) commonly refered) to as ‘ehannel 
country” whieh learured) jnasstands, shrubhinds. alluvial 
plain and dunes. and watercourses, in a subteopiedtury 
elinate (Type Tat Walter a Lien’). 

Moayiitines were sampled at Moomba eump (2806'S, 
IAT i a Cideealpa owatterhule, an ephemeral 
Hrestiwater lc vy UW opel Freeads ancl dead woodlund 
SO ka tihe neeth 77S. OSA Fy and ait two ates 
Ont Toodphain nea Bimbirsst swamp. appresiniately SO kin 
north oF Moomnby carip and 50 kon seutt of (he Coungic 
Lakes syste, The swamp. on the Cooper Creek. i io the 
mile OF oa Foctpliie sipportije Twin shrubland 
(Michleuherkio flonidenta Meissner, nardow (Mapsilend 
deumnneadit A. Being. and sparse Aewete Awooudland, 
Kinibarko swipe Sie bowas ut the northern end al the 
shrublind/floodplin aipprowimutely seven kan north ob che 
Vivowarta oil aad sas cmp (279 8) LAaeoe fy, 
Emibarhe sWwarnp Site 2 vee approximately one kin wipe at 
(he Guin. 

Mosquitoes were sampled ising diy ide bailed iiianice 
Hieht taps! “These trips predominimtly: capnire hist 
seeking ul feniale Mosquitoes. A single (rap was set al 
lodsrone Hur hefsre semen and retrieved at lost one hour 
aller sinrise al wach site on three tsonmmenmes four) 
wecisions front Deveinber |9Os to April 1999. Twa tirther 
{raps were sel Gilgealpy watechole ail Enpbarka swear 
Sih bon U1/101/00. Pour trans were sett Moomba camp on 
T7/1 2/000 i response to heuvy rails. there dluriag 
December (9% mi Prom ba/b2/eo tp 27/12/84), 
Mosquitdes were idenrificd using He keys at Rec an ad! 
dm Russell 


"OT Ties Stree Pah Sa GdeS pers. cuaniin | paged 


Eleven species Were captured in diveiee bane (raps 
(ible 1 Prom December 1998 to April 1999, Gidwealpa 
witerhole aid Moomba camp were characterised! by yery 
low catch mtlinibers. tn contrast, lage aumbers were 
cuptiired at Embarka swap with Cilew aiaidiiavits Skuse 
most ubundint Anupheles annutipes sl Walker and 
Atopheles anichiw Bdwards were cupid in tire 
Humbers periodicully ai the Eribarka swarnp sites. The we 
traps selon TI/L1/99 yielled waly a single, male dvedes 
(Marleave) Marks species 126 al Gidgealpa waterhnte. No 
MUSQHILOGs Were GHUghE at Hivbarka swap Sate 1 or this 
wbthe, 

“Hhups sell Moomba eunp on 27/12/04 cipriced large 
numburs ob Aedes clbvoldeasis Mackerias, with saniler 
numbers of Aedes sapiens Marks, Aedes (Cehlevetatiny 
Marks species SS amd Cy canudirasiis. Raintath ae 
Moomba in iiid-Decenher [990 wis probitbly responsible 
lor The Kirge nuntbers at Aedes mesquitees, whieh have 
desiccation-resistint eves that hotel upon tiandation, ‘This 
trail) makes dedes specivs well adipted to 
environments where ranihill ty eplsadia 

Mosquitoes ditto hima were cuptured (ory bare legs 
from the Knees dower aod on feet using a mieutl opera 
aspinitor und stared fa polystyrene drinking cups covered 
wih nylon meting. All mosquitoes were caught by ihe 
authors (75 be wnt YO Ke respeenvely) silting opposite cach 
orhery Usidg a stile uspinitoy, combining the eateh tom 
bork pairs oF laps. This inethad was Used despite the tak at 
oscuilOes i the Wee CHV arhovirises, Of which bith 
uthors were wore, During the vive this was done 
whenever biting Tmasquitnes were evident Nocturnal 
collections were made on tired Qeensions for ten viinntes 
every howe [rom stinset uno losuitise at Eobarka swan 
Site 2 ahd Mooniba camp (ible 2), Several) other bitin 
Hilehes Commenced sunset and continued for (We 16 tine 
Howes. 

Mosquities were cauuht fram furisins al Bimbarke 
swap Site 20on 9/12/98 LOL Bey l/au, L62o9, 
IS/AMOO Ke VIET /OUL eli antifireseels seux the 
PredoMmiink Bilis species ar this sites AL ofeht bitin 
collvctions of this species (Table 2) peaked! five to six hours 
ater siiser (he A130 to 0230 hb), OF those species not 
suflivicotly ubundkint to present all pighh eateby dani. ve 
vivvoklensiy was caught biting daring the day aid 
Houshode the wiht wheres oda, aeegie vumel ve, 
inadines were only caught during MWe itin Aedes 
Fancrofitanes awards wis Gught baie gt saiser al this 
site, 

At Moomba cup. Ca camiatiiastns and Aa, aninedipes 
wer caunhl ul sunset oon R/P2/98, On DVYI2/M8, Ae 
wldsvuldensis was caupht biting humsdns teeuahaue 
Cho dight hours and was the predominith bung species 
When dn dE WM catelh was pertarmed (Table 2), Cuda 
initia, Aedes sapiens Motks. dedes theta 
Chiylor) and Awe aeiiipey were cauetie i seller pin bet 
HPO OE He HH This site, 


eane 


46 


TABLE 1. Mosquitoes captured by dry-ice baited miniature light traps from December 1998 to April 1999 ane 
December 1999, 


a 


December 1998-April 1999* Dec. 1999" 

Species Moomba Gidgeulpit Embarka Embarka Moomba 

Camp waterhole Swamp Swamp Camp 

causeway Site | Site 2 

EE 
Aedes alternans 0 | 0 0 6 
(Westwood ) 
Ae, eidsyoldensis 0 | 6 18 1215 
Mackerras 
Ae, sapiens () () 0 0 102 
Marks 
Ae. theobaldi 0) 0 i) 0 8 
(Taylor) 
Ae. (Och.) 0 0 () 0 40 
Marks sp. 85 
Ae. (Mac.) 0 | 0) 0 0 
Marks sp. 126 
Anapheles annulipes 8.1. 0) 2 104 289 23 
Walker 
An. amictus 0 0 24 238 0 
Edwards 
Culey annulirostris 6 | 3\4 1789 37 
Skuse 
Cy. australicus | tf] 20 5 0 
Dobrotworsky & Drummond 
Cy. quinquefasciats 24 0 0 0 0) 
Say 
total 3l 6 468 2339 1431 
no. sampling nights 3 3 4 3 1% 


‘ 1998/99 sampling dates: Moomba causeway - 8/12, 12/1, 18/3; Gidgealpa waterhole - 9/12, 12/1, 18/3; Embarka swamp 
Site | - 8/12, 12/1, 18/3, 1/4; Embarka swamp Site 2 - 8/12. 26/1, 18/3 
" Moomba camp was sampled using four traps on 27/12/99. 


TABLE 2. Culex annulirostris ad Aedes eidsvoldensis mosquitoes biting humans for a 10 minute period each hour 
fram sunset (SS) to sunrise (SR) at Embarka swamp Site 2 and Moomba camp. 


SUE nyEI tI EISISISISSISdSSISSSSSSS SSUES SSS 


Time of sampling Embarka swamp Site 2 Moomba camp 
26/1/99* 16/2/99" 27/12/99 
Culex annulirostris Aedes eidsvaldensts 

SS-1 0 0 3 

SS 19 4 22 

SS +1 19 20 9 

88 +2 22 32 | 
SS+3 8 34 7 
SS+4 32 4\ 2 

SS +5 33 58 3 

SS +6 38 63 a) 

SS +7 16 4| I 
SS+k 18 50 8 

SR 15 27 3 

total 220 370 62 


‘ §S: 2100h, SR: 0600h; * SS: 2040h, SR: 0540h; © SS: 2030h, SR: 0530h 


Athowaler bodies eneourtercd chirige Chus study wore 
Saimmied for the presence af mosquite larvae using oa 
aimulind dipprg. techpiqie. Any linvae collected were 
idenlified in the liboraiery using the key proposed by 
Rlisscdl 

Lurval collections returned lime numbers of C4 
mies und Cules caatidious Bobromworsky & 
Drummond al foeutions less thud 2000 nm Fron bP raburks 
swap Site 2 on 26/1/99. 16/2/99, 18/3/98 aim TA99, 
‘These fowmions featured ait abundanee of shallow (less 
than O.Sm deep), clear, trash water Most harvue were 
collected from the fringes of waterways, particularly where 
the bank had heen disturbed by livestock, leaving isolmed 
watler filled hoo! prints. These offen contained [rousanls of 
hwar oat both Cidey species, Despite this, adule 4 
usteions were only collated in small canibers in baited 
lipht traps (Table |) This species is apparently more 
WHHieter te Gvba ited Telit traps! 

On 27/12/90, numerous ephemersl ground pools within 
OL ke tadiis of Moomba ccuop were lowed te conn On 
aunilnesieyaad An aaiipes larvae These wilers: were 
eroatud by canal during (he precoding ten dilys. 


Irs sity dais adil Pour speeies to the List at 


mosquitoes reeorded from SA We. suppres wun Vedes 
(Oc hleratatis) Matks species 85 huve been recorded: fron 
SUvur Sites (Hratiahout the arid zone Wi nord) west NSW. 


PB Wibhan Tertiony Health Servier. Biwi NE) ptr. eum, 


(ny 


‘Lee, Do & Woodhill, AL Re 9d) eThe Anopheline 


Mosquttves of the Auanulisian Region” Unversity af 


Sydney Depuriment of Zoalugy Monogriph No 2 
(University al Sydney, Sytney), 

Pr tiower, AL KR. C1058) THe Mosquifaes of Sortt 
Westerns Astral” SPRPM Service Pobhowtan 7 
(Conunonwerlth Dept oF Health, Sydney), 

Marks, i. N14 1967) Aust. Nat. Hist, $8, 331-336- 
‘Marks. WEN. 11972) Oo Nut, 20, 1h 116. 
‘Ray, BOER (197%) Aust Raxp. Biel, Mud. Sei 37, 197-08. 
‘Kay, 1. TL, Boreham, PoP. & Panning, 1D.) 198s) J 
Mod, Batomal, 22, 520-535, 

Miles, A. Ro Howes, D. Wo C1953) Med, 1) Aust 
1953. 7-12, 

Stunley, NOE C1982) Virw! Diseases in Northers Austealio 
pp. 278-282 dy Maekenvie. 1S. Wd) Viti Diseises ih 
South Last Asio uid the Western Paeatie (Aeaderue Press. 
Sydney. 
“Merritt, A., Phillips DLA. Carney, 1. & Whelan, PL 
(100K) Comin. Das. tatell, 22) 13-104 
Wrderson, S, GX Baple, ML (1953) Meu J Aust, 2953 
WTRAY |, 

Bartow. BOA. TYSS) Brunonia d, 347-382, 

Walter, HW, & Lieth, Me (1967) “Kiinadiaeraimi — 
Welrarlas (VERB Gustav Vischer Verkig. Jem), 

finns De bod Falk R.P.(1979) Ball, Soe, Veetor Beal. 
4. bey, 


CRAIG Ro WIHALEAMS jind MICHAL J, 


Ww 


south-west aqd central Qld and from Albee Springs WP 
dades (Macleaved Marks speeies 126 Has been found 32) 
km tothe cast at Cinnamolke Qkh T5Oki ta the orth 
west al Alice Springs. NT and near Balzo, WAT! This is 
aise rhe first record of Av aiietis trom SA. previously 
revorded Tram Charlotte Waters NP 0550) kin Wwest-north- 
west)!, Cunnamulla Qld! and Charleville Qhe, indicating a 
witle runge Throuehout arid) Australia. Prowiousty, ae 
culsveldensis bas only been recorded int SA fron 
Innomincka, 60 kin cast-seutlr east of Pumewarra canip’, 
Twelve mosquite species bawe (hes far been wentified Cran 
(his pan ob SA, 

This sitaly has extended our knowledge al musqeaite 
invidence in the Cooper Basin of SA, High numbers of Ca, 
AM Tiostis. a Vector af Ross River Virus. Murray Valley 
cnvephalitis und Japanese cnvephalitia!. are of purteuthir 
interest, Together with an abundance of feral pigs ind 
dalive birds (reservoirs for tosqliloeborme  unce. 
phulitides), high biti yates by Cu. eadinasnty my 
render the north-east oF SA vulherble to mosquite burine 
vir disense, 

The logistical support of Santos Lid and the field 
asarstinee Of L bhidsen. Ce Vanstone. Bo Hall Po Caaeh, A 
Richardson and W, Phall is greallyappreeiited, Dr B. Whelin 
(Medical Entomoloey Branch. Terrlory Health Seraces, 
Darwin) vontirmicd mosquita identifigations and identified 
the Aedes (Mupleaval species [26-specimens. This work 
was undertiken by CRW as part ala PAD programme under 
The supervisiol al MEEK, CRW was in receiphota Wiiversity 
of South Australia Postpradagte Ayana 


Lee. Dd. Hicks, MM. Griffiths, M., Russell, R. CL & 
Marhs, B.S. (1984) The Culicids of the Austobesion 
Rewion Vol, Monoyraph Series. Eatomulapy Monoerapl 
No. 2° (Australian Government Publishing Service, 
Canberra. 

‘hee, D, 4h, Uieks, M, My, Debeotam, Mo Lu 
Crriffiths, VL, Marks, B.N., Bryan. AL & Russell, R. 
CL. 1089) The Culieidae Obie Avstrahisian Region Vol. 
7. Monograph Series. Entomology Mosaeruph No, 2" 
(Austrilian Government Publishing Service, Cunborras, 
“Russell, ROO, (1999) Mosquitoes and) Mosquitorborne 
Disease in Soullt Bastern Austratit (revised editions” (Dept 
of Medical Entomeulogy; Westmead Hospinal and 
Liniversity of Sydney. Sydney). 

“Debenham, Veo. & Hicks, Me Me. ClO89) “The 
Culeiie ofabe Austiihishin Beeian Voki) Moregraply 
Series, Bulonelogy Mohogeiph Neo 27 CAUsteulice 
Cavernimvent Publishing Service, Cunberray 

“Taylan FO. (1993) “Mosquito Liternicdiary Hosts ol 
Disease on Austratla ane Now Caines SPIEM Service 
Pobhiowtion +" (Commonwealth Dept of TMealtity 
Sydtey a, 

"Mackenzie, J.S., Heoom, A, K., Mall, R.A. Johansen, 
C. AY Lindsay, M.D, Phillips, D, A,, Rilehie, S, A,, 
Russell, ROCO& Smith, DOW. (1998) Chinn. Dis. Intell. 
22. 3-110), 


KORKKINN, School ol Phannuey & Medical Seteneos, 


Liniversity al South Australia Nowh Terrace Adelaide SA 5000, 


RANGE EXTENSION AND NEW HOST RECORDS OF THE 
ECTOPARASITE PSEUDOSTEGIAS SETOENSIS SHIINO 1933 
(CRUSTACEA: ISOPODA: BOPYRIDAE) 


BRIEF COMMUNICATION 


Summary 


The bopyrid ectoparasite, Pseudostegias setoensis Shiino, 1933 is here reported as 
occurring on Clibanarius taeniatus (H. Milne Edwards, 1848) and Clibanarius 
virescens (Krauss, 1843) collected from South Cooee Bay (23° 08.5’ S. 150° 45.7’ E) 
on the east coast of Central Queensland, Australia. This represents the most southern 
and eastern locality reported thus far for this parasite. Clibanarius taeniatus and C. 
virescens are new host records for this parasite. 


Trinesucions af the evil Nec ofS. Ase (2U00), $240.1, 9, 


BRIER COMMUNICATION 


RANGE E) 


TENSION AND NEW HOST RECORDS OF THE ECTOPARASITE PSEUDOSTEGIAS 


SETOENSTS SHINO, 1933 (CRUSTACEA; ISOPODA: BOPYRIDAE) 


‘The hopyrid eeioparasite Myerdosvregiay yerenats SHO. 
1949 is here reported as ovcurring on CWbararinn faemutis 
(TL Milne Relwards. W848) ail (bananas virescens CRISS. 
1H44) collected [rom Senth Coave Bay (29° 085° S, 1a0F 
45.7 Gyn the east eoust of Central Queenstand, Australia, 
This represents (he most southerth and eastern focwlity 
reported thus ir for this parasite, Clibenerias ennns: and 
© péreyeews are mew host records for this parasige 

Preudostoaians setoensiy fis heen vecorded: as being 
Palsitie On Herniih crabs of the wentls Cobadearisy, oily 
Previnusly. Po setvensis has been reported from Seto, Japan 
Where (he prirusite wits reported yn Obata aharay! (de Phin, 
TRAO), Cowon where if parusitiseu ¢ —yirielafay” Data, 1R52- 
Hong Kong. oC. binacuhiins wand CO. ransont’ Forest. 1952 
ane Phoker Thailand. jifesting € pace nasi de Mun | RRS. 

‘Two species Of CViearartie ae common in rocky intertidal 
Areas of Central Queenshind where de-pool lemperiures 
ty Caveed A> CO cluiriestintner mantis. Shallow pide- pools 
‘lthe cofleeiion sites ul both C. taenicites ale, virescens are 
Most ob coral qubble and/or sand substrate. lined with 


Tay 7 Viewer Hie bopyid parasite Pxeacdesteuieay selon sy 
\ showing stipe and sue On the dorsal i bidlominal aspeer 
ofthe bermil ech Cuban igs denen, Note head region 
(i thtmae lems (. theracie segments (sp and) yeatral 
stirlace (VEU parasite The male is characteristically func 
benenth the lomethued ehdopedites Ce) ef she pleepota 
Pieopyehi aire nor disdimer ithis sew, Sele barb iim, 


‘Stiling, S. ML 132) Mem, Collese Sen, Ken fonpen 
Univ (By 8. 240-400. 

Shino, 8. MLUI9Sky Rep Fish. Mie Chiversity 3 27-73, 
Markham. 20. (1982) Bopyrid Isopads Parasitic on 
Decupadl Crostaveuis in blog Rone and Southern Chin 


imiefoe and macroalpae unel surrounded by unstable toeks- 
Pregedusieway selena wits Tound (o> parisitise both 
fcematiiy nel ©. vireseeds colleeted from South Cawee Bay. 

The first specimen (Queenshind Musein OM WIA187) 
was collected on LS January, 98 from the abdomen of ©, 
Jeeuiatis and was later wentihed by EL Markham asain adalt 
fommale and file Po yetoesists pain The small, worm-like inte 
was observed on the ventral, abdominal surfice of (le laeer 
fermale in what Shine! describes wy a mpsuphal-like cavity 
produced By lamelhitec cndapoites of pleopacay. while the 
female was clinging with its dorsal side elosest to the dorsal 
aspect of the hermib eral’s abdomen (hig. Ch. 

Cullechons of talefemale Poosefaesis puis rom © 
viresceny were also cade on, LE October L998 (OM W2s043), 
5 November, 199% (OM W255). 1S November [99% (OM 
W25090), 8 October 999 Ceravid femiiless (OM W25}00 
and QM W510), 22 Novenbe, P2000 Lepavidl fennvale) 
(Central Queenstand Unwersity Maseum COUM 271 190 and 
personal collection), 14 December, 1999 (OM W25103 snd 
Smthisanin LISMN 200208) wad 3b ary. 2000 (anid 
Temale) (OM W2SL001On 2 November, 1998 wsoliairy mule 
PF oseTnenisis (OM W25094) was collected sigh) posterior fo 
(he carapace oF the branchiy region on a 0. virescens 
specimen, Colleetions OFF serrensiy paits WONT OC” faerie 
were asormide on 1 E Mareh. 1990 (OM W2S007). 15 March. 
100 (OM W25008), TH August, 1990 (OM: WZs000- gn 
peTaonal collechoo) | Nowerbep, 100 COM WIS 102 jane 27 
Juruiry. 2000) (Austra Museum P58a45). A single 0 
serecusiy Male/lennile pair, collected |S September, |Y98, was 
used for SEM work The inedence nile ob Po vefeersty in ay 
sample oF 387 hermit cribs collected henveen 16 Sepreniber. 
1004 and 25 Qetaben MYO Was 1.6%. On destelled leet 
erths the opaque, creaim-voloured parasile wus easily seep. 
covering Up te one fall the toral lena of the ahdnminal 
Saument oar the onih and micwsueing as much as % inn mn 
fengit, Parasilised hermit cribs did net display: obyviousty 
Aeron behaviotir Grong ode panisThacd Ones, Hoe (idl they 
show visible Signs al abdominal puneture nar physical 
demriovativn before orator the parisites were remaved. 

With the eX ceplion Ot tie apart frat the aorthecustern 
Indian Ocean (Phikery) the previous cael cumenr records ot 7! 
yefensén indicale o Western Prcihic uistbution fron Tapa bey 
cast Austrilia fOr this species. 

Sineerest (lwaks ce extended to J) Wharkiiin ol (he Arch 
Cape Mining Laborutory. Arch Cape Orenon USA: Tot his 
Wen fealion Of ie purine 


pp. 225-391 fu Morton, BOS. & Tsengo CK. (EUS) “Proe: 
Virst Tater) Mar Biol Workshop: The Martie Flora and 
Fauna of Hong Kong and Souther China, Hone Bors ~ 
(ond Rone Gaver Press, Hone Kane) 

Markham, J. ©. (1985) Zool Verh. (Leiden) 224. 5164. 


STUPTIEN ¢ DUNBAR und MIKE COATES, School ar Riologicg! and Gavironimental Seenaes, Cental 
Queenshind Uriermsity, Rocklrnplon Qld 4702, Bemail) dunbarst@ ropazcqtedtuau 


OBITUARY 
ALAN FRANCIS BIRD, B.Sc (Hons), MSc, PhD, DSc 


11.11.1928 — 13.xii.1999 


Summary 


Alan Bird died suddenly from a heart attack on December 13, 1999, leaving a large 
void for his family, friends and scientific colleagues. Alan is perhaps most widely 
known as an internationally renowned nematologist but he also was naturalist in its 
most traditional sense. An obituary emphasizing this first facet of Alan’s professional 
career has recently been published (J. Nematol. 32, 1-3, 2000); here I will focus a 
little more on Alan as a naturalist, for it was in this guise that he is known most 
widely to his friends in Australia. 


+ 
“ 


av 
“4 


ALAN FRANCIS BIRD 


PhD, DSe 


BSe (Hons), MSe, 


OBITUARY 


ALAN FRANCTS BIRD, BSe (Hons), MSc, PhD. DSe 


1d. 1928 ~ 13.411, 1999 


Alun Bird died suddenly trom a heart atack on 
December 13, 1999, leaving a liree veid for bis 
fimily, Hiebds and scientific colleagues, Alun is 
perhaps most widely known as an internationally 
renowned nematologist but he alse was naturalist in 
Hs most traditional sense. An obituary eniphasiziniy 
his first favet of Alan's professional career his 
recently been published (7, Nemeth 32, 1-3, 20004, 
Tere Pwill locus a Hille more on Alan as anaturalist, 
for it Was I THs wuise that he ts known mosh widely 
Lo His Friends in Australian. 

Alan Bird was bom on tT) Pebruiary, (924 at 
Seremban in whit were then the Federated Maley 
States. tn 1937 he travelled to Northern teland. and 
wis errolled us a bowrder in Gloucester House, 
Roniskillern which was the Prep. School for Portora 
Royal School which he subsequently attended. 
During this lime. Aki developed a passion for 
Ravby Union football hat stayed with him tor bis 
entire ile. wid indeed. one of the list conversations 
we had Was over The phone. just aller Australia wor 
the Rugby World cup Git was about 3am in Auseralis 
al the time!) 

Alan trivelled to Perth tn 1946, and enretled.as an 
agriculiiral science student atl the University of 
Wester) Australia where he seen to have spent 
inch of his time playing rigby. boil for the 
Vhiversity (and wets awarded a ich coveted “blue") 
and also far the State teank His ragby career i WA 
neHided a match aeaist the New Zeahinl All 
Blacks, T suspect a rather Irighlening: experienee Por 
uw little seruny hall Akin swiehed disciplines ane 
warned a BSe (llons) in Zoology in 1952. with a 
research project on blow Mies that heeessiiated hin 
Waitin a small Mock of sheep which graced. 
tethered to tees on the UWA Cirounds, That same 
year he ioved to Adehude jind begun to work on 
wim parasitic nenmudodes for an MSc. whieh wis 
eranied hy the University of Adelaide im 1955, He 
commimmcd: to pkiy rugby, for the University and tor 
the State of South Australia but had now developed 
Hhe oshills fo halanee sport with acudenic 
Helievement: his thesis on Phe culigle und 
eashvathing, mechanism of third stage intecnve 
stronpyle hirvie’ resufled in four publications, one in 
Nature, one ta Sedan and Iwo in Bypeninental 
Purusiiolipy. 

Newly fumied toon, Alin went ta Seathind in 
195d 10 work Gua PHD which was awarded by the 


Zuckermun (USA, 


University of Edinburgh in 1956, During (hose lwo 
yeurs. Alin also taught full time as an Assistant 
Lecturer in helminthalogy and nematode physiolowy 
a the University of Edinburgh, His ability to prodiice 
a PHD thesis (on “The nematode cuticle’) and 
publish four additional papers in what must be close 
fo record time can be athributed to two lactors Capit 
from Alan's inherent ability. Hirst the iiteleecual 
environment in the Zoology Department at 
Edinburgh in the mid-1950s was of the highest 
culibre, and Alan's supervisor, Peer Mirchell PRS. 
subsequently won a Nobel prize. Second were the 
pressures: inflicted by the poverty-level sahiry then 
paid 16 Assist Lecturers, [bya as a Phl studerst 
in Edinburgh that Alan began to use the (hen new) 
electron microscope and microscopy remained an 
essential, although not-exclusive tool for his entire 
urea. 

Alan and Joa retuimed to Australia in 1957, and 
Alan began lo work for CSIRO Division ul 
Horricullural Researeh. Fest at Merbein but tron 
IO58. in Adelaide. In 198d he moved devoss the car 
park from the Division of Horticulture lo the 
Division oF Soils. During tis second Australian 
period Alan publisher an additional (14 papers. By 
any measure, This is an impressive seientitic 
uehievemien, Even more striking is that fet thal on 
100 of his papers, he was the first or sole jultior, 
Significantly, fis co-nulhors oflen were sabbatical 
visitors drawn to Alan’s lib: Drs 8. D. Van Gundy 
(USA, 1965-66): MOA, MeClire (USA, (974-75): V, 
H. Dropkin (USA, 1976-77); B.S. Stynes (WA, 
1977-78): D. L, Riddle (USA, 1983): C. Preston 
(Wales, L986); K. A, Wright (Candi, 1986): BoM, 
IVER): Gh. Wo Yeutes, (New 
Zeulund, }993). This fist whiel spans some 30 years, 
indigttes for just how long Alia wits at the fore lord 
of Nematology, and is all the more striking for the 
diversity of stib-diseiplives it encompasses 
(incliding, homatode ecology. nemalodle-nicroahe 
internetions., chissicul plunt-Nemarolagy, ulti 
structure und hebaviouwr). Akws  titernatioial 
enllaborations extesded Further thraueh bis own 
sabbatical travels. with stints at ihe University of 
Leeds (UK), Sagi Universiny (Kyushu, laparn) asa 
Visiting Professor, and tis 4 Regents’ Leeturer (he 
first Tor a hematologist) at the Chwersily ol 
Califonia-Riverside. 

Alun’s move from Torticutiure ta Soils marked a 


wal 


vhange in emphisis uway fren: pluni-Nermatology 
und he became inereasingly interested in biceantral 
of parasific nematodes Using, other nematodes, Fung 
and hacieria. He was parlicalurly interested ir the 
surface Gout ol nematodes and dhe means and effects 
of adhesion of parhovens to i. He also beeume 
inferested in nenutedes iO the soil and those in rivers 
and freshwater lakes, art interest which he retained 
und bis eleath, 

Alan formally retired as a Chie? Research Scientist 
in 1993, althouwwh he remained ain Honorary Post 
Relirement Fellow but. in fact, Nis retirement was. i 
name only, His study at home had been converted to 
uosiitall laboriery (eomplere with nicroscopes), ut 
darkroom was bojlt ina corner of the cellar and the 
cars were obliged ty slmre space with growth 
chanibers. Soon after his retirement he wis awarded 
u grunt to study gematedes in likes and rivers in 
sourhern Australia and this work resulted in several 
papers, During Is examinition of nematodes in sail 
trom wheat fields he isolited a tardigeade, whieh led 
lou slight digression from nematodes and resulted in 
(hree papers. Afler his retirement. Alan published ten 
papers and one chapter und had (wo chapters i 
press, AL the lime of his death, Alan had received o 
yrint to identify nematodes colleeted trom every 
freshwater budy in South Anstralia anc was the Pl 
(with Mike Hodda, CSIRO Hintemology) on another 
award to campile a pricheal pretorial key ol 
nematodes. 

Throughout his career Alun received yarns 
awards, including bemg made a Pellow ob the 


Sociely OF Nematologists (M83), and bemy 
appointed oan Honorary Member of — the 
Helminthologival Soctety of Washington in 1997, In 
}991, he was awarded the Vereo Medal of the Royal 
Society of South Australia, and shortly before his 
untimely death Alan was made an Honorary Bellow 
of the sume sveiety (October 1999), Alin was named 
a Fellow of the Australian Saciety for Parasitology in 
1903, Flowever, the honor that gave Alan the mest 
salistaction and the one (hat, perhaps, best reflects 
the merit ol his collective research career, occurred 
in 1975 when he was awarded the degree of DSe 
frou the University of Edinburgh. 

In bis CY. Al fisted microscopy and: history as 
his hobbies (although he probably also should: have 
included wine-tasting). Although he certainly 
pursticd the former, he didn't aetually wetinvolved WH 
the latter, apart front beyne an avid reader oF history, 
Joining the SA Historical Society was something. thi 
he offen talked about but researeh always mamaged bo 
tke a front seat. Shortly before he died. Alan and T 
completed e chapter on plant nematodes fora general 
Nenmalology text. Hostruck me as we were writing 
that it would be possible lo produce aw fiirly 
coniprehensive article citing only papers by Alan 
Rird, Writing together was a surprisingly enjoyable 
expericnce and one that rest sadly, P won't have the 
Opportunity to do again, Aka is survived by his wite 
Jean and two children, Mary and me; we three hive 
much Lo be proud of in Alan. 


DAVID Mckk. FIRED 


Gibtivgraphy 


(95) The cuticle of permatode larvae, Nariie (Londen) 

174, 362, 

(955 Impertance uf proteases us factors jnwelwed: in the 
esha mechimisn of infective mermtode larvae 
al sheep. Arveneu PIN, HT, 

1956 Chemie! composition of the aneniitode cuthele, 
Dbservations OF the whole enticks aye Paranal ¥, 
IMS. 

1956 Chemical composted af fe cuhele al third stiyre 
Hemmatode larvae, Toi VW) 440-457 0 Walh WP. 
Rogers), 

1957 Chenival composition ef jhe remade colict: 

Obsevvatiiis an iitdividual layers sand extracts Cron 

these fivers in Asevhis dadirtomeley comieles (nah O. 

HAS. 

Phe strierure af thre eiitiele nit a veqrin Ganiylendes 

war, vals eidsttatapy AT 318 32s 4 Wath Is 

Dowtschy, 

(OSS further observations on Ue strucHin’ af tefiatode 
curtele, (ord. 4B, AD 47. 

1OS% The adulr lente eurele ind eg sae ot the eins 
Mididitanne Celli, 1887) Nemarofoyicg IE 205. 
22. 

a4 Develapment oof the fou kaor  nennirudes 
Mehudiete Javarieon (Prenky ak Mefiideayia 
digi Chitwood athe torte. Mid ae Ay 


1YS7 


1959 “The attretiveness of roars roe the pie parasiic 
nematorles Maviidesivire pavanice and MW layla Hed 
4, 329-335, 

1960 The effcerot seme single clement deferencies On ite 
erowihoal Midurdoyvne pavediicat, Hd, VOTBBS 

L960) Additional notes on the atlruchivenvuss al roone fa 
plant parasitic nemmlodes, (bid, Y, 217. 

1a) Crow oF a nematode in tomate planks prowe oe 
sodlinmertefieient water culuuresy, Were (Landon) 
189, 4/8 419 1 With Rb, Brownell), 

(O61 The ultvastraeture and Histochemistry ob nediitente- 
induced a ell Biophys. Binchem. Cyid VW 
TONTVS. 

962 The jaducement of aiint cells Wy Mehul 
avant, Nenedologicg By 1M, 

M62 Respiririon studies on Meterdeyetetudiced: salts a 
tomate roors, (bid. 8, 261-200. (With A, Millerd). 

(962 Ovienuition wh the lorie of Muloidae vite pavanecer 
rolalive to rieitss (/vid 8, 275-287, 

6a The growth ol! Weloidoiyne in Prunes persica. dul, 
9 542-500. With dF Borden & LM. Pishert 

Mork Serolowicul studies an the plint purtsite Homie. 
Mafoitawiye juvaniod. Lap Paitasital, tS. 350-3600. 

(Qh) Limbeddine andl sami sarah qemutodes for 
electron microscopy. Vatuee (Couto) 2 1 ato 
(all, 


19AS 


Was 


1965 


1906 


L906 


OG 


}967 


(yo? 


1967 


OGM 


ulate 


one 


160 


1Un4 


Rea) 


1070 


uy] 


| 


(72 


1972 


172 


172 


(rrastructie of the cabele and) d& duration i 
Meloideuvine pavanieu. Nenuttolosiod WW. 224-230 
(Wilh G. To Ropers! 

Ultrastructural and histochemical studies oF the vells 
prodtieing (he velarinis mutria ti Mftefoatifees ye. 
Hill WA 241258, With GE Rowers), 

Phe iifldehice of teniperature on Medaidagvne laple 
and MW. jae. toid TP IS)-349. OWitl HER. 
Willie ). 

Usterases inthe genus Mefoidogsnie, Wid, 12, 359-461. 
fume observations Gn exudates From Mefeidauiune 
harvae. Hie. 12. 471-482 

The ehlect of aiiinetabotites on the arewth ol Wedis 
dawvie javeniou tid 12. 637-680, (With Rd. MeGuire), 
Neing ad stirvation in daryae of Medaidoa ye 
java aml Pylenehats  yenripenetrans. 
Phi topatilogy S7. 359-571. (With S.C. Van Gundy 
SUR. Walltee). 

Chinges assacined with parasilisty ty nemetates. 1. 
Morphology aod physiology al prepurasie url 
pirate hurvite ol Mefonteyie jewaton 
Pravin 33, T6877. 

Changes associated with parasitism in neneitodes. 1. 
Histochemend! ind) mictospecttophatanmende 
wilyses Ob prepirasitig sand) parasitic. hive af 
Meloidowyae javgmeg, thid 33, (262-1269, (With 
W, Saurer), 

Chinges ysseciatcd will) panisiisa i nemauwles, 
U1 Ulirastenctire of the ego shell, Kurvall cuticle. and 
coments Of the subventrab esaphawedl vlads i 
Meloidewin java. wih some observations of 
hatching. Med. 34, 475-484, 

Chnges assogmimd wilh parasitian Te memes 
IV. Cytochenneal stiles dn the caput oP dhe 
Jorsal vsophgeut phind of Meloidea vie jeveien 
ond Gr exidanons foi We buccal stylet. fait 54, 
879-890) 

Changes wysaciuled wilh parasitisnr in nematodes, V, 
Wirastructire of dhe stylet exudation and dersat 
ssuplgcal gland contents ol emule Meloiddevne 
javaiucae Had, 38, 337-345. 

The willdence of tobacco rife spor viries and labaece 
Mosdie vir On the srowth oF Adelajdowyne farwyce 
Newuifosied 0S, 201-209, 

Skeletal oo structites amd fleguiment oF 
Acarthocephsila and Neruitodd pp. 253-288) 
Hlorkin, Me & Scheer Bo oT. (Eds) “Chermacal 
Zrolowy” Vol TE Agadenic Press, New York, (With 
I, Rind), 

Chenneal eoolowy of Acanthocephala sd Nematoda 
pp. 361-502 dai (With PER. Wallace) 

The etheeh ab nitrogen deticiiey on ihe grawllh al 
AITeidoayie peeved aL Teen population toy cls 
Nenidtolosivad 4d, (3-20. 

Speckilizet adaprtions OF penatodes lo punisilsn 
pp 35-44 /y Zuckerman. BL MEY Man, WE & Rohde, 
ROA (Rds) "Plint Parisitic Nemmaltudes'! Vol, 2 
(Acidemic Press, New York & London). 

“Vhe struchire of Nermatudes” jNeaderme Press. Sew 
York |, 

Quintitanive siidies un the prowih oF syneyvia 
inciecih i plaais by raat Kriot nenntddes, der J 
Parasitol X NSTI, 

Chrites He the UT struteie OF Hie gelatinous anatens 
of Melaidogvoe yaewuca dann delivdraten. 7 
Verdot 4 166-109 OWith AL Salt ky), 

Influenve of lemperuture on embryogenesis in 
Mehudoe vie qavaiticute Hbidt, A. 200-214. 

Cell wall breakdowo dite the fornia of 
syneytia Pidiceth Th plants by coor keer mento, 
far fd Pavsnile?. dale 


)O73 


)O73 


\v74 


1974 
jy74 


1975 


IO7s 


1975 


1976 


1976 


bath 


870 


1977 


\O77 


178 


JOTR 


179 


170 


1979 


lu) 
Ly) 


S35 
Observunions on ehroamdsomes vind nicleal) ta 
syncytia Mduiced Ay Weltarne fovaiiod. Physiol 
Plant Pail, 3, STAM, 
The influence of nematodes no photesyathesis i 
tomaterplants, (ile 3525-520 (With BOR, Loveys) 
Suppression of embryogenesis and harehine i 
Miloidugvie javaiieu by Uhernd stress. A Nevin! 
6, 95-4), 
Phint response to root-knot nemamde, Aan. Bey 
Phytopatiolauy V2. 69-X5, 
Ultrastruetaral changes an the nenvitode Adeline 
winced dssacioted with anliydrobiosts, 0 Oe trastene d 
Rey AR, 77-189, (Wilh M.S. Bullrose), 
Cellulase secretion by second stage faevue al she 
rool-knor neniutode (Meloidueyne  panvaiied), 
Meareelia 38. 1OS-169. With Wo S, Duwiaran & J, 
S. Hawker). 
The incdrporation of photusyathates by Melody 
fevenivd, A Nematol. "7 Lbl-tha, (With Bo OR, 
Loveys), 
Symbiotic felutionships betveea nematodes ani 
plants pp. 331-371 fi Symbiosis" Symposia ob the 
Sociery for beaperimentil Biolony. Ro, NNEX, 
(Cambridue University Press. Cunibridee). 
The lylenchic (Nematodes egg shelly sirucrure, 
composition and permeability, Marcyiiedowy 72. 14 
Js. (Wille MA McClure), 
The tylenelid pNematoda) egg shell: formation ob the 
via shell ly Melatdowsie pevann a [biel 72, 2-44). 
(With MAL MeClune), 
The development and ornsainizarion ef skeletal 
Structures TH nematodes ppo LOT-LAT me Croll No A. 
(hd "The Orginrasiton of Neniitodes” GAeidenie 
Press, New York). 
Cuucle formation und moulling iq the cee of 
Meloidoayne tuveniva (Nemavodiah, Parasia 74, 
149-152, 
The momhology ofa Coryuehiete ri sp. punasite 
on ginal eve prass. Plywapatiduey 87, S280. 
(With BoA, Stynes). 
The efleer of various concen(iatians ol soci 
chloride on (he host-parisite relaltornshipy ofthe rout 
knot nennirode (ATeliiddegene peed) cand 
Avyheury (Gye de my vac. bee). Menceltig 40. 167- 
175. 
Obscrvations an ervstuls found in time rbestiau cells 
ol Heemenchis coder and inthe intestinal liner 
Of Ostertaai catenins. Hine F Prariayltad 8 OQ} 
(With Bot Waller, KML Drasth & (7 Mayon 
Roo kno) nematodes i Australia, CSIRO: Division 
OF Horticultural Réseareh Teehiica) Paper Nee a, | 
ace 
Phystolowieal onclk roorphotogiit studies on 
SGETCHON OF u probinecurbohydrie complex by 
gematode, Mie Poaraitreh &, 225 232 (With Vo 
Dropkink 
The growth at Meleaidogvne peundiye ve Sonne 
Australian. fative plants. Sea i ds So, Wilh dP 
Milli). 
Acmethod al disinguishing between livin and clei 
Wenndtodes by enzymiitionl indiictd Mies cies wf 
Nennital. VL WS. TOS, 
Morpholowy unl ulirastraelene pp. Sosa fy 
hamherti, Foo& Tuwlorn ©, TE, (Gus) ‘Koon-knq 
nematodes (Wtefordogyne species)" (Acwderuie ress, 
London) 
Histopatholoes did plysiolowy of svaeyin pp. 19S- 
WT dial, 
Clivesiuenuce Gf the Gail reper al the secund stage 
PrepPUrasilic Harv ot He cue Kiet megiatode. fn 
Paras YAS E AT 


+4 
HARM) 


1YK0) 


INs0) 


1O80 


91 


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he 


10M? 


(O83 


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[ORS 


[Ma 


(ed 


Med 


los 


Wad 


{URS 


1Ona 


(he influence of ight quality Qi the growth: aed 
fecundity of Melanouvie infesting tormiito, Nemaral. 
miedif 8, 59-06, (Wilh M, Ro Sauer RN, Chapin 
& 1, RB. Loveysi 

The denarde eutlele and ths surkiee pp, 2EA-236 far 
Zucker BoM.) Edo “Nematodes as Biologieul 
Maiels) Vol 2. (Aeadenic Press, New York. 

The involyement of cytokinins Ta a) fasteparasire 
rehunionship) belween the tomato Lycopersicon 
everdeuman arta nematode (Mefoiidee vey faveaniod), 
Pivayitedauy 80, 497-505. (With B. R. Loveys), 

A camparisab of nematode dnd bieteria-colanized 
walls jndyeed by Angatie eusostes bedi bogey 
Mhvropaltietogy 70, 104) 1098, (With BL AL Stynes 
& WW. 'Thomson, 

‘The fife evehoot Angin aerosis: enbryosenesis, 
din. Pavasinel, WW 23-34. (With BLAS Stynes). 
The tife cycle of dagiind agrestis: pest embryonic 
growill of the second shige larva. Mid) Wa, 244-250), 
Wilh BLA. Siynes), 

Kilcem oF merhods ut killing. Tis tigcang prrounting on 
mensaremenia of Aid gees Nearer ites i 
26, 467-71 With BOA, Stynes) 

\eenie earostia, the veeror ol cna) rye pres 
toxteily ft Australia, ai? 26, 475-490. (With BL A 
Smyiest 

The fife eyele oF Aneuine ugrouis: develypment in 
hose phan died Pees Weds dae Wate ba, 
Atvnes 
The Ai Coryathrienii dsscciatlon py, Ws- 
$28 fy (Auekernitn, By M, & Rohde, BR, A (Pus) 
“Plant Panisine Senmirmes," Vola, (Acidemie Press, 
New York) 

evelopment al walls iidieed in adi eigidin hy 
Anite cpbostty. Plwaputalety 72. 340-Fdé. 
(Wyll) By A, Styfies) 

Detection vid ulfredruetial daseription ata faeval 
inuult an rhe eee oF Garin phy lhipre do Nematal, 14, 
JOR IOO, OWE CL Onn, 

Growth and MoulliAg in erates: Chamges in he 
Winensions and morphology af Rervlenehiulis 
reniforndy Gong shud to Huish ob mowtiing, ttf 
wreniiols 13, 201-200, 

Changes in he siinensiens oF the oesuphigent glands 
iy veoOL-RNOL menuiitiles dure dike onset al 
pitasitisin, Jol. ba, Fda, 

Hie patie at the ifestival vesiohe vie nenaaiades et 
jhe family Steinernemabdae. fhid) Ta, 584-006. 
(Wath Rh Akhurst) 

Development of annual ryeerass losieity. tase 7 
Awrie. Kes, $4, 654-660, (Wilh BL A. Stvrtes) 
Nenuanilit py, 207-283 Je Bereier-thuin 4, 
Matolisy, AG) & Richards, KS. (Eds) "Totoay of 
the Integument! Vol LoSprinmer Verku. Berlin 
Crowih aid Moulting in nematodes: rmianling sel 
development af the hitehed larva at Beeenediitis 
rented. Parasiteliaay 89, TOF TIM, 

Efeet of alachmenat of Grr yirehecder sian ctu) at 
movement oly gerostd’s uve. dat 0 Presto Da 
SUAS TL. (With Dd. Le Biddle), 

‘The ilicrce oF roar knoe nematides (| Mefloidnweyne 
Javiiog) On vtelt and rm weliyily @l suerose 
synthase Hi TVePse Ti foors col) ceephants 
(Solan nudongena). Pliystel, Plan. Path 38. V9- 
ANZ (Wil WP liusseny), 

Responses of igi apbesrie ta delenent and 
anesthetic frealinenth PE Nemetol. 17, bOA-108. L With 
DL, Rile), 

Vie ature of dhe adhesion at Cae yueedwerden ian 
ratte roothe euriehe of the tfeetive baw al 
Anu avant, died Beaesemn 1S, MW SOS, 


JOKS 


(ORS 


(Ont 


[986 


[YhG 


OKT 


VON7 


|OR7 


1O8S 


IONS 


LORS 


LURK 


JRO 


1989 


JOK9 


LORO 


189 


LO) 


ron 


{UO] 


1ou| 


[ayo 


Corynetos ios and pemutodes, Myrinitelayy OL, (bu 
176, (With M. Vo dawo & PAL Cochin, 
Responses af the plant parisitie  fenmitaces 
Rotvien hulis venifarinis, ARO agtostiy und 
Meloideeyne javanica ly chernent aithnetints. (bid 
9 185-195, (With DL, Riddhe, 

Observyy tions on the tise OF (sect Palast ie Hemarides 
dus drei at bielogical vontrol af riot-knet neni 
lodes. fiat J Pavusiol 16, 30-516. (With. Bird 
Alehiment ol Pisverime penetrans apetes ta the 
cuticles of roet-Knot nenitodes, Reve ve 
Nenratoluvle 9, 251260, (WalLG, Ry Stipling & AB. 
Cukurs. 

The tiflienee af the aetinamyeete. Posteurie 
peneienis. OW Te Hosteprisite nekitlonship ah the 
phint-pirasitic nematode, Mefoidowyne jeavailicn 
Pnasinihesy 93, AT(-380, 

Moning of paniaitic neemodes, (inh Parasitol, V7, 
233-239, 

Physiologicn) and iorphalogicul changes dssoe ited 
willrrecowery Hor aiubrowis 1 the dauer Rilvikot thc 
neratode Anwaine genesis, Papeastiofowy 9S 10% 
133, (With C) M, Preston), 

Adhesion of eonidie of the fungus Oifapleapena 
dlopecwer tothe cuticle al the nent Aged 
Hinata, The veetor in anual ryegriiss toxicity, (ie 7 
Parasital, V7, 1231247. With A Co MeKays. 

The influence of Pastearia penetrey in Feld soils on 
the reproduetion of root-knot nematodes, Kevite de 
Néematatowe W758 (Wil PG. Brisbane), 
Atechitiqie tor staining the endaspares ait Peston 
penelrans. lhid. TR, 304.305 

Acrole for the “excretory” svstcni Ti seecrmentear 
neomtodes, A, Newetel 20, 499496, (WiLL Boni 
& A, Brcic), 

Cuticle printing of nemudodes. dard Harastral, 1, 
MOUNT I. 

Observations on lpliedevichoitas dividras Massey. 
1974 feeding on Tonal pathogens of whear th 
Australia, Revue de Nemetolosie 12, 27-44, (Watt 
Bird, R. Forluner & Re Moen), 

Nematoda and Nematormorpha pp. 219950 fy Adivodt, 
KG. ke Adiyods, Rina Gy. (hds) “Reproditetive Biglogy 
of Tnvertebrates!, Vol, 4, (Oxford and TBE Publishing 
Co. New Delhi). With R, L Sommerville 

Factors alfeeting the adhesion of miero-orsinisits 
the surkices oF planepwasine memaruides. Mate 
Vitelaus YB, (55-164 (With 1. Bomig & A. Baer), 
Studies on the surface cont (alycaeiyx) of the date 
larva al Aven aerosis. tat, 2 Peanasitel. WW, 245- 
240. (With BoM, Zuckerman). 

Composition of high molecular wenlt excretions 
secretions (romp iufective larvae af Medlin 


revatio: 2 Neri, 20, 477482, (Willi A, Bacic de 


¥ Pelion 

Studies On the propeities wl Tig spetes uF sen 
nopulutions of Paweneia pene Lf Livers Carhet 
S560 178. (Wil BG. Beisbane, 8G) Met line & 
ROW. Kinnber, 

Virwl striiie oF vlycopferen) seerehad by mfeehive 
Ihivd ate Eievae OF Meer hdis alten tity pela fey 
exsheathment, fat, 2, Pavestiod 2h 614-623, 
Observations on erystulluld bodies in the pavuite- 
colon of Avmabeiies feptepiltatios. ds Nera, 
23, 30-47. (Wits. G. MeClure & We to Nichols, 
"The Struciem pf Nenuertes” Jed Petey (Aparterme 
Press. San Diego). (With J, Bir). 

Mie nematode Futon al rhe Mirra Riven estiany: ile 
effects at fhe barrages across is diouth, yer 
hielowia 234, 47-101 (Wath WoL, Sieholis Tow, 
ficech & A, ©, Stew), 


1993 


1993 


1993 


1993 


1904 


1994 


[995 


LyO5 


19906 


Association of bactertophage particles with toxin 
production by Clavibucter faxicus, the causal agent 
of unnual ryegrass toxicity. Phytoputhology 83, 676- 
OX!. (With K. M. Ophel & A. Kerr). 

Feeding of the nematode Acroheloides nants on 
bacteria... Nematol, 25, 493-499, (With M, H. Ryder). 
Effect of  Acrabeloides  nanus  (Nematodar 
Cephalobidac) upon the survival of Pseudomonas 
corrigata (Bubaeteria) in pasteurized soil from 
Kapunda, South Australia. Trany. Ro Soe. S. Aust. 
117, 179-182. (With M, H. Ryder). 

Morphology, oviposition and embryogenesis. in an 
Austrahan population of Acroheloidey nanus. J 
Nematol. 25, 607-615. (With P, De Ley & J, Bird), 
Some observations on the influence of agricultural 
practices on the nematode faunae of some South 
Australian soils. Maidan. appl. Neniatel, V7, 133- 
145. (With G. W. Yeates). 

Studies on Aprutides guidetiii (Nematoda: Seinuridae ) 
isolinted From soil at Northfield, South Australia, Treas, 
Ro Soc. S. Aust, 118, 261-266. (With G. W. Yeates). 
Chitin in’ Meloidogyne javanica. Fundam. appl, 
Nematal, 18, 235-239, (With P. G. Self), 

Studies on Eiohrilus heptapapitllatus (Nematoda: 
‘Tobrifidac) the predominant nematode inhabiting the 
bottoms of Lakes Albert and Alexandrina, South 
Australia. Trans. R. Soe. S. Aust. Lb9, 133-141, 
Studies on the sojl-inhubiting tardigrade, Macro- 
hiotis ef. pseudohufelandi, from South Australia, 
Thid 120, 147-154. 


1997 


1997 


1998 


1999 


1999 


In Press 


In Press 


SS 


Composition of the stylets of the tardigrade, 
Macrohiotus ef, pseudohiufelandi, (bj. 121, 43-50, 
(With S. G, McClure). 

Studies of the eggs of Macrohiatuy cl. psende- 
hufelandi (Tardigrada) from wheat fields in South 
Australia. ffi. V2, 51-57. (With S. G. Me 
Clure). 

Introduction to functional organization pp. 1-24 /n 
Perry, R. N. & and Wright D. J. (Eds) “The 
Physiology and Biochemistry of Free-living and 
Plant-parasitic Nematodes” (CABL Publishing. 
Wallingford). (With J. Bird). 

A comparison of some microinvertebrate assem- 
hlages in southern Australia. Trany, Ro Soe. 8. Aust 
123, 69-75, 

Observations of some nematodes from Kangaroo 
Island, South Australia, including the description of a 
new species, Hemicyeliophora flyvialis (Vylenchida: 
Hemicycliophoridae). from Rocky River, [hid, 123. 
121-131, 

Plant parasitic nematodes /7 Kennedy, M. W. & 
Harnett. W. (Eds) “Parasitic Nematodes. 
Molecular Biology, Biochemistry and tmmun- 
ology” (CABL Publishing, Wallmgford, UK). 
(With D, McK, Bird), 

Surface adhesion to nematodes and its 
consequences /n Chen, ZX, Chen, S, Y, &. 
Dickson. D. W. (Eds) “Nematology, Advanees 
and Perspectives” (Tsinghua University Press. 
China). 


OBITUARY 
PATRICIA MARIETJE THOMAS, BSc, MSc, AO 


13.iv.1915 — 16.xii.1999 


Summary 


Patricia M. Thomas was born in Melbourne on 13 April 1915, the elder daughter of 
Sir Douglas and Lady Mawson. Her father was Professor of Geology at the University 
of Adelaide. Her secondary education was undertaken at Woodlands Church of 
England Girls Grammar School, Adelaide, where she completed her Leaving 
Certificate in 1932 in English, French, Geography, Geology, Mathematics and 
Biology and matriculated in 1933. She graduated from the University of Adelaide 
with the degree Bachelor of Science in 1936 and completed a Masters degree in 
Zoology, under the supervision of Professor T. H. Johnston, in 1938. The subject of 
her research was “Studies in Australian Nematoda”. She subsequently held various 
teaching and research positions at the University of Adelaide until 1946. 


PATRICIA MARIETJE THOMAS 
BSe, MSc, AO 


OBITUARY 


PATRICIA MARIETJE THOMAS, BSc, MSc. AQ 


TRavd9ls - beni 1999 


Pairicia Me Thomas was bor in Melbourne on 
13: Apnl (91S, the elder daughter of Sir Douglits ind 
Lady Mawson, Her father was Professor of Geology 
at the University of Adehude, Her secondary 
cilication was dindertiken at Woudhinds Churgh of 
England Girls Grammar School, Adelaide, where she 
completed her Leaving Certifieate i $932 in 
English, Freneh, Geography, Geology. Mathemalies 
and Biology and miatrieulided in 1933, She graduated 
fron) ihe University of Adelaide with the depree 
Bachelor of Science tn M36 and cemplerd a 
Masters dewree ain Zoology, under the stpervision of 
Professor T. TH, Johnston, in 193%, The subject of her 
research was "Studies in Austealian Nematoda’. She 
subsequently held various teaching and rescarel 
positions wethe University oF Adchude anant 96, In 
(947, she (armed Wor M. Thonus, then a leeturer in 
marine blolowy atthe University and fram T8St! te 
JO8O eeupred a variely of parltime posivions within 
the Deparment of Adology, including Junie 
Ruscarch Fellow, Research Assistant and Technical 
Officer, which she competently combined with the 
obligations of raising a family oF (hree sons while 
vise eoyaging in research, primarily on parasite 
nematodes. In 1954 she undertook a period of study 
overseds, vibiting the lihoratory of Dr Sehuunnuns 
Stekhoven, an ennient student of tree-hying 
nenutiWes, al Deventer in Holhind and pubtished 
WIT Tima senes af papers on free-living: nematodes 
ay well us oo mermithids. She uso visited the 
Iystiiite Of Pargsitilogy at MeGill University, 
Montreal, Canadi where she worked under the weets 
of ihe director, Prof TW. M. Cameron, asa Nuffield 
bellow. a posiioo funded by the Royal Saeiety. 

Pats subsequent serene Gureer wiry spent io the 
Zooloey Deparment of the University of Adelivide 
onlil her revrement in Devember 1980. bollowiny 
her retirement from the University, Pat moved to the 
postion Ob Honorary Curator of Hehniallis at the 
South Australi Museum, at position which she field 
until Weheatih forced her lo relinquish iin 1995 at 
the age of BA. 

When Pal began at the musean the calleeron of 
parniside helminths held there wis timited ane 
consmsled primanly al types deposited by TH, 
Jahoston dine bis students. Th cotlaboration with 
Madeline Anwel aid Stud kariornds, Pateniareed! the 
Heh) paricite colleetion of the museum lo the 
pom Where i becume pre-erminent in Austdia, with 


the largest number of accessions and the largesi 
number of primary types: The parasite colleetion ts 
supporicd by an extensive collection oF reprints and 
offer pelted Hiterature available yowhere else i 
Australi aswell as by host-piarasile elalogues anc 
lilerature summaries for cach parasite genus, These 
also are available in ne other Austeahan tistitution 
nor ure they available. dnd probably never will be, 
electronically, During the 1970s, Pat sought, with the 
support ofthe Australian Society for Parasitology, to 
have the vollecrion housed inthe SA Museum named 
the Australian Helminthalogieal Collection aml for it 
a Tom the basis of a pational eallection, Her 
endeavours were successful to a turge degree and 
resulied in fuoding from the Australiin Biological 
Resources Study fo catalogue lally the parasite 
collechon records as well as funding from the 
Australian Society lor Parasitology for cernputers 
and the development of an electronic database, 
Pirally. in LD9Od the muscu appomted ts fest fall 
time curator of helminths, ullhouwh thas posaion is 
currently vaciiol, The contribution of Pat Thoms and 
her colleayues to the diseypline Of parasitology an 
Australia has therefore heen quite uniquc. 

Pac’s contributions to Science were both signilicant 
and varied. Ao large coniponent of her formal 
employment at (he Laiversity Mvelyed ruining huge 
practical classes lor first year students of zoology 
who probubly viewed her with eorsiderable fear us 
something oF a chsciplinaritn. Those who propressed 
in their Studies a! vooloey undoubtedly wot to know 
her a (ite more intimately and discovered a curing 
person. with on enquiring mind. who was not only 
Wiberesten i there czowlocieal studies but also tn the 
progress af Weir personal development. They alse 
Sulmed cn aipprecition ot the depth of hes 
COTTE AL TG Serenihie research, 

The manilestitions. of her Commitment to the 
wubvanec ent of science were diverse, She weled is 
editor, Joflowing the death of Prot. T. Thirvey 
Johnston, for miany of the series of scientific reports 
(on Zoatogy and Botany) resulting from her father's 
expedifions co Adtarcticd, Phese were published its 
the Buitish, AusStralnun iad New Zealand Antaretic 
Research Expedition reports and covered botameal 
and zoologica) explomarions dy Aptaretica, betweeti 
(929 and (YS), The first of Wiese wats published in 
1937; Ihe most recent issue of [he series (Porileria) 
wis published in 1076, Volurme 3. on the traumnniats, 


aN 


is expected to be published in the near Tube and is 
likely io be the fastof the series. 

Pat Thomis's most significint scientific lepuey 
undoubtedly lies in her publications, the majority: ol 
which were published winder her maiden name (PM, 
Mawson), Nurnberg, i excess oF 100. her pupers 
mide i Siiifieant contribution to the study ol 
parasitic fenntodes i Australia, a impact whieh 
was highly respected both withim Australia and 
overseas, The papers published from her Masters 
(hesis during 1938, 1939 and 1940) probubly 
represent (he most significant single contribution to 
our Knowledge of the nematode parasites of 
marsupials made to dale. They indiewted that an 
erobmously diverse Latina Of nematode pumisites. was 
present in Australian marsupials and thereby laid the 
foundations for subsequent studies, which are still in 
progress, Later papers covered the nemiatode 
parasites of frogs. fishes, lizards. snakes. eutherianns 
und tarsupials and, particularly, birds. Numerous 
papers of (his Hitter topic culmingied in L986 in the 
publication (in conjunction with Madeline Angel ane 
Stan Filmonds) of a definitive cheeklist of the 
helminths af Austin birds. a monograph whieh 
remains at essential resource for Australian 
pdrastologists, The parasitic Hernntode fauna of 
Austriliin vertebrates is unique and extraortinartly 
diverse anc) Pat's papers represented the first teal 
utlempt Lo document it comprehensively. While the 
enterprise of documentation continues. tind will 
undoubtedly eontinue tor some considerable line. 
her collected) papers represen) probably the most 
sannilicant advance ode va this are tee cme. An 
outstanding feature of her work was that she studied 


dl published oo virtually allad the kaawe orders of 


purasite nemilodes,. Furthermore, she crossed the 
gull which divides many pentmatelogists ie. the 


division between parasitic and free-living forms, 
Few nemuatologists hive published sinificant series 
Ol papers in both areas of research but Par was one al 
this group. Todi. ian ert of iitense speciilisation, 
such u broad and comprehensive approach is almost 
Hcoutprehensible and, therefore. probably hus net 
achieved the recognition which i deserves, 

‘at's achievements have bee recounized int 
number of ways. She wis awarded the Vereo Medial 
of the Royal Sociely of Soul Ausiralia in (974 for 
her setentific publications and heeame the frst Cand 
ui! 1999 the only) Worn to be made a Fellow ol! 
the Australian Society for Parasitology, Ty (994, Pat 
was awarded an Order of Australia lor her 
ecortibutions ty seenee in Australi, Apu fram her 
UeHVITIGs ds pansiolagist, Pat was ulso a member 
of the Tandbooks Committee (Tandbook of the 
Flora and Fauna of South Australia) for more tha 
Iwenty years and uw very netive member of the 
National Patks Commission. She was also a Counell 
Member of the Royal Society of South Australia 
from 1977-1992 being Membership Secretary fren 
IMH2-1992, 

Her passing will be noted with regret pot only by 
colleawues in Australia butalsa overseas, piuticulirly 
in Prange, Britain, Poland, the United States sind 
Canada, countries fa whieh she estiblished long 
Siunding scientific Collaborations and friendships. Aer 
contribubions (oO parasitology were highly regarded, 
but she will also be remembered for her extremely 
senerous hospitality, her culioary skills. her forthright 
manner, her sense of humour and, in particulin, for her 
gencrosity in sharing seientifie material, 

at wats predeceidsed by her husband Mor and is 
survived by her sons Gareth, Alun and Ernlyn 


LAN BEVERIDOT 


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Mire lrecliving nenitodes tram South Australia, 
Trans, KSow S Aust. 80, OR TOR, 

Some nemitodes from fish trout Heron tsar. 
Oucenstin, (id SO, (77-178 

Pihiriid nemareides from Canali birds. Cone ob 
fot AS, TG2Y, 

Some nefmatode patasines Crom Avastin hosts. 
Tins RN 8 Aust, 82, 15)-162, 

Two new species of (richostronsyle nenmatares: [rom 
(he Australitn pouched touse. Plascoeale flavipes 
(Warerhouse), Purasiivfauy SQ, 425-124, 
Nematodes belonging to the lrichostronyy lide, 
Subuluridae. Rhabdiasidae and Trichuridae from 
bandhicoors Auv_f. Aent &, 261s, 
Trichosironeyles from paderts in Queenslandk with 
comments an the genus Lenedsidate (Nemtacdky: 
Heligtmosonmaridae | dive & TUB 26, 

A ie spevics and some records if tie peries 
Claaeind UNemanwla: Strineyloidea) tron Western 
Aurelian. Dra ®oSee So dust BE SIEN3, 

A nole on fhe aecurenee oF oesophageal teeth in 
specivs ef the genus Cluaeiia (Nematodiic 
Strommy hotles). (ai 8585-89, 

Two tnichostronayle nematades fron ae mrmoser, 
{hid S®, VST-TSY 

Sane Nenniloda (Strongylingy woul Ox yuri) fron 
ATTOOS OMe npr spp.) Com custern Austrutia, 
Phirusiiilivey S34. T4782, 

Noles oo some species of Nenmairyht (ron kane ies 
ad wallitbies. inelidity a new wenus andl three Hew 
species, dhal, BS, 145-192. 

Three species wih the Qenus Geopentia Chita 
(Nematoc), Spicweani fron Australian bitdy. fail, 
56, 7IS-718, 

Miscelhitiea temmtoceilowion, WI, Ee tiaeaneentes 
HUST HEN sp. TY Eron Semis iy creisale aiteleon i 
South Australian Zoe, Hise, 4, VTS. With WG 
Tu liny. 

Praiivaneme (pos Sen. av. ap. Hoy. (Nematoda: 
Scurmloidea: Inalisonematinae. sablank nove tren 
in Australian bird. Parasalapy S87 | 73. 

Two species ar Memories eSpiranda: Spuviridact 
from Sustration dhsyurids. had S874 78 
Nematodes from Australian waders Are SR. 277-405, 


a 
1908 
LO6K 


L068 


1969 


1970) 


1971 
197] 


1972 


1972 


1472 


(O74 


1974 


1975 


197 


1977 


1977 


177 


1O77 


17S 


Anew genus, Maiichdbandia (Nematoda : Spirurinae) 
from coracilorm birds in Australia, (bid. 58. 741-744, 


Hahronematinae (Nematoda: Spiruridae) trom 
Australian birds. (id $8, 745-767, 
Helminths from some lizards, mainly from South 


Australia, Travis. Ro Soc, S. Aust, 92, 50-72. (With 
L.. M. Angel). 

Some nematodes from Australian gulls and terns, 
J bish Res. Ba. Can, 26, | 103-1111. 
Skrjabinoptera galdmanue a. sp. (Nematoda: 
Physalopteridac) from an Australian agamid Heard, 
Trans. R, Soe. 8 Aust 94, 223-225. 

Two new species of Rictularia (Nematoda) from 
Australian rodents. fhe. 95, 61-64. 

Pearson Psland Expedition 1969-8, Helminths. fied. 
95. 169-1823, 

The nematode genus Miavvechonia (Osyurati: 
Cosmocercidac) i Australian reptiles andl [rags. 
hid. 9, LOL LOR. 

Three new species of the geaus Cloueli Linstow 
(Nemarada: Strongy lity) from macropod marsupials, 
Thid. 96, 109-113. 

The genus Acuaria Bremser (Nematoda: Spiruridit) 
in Australia, (hid, 9, 139-147, 

Amidastumatinue (Nematoda: ‘Trichostrongyloidea) 
fom Australian mursupitls and monotremes. Hid, 
97, IN7-279, 

The genus Pororestronayias Johoston and Mawson 
(Nematoda: Trichonemiutidae) from macropod 
marsupials, Mid 98, 135-138, 

‘hwo new species of the genus Clouesied (Nematoda: 
Strongy lida) from the Gummi Macrapis engeii, 
hid, 99, 39-42, 

Woodwardostrongyins  obenlorfi new species 
(Netnatodas Amidostomatidae) fran) kangaroos, (id, 
100, 12]-124, 

The genus Cyelosironeyiias Johnston ond Miuwson 
(Nemittoda: Trichonematidue). (bid. LOL, 1920, 
Cloacine cornia (Davey and Wood band CO) cubation 
sp. noy. CSemutoda: Cloacininae) from miacropods 
from Papua. Tavitiie de Bivlogia, Publicaciones 
Especiales, ¢. Excerta Parasitologica ent memoria det 
doctor Eduardo Caballera y Cahaltera pp, 455-458, 
Revision of the genus Mucrapestroneyiis and 
descriplions of three new genera: Popovastrongylis, 
Dorcopsinema unl —Arundelia — (Nematodic 
‘Trichonematidae), Trans. R, Soe, 8. Aust, WE, 51-02. 
The genus Micruerameres Travassos (Nematodi. Spir 
uridiadin Australian birds, Bee §, Anat, Mais, 17239259, 


A taxenomig revision of the genera Maer 
posmongyloides Yumaguti and Maramacrepe- 
svrangvlns Johnston and) Mawson (Nematodi: 


‘Trichonematidae) frony Australian marsupiils. Ais, 
J Zeal, 26, 763-787. (With 1, Beveridge), 


197K 


1078 


OTS 


1078 


197K 


1978 


1978 


1974 
19749 
1980) 


Pa) 


19K) 


1982 
1983 
18S 


1986 


OSG 


1oou 


Mitrapienla vevdromi oe wm sp. CNematodas 
Stroneylidac) trom a western Australian kangaroo 
Trans, RO Saue 8. Aust. 2, 113-015, 

Macraponema (Nematoda: Trichoneniaiidie): a mew 
genus Tron macropod mitrsupiitls. Jit. Parasol &, 
163-166, 


A new genus Adeloneme (Nemmroda: Oxyuridie 
from Australian phalingerid marsupils, Praia. ht 
Soe, So Aust, 102, 223-226, 

Nematode parasites of the Kangaroo Island walluby. 
Macropus eagenit (Desmarest). 1, Seasonal ane 
geographical distribution, fbi. 12. 9-16. (Wilh 
L, RK. Sales), 

Nematode and other helminth parasites of the 
Kangaroo Island = wallaby, Maeraypuiy  eugenir 
(Desmarest). 2. Site selection within the stomach. 
Thiel. 12, 79-83, (With L. Re Sales), 
Cross-transmission of sirongyle nemiutodes benween 
macropods and domestic stock, Ast vet, 54. 181- 
182. (With L. R. Smales}, 

Parasites of Australian native miniinuls. Buel Alir. 
Mammal, Soc. 3. 8-15. (With D. Me Sprat & 
Beveridge, R. Domrow, D. Kemp & M.D, Murray). 
Alacastama new genus (Nemkiteda:  “Tricho- 
nemitidae), Trams AL Soe 8 Auer, 103, 124-126, 
Some Tetrumeridae (Nematoda: Spiruriday Tron 
Australian birds. hid, LOR, 1776184, 

Some strongyle nematodes (Anidosteninnn spp) 
from Australian birds, (hie 1040-12, 

Beverilgea new vents CNergatoda Strongylida) 
from the agile wallaby from northern Australia. (ie 
104, 81-82. 

On some oviparous filurial nematodes mantty front 
Australian bids, Rec. 8. Arst, Mits, TR. 205-284, 
(With O. Bain). 

Some Acuariinue (Nematoda) trom Austealian birds, 
Tratis, R. Soe. §. Aust. 106, 19-30, 

On the status of some nematode species fron 
Australian birds. (hid, 107, 247-248. 

Nematades (Avearidia species) front some eiphve 
and feral parrots. $. Aust, Ornithol, 29, 10-191, 
Redeseription of Temameres certo (Leidy, PakO) 
Nematoda: Tlibronematoides, Travis. Raa 4 Auer 
110, 77-79, 

A checklist of helminths tram Australian birds. Ree 
§. Aust, Mus 19, 219-325, (WiLL. M. Angel & Sd 
Rdmonds), 

A checklist of helminth parasites oof Australian 
repriles. Ree. S. Aust, Mas Manage Ser 8. Ot, 
(Wah S, Pichelin & M,N, Tlateliisan) 


VOL. 124, PART 2 
30 NOVEMBER, 2000 


Transactions of the 
Royal Society of South 
Australia 


Incorporated 


Contents. 


Reed, E. H. & Bourne, S. J. Pleistocene fossil vertebrate sites of the South East 
region of South Australia - - - - - - - 

Brown, S. P. & Wells, R. T. A Middle Pleistocene vertebrate fossil ae 
from Cathedral Cave, Naracoorte, South Australia - — - 

O’Callaghan, M. G., Davies, M. & Andrews, R. H. Species of Raillietina 
Fuhrmann, 1920 (Cestoda: Davaineidaec) from the emu, 
Dromaius novaehollandiae - - 

Cribb, T. H., Daintith, M. & Munday, B. A new blood: fluke, Fein 
forsteri, (Digenea: Sanguinicolidae) of southern blue-fin 
tuna (Thunnus maccoyli) in aquaculture - — - - 

Kolesik, P. & Cunningham, S. A. A new gall midge species —Binisea! 
Cecidomytidae) infesting fruit of punty bush, Senna 
artemisioides (Caesalpiniaceae) in Australia - -  - 

Goldberg, S. R. & Bursey, C. R. Intestinal helminths of five species of Seat 
lizards (Sauria: Scincidae) from Western Australia -  - 

Walker, S. J. & Goonan, P. M. Re-evaluation of the distribution of Geocrinia 
laevis (Anura: Leptodactylidae) in South Australia -  - 

Davies, M. & Burton, T. C. Redefinition of the Australian frog Limnodynastes 
depressus Tyler (Myobatrachidae: Limnodynastinae) -  - 

Beveridge, IL, Campbell, R. A. & Jones, M. K. New records of the cestode 
genus Pseudotobothrium PEPER Otobothriidae) 
from Australian fishes  - - 

Nicholas, W. L. & Hodda, M. Dorylaimus baylyi sp. nov. (ar vineniaae, 
Dorylaimida) a nematode collected from sediment in a 
freshwater rock-hole in the Northern Territory - = - 

Tyler, M. J. & Davies, M. Developmental biology and larval morphology of the 
frog Limnodynastes depressus Tyler (Myobatrachidae: 
Limnodynastinae) - - - - - - 


Brief Communication: 
Clarke, R. H. First record of the Southern Right Whale Dolphin, 


Lissodelphis peronii (Lacépéde, 1804) (Odonoceti 
Delphinidae), from waters off South Australia- - - - 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 


61 


9] 


163 


169 


177 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL, 124, PART 2 


PLEISTOCENE FOSSIL VERTEBRATE SITES OF THE 
SOUTH-EAST REGION OF SOUTH AUSTRALIA 


By E. H. REED& & §. J. BOURNET 


Summary 


Reed, E. H. & Bourne, S. J. (2000) Pleistocene fossil vertebrate sites of the South East 
region of South Australia. Trans. R. Soc. S. Aust. 124(2), 61-90, 30 November, 2000. 
This paper provides a summary of the Pleistocene vertebrate fossil sites of the South 
East region of South Australia and builds upon an earlier paper by Williams (1980). It 
also provides the first detailed review of all known Pleistocene faunal sites of the 
Naracoorte Caves World Heritage Area. Each known site in the region is listed with 
details of the site and faunal assemblage, fossil collections made from it and 
references to previous literature. The representation of the major vertebrate groups in 
the Pleistocene sites of the South East and the level of scientific attention they have 
received are discussed. 

Key Words: Vertebrate palaeontology, caves, South East, Naracoorte, South 
Australia, Pleistocene, Quaternary, vertebrate fossils. 


Transactions ef the Revel Snetery of 8 Aust 12000), 124(2),.01-90, 


PLEISTOCENE FOSSIL VERTEBRATE SITES OF THE SOUTH EAST REGION 
OF SOUTH AUSTRALIA 


by BE. EL. Reto & S.J. Bourn’ 


Summary 


Rebb, BH & Bobrpn, S.J. (2000) Pleistocene fossil vertebrate sites of the South East region of South 
Australig, Trans. R. Soe. S, Aust 124(2), 1-90, 30 November, 2000, 

This paper provides a summary of the Pleistocene vertebrite fossil sites of the South East region ot South 
Australia and builds upon an earlier paper by Williams (1980). Ht also provides the first detailed review af all 
Known Pleistocene faunal sites of We Naracourle Caves: Worl Aeritige Area, Buch known site in Lhe region is 
Tisted with details of the site.and faunal assemblage. fossil collections made from it and references mm previous 
literature, The representation of the major vertebruie eroups in the Pleistocene sites of the South Bust and the 
Jevelol serendtie atiention they have reveived are discussed, 


Khy Worbs: Vertebrate palacoitology, caves, South Bast, Neracourte, Saath Australi, Pleistocene, 


Ouaternary, vertebratys lassibs, 


Introduction 


The South Eust region of South Australia (Fig, 1) 
is predominantly a kurst terrain characterised by 
features such aS dolines, caves and cenotes 
(sinkholes), The Oligocene to Miocene Gambier and 
Naraceore limestones of the South East ecantain 
humerous eaves, with more dian 170 haying been 
recorded forthe Upper South Bast and more than 400 
in the Lower South East (Lewis 1979!> Matthews 
1O85; K. Mott pers, comm. 1999). Many af these 
caves contain skeletal malerial of Pleistocene 
vertebrates. “These sites have received much 
scientific attention and will he the main focus of this 
paper, Wihams (1980) published the first catalogue 
Ol Pleistocene vertebrate fossil sites of South 
Austria. but listed only a small number of sites for 
the South East. Palacontolowieal research in the 
region has beet steadily inereasing since 1980, 
paruicularly on sites in the Naracoorte Caves: World 
School Ot Biologie Sclenves, The Flinders Ciniversity af South 
Australia GPO Box JOU) Adehude SA. SOOT, E-ajail 
Ne dod Hindet’s cathau 
Numecourtle Caves Conservation Park, PO Bus 134 Naniaorte SA 
SI71- 

‘Lewis, 1D, 1974) South Austealiin Caye Reference Hapdboub, 
Oceasonl Paper Noo 3" (Give Exploruiiuin Grow of South 
Auistoilid, Adelaide 1. 

Nbwros, ©. oy, (1088) WY faphonomic dnd) paliedeealogreal 

amilyois vf the Civeen Waterhale (SUKI a submerged Tale 

Pleistocene bone deposit in the Lower southewst oF South Austria, 

Boe (Hons) ‘Tears. The Plindecs Gniversity of South Australia 

(unpub) 

Brows, S, BR (IY98) Ao wenlogiceal pod palteantologival 

examin oof the Pleystcene Cathedial Cave tossil 

awweuintanoan, Nardeoorte, Suuth Austnilia, BSe (Hows) Thesis, 

The Flinders University of Swath Australia (unpub. 


Heritage Area and surrounds. Thus, with further 
reseurch, Ongoing cave exploration and, most 
revently, Vineyard development new cave sites have 
been discovered highlighting the need Lo review the 
fossil sites of the region in depth, The currert paper 
builds on Williams” (1980) study and includes sites 
ihat were only under preliniinary investigation at that 
lime. sites omitted by thir author and those 
discovered and investigated more recently by the 
present authors add the palaeontological research 
team at Flinders University. This study originated as 
part of the PhD studies of one of us (EH R). 

The mujority of the sites discussed in this paper are 
incaves, Various modes of bone accumulation have 
been suggested, including natural traps and predator 
aecumulauons (Sauth 1971, 1972: Pledge 1980a. 
1990; Wells eral. 1984: Batrd LOS; Newton LO8h*: 
Barrie 1997; Brown 19987, Brown & Wells 2000: 
Morurty ef af. 2000), Many of the sites display 
multiple and overlapping aecumulation modes. Less 
common in the region are surface sites and others 
sack us the accidental finds where drilling of bores or 
construction works have led to discoveries (Wells & 
Pledge 1983). Several ol the fossil deposits in the 
region have been extensively researched, such as 
those of Henschke’s Fossil Cave (3091, 5097). 
Green Walerhole Cave (5L&1) und the Victoria Fossil 
Cave (SU 1) in which research hus been continuous 
for almost 30 years (Smith 197), 1972, 1976; Van 
Tets & Sinith 1974; Wells 1975: Wells eff. 1984; 
Moriarty er af. 2000), Other caves such as Wombat 
Caye (5U58) have received little more than 
preliminary investigation, while others have only 
heen surveyed und fossils identified ia sim, e.g 
Rabbit Cave (S066), Some of these cave sites po 


(2 E. H. REED & 8. J, BOURNE 


, 


VICTORIA 
N a7. 
15-24 
UPPER SQUTH EAST 4-14.25 
osquito Creek 
Bool Lagoon 26 | aa 
28 | 
o92/ | 
—_ LOWER SOUTH EAST 
vo i 
/ 31-32 0 amt Burr f 
33 ( 
{ 3435 
\ 3637 «3839 
do. 7 Mt. Gambier 45-49 % 
. 414942 
43 Mt. Schank 
\ J 44 


SouthEast ©” 


km 


Fig. |. Map of the South East of South Australia, with sites marked by a Number corresponding to those mentioned in the 
iext. Divisions of Upper and Lower South East sub-regions are indicated. 


longer exist, due to land development and others are 
yet to be fully explored. This paper ts up-to date as 
of 31 July 2000 but research in the region is 
continuing, particularly in the Naracoorte Caves 
World Heritage Area, 

This paper is not intended for use as a 
biogeographical database but simply provides faunal 
lists for each of the sites, along with some 
background information. Due to differing 
chronological sequences, it should not be assumed 
they ure contemporaneous, The main aim ol the 
paper is to report on new sites m the region. ta 
highlight their significance and to provide more up to 
date faunal lists, particularly for sites within the 
Naracoorte Caves World Heritage Area. 


The Naracoorte Caves World Heritage Area 


The presence of bone material in the caves of 


Naracoorte was recognised soon ufter the discovery 
of Blanche Cave in 1845 (Wells & Pledge 1983), The 


first significant work on vertebrate fossils from the 
region was carried out by Woods in 1857 and 1858, 
and recorded in his book “Geological Observations 
in South Australia’ (Woods 1862), Later, yertebrate 
fossils were reported from Specimen Cave by 
Stirling (Stirling 1908, 1912: Wells & Pledge 1983), 
Very litthe  palacontological reseurch — was 
subsequently undertaken in the region until the 
1960s, when material from Haystall Cave and 
Henschke's Fossil Cave was investigated (Merrilees 
1965; Pledge 1980c; Barrie 1997), The discovery in 
1969 of the Fossil Chamber in Victoria Fossil Cave 
(then known as Victoria Cave) and subsequent fossil 
discoveries in other caves of the Naracoorte Caves 
Conservation Park, led to an upsurge in research 
activity in the region und a growing awareness 
among the scientific community of its importance. 
The significance of the Pleistocene fossil deposits 
of the Naracoorte Caves Conservation Park was 
recognised internationally in 1994 when the Park 
wus inscribed on to the World Heritage List. The 


PLEISTOCENE FOSSIL SITES OF THE SOUTILLAST il 


Narnicoorte Caves deposits, together woh 
Riversleigh in Queensland. form. the Australian 
Fossil Mammal Sites. ‘The Pleistocene fannil record 
at the Naracoorte Caves is extensive, the caves 
Inivinw acted as pillall iraps and owl roosts, 
collecting examples of the fauna of this small 
gedgrdphic region over atleast the last 400,000 yeurs 
(Ayhite ete. J998: Brow) 1998: Brown & Wells 
2000; Mortarty ef af. 2000), Within the World 
Heritage Arew J1 of the 26 caves have yielded 
yerrehrale bane material When combined with 
recent climatic apd geoehronologioal work, the 
potential of the bone material for resolying 
pulucoccologicul and other contentious issiies, such 
as ihe timings of the megafaanal extinetivns. is 
vonsiderable (Aylifte er a 1998; Moriarty er vl 
20), 


Materials and Methods 


Tic Hist of sites and faunas provided in this paper 
has been compiled from the collections and records 
of the Sooth Australian Museum and. the Flinders 
University vertebrate pulaeontolozy labarutory, 
vurrept resedreh. published literiuture. personal 
communication with researchers studying sites in the 
trexion and held research by the authors, The 
Jocauoos of the fossil sites discussed im this paper wre 
shawn in Fig. |. Their nunibers correspond (9 site 
Himbers given in the Usts of sites and faunas 
provided and in Table 3, The format is similar co that 
of Willams (1980) but additional information, 
melding site details and carrent research is provided 
for each site, Cave numbers Ui.c, Cave Exploration 
Croup of South Australia CEGSA registration 
numbers) follow thase of Lewis (19794. Matthews 
(1985) and current CEGSA recerds. For these 
Mienbers 5” ydicates the state of South Austratia 
and “Uae “Lh rpper Oy lower South East sub- 
reson. The division inty Upper and Lower sub- 
regions used in this paper (Fig. 1) conforms with the 
CEGSA divisions for cave Jocutions. Within these 
sub-regions siles have heen grouped uccording lo 
distiict, determined by the au(ors as encompassing 
wi approximidely 25 km radius of the migor 
townships Of the South bast. The Naracoorte Cuyes 
World Heritage Arca is presented separately fron the 
Sunveorte distdet, Site names follow Waillians 
(1980), CEGSA revords and the published literature. 
The storage locution of fossil collections fram eich 
sie is also ineluded, as are the sodrees. for the 
InfopMAOn presented. Sites under investigation by 
the wulhors are identified, 


Watanws QOL) PR) Mie late Pherstoecene of Wie Flinders sieved 
Mi Fatty Moenees PADS Theo, Pte TP Tieteee Criveesiny od Seanib 
Nostale canpub | 


Systematics 


Checklists of faunas represented in the Pleistocene 
deposits deseribed in this study are presented ih 
Tables | d& 2. Tuble 3 shows the distribubion af 
species hetween tie sites presented in Ute win (ext. 
Phylogenetic order tor marsapials fullows Aplin & 
Avoher (1987), Robinson ef af (2000) ace followed! 
for placental mamnuis, replies. amiphibiins and 
birds. Names. tuxenomic authorities and 
distributional dala were taken primarily fron 
Robinson ef af, (2000), with some additional 
informabion taken rom Strahan (1995) tor 
mammals. Cogger (2000) tor repiles and 
amphibrims und Pivzey & Kaight (1997) for birds. 
Names and authorities for fossil species follow 
Archered af (1984) lor marimals, Baird (F985). Vien 
Tels & Smith (1974) und Stirling & Zietz (1896) for 
birds. aud Smith (1976) for the fossil reptile 
Wonamhi nracemieisis, Relerences for authorities 
for Hames published subsequently are ineluded in the 
Relerences section uf this paper, Distribubonal anc 
survival status changes are indicated for cach species 
in the list of sites and the lanl checklists (‘Tables | 
& 2). with #4 referring to species which became 
extinet during the Pleistocene, referring to 
historically extinct species. jind * indicating those 
species Which are locally extinet, of currently not 
found in the South East reg@iin. 

Fauial names used inthe faunal fises confor with) 
current usage, Nomenelatural changes that affect 
species included in this paper are summarised in 
Table 4. This table [sts the current name (is used an 
this paper), the previous. pane as fl appeared in 
earlier publications for Pleistocene sites of the South 
Exst und the relevant references, Changes in 
identification of fossil specunens are noted i the 
Falinal lists With appropenute yefererices giver, 

Williams (1982") provided revised diagneses tor 
the genus Dipreredoan. Ve fisted *+Piprotoitan 
australis and *¢D, optetun ats separate speeies. The 
identification provided for site 32 conforms with 
Williams” diagnoses (R. Wells pers, comm. 2000, 
Sraith (1972) identified “Aytechiniy stnartie Prous 
site ta, Subsequent work hie changed the concept ol 
the modern species OF Tel, Adearih ancl papulalines 
formerly included in FA. seed? actually coniprise 
Iwo siblings species, 7A. anartin aid “AL auiley 
(Dickman e7 af. PORR: Dickowit 1898). On ihe basis 
of modern ranges (Strahan 1995). any identification 
at SA, srueirtii Crom Pleistocene tossil deposits af the 
South East is likely te be the newly recognised "A 
auilry rablrer than the true A. sieearsiv, 


Results 


The following list of Sites und fats provides a 


O4 EH. REED & S. J. BOURNE 


TAaLe |. Checklist of amphibian, reptile and bird species identified or tentatively identified from Pleistocene fossil sites 
of the South East of South Australia. 


CLASS AND ORDER FAMILY AND SUB-FAMILY GENUS AND SPECIES 

AMPHIBIA 

ANURA Hylidae (Tree frogs) Litoria ewingi (Dumeril & Bibron, [841) 
Myobatrachidae Crinia signifera (Girard, 1853) 
(Southern Frogs) Geocrinia luevis (Giinther, 1864) 


Lintnadynastes dumerili Peters, 1863 
Lintnodynustes tasmaniensis Giinther, 185%. 


REPTILIA 
TESTUDINES Chelidae Chelodina longicollis (Shaw, 1794) 
(Side-necked Tortoises) *Emydura macquarti (Gray, 1830) 
SQUAMATA Agamidae (Dragon lizards) Pagona barbata (Cuyier. 1829) 
Scincidae (Skinks) Egernia whitii (Lacépéde, 1804) 
Eulamprus iympanun (Lonnberg & Andersson, 1913) 
Lerista bougainvillii (Gray. 1839) 
Tiliqua nigrolutea (Quoy & Gaimard, 1824) 
Tiliqua rugosa (Gray, 1825) 
Varanidae (Goannas) *Varanus gouldii (Gray, 1838) 
“Varanus varius (White, ex Shaw, 1790) 
Mautsoiidae “| Wanambi naracoortensis Smith, 1976 
(Madtsoiid snakes) 
Elapidae (Elapid snakes) Notechis scutatus (Peters. 1861) 
*Pseudechis porphyriacus (Shaw, 1794) 
Pseadonaja wuchalis Ginther, 1858 
AVES 
STRUTHIONIFORMES Casuaridae Dromaius novaehollandiae (Lathan, 1790) 
(Cassowaries & Emus) 
Dromornithidae ++ Genyornis newtoni Surling & Zietz, 1896 
(Dromornithids) 
GALLIFORMES Megapodiidae Leipoa avellaia Gould, 1840 
(Megapodes) ++ Progura naracoortensis Van Tets. 1974 
Phasianidae Coturntx pectoraliy Gould, 1837 
(Pheasants, quails & allies) Coturnix ypsilophara Base, 1792 
ANSERIFORMES Anaudae Gen. et sp. indet. 
(Geese, swans & ducks) 
PELICANIFORMES Phalacrocoracidae Phalacrocorax melanoleucos (Vieillot, 1817) 
(Cormorants) 
FALCONIFORMES Accipitridae Accipiter Brisson, 1760 sp. indet. 
(Osprey, hawks, eagles & allies) Aquila audax (Latham, 1802) 
Faleondue (Falcons) Fulco berigora Vigors & Horsfield, 1827 
GRUIFORMES Rallidae *Gallinula mortierii (Du Bus, 1840) 
(Rails. crukes & allies) Gallinula tenebrosa Gould. 1846 
Gallirallus philippensis (Linnaeus. 1766) 
TURNICIFORMES Turnicidae (Button-quails ) Turnix varia (Latham, 1802) 
CHARADRIIFORMES Pedionomidae *Pedionomus torquatus Gould, 1840 
(Plains-wanderer) 
Scolopacidac 
(Sandpipers & allies) 
Gallinagoninae Gallinage hardwickii (Gray, 1831) 
Tringinae Tringa glareola Linnacus, 1758 
Calidrinae Calidris ruficollis (Pallas, 1776) 
Burhinidae (Stone curlews) Barhinas grallariuvs (Latham, 1802) 
Charadriidae *Charadrius australis (Gould, 1841) 
(Plovers & dotterels) 
COLUMBIFORMES Columbidae Phaps chaleoptera (Latham, 1790) 


(Pigeons & doves) 


PLEISTOCENE FOSSIL SITES OF THE SOUTH EAST as 


PSITTACIFORMES Cacatuiday 


(Cockatoos & cockutiel) 
Psithacidive (Parrots) 


CUCULIFORMES 
STRIGIFORMES 


Cuculidae (Cuckoos) 
Strigidae (Typical owls) 
Tytonidae (Barn Owls) 
CORACTIFORMES Alcedinidae (Kingfishers, 
bee-eaters & rollers) 
Acanthizidae ( Bristlebirds, 
thornhills, scrubwrens & 
allies) 

Meliphagidae (Moneyearers 
& Australian chats) 
Orthonychidae 
(Chowehtllas, quail-thrushes 
& allies) 

Dicruridae (Monarchs, 


PASSERIFORMES 


drongos, magpie-larks & allies} 


Aruimidae (Woodswallows. 
buteherbirds & allies) 
Corvidae (Crows) 
Hirundinidae 

(Swallows & martins) 
Estoldidae (Grass-linches } 


Cacatue tenuirostris (Ruhl, |820) 
Callocephaton fimbrianun (Grant, (803) 
Calyproviynchus banksi (Latham, 1790) 
"Culprovhyachius lata (Temminek, (807) 
"Pezoporus wallivus (Kerr, 1792) 
Plarveerous Vigors, 1825 sp, indet, 
"SCenrropus colossus Baird, 1985 
Ninox novacseelandiae (Gmelin, 1788) 
Nyro alba (Scopoli, 1769) 

Wro novaehollundiag (Stephens, 1426) 
Docelo wovueviinede (Hermann, 1783) 


Dosvornis broadbenti (MeCoy. 1867) 


Matorina inelinoceplala (Latham. |802) 


SOrthonys hypsiopliis Baird, (XS 


Gralline cyanoleuce (Latham. $802) 
Gurnavhing tibiven (Latham, 101) 


Chrvus Linnueus, 1758 sp. indet. 
Hinundy neorena Goull 1842 


Gen. et sp, indet. 


Invomplete identifications are included only if they represent the only entry representing the family or genus concerned, 
*F indicates species extinct during the Pleistocene, ** indicates historically extinct taxon, * indicates Gixon no longer 


vecurs in the resion. 


catalogue of all known Pleistocene fossil vertebrate 
sites und faunas of the South Bust. 


Sites and Faunas of the 
Upper South East region 


Kingston district 


1. BLACKFORD DRAIN 

LOCATION: 21 km NE of Kingston. 

SITE DESCRIPTION: Fossils were uncovered in the 
north side of the creck-bed duting bridge 
construction in 1954 1n a bed of “waterworn stones” 
ula depth of approximately 3.5 - 4.5 m (Williams 
1980). A letter from Ro V. Flint accompanying the 
specimens described the lowest level usu hard stone 
which looked like a flow of black mud, thickly 
impregnated with small white shells” (Williams 
1980) 

COLLECTION: South — Australian Museum 
palucontolory collection (vertebrite fossils), 
KAUINA; 
MAMMALS 
Diprotodontidae: 


+ Dipratodeun sp. indet., 


*“tZyeomalurus trilobuss Macropodidae: Macropus 
giganteus. M, rufagriseus, *}Procoptodon sp. indet. 
*+Simosthenurus aecidentalin, 

REFERENCES: Williams (1980); N. Pledge (pers, 
comm. 2000): South Australian Museum paliae- 
ontology collection records. 


Naracoorte Township 


2. HENSCHKE’S Fossi. Cave SU91, SUIOT (also 
known as Henschke’s Quarry Cave) 

LOCATION: Outskirts of Naracoorte township. at 
Henschke’s Quarry, 

SITE DESCRIPTION: The cave was exposed by 
quarrying and found to contain a rich and diverse 
fossil assemblage. It was excavated by workers from 
the South Australian Museum from 1969 to 1981 te 
salvage material from the eave, Whieh was part of the 
active quarry (Pledge 1990), Subsequent excavation 
was curried out by J, Burrie from 1981 to 1997, 
investigaling an extensive section radiating from the 
location of the carlier excavations (Barrie 1997), As 
quarrying lias continued the cave hus been 
completely destroyed. 


G6 E. H. REED & S, J, BOURNE 


Taner 2. Cheeklist of mammal species identified or tentatively identified from Pleistocene fossil sites of the South East of 
South Australia. 


CLASS AND ORDER FAMILY AND SUB-FAMILY GENUS AND SPECIES 

MAMMALIA 

MONOTREMATA Tachyglossidae (Echidnas or *+Mevalibgwilia ramsavi (Owen. 1884) 
spiny anteaters) Tuchyglossus aculeatus (Shaw, 1792) 

MARSUPIALIA 

DASYUROMORPHIA Thylacinidac (Thylacines) “*Thylacinus cynocephalus (Harris, L808) 
Dasyuridae Amechinus flavipes (Waterhouse, 1837) 
(Carnivorous marsupials) Antechinus minimus (Geottroy. (803) 


*Antechinus stuartii Macleay. 1842 
*Antechinus swainsonti (Waterhouse, 1540) 
*Dasvurus geoffroli Gould, 1841 

*Dasyurus maculatus (Kerr, 1792) 
*Dasvurus viverrinuy (Shaw, 1800) 
*Ningaui yvonnue Kitchener, Stoddart & Henry, 1983 
*Phascogale calura Gould, 1844 

* Phascogale tapoatafi (Meyer, 1793) 
'Sarcophilus herristi (Boitard, 1841) 
"Sarcophilus faniariuys (Owen, 1838) 
Sminthopsis crassicaudata (Gould, (844) 
*Sminthopsis leucapus (Gray, 1842) 
Sminthopsis murina (Waterhouse, 1837) 


PERAMELEMORPHIA — Peramelidae Tsoadon obesulus (Shaw, 1797) 
(Bandicoots & bilbies) *Perameles bougainville Quoy & Gaimard, 1824 
*Perameles gunitt Gray, 1838 
DIPROTODONTIA Phascolaretidae (Koalas) Phascolarctos cinereus (Goldtuss, 1817) 


+ Phascolarctos stirtoni Bartholomai. 1968 
Diprotodontidae (Large extinet 
marsupial quadrupeds) 
Zygomaturinae 


*+Zyeomaturus trilobus Macleay. 1858 


Diprotodontinae **Diprotodon australis (Owen, 1844) 
"+ Dipratodon opranin Owen. 1838 
Palorchestidae (Large extinet + Palorchestes asael Owen, 1874 
tapir-like marsupials ) “+ Palorchestes parvus De Vis, 1895 
Vombatidae (Wombuts } *Lasiorhinus kreffiii (Owen, 1872) 


"Lasiorhinus latifrons (Owen. 1845) 
Vombarus ursinus (Shaw. 1800) 
“+ Werendia wakefield’ Hope & Wilkinson, 1982 


Thylacoleonidae *tThylacolee carnifex Owen, [858 
(Marsupial “lions’) 

Phalangeridae (Brushtuil Trichosurus vulpecula UXerr, 1792) 
possums & cuscuses ) 

Hypsiprymnodontidae *+Propleapus oscillins (De Vis, 1888) 
(Sectorial-loothed rat-kangaroos) 

Potoroidae *Aepyprvmnus rufescens (Gray, 1837) 
(Potoroos, bettongs & *Bertongia gaimardi (Desmarest, 1822) 


rat-kangaroos) *Bettonsia lesueur (Quoy & Gaimard, 1824) 
*Bettongia penicillata Gray. 1837 

*Pororous platyops (Gould, 1844) 

“Potoreus tridactylus (Kerr, 1792) 


Macropodidae (Wallabies, 
kangaroos & tree-kangurous) 
Sthenurinae (extinct | Procoptodon goliah (Owen, 1846) 
browsing kangaroos) 4} Pracoptodon rapha Owen, 1874 
“+ Sunosthenuras baileyi (Prideaux & Wells, 1998) 
“Ss Simosthenurus brownei (Metrilees. 1968) 


PLEISTOCENE FOSSIL SITES OP THE SOUTH EAST 67 


Macropodinae 


Burramyidae 
(Pygmy-possums } 


Pseudocheiridae (Ringtail 
Possums & Greater Glider) 
Petauridae (Striped Possum, 
Leadbeater’s Possum & 
wrist-winged gliders) 
Acrobatidae (Peathertail Glider) 


PLACENTALIA 

CHIROPTERA Vespertilionidae 
(Ordinary bats) 

CARNIVORA Canidae 


(Dogs, foxes & allies) 
Felidae (Cats) 

Otartidae (Eared seals} 
Suidae (Pig) 

Bovidae (Horned ruminants) 
Muridae (Rats and nee) 


ARTIODACTYLA 


RODENTIA 


LAGORMORPILA Leporidae 


Simosthenurus gilli (Merrilecs, 1965) 

ty Simosthenurus maddockt (Wells & Murray, 1979) 
+ Simosthenuras newlonae Prideaux, 2000 
“TSimosthenurus occidentalis (Glauert, 1910) 
“+ Simosthenurus pitles (De Vis, (895) 

“+h Srhenurus cuderyoni Marcus, 1962 
Congruus congruusy McNamara, 1994 
“tLavorchestes leparides (Gould, 1841) 
*Lagostrophus fasciatus (Peron & Lesueur, 1807) 
*Macropus cugenil (Desmarest, 1817) 
Macropus fuliginosus (Desmarest, 1817) 
Macropus giganteus Shaw, 1790 
**Macropus greyi Waterhouse, 1845 
Macropus rufogriseus (Desmarest, 1817) 
*}Macropus titan Owen, 1838 
“Onvehoxalea lunata (Gould, 1841) 
*}Protemmodon anak Owen, 1874 
=+Protemnodon breliis (Owen, 1874) 
*+Protemnodon roechus Owen, 1874 
*Thylogale billardierti (Desmarest, 1822) 
Wallabia bicolor (Desmarest, 1804) 
Cercartetus concinmus (Gould, 1845) 
Cercartetms lepidus (Thomas. 1888) 
Cercartemts nanus (Desmarest, [S18) 
*Petauroides volans (Kerr, 1792) 
Pseudocheirus peregrimts (Boddaert, 1795) 
Petawrus breviceps Waterhouse, 1839 


Acrobutes pyenuaeus (Shaw, 1794) 


Mintopteras schreiberstt (Kuhl, 1817) 
Nyctophilus geoffreyi Leach, 1821 

Canis lupus familiaris Linnaeus, 1758 
Vulpes vulpes Linnaeus, 1758 

Feliy catusy Linnaeus, 1758 
Arctocephalus Geotfroy & Cuvier. 1826 sp, indet. 
Sus scrofa Linnaeus, 1758 

Ovis aries Linnaeus, 1758 

*6Conilurus albipes (Lichtenstein, 1829) 
Hydromys chrysovasier Geoffroy, 1804 
*Mastaucomys fiscus Thomas, (882 
Notomys mitchellii (Ogilby. 1838) 
Pseudomys apademuides Finlayson, 1932 
*Psendomys australis Gray. 1832 
*Psendoniys fumeus Brazenar, | 934 
*Pseudoniys gouldii (Waterhouse, 1839) 
Pseudomvs shortridgei (Thomas, 1907) 
Rattus fiusvipes (Waterhouse, 1839) 
Rattus lutrealus (Gray, 1841) 

*Ratis taineyt (Thomas, 1904) 

Ory ctolagus cuniculiy (Linnaeus, 1758) 


Incomplete identifications ure included only if they represent the only entry representing the family or genus concerned. 
FF indicates species estinet during the Pleistocene, indicates historically extinct axon, * indicates taxon no longer 


occurs m the region, 


x nsoydopsds XTUsMey 


x sypsojaad XMUANJOD 
py x. xX x x SISUALLOOIVADU DAN ON A 
x pIpjavo vodiaT 
x x Japul ‘ds s7mioauay s,s. 

x IUOJMAU STULOAMUAD) 
xX x yp x Xx AVIPUDLJOYAVAOU SMIDUOLG 
Spite 
x yapur “ds vipuopnasg 
x x ‘yo syjpyonu pipuopnas gf 
x x "yo snopiudcydiod s1ydapnas x 
x x x ‘J SAIDINIS SIYIAION 
x x XxX Xx SJSUALMOOIDADU 1QUIDUOA 4. 
x x x yapur ‘ds snunsn, 
p cp ‘Po ge SNLUDA SHUDADA x. 
aa] yp cp MP]NOS SNUDADA x. 
rs) x x x XxX Xx x OS psosns pnbipiy 
5 x *¥ Xx x Xx pajnjo1s1u pnbyty 
oa) x UPPIAUIDSNOG VISMIT 
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a ‘Pp pinging PUOSOg 
x xX cp MADNDIVUL DANPAUWT 
ss} x S1]JOI18UO] DUIPO]aYD 
w saqnday 
x ‘yapur “ds sayspucpouurT] 
% SS SISUALUDUISD] SAISDUXPOUWTT 
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1p Xx SIMav] DIULLIOAN 
Soe pdafiusis DID 
Xe 1SULMA DONT 
suviqrydiry 
BRERA PE Se Ree Oe Ue BOS REESE SSR bh SBE Ss CRA AES eS SaIOdds 

ALIS 


“SIST] [PUNR] 
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S ‘yxa] UIDUL aYl Ul pajuasatd Says ayl Uaaattag Saloads Jo UOLINGLSIpP IY] SiMOYS AjqVI *¢ AVAV I, 


69 


PLEISTOCENE FOSSIL SITES OF THE SOUTH EAST 


x x x 
x 
yoyo 
x 
x 
x 
x 
x 
x 
x 
x 
x 
x 
x 
x 
x 
x 
x 
1p 
x 
' 
x 
9 
x 
x 
x 


XX X 


ae 


x 


snpoydadouayd SNULIDIAY 


SNIDIINID SNSSOPSKYIVI 
IADSLUDS DIPIMS GIVI 4. 
S/RULLUR AY 

DUaxXOAu OPUNALAD 

‘jopur ‘ds smasoy 

UIIIGH DUIYAOUWAD 
DINAJOUDAI DUIIVLD 
snydopisdaAy XAUOYLIO 4s. 
pjpydasouvjalt DULIOUD JY 
MUIGPVOAG SIULOASDG 
IDIUINSADAOU O]AIDG 
WIPUY]|JOYPDAOU OIA 
DqY OK], 
ADIPUD]AISADAOU XOUINY 
SHSSOIJOI SNAOMUIT |, 
Jopul “ds sna4aIAV] 
SNINJOM SnLOdOozag s. 


japut ‘ds snyousysoidajp9 
MUDYID] SNYIUKYAOJAN]D I), 


usyupg snyoudsysolda]py 


wnpiiquaf uojpydaso]|09 


SLUSOLINUAL DNIDIDI 
‘joput ‘ds sdpyg 
psajdozjoyo Sdvyd 
SYDAISND SHLMPDADY Ds. 
SNLUD]DAS SNUTYANG 
DJOALD]S VDEULLL 
HYIIMpIVY OSDUI]TOH 
Sqpoayint sLipipoy 
snyonbso] snuouoipad. 
japur ds xmany 

DIIDA XUAN 
sisuaddipiyd snj]payjoy 
psosqaual DjNUDH 
MAIYAOU DIMU]VY x 
DAOS19G OJ]D 

xppnp pjyinby 

yapur ‘ds sayidiooy 


SOMALOUD]AUL XVAOIOLIDIDY 


yapul ‘ds x70 


E. H. REED & S. J. BOURNE 


Ol 


70 


‘jepur “ds snuysoiwpT 
SUOLUD] SNUIYLOISY 
miffary SNUIYOISDT] 
snaupd sajsayr1ojDd 
[ADD Sajsayrojdd }. 
SNGOP] SNANMUOTKT. 
yeput ‘ds uopojosdiq }... 
wnjpido uopojosdig, 
syp.usnp uopojoidig ds. 
‘Jeput “ds sojamjoospy dg 
IMOIANS SOJIMDJOISDY bse 
SNALIUII SOJIIDJOISDY 
apul ‘ds sajauipsag 
IMUNS SA]aUlD1I x 
A]IAUIDSNO SajaUp.Lads. 
japul ‘ds wopoos] 
S7]JNSaqgoO UOPOOS] 

‘japur ‘ds sisdoysuiuy 
pULINi sisdoyjulUug 
sndoona] sisdoyjunus.. 
pippnnaissp.io sisdoysuius 
japut “ds saprydooings. 
SMLIDIUD] SN[LYAOMVS |... 
TISLLADY SNIN{AOIIDS s. 
aput ‘ds ajpsoospyg 
Y{pIModY] a[DIOIS DY ds. 
DANIDI 3]VSOISDUY x. 
ADULOAN INDSUIN 
‘jopur ‘ds snanaspg 
SNULMIAAIA SNANASDG, 
SNIDINIPUE SN.ANASHGs. 
OAJOIS SNANKSOCs. 
Jopur ‘ds snuijpoayuy 
MIUMOSUIDAMS SNUIYIAIUYs, 
INADNIS SHUNJIAJUY, 
SMUT SHUNIATUY 
sadippf snulyoaluy 


SHIOdd$ 


7\ 


AST 


SOUTH I 


x ‘japul ‘ds aynsojAy 7 
MAAIpAD] [IG 2]DSONY J 
MMOSAIPUD SNANUAYLS 4» 

‘japut ‘ds sMMUayISOUl]S 4. 
Sappd SnANUaY{ISOUNS + 

X SIDIMAPIIIO SHANUPYJSOULS 

aDUOIMa SHNUAYISOUL 

LYOOPPPUL SHLANUIYISOUNS 

1/J15 SHANUAYIJSOULS 4 
JOUMOL SNANUIYISOULNY | 
IAa]ING SNanuaysouns | 
“japut ‘ds vopouiuajorg 4 
SY 2IOL UOPOUWAOL 
SNYAAG UOPOUWAION | 4. x 
YPUD UOPOUWAOL | 4. 
apul “ds uopoldooodd 4... 

x pydps uopoidovo.tg 4 

ypyos uopoidosodsd |... 

DIDUN] DA|DSOYILUO x. x. 


oe 
« 
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tal 
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a 


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PLEISTOCENE FOSSIL SITES OF THE 


bal 


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i 
4 
a 
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x 


x x 


moe 


woo 


i“ 


.o} 
ss 


oe 


mnjjioluad pisuolag 


japut ‘ds sundown 
up Sndodovyy bs. 


snasiisofns sndosony 


IMAL3 SNAOLIDP x x 
snaquns1s sndosovy 


snsoursynf sndosavyy 


Muasna SNdosgV x. 


SHIDIISDL SNYMOASOSVT 
Saplloda] SalsayHOsyT x,y. 
SHNASUOD SNNABUOD 4. 
SHJAIIVPLUL SNOLOIO” 5. 


sdoxqv]d Sno10j0g x. 
japur ‘ds nisuoyag 


AHINSA] DISUONAY se. 
IPADUUDS DISUONAG ¢. 


suasafns snuussdxday x 
suppioso sndoajdod g 4... 
DNIadpNA SAANSOYILLT, 
NALIUIPI OFJOIVINY [4.5 

x IPJAYayYM VIPUL +. x. 
. : ¢ Z f cx SHUISAN SNIDGUIOA 


ca 
uv 


E. H. REED & S. J. BOURNE 


x 
x 
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49 
x 
RRARDRRRAL 
OmMmAIADUMEWH— 


“UOISAL DY] UL SINIIO IBUO] OU UOXE] SAIBSIPUT ,. ‘UOXR] JOUNXA AT[LOLOISTY SayIIpul 


2 WW WW 2 Wa LW Ln Wd 


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SNNIWUNI SNIVDJOIIAC 
apul ‘ds spy 
INAUU SNIDY x. 
snjoasn] SNUPY 

sadiosnf snyiny 

‘Jopul ds stwopnasg 
193 PIALIOYS: SAULOPNAS 
LIpnos sauopnasd x 
snaunf SXWOPNasd ». 

S]JDAISND SKULOPNAS x. 

saploulapodp sKwopnasd 
NPJAYI[IUL SAUWOJON 
SNISNf SAWOIDISDIN 
apur “ds swmoipaAy 
AA]SPSOSKMAYI SKUOAPAH 
‘yapul “ds snunjiuoy 
SadIq]D SNANPUOY x. 
SID SIAQ) 

pos sng 

aput ‘ds snjpydasojo1y 
SMIDI SY}24 

sadjna sadjna 

stupipiuny sndny siuny 
Moadffoas snjiydolovn 
apur ‘ds sniaidommpy 
msdaqiaiyos snsajdoluyyy 
snapusad Salpqo1oy 
sdadiaalg SNANDIag 
snulisasad SnAlayIOpnas 
SUDJOA SAPLOANDIA ». 
Snub SNJajIVIL1aD 
snpida] snjajivo1a7y 

SNUUIIUOD SNIALADIMID 


AOJOIIG DIGD]INA 


SHIOddS 


PLEISTOCENE FOSSIL SITES OP THE SOUTH EAST TA 


TABLE 4. Stmumary of nomenclatural changes related to species included in this paper 


THIS PAPER 


Myobatrachidae 


Crinty signifera 


Pagora barbate 


Fuamprus tympani 


Tilique rugosa 


Conraix vpstlophova 


Gallirallis philippensis 


Brrliiins wrallarits 
Charudriuy australis 


Calyplorhynchus banksit 


Dicrundae 


Artumidae 


Megalibewilia reunseyi 


Thylacims cynocephatus 
Vonnbeatis 
Poterous tridacty lus 


Preudomys apodemoides 


PREVIOUS NAME 


Leptodactylidae 


Ranidella signifera 


Amphibolurus barbaris 


Sphenomorphus Nmpanua 


Trachydosaurus rugosuys 


Coturnix australis 


Rallus philippensis 


Burhinus magnivestriy 
Peltalivas australis 


Calyptorhynchits magniticus 


Grallinidac 


Cracticidae 


Zarlossus ransayi 


Thylacinus major 
Phascoloms 
Potorous apicalis 


Psendomys albocinereus 


REFERENCES FOR THE SOUTH 
EAST IN WHICH THE PREVIOUS 
NAME APPEARED 


Tyler (1977, 1991); Williams (1980): 
Wells & Pledge (1983); Brown & 
Wells (2000); Moriarty er al. (2000), 
Tyler (1977); Williams (1980); Wells 
& Pledge (1983); Wells ev al, (1984): 
Pledge (1990); Moriarty ef af. (2000). 
Smith (1976); Williams (1980); Wells 
& Pledge (1983); Wells eral, (1984), 
Pledge (1990); Moriarty e7 al. (2000) 
Smith (1976): Williams (1980); Wells 
& Pledge (1983); Wells ev al. (1984): 
Moriarty ef al. (2000). 

Smith (1976); Williams (1980); Wells 
& Pledge (1983): Wells et al, (1984): 
Cogger (2000). 

Van Tets & Smith (1974); Williams (1980): 
Wells & Pledge (1983); Wells et al. (1984); 
Baird (1991); Baird et al. (1991). 

Van Tets & Smith (1974), Willtams 
(1980): Wells & Pledge (1983); Wells 
ev al, (W984); Newton (1988*): Baird 
(1991): Baird et al, (1991). 

Newton (1988*); Baird (1991). 

Van Tets & Smith (1974); Williams 
(1980): Wells & Pledge (1983): Wells 
ef al. (1Y84): Baird (1991), Baird e7 al. 
(1991). 

Baird (1985); Newton (1988*): Baird 
(1991); Baird eral, (1991). 

Van Tets & Smith (1974); Williams 
(1980); Wells & Pledge (1983); Wells 
eral, (1984): Baird (1991): Moriarty er 
al. (2000). 

Van Tets & Smith (1974); Williams 
(1980); Wells & Pledge (1983); Wells 
et al, (1984); Baird (1991): Moriarty er 
al. (2000). 

Murray (1978); Pledge (1980c); 
Williams (1980); Wells & Pledge (1983): 
Wells ef a/. (1984): Pledge (1990): 
Griffiths ef al. (1991), 

Williams (1980)- 

Tindale (1933); Williams (1980. 

Smith (1970); Williams (1980); Wells 
& Pledge (1983); Wells eral, (1984), 
Wells & Pledge (1983); Wells ev al. 
(1984). 


i (OUD REED & Ss. BOURNE 


COPPFOTION: Sauth Australian Museum palae- 
Ontology calleetorn Cyvertebrate fossils), 

bAUNA? 

AMPIIBEANS 

Livlidkie: Lieria ewirai: Myobatrachidues Crna 
sismfere, Chea Mi lees, Ldarendvinensres 
NNT en AEN, 

REPTILES 

Chelidue: Chelulina longicollis, cl *Lawvduvit 
me qari, Agamidie: Povons sp. indet. Scincidue: 
Tiliguea nigroliled, T, rugosa: Varanidae, Varaiis sp, 
ch OV park’. Vuranus sp. ch 2 varius: 
Madtsondae: oh Wenamhi nardcoortensiye Elapidies 
Preudonaja sp, indet, 

Bibs 


Casnariidae: Dyrnmains hovaehallandiac 
Mesapouiidie: Progra IMEFUCOOPLEN SEY: 
Phusianidue: Comrie 4 mdet.; Anand: 


indeseribed taxon; Rallidae: *Gallnahe mortiert: 
lurticudae: Turnia vertes Psittaciforiies: farnily 
indef.: Passeriformes: amily indel: Corvielie: 
Corvus sp. inden: Hirondinidae: Airunele: reeena. 

MAMMALS 

Tuchyglossidae: “tMewelibewilia ranisevie, Lach 
glossy achleaius, Thylacinidaes *° 7 rviietniey 
cymocephatus; Dasyuridae: Anivehinus sp. ch A, 
minis, HDasyartis viverrinusy, SPhaxscovale sp 
indet., 7? Sarcophilas lanidtins, *Smirithapsis 
lencopuy: Peramehidae, Iyeodon obeyulus, Peranteles 
sp. OF HP heugatiile. Perameles spelt P warn 
Phascalaretidae: Phasealarctas sp. el Eo emerety, 
Diprotedontidac: &+Dipreteden aptatin, bev e0 
malurus irtlotis; Palorebestidacs Palorchestes 
acael, Vorbutdae: Lasierhinis sp. ck FE krefftii, 
Vibes tersinas: “Thylacoleonidae: biivlecalen 
cortex, Phalanveridae: Trrehosuriy vulpeculis: 
Hypsipryinnodontidae: Th Prepleapas oseitdaiis; 
Polonia: “Aepvprimrnis rufescenis, Bellongia sp. 
eh OB vatmards, *Berangio lesuenr, Betis sp 
uh *B. penteilata. Patorons spo cl! FP. plewveps 
=P tridaciuliy: Macropodidie: ** Lagerehestes 
loparides, * Lagesirophay fasciatus, Marcraypns sp. vt 
Mo vrvantens, FM. erevi, Mo rufieriveus, \M. 
lila. *"Onveheavalea lundta, “+ Procepleden raphe, 
'FPrademnodow roechus, © Stnestienicuy brawnet, 
POS. wall, TSS. nandidewki, OFS. neawlanue, FS, 
necidemalis, V8. pales. *oSrhenuras aidersont 
(= Sileruris atlastandersant ot Pledge |99U, see 
Pridewix [999% Wallabia hicalor: Burramyidae, 
Cercarens nana, Psoudocheinidaes Previdocheinis 
perewiitis,  Petauriye:  Peraideis — brevieeyay, 
Vespentiionidae: Vyctoplilys sp. ef No vealproyy, 


OPI AUK Gr CP99) Syvstennitios and cyatllional) Ue es bel 
heme SubRunih Sthenwiinie. PHD Mesias The Flinders 
Uriwersiiv at Sen Gustelin Cenpub ) 


Moridae: *Contlurus sp. indet.. A yelroniys 
Chrvwogaster, *Mastcomys fuvens, Psendomys sp. 
indet,, Keiras sp. inde, 

REPERENCES: Van Tets (1974); Tyler (1977, 1991); 
Pledge (1977. PORE. 1990, 1991); Murray (1974): 
Williams Cl9O80): Barrie (1990, 1997): Baird 
(1991b); Baird et cil, (991), Geitfiths er al (1991): 
MeNumiara (1997); Pridewux (79997, 2000); Scunton 
& Lee (2000): South Australian Museum 
palucontology ealleetion records. 


3. JAMES’ QuagkyY CAVE 5U29 

LOCATION: Naracoorte township. 

SITE DESCRIPTION: This small cave was. uneoyered 
by yuurrying in 1956, discovered by A, James. 
proprictor of (he quarry, Th contained a partial 
skeleton of Thylaceles carnifex (Daily, 60), The 
wave has -sinee heen destroyed hy quarrying, 
COLLECTION: South Austeuhiin Museum 
palucontology collection (vertebrae Tossils), 
PAUNAL 

MAMMALS 

Peramelidae: "Perameles bareainvilles Vombandie 
Vambatus vrsinas: Vhylicoleonidae: *°7hvleenlea 
canilfer: Potoroidae: +Betroeia valmardi, °B 
lesneur: Macropodidaes Macropuy giventens, MM 
rufavensens. =F Onyehovelea linear. Ft Sline- 
whens gilt 

RELERENCES: Daily (1960); Pledge (1977); Williams 
(1980), South Adstridan Museant palacontolosy 
colleetion records, 


Naracoorte Cavey World Heritage Area 

The Naracoorte Cuves World Heritage Area has 
loll area of wpproxtmately 305 heelures und as 
centred Fl) kim sontheast ofthe Naracoorte township 
There ure 26 caves on the reserve, imainy of whiels 
contin deposits Gf Pleistocene and/or Holocene 
vertebrates, with a particularly rich record of 
marsupuuls, The Naracoorte Ciyes were inseribed on 
to ihe World Herilage List in December 1994 as an 
Australian Fossil Mammal Site (serial pominition 
with Riversteigh, Queensland) for them exceptional 
Hitiraband scientific value, 


4, Vievroria Foss Cave SU | 

LOCATIONS Naracoorte Caves World Hertize Ares, 
DESCRIPTION: Lurge cuve of approximately 4 ki ol 
Thapped passages and chambers. The eave contains 
five known fossil deposits. wih the largest and most 
studied being that i the Main Fossil Chamber. 
which wis discovered in 1960 by members ol 
CEGSA, Other chambers containing fossils have 
heyn found sinve then, All are currently under 
Invesiyalion by Flinders University palacontologpsts 
and the fiwinus identified to dale ure listed below 
Uranium-series dating of speleothems isso) nitvd 


PLEISTOCENE POSSIL SITES OF THE SOUTITEAST is 


With these fossil deposits has placed their age range 
from) Middle to Late Pleistocene (Ayhffe eral. 1998; 
Moriarty er af. 2000). 

COLLECTION: Flinders University vertebrate 
palwontology Collection; South Australin Maseum 
palatontolowy collection (vertebrite fossils). 


di, MAIN bosst. CHAMBER 

SUL DESCRIPTION: This chamber has ao extensive 
hone deposit within a lane sediment cone and fin, 
The deposit has a complex depesitional history with 
multiple modes oF aecumiuation aml concentration 
evident. chictly pital) (rap. predator accumulation 
Givi dud mininalian) and hydraulic transport. 
Litaniiescries dating of fowstone on (he surface ot 
the deposit has provided a minimum age of about 
23 ka (AytMe eral }998: Moriarty er al. 2000). 
COLLECTIONS Flmders University vertebrate 
pahicontolowy collection: South Australian Museu 
paleontology collection (vertebrate lossils). 
FAUNA; 

AMPHIISLASS 

Hylidae: Line ewingi, Myabitrachidaes Crtia 
sinitera. Geocrinivsp, cb OG. laevis; Loninndynastes 
speck de dumertli, Linmodynisdes fersareaniertts, 
Rerris 

Cheldue: *baivdira neque Agamidae: Pergeies 
sp el Po harhatia, Seincidue: Evernia whitii, of, 
Lvtanprus tympanunt, Leite beouganertit, Higvee 
nivoolitea, TE rugesa, Varumdae, Vareniiy sp. cb ove 
roukdiy Varin sp. eh FV veri: Madtsotulae: 
hr Worn nardcoortensis, Mhapidae. Noreohiy sp. 
of Noweutaris, Beatties Wis sp at ME porphyrtae is, 
Premlomier sp. cl, Po ntettalts. 


Lirbs 
Casunieidae, Drones Hovaehullandiae: 
Mesapodiidiwe:  Leipod ecellata, rh Pragura 


noracvoriensis, Phasiuidae, Colurnis pecloraty, ©, 
yaitlophora, Ballidae: Gelfrratias philippeusis 
Vurnietdie: fais verte Din osp. Tadets 
Podionomidac: * Pediononiis HSPs 
Seolupucidac,  Calielris  ruficallis, Cel lttetyer 
hunhwekdi, Tring shares  Chiradridac: 


*Charedrits ciytvatiy, Psittacutae, =Perapeariy 
wallicvua;  Tylonidwe; Tle neveehollandiue: 
Dieruridue: Grafling cvaneleuea: Ariamidae: 


Gynmnorhing this: Cocvidae: Corvus sp. inet. 
“Kallas wellivaiius’ listed by Moriarty ef al (2000) 
appears to he misspelling (Wo Boles. pers. corn 
O00) 

MAMMALS 

Tuchylossidue: ehMevalibawilta rams 
Tavliyelossus vette, Thylacinidae: © Thvloe ins 
oynueephalis. Dusyundae: Anrechiny flavins. ON. 
wedi, AL seatnxeamt. “Denvuriy macula, 7D, 
viverriitusy,  Ningerat sp. ele ON. vitenutete. 
“Phascosih valued, *P rapoeidtafa, Sarcophilus sp. 


funieln, 


ch 27S. daniartuy, Soiathopsiy craysicaudate. &. 
jorhias Peramelidac: dsocdon obesidis, *Perameles 
howarinuille. =P. unanii: — Phascoliretidue: 
Phisweokuretas CTE TRIENS Diprotodonticdae: 
=“ Dipratadeant sp. indet.. *Aygemetirus trilobiy; 
Palorchestidae: +h Palen theses. agdel, Vombatidae: 
“Lasivrhinuy Krelftit. Fhe batitrons,  Viaubectey 
ursinuss ‘Thylacoleommdies FF Tivkiwelea carnifey; 
Phalangeridae: Trichosnrus valeting 
Hypsiprymnodontidae: ch At Prapleapis avciltans, 
Potoroidac: ef “depvpevaniis ciufescears, *Reroneia 
catmundi, Bellunuin sp. ch HB desneur, TR, 
pemeillant, ** Petros planus. * PL tridaceylis 
Macropodidac:  ' Lagarchestes  leporides (= 
Lagoichestes spel. HL conspiciitaniy Of Wells et al 
1Ob4 und Moriarty ef af. 2000. sce McNunnura 1997), 
*Mucrapuy vugenit, M. faiteinosus, My sglecuteis, 
SSM prey ML orufagriveus,  *0M. rite, 
*ePracaptodon poliale (= *tPrecoptedan rapla ol 
Wells ef af 1984, see PrideaX = T899)), 
*e Prafemmearta reeclins, **Sinnsitenurus baileyi, 
FES. browne FPS. wll TS. reedelawki PTS 


1, 

newronae, oTS. decidentatiy, TS. pales. er Sdienturws 
ctnhidervont (= Srhenurus atlas of Wells et a. 1984, 
see Prideaux |9Y%), Wallabre bicelor: Burrumy ida: 
Crrcarrenis lepidus, Co nuaus: Pseudocheiridae; 
Pseudocheinuy peregrinus: Perauridue: Perernctes 
brevieepy, Acrobulidie: eb, Acrehetes pryginaeny: 
Vesperilionidue: Mirfapterus sp. Inder, Muridae: 
*eConttnray albipes, Hydronws  chrysexasler, 
= Musiucimiys firscis, Neinmys sp. ef No uniiwheltit, 
Papeelemoides. *Poaustraliy, Preadoimys sp. cf. &2 
Psotudwis spo ck Fee weuifdir 2 
shoriridger, Rattus fuscipes, “R. laaneye, 
Moriarly er ud, 2000 listed two additonal marsupis, 
“Desvecrenus eristeandea and *Pemarneles teasvtie. 
However no specunens can be located li support 
these idenufications, Which appear unlikely, They 
hive therefore been omitted from this list. 


4h. GRANT HALT. (also known as White Chuinber) 
SITE DESCRIPTION: Exauvations hive yielded bone 
imalerial [rom the sediment floor of the chamber al 
the base ofa urge Galus cone. The bone deposits are 
ussochiled with several levels ol speleothems, 
uranium-serivs dating of which indicates (hat the 
deposits accumulated between about 206 ka and 76 
ka CAyliffe eral, 1998: Moriarty etal, 20001, The site 
is currently under mivestiquiion hy Rebevod Greshurnt 
from blinders University. 
COLLECTION? — Flinders 
paluvoulology collection. 
FALINA, 

REOTIEDS 

Aguimdie: genet sp. inden: Vaarainidaer Vararies sp 
inde, = Madtsovidie: °F Woneabt neaneaartenst: 
Elapidie: pen. et. sp. mae 


University vertebrate 


mM EL REED & SO BOURNE 


BIRDS 

Order mndet. 

MAMMAI & 

Fachyplossidic: Mvehyelossis venleanes: Thyla- 
cmnidues F*Tivlaciis  evnecephalis:  Phasco- 
larciidae: Phascolaretos cinereans: Diprotodontidae: 
Aywemalurus trilabus. VYombutdae: Vonibarttus 
vesinuys Thylacoleonidue: *?Thvlacelen carnifer; 
Potoroidae! *Berenigia penivoillute: Maecropadicdac: 
Macropus giseneuy, M. rufogrixeus, Macrapus sp. 
nidet,, “?Simesthenurus hrownei, @ eS. gilli, & PS, 
vewranae,  Wallahia  biceler; — Burramyidive: 
Cercarterus lepidus, C uenius: Muridae: *Mista- 
COs fusens.  Pyeudomvs apodemoides, *P. 
australis, Psendanye sp. cl 8 vwauldii, P. 
shoriridger, Pseaduays sp. met. Raties fiscipes, 
VR panne. 


40 SPRING CHAMBER (also known as Starburst 
Chamber) 

SITE DESCRIPTION, Bone material has been 
exeavaled rom the sediment floor of this large 
chamber. Although only preliminary work has been 
done on the site, vrainiumeseries dating of ussourated 
speleothems sugeests that deposition bevun before 
327 ka ond upper layers of the deposit accumulated 
heeween 280 ka and 210 ka (Moriaty: er af, 2000) 
The site is clirrently ander investigation by 
palacontologists from Flinders University 
COLLECTION; Flinders University 
pulacontology volleetian. 

FAUNAS 

MAMMALS 

Dasyvuridae; Dewveray sp. ch Fl viverrinus, 
Phaseolarctidae: Phascalarctos rinerews, Phas 
volerctay sp. el TP winnk Vombaridae. pen, eb sp. 
indet., Macropodidae: Macrapus gigeanteus, M 
rufewisens, Macrupus sp, indet. “hSineasthentris 
wil 24S. eee ddenalis. 5S. pales: Muridae: 
Hydroniys chrvsogesten HRartas Wiens 


vertebrate 


4d, Urrer Asp Lower OSSUARIES 

SUH DESCHIPTION: These two vliwnbers in the distal 
part of the cave contin immensely rich bone deposits, 
Discovered in the early 1970s by BL Wright and BR. 
Gulbreath (CEGSA), they reniwin lirely antouched 
as avrelerence sile. To date, surface material only has 
been examined, mostly fa sin No excavation has 
been done in these chambers, A few specimens were 
removed front the aceess tunnel for ident fieakion 
when the chimbers were discovered. Additional 
material has heen identified fn itv by the wathors 


(CPE P OPMOS TD TWestnitron: ash The thers Aterapitis 
UMusapiilin; Mactopia) fond the Viera Fissit Cave depesil 
Nanos BSe Uo heat Te Pinder CIN Orsi) ah South 
Avstiilis Loepiehy 


COLLECTION: — Flinders vertebrate 
palacontology collection. 

KAUNAT 

Bigbs 

Casuariidae: ef. Dramnains noveehollandiac, 
MAMMALS 

Tachyglossidae, yMeealibewilae ramsayi, Vhyla- 
einidaes *°Thvlacinuy cvaecephalus, Dasyuridae: 
Sarcuphitus sp. ob eS. latierius: Diprotodontidue: 
“t+ Zygomanirus trilobas. Thylacoleonidae: °F yla 
coleo curnifes; Mactopodidae: Macropus spool, MW 
gigunienus, MM. rufogrineus,  FPSirestheniurivy 
browet eS. gli, PhS maddaeki, EtS_ ec iedentalls 
~TSthentirns atedersenr. 


University 


de. BUTCHLAND LAKE CHAMBER 

SUT DESCRIPTION: A small chamber adjacent 66 the 
Many Fossil Chamber discovered in the early 1970s 
hy A. Lake und B. Alveres (CEGSA). Bone mater 
Was discovered and colleeted, Additional material 
was collected in 1997/1998 and identified hy one ol 
the authors (2 HR) and colleagues from Flinders 
University, Al bone material within the chamber is 
found within the roek pile on the chamber floor, 
without any sedimentary context, thus providing 
some inleresting preservational features (Moriarty e/ 
wl, 2000), 

COLLRCTION: — Flinders 
palacontology cullection, 
HAUNAL 

REPTIURS 

Vierunidae: Varonts sp. inde. 

MAMMALS 

Thylacinidies |? Tavlecinny cyaocephalas, Phasco- 
Jurchidae: Phisealarctos cinereus: Thylacolvonidie: 
“)Thylacalea cartifex; Macrapodtdae, Meerapis spe 
el, M. filiginosus. Macropus sp. eb MM. rufegriveuy, 
STtoslenirtas sp. cf 8, gilli. 


University vertebrite 


REPERENCHS: Smith (1471, 1972, 19764, Van Tets & 
Smith (1974): Pledge (1977, l980b,-c, 97); Tyler 
(1977, 1991): Wells (1978): Wells & Murray (1979): 
Wilkos (1980); Duwson (1982): Wells & Pledge 
(O83): Wells eta. (M084): Baird (199 1a, by. Bate 
ehh (lO0L): Griffiths ep ai ()991): MeNumarn 
(1997), Aylite ef al. 1998) Pride@auy & Wells (1998), 
Prideauax (1999, 2000), Turner (1999%; Moriirty ¢/ 
al. (2000): M. Hutehinson (pers, corm, 1999, 2000), 
A, Baynes (pers. comm. 2000); R. Cireshain (pers 
cumin. 2000). C. Williams (pers. conn, 2000): 
Finders University vertebrate pulieontology 
collection dutabase; South Australian Museu 
palacontology collection records. 


5, BAT CAVE 5U2 
LOCATIONS Naracoorte Caves World Heritage Ares, 
SITL, DESCRIPTION: Bone mutterial was collected 


PLEISTOCENE FOSSIL SITES OF THE SOUTHEAST W 


from sediment beneath a ledge m the entrance 
chamber by Walsh in L959. The deposit is estimated 
ty be of Late Plenstocene age by fuamal association. 
wlthoughy ih is likely that more recent matovial has 
heen meluded with the colleghon. 

COLLECTION: South  Austrahian = Museum 
palucontolowy collection (vertebrate lossits). 

UAUIN AS 

MAMMALS 

Posyuridae: “Dasyiries maculata, “Daspurus sp. 
indet. *Phevengele tapecula. Sarcophilus sp. ef. 
“S. harrivit, Phalingeridae: Triehosaeny valpeculas 
Potoroidae, “Betroneiae gained’, Macropodidae: 
'TSimosthermmuy browne, *PS. sill Petauridae; 
Peniuris breviceps. Muridae: '* Conrluruy albipes, 
'Mastacoays faseas, Rattisy sp. mauet. 
REFERENCES! South Australian Museum palie- 
onlology collection records, 


6. ALEXANDRA CAVE SUS 

LOCATION! Nuritoorte Caves World Heritage Area. 
STL DESCKIPHON: Bone material was recovered 
fron) sediment when the second tourist entrance of 
the cave was dug out im 1978. Other fossil muterial 
wits discovered during cave exploration excayarions 
of small sediment filled tunnels in the current tourist 
seetion of the cave, Only preliminary investigatien 
of this site has so fur been attempted, 

COLLECTION: Minders University vertebrate: pulae- 
ontology callection, South Australian Museu 
palucontolagy collection (Vertebrate fossils). 
FAUNA? 

MAMMALS 

Doyyuridie: Dewars weenlatis. “Do piverruits. 
Surcuphilus sp. ch eS. lureriviic Phuscolaretidue: 
Phoscolarelos. cinertuss Vormbatidaes Vewibatas 
vesmiuss Thylacoleonidae: tT? Thvlecelee cameo 
Phalungeridae: Trrehaswres valpeculas Potoreidac: 
Bertone sp. indets: Miaeropodidie: Adeenupiy sp. 
eh A eieaiteins, Maur lapiis sp. cl ML rufeerivens, 
Mucropuy sp. inde, '?Preeeploadeon gel. 
“TSatemthenuras  hrownei, tS. gilli 2848 
Hcctdemelis. Wallahia divatar 

REPERENCKS! Pledge (1877); Williims (1980). 
Flinders University vertebrate palaeontolowy 
colleetion duitihuse: South Australian Museun 
palacontolagy collection records, 


7. BLANCHE CAVE SUA, SUS, SUG 

LOCATION: Naracoorte Caves World Heritage Area, 
STEED DESCRIPTION: Fossil material front this cave 
wus Ueseribed last century by Woods (1862), with 
sddivional material having heen collected since then- 
Hestly in the 1970s (notably the Gemvernis 
specimen). A small number of bones has been 
collected from the third chamberol the eave by eave 
explorers. 


COLLECTION:  Plinders University vertebrate 
palucontology colleetion: South Australian Museum 
palacontolugy collection (vertebrate Tossils). 
FAUNA: 

BIRDS 

Casuariidue: Dromainy novaehallandiae: Drome 
mithidae. *+Genverniy newer, 

MAMMALS. 

Pasyuridie; “Denyviruy sp. indet.. “Sarcepliiay sp. 
indet.: Peramelidae: *Perameles beonecinville, 
Phascolurcuide: Phaseolercios elnerens. Thylo- 
voleortidue: "TTiyliealeo carnifev: Phulangeridac. 
Trichoyuruy yulpecula. Potoreiduc: Berteiivry 
gaimondi: Macropudidae: **Lagorchestes leporidey. 
Myernpes sp. cf A gleanrens. ML rufeertseus 
“Onvelhoxalea latata. 'yPratembnvden Predury- 
He Some henustts ailll: Pacudocheimidie: 
Pseudocheirus peregriins, 

REFERENCES: Wools (1862. [866)0 Rich (1979); 
Wells & Pledge (1983); Baird (199Tb): Baird et al 
(1901): MeNamura (1997). Blinders University 
Verlebrale palneontology colleetion dutibase; South 
Australian Museum  paldeontolosy collection 
records. 


& Wer Cave - Stick Entrance SUL0, Tomato 
Enlranee SUT) Gulso know as Tomato-Stick Cuve) 
LOCATION: Naracoorte Cuves World Heritiwe Area. 


STH DESCRIFTION: Excavation wis carried out in 


1997/1998 in the distil fan sediments ofa lurve 


sediment cone im the culrent tourfst cave by 


Minders University researchers, Abundant borne 
maternal wits peeovered. purticukirly small aiunals, 
with Some imewaluunal species atthe lower levels 
ol the sequence, The site is) currently under 
investigation by M, MeDawell (Flinders 
University), Prelimioury resulis saggest a Late 
Pleistocene ta Holocene age for the deposit. Other 
material has been collected fron the cave im the 
pant aud lodged with the South Australian 
Museunr. 

COLLECTION: Flinders University vertebrate 
pulicontology collection. South Australian Maseuin 
paluavontology collection (vertebrate fossil). 
FAUNA 

AMPHIBIANS 

Family inet, 

REPTILES 

Agamidae: gen. el sp. indel; Semmeidue: Huernin 
whit, Viligia nigroluiwan To eiteeses Varanidae: 
Veramis sp. cl OVE veeriis: Madtsotidae: Pr Werenht 
naraceerrenviss  Wdapidaes Netechty  yeurattts. 
» Pseudechis porphyrideus, Prendaneaa uehaliy. 
Biris 

Psittucidues gen, ebosp, mdets Tytonidae: Ayre alba: 
Arlumidaes gen. cl sp. indet.; stride, gen, cb sp 
det. 


is bk HW REED & 8d BOURNE 


MaMMALS 

Thylacinide; °° ivlaeiiaus eynoceplitlus; Dasy- 
wilde Aiechnins flavipes, “Dayvirus weaffrait, 
'D. vivertinus. *Ningeut wwornide, “Phoseogale 
tipourafd. *Sareophilus  hearvish, Sminthopsis 
creasrcondala. Scouring, Peramelidac; *Peraineles 
wun, Phoscolarctidue: Phaseolarcios etrerens: 
Diprotodontiduc, T+ AZyeenanauy trilahus, Thyla- 
colvonidae: }Thylacelee carnifex: Phalangeridae: 
Trichosuruy vilpecula, Potoroidae, “Betronuie 
fesneur, "Patorous  platvopss Mueropodidae: 
Miacropus gigameus. ME pufoerisens. Macrapus sp. 
inden, '?Prvemnoden hreltus, ++ Protenmoedan sp 
indet., *?Stnrasthenneis brawie FPS, wall Hes. 
neavlonae, *TS. veadentalis: Burramyidaes Cerce- 
rrers, coneinnis. Co depridas. Co tariiss Pseu- 
dovheirtdae: Pyeudecheirts pereertiuy: Petautidie: 
Penman brevicepss Vespertiionidies Miniepterius 
seliverbersii, = Mupidae:  'Conilurus — albipres, 
*Masiucanys fiuscits., Notonws wmitchellii, Pre 
emis apodemaites, “PP. australis, VP. funets. 
Pyevudanivys sp. el. #77. geaddil. P. shortridyer. Rann 
fuseipes, Ro lureals, R tnaneyvs 

KEPERESCES: Willaims (1980); M, McDowell (pers. 
comm. 1999, 2000) Flinders University vertebrate 
pulucontology collection database; South Australis 
Mirseunt palacuntology collection reenrds. 


9. CATHEDRAL CAVE SUT2, SUI3 

LOCATION: Narmcaorte Caves World Hentige Area. 
SHE DESCRIPTION: A large chaniber in the distal part 
of the cave has waediment cone deposit containing 4 
large aumuunt of bone material, some in assecialion 
with dated Howstone. Uraniunvthoriunr dating of 
these speleothens sugvests that the material 
uccuniulated between upproximately 279 kaand 159 
hu (Brown (99%) Brown & Wells 2000), Brows 
{Grown 1996%, Brown & Wells 2000) concluded that 
The primary uceomulation mode wus pittall via a 
pow-blocked salihon tube. Other material ineluding 
Hoavlacales carnifes, was collected from other small 
passages inthe cuve by CEGSA meinbers in Maret 
1959 und reported by Diily (1960). 

COLLECTION: Tlinders University vertebtate 
pulacontology collection: South Australian Maseurn 
palacentology collection (vertebrate Mossils), 
BAUNAL 

AMPUIBIANS 

Myobutruchidies Aanraed veteran sp. Tider, 
REPTHIES 

Seineidde: Hfigiil Migesu 

Binps 

Parnily made. 

MAMMALS 

Thylacinidae: "Paver ins OVE pal’: 
Dusyuridice Auechaney flavipes. Antechinus sp. 
mde. Devers mentees. 8D. viverrnhnis 


*Phaseogale cdlura, Saiathopsis inane, Pera 
melidae: Aseoden obesuliy. "Peruneles hougein 
ville, *P. gunnil; Diprotodontidae: “tZAyvemeaneris 
vilobus: Vombatidac: gen, et sp. indet. 7 Leasterdinus 
lanfraus, Thylitcoleomdues e+ 7Ayhiealea camnifer: 
Potorpidac: “Rerrenieia penvillata, “Betlaneta sp. 
indet.. T=? Palarouy plaivepy. TP, (ridactylus, Macro 
podidae: Macrapns elgantens, Mo rufnerisens. 
Meeropus sp. indet..  HpPravepraden gollal. 
ht Simasthenurus  brawaei, 818. all ehs 
ocoidentaliy, Wallabia bicolor; Burramyidae, Cere 
urielux Hanus. Murtdae: Mateos [lwens, 
Netomys netehellii *Pseadomvs australis. 2. 
shortridgel, Pseudonis sp. inde. 

ROPERENCES: Daily (1960): Pledge (1977); Williams 
(1980), Aylilfe ef al, (1998); Brown (1998%, pers, 
comm, 2000); Brown & Wells (2000): Moriarty eral 
(2000); Blinder University vertebrite paluvontology 
collection database; South Australian Museum 
palwontology collection records, 


10. ROBERTSON Cave SLIN7, SUT8, SUT9 
LOCATION: Nirteoorte Caves World Heritage Atea. 
STP DESCRIPTION: ‘The inner chamber ol the cave 
contains a tich bene deposit, particularly of srnall 
mummal renuwins. Megataunal species have been 
found atthe lower levels of the sequence sugsesuiig 
a Pleistovene uge. The sites is currently tinder 
Tyestigution by M, McDowell (Flinders University, 
COLLECTION; Flinders University — vertebrate 
palicontology volleenon, 

VAUNAL 

AMPITLBIANS 

Pumly indet, 

REPTILES 

Agamidae: gen. et sp. indet, Sermudiae: Beernia 
whit, Tiliqna nigraluled, To riagesxa, Varanidae: 
Vardiis sp. ef MO varivy, Elapidae. WNelechis 
Seutumis. "Pseudechiy porphyrins. Psedanafe 
nuchealis. 

BIRDS 

Fuamily mies, 

Parttucidite: gen. ob sp. idets Tytonidae: (rer ad/ees 
Artiumidae: ven. et sp met; Estrikilidae: gen. et sp 
indet. 

MAMMALS 

Dusyuridie: Amechinis flavipes,  *Duxyurus 
veoffrati. TD. vivertinis FNingant yvonne. 
*Phoscogale tapoutaja. Srmuthopsis crassieaudate. 


S. mmurine: Pecamelidae: lyeedon obesity, 
*Perumeley enomiits Phascolarctidae: Phescoleretes 
CPMCT ONS Vombatidie! Vomibatus HESTDUN, 


Pholungeriditie: Trielesuris pulpecnla: Potoronae, 
“Bellonula  leynenr,  **Pearerouy —— pleatvapiss 
Mieropodidites Mecrapuy efeantens, My rufugriseny, 
“FProtemnedear sp. inde SU Stnasrhenieries 
ucvidenulis, Birramyvidue: Cercedtens leptin. ©, 


PLEISTOCENE FOSSIL SITES OF THE SOUTIEEAST zh 


nami, Pseudocheinidae: FPefaurides valgus, 
Msendocheirns pereurimis, Petaoridue. Pernrus 
brevieeps, Vespertiliomidie: Minloptertis seliet- 
hepsi, Muridaes **Contfarus albipey, Hydroniys 
chrysovasier, “Maxticomys fuscix,  Neneuiys 
mitehellin, Pvewlonys upodenmides, “Po austrelis. 
*P. fiunens. Pyeudontyy sp. el BTR gould P. 
Shoriridver, Rutty fuscipes, Ro dutreoliw, *R- 
HWnneyy, 

REPERENCES: M, McDowell (pers. comm. T999. 
2000): Minders Liniversity vertebrate pulaeontology 
collvetion databuse. 


Il. Fox Cave 5U22 

LOCATION, Naracoorte Caves World Heritage Area. 
STrE DESCRIPTION: Fossil material has been 
recovered. from this large cave, with excavations 
being conducted by reseirehers from the South 
Australiins Museum and Plinders University. The 
deposit consists of numerous bones contained in the 
sediment floor of the cave und lurge sedinient cone. 
COLLECTION: — Flinders University vertebrate 
paleontology collection, South Austealiin Museum 
paluwontology collection (vertebrate fassils). 
PAULINA: 

Bikios 

Meipodiidie: Pravira iareceortensiy, Alee- 
dinate: Daeelo sp. cl DL nevaeguinede, 
MAMMALS 

lachyglossidae: by Mevelihgwilta renusivi, Tacly- 
wlossas cdeulears, Thylicinidae: **Thvlacrins 
cynecephalws: Dasyuridues *Dayyvuruy otaeulelity, 
°D. viverrinis. Sarcophilus sp. el tS. laniarins: 
Perametidac: hyoudon abesuley. *Peraneles eanniiz 
Diprotodontidie: “4Zvgenaturus trilobux; Vor\ba- 
Hidde: Vombaris Wsitus: Thylicoleonidae: 7 iile- 
cleo carnifex: Phaulungendae. Trichosirius vulp- 
evita; Potoroidae: *Bertongia gaimardi. * Porras 
tdeetyiny: Maecrupodidiue: Maeropus sp. ch MM. 
vigentents. Morel MM rufovriseus, Macropus sp- 
of A titan hPreceplodon woliah, °° Stme- 
sHenurus browned, FES gilli, @T8. eecidemnilis. 
Willubta hicater. Pseudoehetidae: Pyendenhelruy 
pereertniss Muridae: TMestaceuyy fiiyeus, Ratiay 
sp ck Ae latreetiny. 

REFERENCES: Pledge (L977. 1Y80e): Murray (1978): 
Baird (lMO9lb) Baird ef ed (l99OT Flinders 
University vertebrate palacantolawy  calleetion 
ditibase; South Australian Museum pialaconlology 
collection records 


P22 LIN-SAMED Cave SLAY 

LOCAMON: Nuraceorte Caves World Lleritige Area. 
SVT DESCRIPRION: A stnall cave witha solution pipe 
enirunce and very bithe cave development. Bone 
material iscontiined wilhliia sediment! cone beneath 
a blocked former sefutton pipe entyanee. No 


excavations haye been conducted in this cuve but 
identifications were made from a smulh amount al 
material Collected by the authors fon areas that had 
been previously disturbed. 

COLLECTION: Flinders University vertehnile pale 
ontolopy collection, 

VAUNA? 

REPTILES 

Varmidac: Vareniay sp. indet, 

Bikps 

Megupouiidaes #7 Prasad naracaartenyny, 
MAMMALS 

Thylucinidae; =" Thylacinas exrocephealus, Yorbi 
tnlaes Vowbeadies ursiniis; Macropodidae: MWucrpus 
sp, indet “*Simosthenuris gill 

RELERENCES: None. 


13. WomBAT CAVE SUSS 

LOCATION: Naracoorte Caves Worle Heritage Are 
SITE DESCRIPTION: The second chamberol the eave 
was discovered in the carly 1970s, with a sitall 
collection of bene material being lodged with the 
South Australian Museum, Additional mitterial his 
been identified (7 sit on a rubble slope adjacent tou 
large area of speleothem development. much of 
which has formed over what was probably a cone 
beneath w solution pipe. The site is currently under 
investigation by one of the authors (FER). 
COLLECTION: — Flinders University vertebrate 
palacontology collection; South Aastraltiin Museuin 
palaeontology collection (vertebrate fossils). 
FAUNAS 

BIRDS 

Mesapodnidae: 'yPragunen naraceortensts. 
MAMMaArLs 

Dasyuridae: *Dayvaruy viverviniy: Peramelidive: 
*Perameles gunnii: Vombatidae: Vembuitts ursinus: 
Phalangeridae: 7richoswrus  vilpecula: Macra- 
podidae: Macrapuy sp. el Mo giganteus, Mo rufi- 
griseus, © Protenmodon sp. inde, Fe simiosthenwrus 
brownei. TS, villi, **Sthenuruy andersani, 
REPERENCES: Baird (1991p): Batrd er af (1991): 
Flinders University vertebrate palicontologs 
collection ditabase; South Austrian Museun 
palaeentology volleetion records. 


14. SANT PUNNEL CAVE SLIT2 
LOCATION: Naracoorte Caves World Hertaze Area. 


SITE DESCRIPTION: Bone malertul wis collecteil 
from surface sediment in the L97Os. connest 
unknown 

CoOrLecTion: Flinders University vertebrate 


pulacontologey collection. 

FAL NAS 

MAMMALS 

Macropodidae; SMecrapus vugenit  Sinieathe 
nurs maddacki. 


SO OH. REED & 8, J, BOURNE 


REPERENCES: Flinders University yertebrate palac- 
ontology collection database 


Orher Naracoorte [district] cave sites 


15. BRown SNAKE Cave SUI 

LOCATION: Naracoorte Forest, Forestry SA. 

SITE DESCRIPTIONS Bone material collected by 
CEGSA members, context unknown, 

COLLECTION: South Australian Museum patie 
ontology collechon (vertebrate fossils). 

FAUNAS 

MAMMALS 

Macropadidac: 77 Simosthenurus gilli, 

REPERENCES: Williams (1980): South Australian 
Muscom palicontology collection records. 


Io, HAYSTALL Cavii 5U23 

LOCATION: Private Jand 

SITE DESCRIPTION: Bone material was excavated by 
W. Rouse and ML R. Wallis und N. Pledge und R. 
Callen, from within the cave duping the 960s, with 
ibundant hone material discovered in the slape and 
fan of adarge sediment cone. 

COLLECTION: South Austruliia Museum palie- 
ontology calleetian (vertebrate fossils) Some 
identifications of material in site were made by the 
authors. 

PALNIAT 

RePiiL Fs 

Seincidues Tilique nigreluteds Madtsoiidie: +] Won- 
AT meracaertensiny 

MAMMALS 

Tachyglossidue: Tachy gloss tts weuleats: 
Thylacinidac: ' Thylaejans evnecephilus, Dasy- 
uridues Amechinuy sp. mdet., *Dasvyrty viverrtiitis, 
“+Surcaphilus faniarius; Peramelidue; lyeodon 
dhesuluy. @Peraumeles gunnity Phascolaretidae: 


Phascolarctes sp. indet Diprotodontidae;, *FZvyee- 
maturuy (rilebus; Vombatidae: Vonibenis asinus: 
Thylacoleamidge: 4 7hylicales earnifer: Potoroidac: 
=Aepypryputus rufescens, *Beronuia gutinaredi, 
= Polorouy plabvops, “P. tridectyluss Macropodidae, 


Pseuducheiidue: — Pserdarherris — peregrinus: 
Murnlae; *Mustectpays fayetis, Ratiny lutrealas. 
REFERENCES? Merrilees (1965), Pledge (1977, pers, 
comm, 2000) Williams (1980), South Australian 
Muscuin palucentology collection records. 


17. UN-NAMEID CAVE SU28 

LOCATION: Narth of Naricoarte (ownship. 

Sut BPSCRIPTIOGN! Small cave 40 mt from Woe 
Cave (SU26). Bone snaterial Was collected during 


cave exploration, context unknown, 

COLLECTION: South Australian Museu) palae- 
ontology collection (vertebrate fossils), 

FAUNA: 

MANMALS 

Maeropodidie: Macropus sp. indet, 'Simosthe 
nuruy brevwnel. 

REPERENCES: South Australion Museany patie 
ontology collection records, 


18. Specimen CAVE SLI35 — false known as Fie 
Caye) 

LOCATION: Private land, 

SITE DESCRIPTION: A-solution pipe leads down toa 
large chamber with bone deposits tn sediment 
associated with flowstone kryers. Stirling reported 
malerial of extineL marsupials, including 7iyvlacoler 
carnifex. trond the eave in J908 (Stirling, L9O8: Wells 
& Pledge 1983). Another report by Surling in 1912 
tnenboned 7. cumifes yaaterial Tram the Naracoorte 
Caves which had heen presented to the South 
Australian Museunt by W. Redden, the caretaker of 
the Ciuves. The nume of the site from which this 
material was collected did not appear in the report 
However, ik is most likely ta be Specimen Cave. 
COLLECTION: Plinders University vertebrate palace 
vontalogy colleeuon: South Australian Museum 
pakiwontology collection (vertebrate fossils). 
VAUNA: 

REPTILES 

Seimeidae, Whyne rigesa- 

MAMMALS 

Thylacinidae: VET Ay leciius evMnace plein: 
Dusyuridae: *Duyveruys sp. indet.. Sarcaplilis sp. ef 
48) Janiarius. Sminthopsiv sp, inet; Peramelidie, 
*Perameles sp. indet.: Vombatidae: Vonbatis 
ursinus: ‘Thylacoleonidue: ')Thyvlacelee cornifes. 
Potoroidae: *Beploneia penteiflai, "'Patoroaus 
platrvaps: Macropodidae: Macropus giganteus, VW. 
rifugrisens. Mactopus sp. ef SM. titan, Macropus 
sp. indet.. Protemmodon spo ck TP. anuk, TTP 
roechius, =) Sunosthenurus baileyi. FS, browned. YS, 
gilli, “4S. maddacki, M48. aveidentalis, *Thylogule 
sp. indet. 

REPRRENCDS: Stirling (YOR, OD): Pledge (1977); 
Willins (L980); Wells & Pledge (L983), Flinders 
University palaeontolovy collection database; Sout 
Australian Mirsennt palaconfolozy collection 
records. 


19. RaBBir Cave SU66 

LOCATIONS Private lund. 

SITE DESCRIPTION: This small cave has hones 
evident in sfteon the surface of he sediment Moar at 
a osmiall distal chamber, No cacwwation fas beer 
conducted jn the cave. However. identifications of 
maternal we sTit were made by the authors 


PLEISTOCENE FOSSIL SITES OF THE SOUTH EAST ¥I 


COLLECTION: None made 

PAULINA! 

MAMMALS 

Muaeropodiidee: Maermpuy sp. ef ML fitltuitasies. 
Somosthenuriy spo eh. OS. brovated, Simantheniras 
sp. ch FES. ill, 

REMPRENCES: None: 


20). PosstM CAVE 5U81 

LOCATION: Private lund. 

SITs DESCRIPTIONS Plentiful bone material apparent 
willitiat large sediment cone and on the sediment 
surface, particularly the distal fan and beneath rock 
ledges. Thus fin ooly preliminary jovestigations 
have heen made. wilh a very small umount of 
material collected for identifieation. The site is under 
further Investigation by the aulltors. 

COLLECTION: Flinders Universily vertebrate palac- 
ontology collection. 

LAUINAL 

MAMMALS 

Mieropodidae: Macropus spo el ME fuliginesus, 
'"Prarennodon Drefis. }Sumastienurus brawner. 
REPERENCHS? Prideuuk (1999); Plinders University 
vertehrale pulaeontology collection ditabase, 


21. CaBLh CAVE 5SU125 

LOCATION? Private fund 

SUPE: DESCRIPTIONS The cave was discovered by 
workers laying cables in JO81, hence the name. A 
small solution pipe entrance leads lo a steep talus 
cone, the distal portions of which contain abundant 
bone material within the sediment. Sone preliminary 
collection and identifications have been made in 
disturbed areas, 

COLLECTION, Flinders University vertebrate palaec- 
ontology colleeuon, 

PAUNA: 

MAMMALS 

Phylacinidae: “*Thiyvlacinus evaece pleats, 
Peramelidae: (saodon obesulus. “Peraneles sp. 
indet.. Vombutdae: Vormberiy trsinus, Thy la- 
colemmdae: *F7iwaralea carnfex. Potoroidae: 
*Retongia  lesueur, “Bettongia sp. indet.: 
Miacropodidae: Maecrapis sp. cl MM. eivaateny, eM, 
vidi, Macropis sp. indet.,  tSfiosthetiurus 3p, 
indet. 

REFERENCES; Flinders University vertebrite pulac- 
ontology voflection datibiase. 


22. SOS Cavir SULR2 
LOCATION: Private lund 
SITE DESCRIPTION: The entrance to thiscave opened 
Up naturally in YSS. Some bone pater fas beer 


SHETOS OME (LYYST SOS CUE ESL IDI) Che Layleriin Gireap 
mt Sauk Maiti News, a. is-55 


collected by members of CRGSA und taken to the 
South Australian Muscum for identification, ‘This 
matefhal inchided an almost complete skeleton of 
Thylacinus eyneeephales and the holotype of the 
extinet Wallaby Congriuis comers. 

COLLECTION: South Australi Musetun patie- 
ontology colleetion (vertebrate fossils), 

FAUNA? 

MAMMALS 

Thylicinidae: EMP ylereinus evnecephalts: 
Macropodidie: Congruus congruns, © Simosthen- 
Hrs Hewiande, 

REPERENCES! McNamara (1994); Sefton (198%); 
Prideuus (19999) 2000); South Australian Museum 
palacontology collection records: 


23. Buekribor Cave SL) 69 

LOCATION: Private land, 

SITE DESCRIPTIONS Small cave uncovered duno 
vineyard preparauon in early 1999 and subsequently 
filled in within 72 hours of its discevery, The 
authors and colleagues were contacted by members 
ol CEGSA to jovestigate (he sile which was found to 
contain significant fossil material A salvage 
excavalion wis undertaken during the night to 
prevent the complete loss of the material and 
information. AIL obvious bone material and most 
sediment were removed fron an approximately +n’ 
area tod depth of approximately 50 em. No. other 
material was visible. Preliminary taphonomiv 
analysis Suggests that this small cave muy have 
acted as a den for carnivores. colably Tasmanian 
Devils and Thivlaceleo carnifex, Bone material from 
the site 1s currently under mvestigation by the 
authors. 

COLLECTION: Currently held by the authors, 

FAL NA? 

REPTILES 

Elpida: Norechiy scalars, 

BIRDS 

Cusuaridaes Deromaiuy noveehollindiae, Mega- 
podiidac: ef "+Progura naruceortenyty. 

MAMMALS 

Tachyslossidue: oy Megalibuwitia raniwvevis Vhyla 
enidie: |" Tivlaciny eviacepheatis, Dasyuridie: 
*Dasvirns maciatis, ©. viverrinus, Sareaphilns 
gp. ch NS latedetrvityy Peramelidaes Lyooedan 
obestlus, Perameles sp, ch HP, suamnil, Palorchestidacy 
“F Pularchestes usael, Vormbatidae: “Lastarhinus sp. 
indel. Thylucaleonidac. “'ivlecelea carntfer: 
Potoroidue: ~Berengia gaomard!. &B. lesueur. *B. 
penicillin: Macropodidie: Meeropus sp. et M 
filiginosus, Macrapusy sp. cl Me piganreis. *M 
wrevi, M rufouriseus, OM. fii, Macropis. spy 
inde. Protentitiodan sp. och PHP, anek. Sine 
Vihenuris sp. indel. “TSihernuris anderen, 
ivlogale billardierii, Wallubia bicolor: Muridas 


n2 LL REED des. J. BOURNE 


Notamys mitchellii, “Pseudenys australis, OP. 
vouldii, Po shortridgei, Pyeudantwy sp. inde, 
KEPERENCES: None 


24. CRAWFORD’ S CORNUCOPIA CAVE SUTTI 
LOCATION: Private land. 

STTE DESCRIPTION? Sinull cave recently uncovered 
during vineyard preparation in mid- 1999; 
subsequently opened up by machinery, A small 
sediment cone contains very Fragile bone miterial 
will) a lower, eemented layer with numerous cranial 
and. post-eranial elements, seme i articulated and 
associitled states. Investigation of the site hy the 
vuthors hus begun, 

COLLECTION: Currently held by the authors, 
FAUNA? 

REPTILES 

Chelidaes gen, et sp. indet, Elapidae: gen. et sp. 
indet 

Birps 

Cisuartidue: Dramains novdehallaundiae: 
Megapodiidae: Progura sp. cf STP naraceortensts. 
MAMMALS 

Thylacinidaes **Thvlacinus cynecephalus; Daisy 
uvidite: Sereephilis sp, cf FS. laniarius, Pera- 
melidae: Aooden obesulus, *Perameles sp. indet.. 
Phascolarctidae: Phaseolarctas cinereus; Vomba- 
tidus: Vemfatny  ursinuys — Thylacoleonidac; 
“+7hivlaeoleo carnifex: Potoroidae: *Bellongia 
levuewe, Mueropodidae: Meeropus sp. ck MM, 
vicaitens, Mo rifoxriveny, &+Protemnodan hrehius, 
*FSimosthenurus baileyi, eS. hirawynes, 24S. willl. 
eS maddeacks, 78. eecidentalix, Wallabia bicoler, 
Muridite: ven, et sp. indet. 

REFERENCES: G. Pridcaus (pers. comm, 1999), 


25, CHhESH AND Purry Cavs 5076 

LOCATION: Private land, 

SUE DESCRIPTION: Material collected from the cave 
in 1967 by G. Langeluddecke. and lodged with the 
South Australian Maseum; context unknown, 
COLLECTION. South) Australian Museum palae- 
ontology colleetion (vertebrate fossils). 
FAUNAt 

MAMMALS 

Maeropodidac: Mucropus sp. 
WP Simasthentrus sp. ineler 
REFERENCES: Sowh Australian Museum patie- 
ontology collection recorils. 


indet.: 


26, COMAUM FOREST CAVE SUT (also known 
as Coma Quarry Cave) 

LOCATION: Comaum Borest, Forestry SA, 

SIP DESCRIPTION; Bone materi! excavated Tron 
the cave by the South Australian Museum in {he 
durkly LOBOS. 
COLLECTION: Mutscun 


Sout Austen 


palucontology colleetion (vertebrate fossils) 

FAUNA 

Reptiles 

Chelidae: ven. ut sp. indet. Seineidie. fliyaa 
PUBOrsel, 

BikDS 

Family indet. 

MAMMALS 

Tachyglossidae: + Megalibgwilia PISA, 
Tachyglossus aculeanse: Thylacinidae: © Thylacinns 
cynecephalus, Dasyuridae: Antecluitis sp. indet,, 
*Dasyurus maculatus, “DPD. viverrinuy, Ft Sarcoplilas 
faniurins, Peramelidae: “Percemeles — guanil 
Phascolarctidae: Phieweolarcios, cinereus. Vomba- 
tidae: Wonbetns isos, “tWarenedia wakefieldi: 
Thylacoleoniduer 'YThvlaceleo earnifes: Poto- 
roidae: *Bevongia gaunardi, *Potoraus iridacivlias, 
Macropodidae: Macrapuy sigameny, *°M. grey M. 
rifaurixens, °F Protemhodent sp. indet., *'Simesthen- 
mruy brane. “FS. will, ES. meuldacki, eS. 
newnae, “TS. oevidentalis, *7S, pales, | Sthenurus 
anderson. Thylogate billardierii. Wallabia bicolor: 
Burrannyidae: Cerearretiy nanis: Pseudocheimulite: 
Pseudocheirus peregriius, Muridae: gen. et sp 
indel, 

REP BRENCHS: 
Austriliitn 
rewards. 


Flannery & Pledge (1987); South 
Museum  palueontology collection 


Sites and Faunas of the Lower South East 
region 


Penola district 


27. PENOLA 

LOCATION: 22 kin NNW af Penola. 

SITE DESCRIPTION: Bones were discovered duriny 
the sinking of a well ou the edge of a swamp in the 
mid-nineteenth century. 

COLLECTION: Whereabouts of material unknown, 
FAUNA? 

BIRDS 

Droniornithidae: “+Genyanis sp. inde. 
REFRRENCES: Woods (1866); Stirling & Zietz (1890. 
L900); Rich (1979), Williams (1lO8O), Wells & 
Pledge (1083); Baird er al (M991). 


28. MONBULLA CAVE 5SL5 

LOCATION: Monbulla area, west of Penola, 

SITE DESCRIPHON? Bone material was collected in 
1975 unl 1992 by cavers, from a low passage in the 
entrance chamber of the cuve on the surfiee of the 
cave floor, The presence of Simosthemirin browiel 
material from the eave indicates some Pleistocene 
maternal. 

COLLECTION; South Australian Museum palace 
ontology collection (vertebrate fossils) 


PLEISTOCENE POSSIL SITES OF THI SOL TH ILAS'T ns 


PAUNA: 

REPTILES 

Scincidac; (iiqna rugaya. 

MAMMALS 

Vormbatidae: Vonibates resins: Macropodidue: 
“tSimasthenurny browne: Peliduey Feliv vatus: 
Bovidae; Ovis aries, Muriue: ’Conluruy albipes. 
KEPERENCES: South Australian Museum pialae- 
ontology collection records. 


29. UN-NAMED CAVE S1.122 

LOCATION: Newr Penola. 

SITE DESCRIFTION? Bone tmateriul was collected by 
bh W, Astin, and presented to the South Australian 
Museum in October 1970, 

COLLECTION: South Australian Museum pale: 
ontology collection (vertebrate fossils). 

PAUNAL 

MAMMALS 

Mieropodidae: Simoytienuras sp. cf. 8. gilli, 
REFERENCES! JJ. MeNamart (pers. cont, 1999); 
Soutle Austrian Museum pulacontolagy collection 
records, 


Millicent district 


30. Mv BURR CAVE 51.69, 5L70 

LOCATION: Mi Burr Forest, Forestry SA. 

SUL DESCRIPTION: Bone material collected by 
vavers duting exploration; conmleat unknown, 
COLLECTION? South Australian Museum pulae- 
ontology collection (vertebrate fossils). 

PAUNAS 
MAMMALS 
Macropodidae: 
entalts. 
REFERENCES? Walliams (1980): South Australian 
Museum palacontology collection reeords. 


Ee Sunosthemirus gilli, OFS. veeide 


3], UN-NAMED SITE NEAK MILLICENT 

LOCATION: Unknown, 

SITE DESCRIPTION: Fossils found ta peat matrix att 
a depth of approximately 2 mo suggesting un aneient 
swamp aceumulition, “Phe tod was reported in 
Waterhouse (1882). 

COLLECTION: Whereuhouts of material tinknown. 
bAUNAT 
MAMMALS 

Di protodontdie; 
retinas: Eritedies 
REPERENCLS: Waterhouse (]&82> Williams (1980), 


PZ yen 


SF Diprolodon sp. imlet.. 


32. LINN AMEDD SIRE 

LOCATION. Prive lind, Millicent ares, 

SEM DESCKIVLION: Bollowii excavation ab a mew 
dai ta Mareh 2000 the Lindowoer collected hones 
Fron ple of sediment discarded during bullloziny 


The landowner brought the bone miterial ta the 
attention of the authors whe identified some of the 
elements as belonging to fiegafaunal species and 
others lo macropodiids, sheep and pigs. In order to 
determine the stratigraphie pesition of the 
Memifadnal elements, and the extent of the deposit. 
the authors partially drained the dat, whieh had 
heen filled, and searched for more bone material, 
Bones of diprotodontids, sthenuring kangaroos and 
other macropodids were collveted from a thick, bhick 
organic mud matrix ata depth of approximately 1.5 
m below the land surface. No introduced species 
were found at this level, therefore their presence in 
the inatermal collected by the landowner from the 
discarded sediment suggests mixing of material 
during excavation und dumping of sediment. The site 
represents 4 swamp accumulition and is currently 
under further investigation by I, Wells, the authors, 
und colleagues trom Flinders University. 
COLLECTION: Flinders University vertebrate pul 
ontology collection, 


FAUNA 

MAMMALS 

Tachyglossidae: laelivglossis ceuleats Diprow 
dontidae: *tDipratadan ausiralix, PAY Roman ys 


rrilobus: Macrope wlidhies, "eMC ropuy 
Macrapuy sp. andel., Sthenuruxs sp. cl 4 
andersant, Suidae: Sus sero: Bovidue: Ovis aries. 
REKERENCLS! R. Wells (pers. comin, 2000). 


Mount Gambier distvict 


33. CHENCOE 

LOCATION: 22 kin NW of Mt Gambier. 

SITE DESCRIPTION: The preservation of the fossils 
(he. white bone with red sediment adhering). is 
sugeestive ol a cuve deposit. passibly Gleneoe West 
Cave (5177) or Glencoe Lust Cave (51.108). Purther 
informaviun is unavailiible, 

COLLAETION: South Australian Museum palae- 
ontology collection (verlebrate fossils). 

PAUNAS 

MAMMALS 

Diprotodontidae: gen. ct sp. Jadet. Macropodidae: 
Macropus sp. inden, *eSimesthenurtiy gilli, ers. 
occidentalis. 

REFERENCES! Tindale (1933): Williams (1980), 
South Austhalint Museum puluconlology collection 
records. 


4h. TANTANOOLA CAVE 3LI2 

LOCATON, Near Tantunauha. 

SITE DESC RIPTLON: Bone tier tis heen colleeted 
from at sediment-lMoored tunnel und iv breecia in the 
CHEPeNE fouiPiSE Cave. Benelt sediments und sea-shells 
partially filled the cave CN. Plealee peeps, comm 
2000), 


M4 IH REED & S. 1, BOURNE 


COLLECTIONS South Australian Museum palace 
ontolowy eollection (vertebrate Fossils). 

PAUNAL 

MAMMALS 

Dusyuridae: "Dasyurus spoindel., “Sercoplitiy sp, 
indet., Peramelukie:  Jyoador sp. inde. 
Diprotodantidsie: “FAvgemenirny Wilabus, Yomba- 
lidac: Vorrbarus arsfaas: Pholunveridae: Triehaxurns: 
vidpecila, Muacropodidae: *FPratemnedon roechtus, 
“+ Sinosthennrus pili, O48. oecidentaliv; Olaridie: 
Armuephalay sp. indets. Moridues Aydraniys sp. 
inden. Retys sp. Inder. 

REFERENCES: Tindale (1935); Williams (1980): 
South Australian Museum palaeontology collection 
records. 


35, TINDALE Ss Cave “EY SEIS 

LOCATION? Taintanoola. 

SITE DPSCRIFTION: Bone material wis collected by 
cavers and presented to the Sourh Australian 
Museu, 

POLLECTION: Soudh Australian Muscum pulue- 
omology collection (verlebriile fossils), 

LATINA: 

MAMMALS 

Macropodidae, “hSiiosteninns gilli. 
REVIRENCTS: Tindale (1933), 0 MeNamura (pers, 
comm. 1909). South Australian Museum 
pulaeontohiey calleetian reeurds, 


36, Morcians Cayk S5L34 

LOCATION: ‘Tunlanoola. 

SETH DESCRIPTION: Material was collected in 1958 
by B. Daily: context unknown, 

COLLECTION: South Australian Museum patie: 
ontology collection (vertebrate fossils), 

AUNA: 

MAMMALS 
Mawapodidie: 
SSiimostheaniilus ga. 
ROPERENCES: South Austrahan Muscum patie- 
ontelowy collection records, 


Macropus rufogrivets, 


97. GREEN WATERNOLE CAVE SLATE Gilso Known 
iby Fossil Cave) 

LOCATION: 22 km NW of MU Gumbier. 

SITH DESCRIPTION! A water-filled cave. with fossils 
discavered by divers on the surface ol a rockpile. ut 
wadepth of (5 0. Collections were iude by divers 
wluring the piel to late M96Os and 1970s and taken to 
ihe South Austratian Museant and Australian 
Museum, Exjensive collecting trips were organised 
by ROT Wells i 1479. Tr bus been suggested Tat the 
probuble aecumulation mode was drowning wl 
animals that (ell inte the cave, trying to use Tas a 
diioking water source (Pledge 19S8Qu: Rewsin 
INES), 


COLLECTION: ONustralian Muscuin palacontology 
collection: Flinders University ‘vertebrate palite 
ontology collections South Australian Museum 
palacontology collection (vertehrate Fossils). 
PAUNAS 

BIRDS 

Phastanidae: Conanix sp. tndets Aceipitridives 
Undeseribed taxon: Paleonidae: Palen sp, ef be 
berivara: Raliidae @Gallimlla mortierti, Gallinullet 
sp. ch G. fevehroasa, Turnicidae: Tanke varia; 
Burhinidue: Burhinus sp. ch Be gralfariins, 
Columbidue: Phaps chaleuplera. Paps sp. inde; 
Cucathidae: Ceca teniirayeris. Callacepletan 
Jimbriatiun, *Calyproriwnehus banksil, Co batho, 
Culyprovlivnciiuy sp. indel., Poitiers: Plaiyoerens 
sp. indet.; Cuculidae: *?Centropus colossus, 
Strigidae: Nines novdeveelandiae Aleedinidae: 
Develo pavaegiiinedae: Acunthizidie: Dayyarniy 
broudbenti, Meliphasidae: Manoring imeldiia- 
cephulas Oribonychidae: *FOrihonye hiypsilophus: 
Corvidue: Corvus sp. indets Pirondinidae: gen, eb sp, 
indet. 

MAMMALS 

Thylacinidie: *7Thylacinis evnocephalins Dasy- 
unidace: *Dasyeas maculains, *Sarcophilay xp, 
indet., Perumelidue: Jseedon ehesitliss Phaseo- 
lurctidae: Phasealaiities cinereus: Vombatidie: 
Vombaluy urstuis, Thylicoleonidae: 9 /ilacalen 
carnifex: Phalangeridae: Trichosuray vulpeculay 
Hypsiprymnodontidae: Ff Propleopus aseithins; 
Potoroidae: *Betongia penieillata, Pororous sp. ol, 
“P, midactylus, Macropodidue: Macrapus sp. cl. Me. 
wiganteus, 77M wrevi, M. rufouriveus, “TAR tina, 
Macropus sp. indel,, Protemnoidan spol. 84 P. anak, 
“+Protenmnedean sp. indet. #1 Simasthenurus gilt, 
ATS. daddocki, SUS. newionee. OPS. cee telentelis, 
Wallahie bicolor, Chiroptera: lamily indet; Suidac; 
Sy scrofa, Bovidaes Oviy aries; Muridue: gen, et sp, 
indet, 

Newlin (198K) dlisted Metcrupus rufus for the 
deposit, However, no specimen cin be located by the 
iuithors to suppert this identification, which appears 
unlikely. "This species has therefore been omitwd 
from the fist, Newton (198S8%) also listed * 7 Siennris 
stirtinet. whieh was later found to be a 
misidentification (G. Pridewus pers, comm, 2000), 
REVERENCHS: Wells & Murray (1979); Pledee 
(19800); Willams (M980): Baird (198Si; Newlou 
(M9887); Baird (M9la. bis Baie er af (LOUt 
Pridegux (1999% 2000). Flinders Universiry 
pakiwontology collection dulubase; South Austeatiut 
Muscuin palacontology cofleetion records. 


38. WANDILO FOREST CAVE SLA05 

LOCATION: Mount Gambier Forest, Foresiry SA. 
StH DESC RIPHON: This small cave was discovered 
in (997 by members of CRGSA. Numernts bones 


PLEISTOCENE POSSIL SITES Ob THE SOUTH EAST a3 


were obvious on the surface of the lloor sediment, 


und within the sediment eone, A small number of 


bones was collected by CEGSA und taken to the 
South Australian Museum for identification. The 
authors and M,C. McDowell vistled the site with 
CRGSA in August [998 whee further 
identifications of some fossil material were nace 
NLA, 

Collection: South Ausuiian Musetim palacontology 
collechon (verlebrale fossils), 

bALNAT 

MAMMAUS 

Phascolaretidic: Phrisealeive tas CTNETEUSY 
Diprotodontidie: ef VZyaenndariy (rilobass Yorba 
lidae: Vommbantes vesmiuss Thylacoleonidae: #4 Thyla- 
coled cartifes: Potoroidae: *Patorouy wideerylis; 
Macropodidae: Maeropus elgcntens, MA. rufauriseus. 
“1 Procoplodon sp. indet. *Simesthenurus browned, 
REPERENCKS: Reed (19982 South Australian 
Muscum palacontulogy collection records. 


39, WANDILO CAVE SL74. 

LOCATIONS Mount Gambier Forest, boresiry SA, 
SITE DESCRIPTION: Bone material was collected by 
cuvers dure exploration in 1992 and taken to thie 
South Australian Museu. 

COLLECTION; Soudh Astra Museum 
pulacontolagy collection (vertebrate fossils), 
TAUNAT 

MAMMALS 

Vombatidac: Vainbatus visiiis: Phalangeridae: Triche- 
yuri vnlpeciulen, Macropodidiie: Maeropas sp. cb ve 
vinantens, Macrapus sp eh ML rifoyrisens, “VSinias- 
Menuray meddockt, “FS. occidentalis. 

South Australian Musenimn palicontology, collection 
rewards, 


40, Moorark 

LOCATION! 3 kat south of Mount Ganthier. 

SITE DESCRIPTION: Probable cave deposit, comext 
UNKNOWN, 

COLLECTION: South Australia Maseum paithae- 
ontology collection (vertebrate Mossils), 

bAUNAS 

MAMMADS 

Thylteoleonidae: th Tivlacelea camifex. Miacro- 
podidde: Macrapay vivannens, Macmapuy sp inde, 
+e Somosthenurus pales. 

REPERENCHS: Pledge (1977), Williams (1980); Sauth 
Australian Museum palicontology collection records, 


41. KinsbyS Hou: 5L46 
LOOANON! 3 kin south west oF Mount Gambier, 


1 Ron, EH, CHK) A Spressine” engoyeeny with: some-surruusty 
tomy Trssils ie SLAG, Wandile: Cave Laydeeerten Coreup of South 
Vaneruilee New 43, 20 10M, 


SIM DESCRIPTION: Bone material was found in the 
Hill froma small solution tube exposed by excavalion 
i TYSS OF a ralop to the water level in the sinkhole 
COLLECTION, South Australian Museum pide 
ontology vollection (vertebrate lossils). 

PAUIA? 

MAMMALS 

Tachyglossidie: Megulfhgyrifia sp. eh SM, 
ramayl, Thylucinidaes ©" Thvlaeciius cvaocephulus: 
Dasyuridaes "Dasyeeriy matculatia, Sarceplilis sp 
indet.; Perumelidae: Jyoedon ebesulas, *Perameler 
boreemille, “Po wana, Phaseolarctidae: Phaycel- 
arctos -cinerens, Diprotodontidie: + 4ygerelirny 
Irilabus: — Vormbaticae: Vener erydines: 
Thylacoleonidae: “TThylacelea carnifex: Phatan- 
weridae: Srichayurus vialpecuta: Potoroidac: *Beite- 
ngia lesueiur *Potorints triductyiis:, Macropadidae: 
“thagerchestes leparudes. Macrapus sp. indet.. 
=7Prvtenmadan sp. indet. © hSimevthenurny silli. 
a8 newtonde, *'S. acckdentalix, = !S. pales, 
=F Sthenuruy anderyent Burruinyidae; Cercarieluy 
ap ch C. pans; Pseudecheiridae: Pyendocherrits 
nere-grinus: Chiroptera: Tnily indet.: Cuntdae: 
Canis Hipus familiaris, Bovidaes Owls aries: 
Muridae: ’Canitiras allipes, Mas taconis (uses, 
Ralius sp, indet; Leporidie: Orvelolagis cuniculi. 
REFERENCES: MeNamara (1997): South Australian 
Muscuin pulaeontology collection records 


2. Simpson's Hone SL42 false known as Jen- 
eighty Sinkhole) 

LOCATION: Near Mount Gambier. 
AME DESCRIPTIONS Bones discovered by divers i 
the flooded section of the cuve. 
COLLECTION:  Plinders University 
palacontology collection. 

FAUNA? 

MAMMALS 

Diprotodontidae: *+Diprotodon sp. todets Macro 
podidae; Mierapuy sp. indet. ?) Prateanieden 
reechus: 

REFERENCES: Flinders Universivy ptieontalaps 
collection dutubase, 


verlebrile 


43..G0UL DENS HO 518 (also Known us Gouldens 
Hole Cave) 

LOCATION: Several Kilometres west of Mi Sehank, 
SITE DESCRIPTION: A sniull tunnel on SSE side ol 
the cenote (Gowldens Hole), was uncovered by a 
Korner digging an ueeess rump ta the water, The 
tunnel floor was covered with sill containing fossil 
materia! ded hones of modern vertebrates, The site 
was oxcavaled by researehers from the Soult 
Australian Maseurn in 1982, Pledwe (1991) suaeests 
(hat the bones were probably derived from a filled 
entrance firtber up-slepe in the tunnel ane reached 
ihe lower extremiry ef the tunel by winer 


sO, Eth REED & S41. BOURNE 


winnowing. He deseribes it as a “reworked, mixed 
assemblage” (Pledge 1991), 

COLLFETION: South Australian Museum palace 
ontolagy collecuon (vertebrate fossils). 

FAUNA! 

REPTILES 

Vamily indet, 

Bins 

Phulacrocorameidae: Phalaeracorax melanoleucas: 
Accipitvidae, Aguile uuday. 

MAMMALS 

Vachylossidue: Mevalibgwilia sp. cb. “tM. resent. 
Tachyelossas acileatis: Thylacinidae: + Tiyvlacinus 
cvnecephaliys Dasyuridae: “Deasvaris rereidlattes. 
'Dasyurus sp. indet, Sarcoplilus sp. indet.: 
Peramelidac: dsacdon sp. ef 7. abesulius, Peraneles 
sp. ef "RL gumity Phascolarctidae: Phescolaretas 
cinereus: Palorehestidde: Palarchestes spo ct PtP. 
parvin: Vombatidiae: Vembeluy ursinas, Phyl 
voleonidae: *)Thylucolea carnifexy; Phalangeridae: 
Trichosurus vulpecula: Potaroidaes *Berongia 
gatmardi, “Peteraus tridactvlus, Mucropodidae: 
Macropus givanieus, Me rufauriseus, ML titan, 
Macropay sp. mde. *tPratenmodon brehus, &*P. 
raecluis, *YSimasthemirus gilli, *78. maddacki, #78. 
newtonae, “AS, occidentaliy, ?Sthenurus andersani, 
Wallubia bicolar, Pseudocheiridae: Psendocheirus 
peregrinus, Candace; Cuniy lupus familiaris, Vulpes 
vulpes; Felidue: Feliy cats, Bovidae: Quix arivs, 
Muridae: "Mesrconivy fuscus, gen. et sp, indet, 
REFERENCES: Pledge (1991); Baird (199Tb), Baird 
ered, (1991), South Australian Museum palic- 
ontology collection records, 


44. TAN KSTAND CAVE SLGO5 

LOCATION; 3 kin west of Mt Schank, 

8b DESCRIPTION: Bone material was collected 
from the drowued part of cave, situation unknown, 
COLLECTION, South Australian Museum pulae- 
ontolowy eollection (vertebrate lossils), 

PAUNAT 

MLAMMALS 

Macropodidae: bi Simesthenucuy gilli, 
KEFORENCOS: Williains (lO80); South Australian 
Musciun pulaeontology collection records. 


Moun Ganbier Township 

45. LIN-NAMED CAVE 

LOOAHION! Derrington Street, Mount Gambier 
{howy) 

YIPE DESCRIPTION: Cave exposed by earthworks for 
wsewer trench in 196A, 

COLLNOTION: Soul Australia Museuin 
palweonlology eollection (vertebrile Tossils), 

PUN AT 

NIAM MALS 

Peruimelidie: "Perdoiele ¥ sp. inlets Phascaliretidie: 


Phaseolarctos sp. indet.. Diprotodontidaes '*+ 
Zvvomaturuy tilobus (= Netotheriium of Willkanis 
1980); Thylacoleonidue: *?7hyvlacelea carnifen; 
Potoroidue: “Berongia sp. iidet: Macropedidie: 
“+ Simosthenurus browne, &+8. alli, *78, meeleloeki, 
“+ Sphenuus anderson Pseudocheiridae: Hsetdo- 
cheirus peregrinny, 

REFERENCES: Pledge (1977): Williams (1980). South 
Australian Museum palacontology collection records. 


46, UN-NAMED CAVE 

LOCATION: Mount Gambier Clown): other details 
unknown. 

STTR DESCRIPHON: Unknown, 

COLLECTION: Natural (History Museam (London), 
TAUNA: 

Binns 

Dromornithidae, *'Genyorniy sp. indet. 
REFERENChS: Stirling and Ziets (1896. 1900K Rich 
(1979): Williams (1980); Baird ef al (1991). 


47. UN- NAMED CAVE 

LOCATION: Grey Street, Mount Gambier (lowe). 
SITE DESCRIPTION? Cave exposed by exciuvalion. 
COLLECTION: South Australian Museum) palae- 
ontology collection (vertebrate fossils). 

FAUNA 

MAMMALS 

REFERENCES: Williams (1980), South Australian 
Museum palacontolowy collecuon records. 


48. ExGkharscur Cave Sito 
LOCATION: Jubilee Highway, 
(town), 

SITE DESCRIPTION! Bones have been recovered by 
divers in the flooded section of the current tourist 
cuye. The bone mitterial wis identified py pile 
ontologists from Flinders Universily, 

COLLECTION: Fngelbreeht Cave manugement, 
FAUNA 

MAMMALS 

Vombatidac: Vewnbertiy ievinis. Maecropudidae: 
Macropus sp. cf MM, eigaiitens, Proteninodon sp, ot 
"YP. brehuy, 

REFERENCES, None, 


Mount Gambier 


49. Tih BLué LAKE 

LOCATION: Mount Gambier. 

SITE DESCRIPTION: Bone material has been found in 
A solution cavity dude excavation of aw qecess 
tunnel aba depth oF bernween +f rand 42 m fron the 
lower pump slalion entrance, 

COLLECTION, South Australian Museuny palie- 
onrotogy wollection (vertebrate fossils). 

HAUNAS 

MaMMALS 


PLEISTOCENE FOSSIL STTES OF THE SOUTH EAdt nY 


Micropodidites |) Simastenurny sp. met. 
REPERENCRS:; Soul Australian Museunr pualie- 
oitlolugy collection records. 


Discussion 


As the date presented ubove clearly Jemonsuute, 
the South Hast of South Australia holds wsignificunt 
yocord of Pleistocene sites add) vertebrate: fossil 
launas, The numerous cave systems and other sites 
the region have uecumuhued vertebrate remains over 
an extended penoiu of dime und with increased 
puliwontolagical researeh i) the region more sites are 
being discovered, Recent improvements io 
veochronolovical techniques have enabled researchers 
fo Concentrate on developitg chronulagies for 
several of the Sites in the remo, particularly those 
willing the Naracoorte Cuyes World Hentage Area 
(Ayitiie & Veeh JOSS: Ayliffe ef af. 1998: Moriarty 
er al. 20001, Current laphonomie research being 
varhed Gut on various deposits within the World 
Herilawe Area vind its suyratinds is allowing ts 10 
piece lovelher (he accumulation history ol many of 
these deposits and to determing their repre- 
semlutveness aud osailability far use ta 
paldcoctalawical reconstructions, Onby wath a 
thoroueh knowledge af the faunas, taphoneny, 
eenlogy und chronglogies of these sites can valid 
palueoecological analyses be made 

The distdbutions of taxa between the sites are 
summarised’ in Table 3. The ditt reveal some 
Interesting pattems the faunas represented aad (he 
level of seiendlie attention that they have reveiverh 
There las been yery Title research done an the 
amphibian tossils from the region since Tyler (1977. 
VOL) warked on material from cave sites at 
Nanieoorte, Pere is pow more Material available 
anda review of this group could reveal more speeies- 
Ascallutthe species listed by Tyler (1977 L991) are 
Still living in (he region today, the frog assemblages 
could be very tselul ty pakwenecolowical 
reconstructions. The lossil reptile luunas hive also 
reecived litle attention sinee the 1970s. with the 
esveprion of the work of Banie (1991), Willmuns 
(990 and Selon & Lee (2000). The varanids. 
wsanics aid) claprls all reqaire further research. 

The fossil bird faunas of the region have received 
Some allention (Van Tebs l974: Van Tets & Smith 
IPG Rich [O79 Bail (985, 199Ta, by Batra er uf 
IYO M, MeDrwell pers. como, 2000) but farther 
Hivesnaunon of materil recovered in recent years, 
particuliuly Trop the Naraeoerte Caves Wark 


SOWIE ELAN © CLUS Tabet | LOAN a Ee eS h Pht) AX ahasyt 
Ory AP Oe Petey pais, WS) CMS) Phese) Phe EE versity 
OW Adhedisitle Garqealed, 


Herituge Area, is required as it may reveal more 
species: Buird (1985S. 19914) did extensive work on 
avian taphonamy and desenbed different modes of 
accumiulauion for bird reniams. A compirtson 
between sites, purticulady the Main Fossil Chamber 
in the Victoria Fossil Cuve (site 4) und Green 
Wauterbole Cave (site 37) which have the hugest 
fossil bird assemblages from (he region. reveals quite 
different species conipositions. probably related to 
different accumulation modes. Further research on 
the bind faunas muy reveal sane other interes 
luphonomic biases. 

Some research hus been eayried GUL-on the fossil 
small mamol Giunas ob the region (Smith lY7y, 
1972) M. McDowell pers, camm. 1999) 2000: A. 
Buynes pers. comm, 2000). Recent work in Wet 
Cave (site A) und Robertson Cave (site 10) (M, 
MeDowell pers. comm, 2000) reveuls ussemblages 
composed primarily of sniall mammals (see Table 3}, 
perhaps dermved mainly front owl pellets cather tran 
wopilfall trap, which Has been suggested as the muri 
mode of wenmulition in many other clive sites, 

Reinvestigalion ob some of the tossil small 
mammal material may be required to confirm seme 
cho heations, One example is tie fossil Anwesinaes: 
mitertal, Pour spevies of Amechiney have beer 
Wentified in Pleistocene faunas from the South Bast 
(Table 3); of these species, Al flavipes and A, 
minimis ace still Tivinge in the region (Strahan 190s; 
Robiasan ef a. 2000). Twa species, Fa. viene (seu 
Systemilics section) and “AL swernsenti, are not 
found in the South East todwy (Strahan 195, 
Robinsun et al. 2000), and are only listed for one 
Pleistocene site (site -ha) i the region, as is vA. 
miniuidy (site 2). These four species may hive been 
part Of a more diverse fiund during the Pleistvenc 
of allernarively, onc or mare of (hese may represent 
misWentificauon. The fossil Ardee/inus mualerial 
trom. the South East needs further work to resolve 
such issues. Another group that has been very tittle 
studied ts (he fossi) bats, Phete ave Gurrently aroun 
Hl species tiving in the region (Robinson ay er, 
2000). oF whielr at east five ure known te. inhabit 
caves, veronly ovo species (ATAaprerns schretbersiy 
wd Nyewphilus veoffray) have se tar been 
identified from fossil deposits (see Table 3)..As owls 
are still active predatars of bats in the region and 
have been accumulators of soll mainymitd remains ty 
the past. amore intensive study of (he plentiful fossil 
small qoummal material (particulsrly Tron the 
Naracoorte Caves World Teritage Area) should 
reveal more bat species. 

The fossil large mammals (25 Ke live weight) have 
received More attention than other grdups in (he 
region (Daily b960Q, Mervilees 1965: Pledge 1977, 
(9808, 0 1990, P94, Murray D978, Wells & Murray 
1870, Wells ef af, 1984: Flannery & Pledge {9S7, 


th EO REED & $1 BOURNE 


Griffiths e¢ ud. POL: MeNamary 1904) Brows LOO8” 
Podesuxs & Welly 1995. Prideaux 999° 32000, 
‘Torner 1999°), The sites af the South East contin a 
rich record ol the extinct megafauoa, particularly the 
sthenuring kunguraos (Merrilees 1965; Wells & 
Murray 1979; Pledve 1980u. Prideaux & Wells (998, 
Pridequx 19999, 2000). The mucropodids are the 
dominant gronp will 2o-speeies represented; of (hese 
I@ beoame extine) during the Pleisfocene. three 
Witully extner following Burapean setdlement and 
three locally extines (Table 3) Currently there are 
only (ourspecies found i the pein (Robuiser eral, 
2000), The large diprotodontits ure sparsely 
represented in jhe deposits with 9? Ayewomatariys 
wilabus he most commonly found (recorded i 15 
sites). Palocchestids are particularly rare These may 
represent real abundances in the ancient faunas or 
taphovemie biases related) ta the modes oof 
uecumulation. GF the huge naman carnivores, 
*eTivideales earnifex, is well represented iw the 
region (reeorded frora 24) sitesi, A Cvlderins 
cyrneuplalas moderately well represented (19 sites) 
wnd the devils (Surcuphiies spp.) represented in 17 
sien (Table 3) 

Milterial representing species for which there 1s 
oily a sitle record from the region may: fequire 
curerul ome-investigalian lor ensure that the 
idenriticuuons ure conmect (sec the note regardiny 
“Dewees envatonde aad “Peranelesy tine Vor 
site hy). However, single specimens of EC renin 
coneruns UMeNamare P9Od) aid“? Warenredpe 
Wwekefehli (Flinnery & Pledge 1987), indicale such 
reconls cin be reliable MeNumara (1997) has alse 
Wighhehted the need for re-aivesigabon of some 
iutecial With his work on the small mueropadids of 


South Austria, Closer examination of (he wealth ob 


fossil material from the South East has the potenthil 
lor rosalvine ather issues such us the taxonomic 
stinis of the devils (Sarcaphilis harvivd and '8. 
felines), dnd the question of whether these 
refiesent distiner species or suh-species, or the 
extol species popresents a dwarted form of the 
lurger 778. lonieriny (Murshall & Corruccini 197%: 
Dawsin 19822 Werdelin 1987). Both have heen 
recorded. dni siles in he South East. Stmilivly, the 
distinction between the koalas Phescoheretas 
Pliers and FP stivtent Was heen the topic ot 
discussion (Archer & Mand (O87), ancl beth species 
are Tisted for Pleistocene sites of the South Eiest. 
Collection biises ean be reduced hy employing 
thorough eseavalion methods and reducing (he 
number of specinens simply collected from sites 
Without rewind to pravemimee. Examination of the 
itd Sumimutised in Table 3 reveals a great muniber ol 
sites with practically ne record ol amphibian. reptile 
Kind or SHmilh avimml specles tine this. yay be 
refuted fo collechon biises where janily hare bone 


milenal has been eolleated. Exceptions to this (ine 
sites where recovery of small specrmens is dil feult 
(ew. Green Waterhole Cave and swarnpy siles), 

Au interests feature of the South Mast is rhe 
yaition in the “types” oF Sites (eat. caves, Sinkliales 
und Aawenps) within the revion. AlLof the Pleistocene: 
siles in the Nuracoorte arew. and much of the Upper 
South Bitst. are cave sites. Swannp sites are found tn 
the Penola und Millreent ares. The Mount Garber 
aed Has cave sites, but alse has sinkholes which tire 
tinue to This area singh dre not found. tir the bopper 
Sout Bast. Obviously the wpe wad morphological 
features of a site Ive a direet Infldenve on. the 
species represented in the deposit und this highlights 
{he imporkunce of taphononiic research. 

Within the South Bast region there has heen a has 
iowards sites of the Upper South East. particularly 
the Siles around Naracoorte. This is probably due ta 
the mueh higher level el yeseareh ane ex plaratian 
conducted in this area, This, further tivestigation yl 
ihe Lower South Basi region is un important rest 
slep to understinding the vertebrate palicoitolagy ol 
the region as 2 whole, 


Acknowledgments 


The wuthors wish to thunk the relerees N. Pleuge 
und paruicularly, AL Baynes for helping greatly to 
Improve the paper, RB. Wells and G. Prideaus for 
commenting on an early drafl and for continued 
guicinee with the project, Speci thanks go co M, 
MeDowell for help wath excavation, site surveying 
dnd for allowing Us to use his extensive Speeies lists 
for Wet Cave (SUT0) wid Robertson Cave (SUL7 
SU 1K, 5019) and for identilying rodent rmiterial 
Hronrather sites at Nareoerte. We would partiva larly 
like to thank NL Pledge, Jo MeNumara and M 
Heotchinson of the South Australiin Museun foe 
assisti with information and providing aveess to 
the Museunrs Palaeontology collection, the siaff of 


ihe Naracoorte Cayes World Heritage area 
(purtivularly the Cave Guides) lor contiiied 
cnthusiasin anh assmninee with the research 


promramime at te Caves, G, Prideaux and 8. Brywir 
lor issishingce with sre surveying: aod identifention 
of yulerial L. Euston (previousty Turner) for 
invaluable help with the identifieating of 
mawrapodids Cron Naracoorte. Eo Bailey for help 
With sile and speaiinen relerenees eontaned ah the 
Mlinders University Vertebrite — Pahteontotiey 
collection und Po Duenke for assistinee with the 
Jocation OF specimens in this colleetion. Thanks jure 
due to W, Boles of vie Australian Muscunp and J, de 
P, Bourne of Bool Lagwon for assistinge with current 
bird names. We would also like to-thank members af 
OPGSA for assistance with (he locaron of sites ane 
material L. Dawsen. Cor fput on the mderial fron 


PLEISTOCENE FOSSIL SEPBS OF THE SOU TH EAS | au 


the site SU169 and helpers and students thom 
Minders University. Thanks to 5. Hayter for helping 
with excavations at Millicent, aud tothe following 
lindowners for providing ugeess fo caves und sites 
on heir propeny. for vikings an active mnrerest in 


palucontologieal rescureh in the region und fin 
reporting the location OF new sites — PL Bird. bh & S- 
Crauwlord. Po d& Ro Freckleton, J. Hole Ro & D. 
Hooper, Re Hunter & family, B, & B. lolisom Re & 
R. Menvel and DO & S, Williamson, 


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A MIDDLE PLEISTOCENE VERTEBRATE FOSSIL 
ASSEMBLAGE FROM CATHEDRAL CAVE, NARACOORTE, 
SOUTH AUSTRALIA 


By STEVEN P. BROWN*® & RODERICK T. WELLS* 


Summary 


Brown, S. P. & Wells, R. T. (2000). A Middle Pleistocene vertebrate fossil 
assemblage from Cathedral Cave, Naracoorte, South Australia. Trans. R. Soc. S. Aust. 
124(2), 91-104, 30 November, 2000. 

Located in the Naracoorte Caves Conservation Park, Cathedral Cave represents one of 
the more fossil-rich vertebrate sites within the region. An analysis of the geology and 
palaeontology of the fossil assemblage, coupled with U-series dating, has enabled a 
reconstruction of both the accumulation modes and the proximal environment 
between about 280,000 and 160,000 years ago, during the Middle Pleistocene. A 
pitfall trap is suggested as the primary mechanism for collecting animals whose 
remains became incorporated in the deposit. The fauna indicates an environment 
dominated by large herbivores inhabiting a grassy open forest or woodland with little 
suggestion of aridity. 

Key Words: Naracoorte, Cathedral Cave, fossil assemblage, U-series dating, 
taphonomy, Middle Pleistocene, pitfall trap. 


Tremascretions: af the Bevel Sect al Se Mest (200, LAG 2 OTN. 


A MIDDLE PLEISTOCENE VERTEBRATE FOSSIL ASSEMBLAGE FROM 
CATHEDRAL CAVE, NARACOORTE, SOUTH AUSTRALIA 


by StrvEN P. BROWN" & Robekick T. Wes’ 


Summary 


Brows. 8. Pod& White. Re T2000), A Middle Pleistocene vertebrate fossil assemblage fron Cathedral Cave, 
Numacoorne, Sourh Australia, Tua. & See, S. Aur 124 (2). 41-104, 30 November, 2000, 

Lavated in the Nareoorts Caves Conservation Park, Cathedmil Cave represents ne of the mare fossil-reh 
vertebrate Sites Within the region, An dhalysis of the geology and paleontology at the fossil ussemblige, 
coupled With U-sevies dating, fas enabled a reconstruction of bath rhe vectimubition modes and the proximal 
cuvironiment bebween about JRO.000 vad F60.000 yours aoe, during the Middle Pleistocene: A pilhall teapy is 
sumeested as (he primary mechanism lor collecting animals whose remains became incorporated in the deposit, 


The laure iidienies in environment dominated by large herbivores inhabiting 


Wilh Tite stigwestion af ariatity. 


2a gritssy ypen forest or woodland 


KEY Words: Naticnorte, Cathedral Cave, fossil assemblage. b-series dating. laphonemy. Middle Plerstovenc. 


milla (rap 


Introduction 


The Oligo-Miopeene limestone (Naracoorte 
Member of the Gambier Limestone) underlying 
much of (he Naracoorte region conltins an extensive 
system at caves well Known for their fossil content 
‘The best studicd ure the richly fossiliferous deposits 
in Victoria Possil Cave (ee, Smith l97l, 1972, 
1976; Van Tets d& Smith 1974, Welly 1975; Tyler 
1977: Wells eral JO84), However, Tiltle is known 
about the fossil content of other caves in the rexion, 
Ongoing geochronological und palueontologicul 
reseiurcht (Aylifle & Veelr loss: Ayliffe ev af, 199M: 
Moriarty ef ad 2000) has encompassed many tnew” 
fussiliferous cave sites, The Fossil Chamber i 
Cathedral Cave is one such site (Mig. 1). The present 
study presents a detailed geological, laphonomic 
wd Faunal analysis of the Cathedral Cave fossil 
deposit. 


Materials and Methods 

Geology: stratigraphy ane sedimentology 

Acsurvey datum wis established on at limestane 
block mn close proximity to the sediment cone tn (he 
Fossil Chaunber (Fig. 2). The sediment fil was 
systematically probed with a Tom long 10 mm 
diameter rod te locate subsurface limestone blocks, 
A 75 nin diameter soil aueer was then used fo 
simple the sediment tow depuh of 2m avoiding the 
buricd blocks. Auger holes were spaced ab Som 


Sebel oF Blohwwical Scenes Pliers Linecesily oF South 
Australian, GPO Box 2h Adehude SA SOUT fe rai 
steyel brown llinders eultiuu 


inferwals across the sediment (il, Each bore hole was 
sampled al [0 en inerementis and a subsurlace 
stritigraphy constructed, ALL depths ta datum were 
measured. A sample of sediment from each LO ci 
interval was phiced joto a cleat shap-loek plastic bie 
and Jubelled with the auger hole location and depth, 
Samples were returied to The Flinders University of 
South Australia and scored for sediment colour 
(Munsell colour charts), clay content, caletuiy 
curbonale content, grain size and sorting (following 
the procedures of Day (965), and grain morphology 
(McCollough sand eauge), 


Palucantolosy 

Access tothe Fossil Chamber was vin a dug crawl 
Wiy approximately 120 m in dength (Pig. 1). 
Excavation of fossililerous sediments was carried out 
over a three-month period to April L998, Two pits 
were excavated within the Fossil Chamber (Mig. 2), 
Vhe furst (Pit Ad measured 28m 8 1.5 140.76 mand 
the Second (Pit B) measured 2.6 mx 2) mx 1.04 my, 

Sediment was carefully excavated along the 
bedding planes using towels. dental pieks and 
brushes. Exposed fossils were lett i yd while the 
following dite were collected: the depth below, 
distance Lo. and direction from datuniy the dip and 
bearing of the specimens, A sighting compass, lipe 
measure and a tine level were used to accomplish 
this. Specimens were them remoyed front the 
sediments and trinsported ta the laborwtory where 
they were cleaned. and stabilised with a polyvinyl 
butyrate (Mowital’’, Hoechst). Due to the difficulty 
of removing sediment trom the eave th was apy 
screened on sile using 3 mm mesh sieves, 


92 S.P. BROWN & R.T. WELLS 

Taphonomic analysis classified as irregular perpendicular, crenulated, 
Each specimen was identified where possible to spiral and compression following Lyman (1994) and 

species or genus and element type. It was then = Marshall (1989). Characterisation of weathering 

examined for breakage pattern, presence of predator stages follows Behrensmeyer (1978). 

marks, abrasions, weathering and any other Following Andrews (1990), all large mammial 

noticeable surface modifications (root etching, skeletal elements were counted (Ni) and their 

burning, colour alteration). Breakage patterns were — relative abundance (Ri) calculated. The relative 


Entrance 


LS 


7 

} 

| 

| 

S. Aust | 
we 

i 

! 

' 

j 

i 

i 

t 

' 

1 
ms, 
bas 

! 

i 

1 

i 

i 

ei 

Naracoorte i 


Fig. |, Map of Cathedral Caye, Naracoorte, showing the position of the study site, Modified from Cave Exploration Group 
of South Australia surveyed map of Cathedral Cave. 


FOSSIL VERTEBRATES FROM CATHEDRAL CAVE, NARACOORTE Va 


abuninee counts were bused on the rekitionship 
hetween the Niv the expected numbers af each 
clement (Ei) within a& complete skeleton, und rhe 
minimum number of individuals. (MND, This is 
SUITES ats 


Ri% = _Ni_ x 
(MINI) Ei) 


The MNI Was prodticed by suimming the most 
ubundant skeletal clement refenable to the Gixon in 
question and dividing this by the number represented 
iv a complete skeleton. These results were then 
totalled ly vive an overall large manimal MNL 
Species MNE were biased primariky on the number ob 
craniodental specimens. as these contain the species- 
specie characters, while MNT yialues for genera 
were based On elements, ustally post-cranial, whieh 
could nol be identilied fo species level, En wis 
culculated by multiplying the MNI by the number of 
cuch clement type present in a complete skeleton, 
The Raat Kangaroo, Macropas rufus (Desmavrest, 
1922) was used as (he compurilor skeleton lor a 
culculations. To enhance accuracy, caudal vertebrae 
were excluded from these calculations. as there is 
considerable variation in the number of Gul vertebrae 
herweerl species. 


Results 


Cove sedtnnenys 
Sediments accumulated as a simple cone beneath a 


=. Limestone 


@ee, Spelcothem 


zm 


Rie, 2 


solitary rool entranve flow choked with sediment, 
limestone blocks and colette formation. Within the 
come, Tour sedimentary units were ensily identiticd 
(desiginted 1.2, 3 ind 4) on the bitsts of colour All 
UNIS Consist ol quarty sands with variable quantities 
uf admixed clay (Table 1), Carbonaceous material is 
ubundant in Units | and 2. sparse in Unit 3 and 
ibsent from Unit 4. Vertebrate fossils were recovered 
from wt units with the exeepuion of Onin. Units t. 
2 and 4 are cogtinuous throughout the Fossil 
Chamber, Unit 3 is restricted (o the distal regions of 
the sediment cone. All sediments have similar 
characteristics of grain sree (Pre. 3) and shape, and 
clay content. Sediment colour was unique for each 
unit, 


Geachronaony 

Three U-sernies dates on calene deposits 
mierlayered with (he Jossiiferous sediments i 
Cathedral Cave were reported by Ayliffe er al, 
(1998), These ages vlone with din additional date (CC 
FOC BS-+) obtained during this study are presented in 
Table 2. A small quantity af bone wis found 
deposited within (he Howstone layers. All sediment 
units (bh to 4) lie beneath the CMowstone stricture 
(CCKC FS-2) dated wt 159.2 + 2.2 ka. A flowstone 
(CCPC PS-4) dited at 279.2 + 7,2 ka hes below Line 
3 while overlying Unit 4. ‘The stratigraphic 


relationship between (he sedimentary units and the 
dated Hlowstones is shown in Fig, 2 


Sochion of (he Cathedral Cave Posstl Chiunbers showing the straivraphy, cited speleatheris, the position al the 


eXeuvalion pits und the lovation of the ulituin. Verlebrite fossils were recovered fren sedimentary Lanits 1, 2 nda, 


4 5S. P. BROWN & RT. WELLS 


Palaeontology 
ASSOCIATION AND ARTICULATION 

During excavation it was evident that some bones 
were cither in association or articulation. 

All of the associated specimens were from either 
extinct (Preceptodon, Sthenuruy) or extant 
(Macropus) species of kangaroos. Based on MN] 
calculations, parts of five (3 Sthenurus spp. | 
Pracoptadon sp. and | Macropus sp.) individuals 
were found in association representing 2.8% of the 
total number ol specimens recovered, 

Figure 4 shows an articulated specimen i stfu. 
Articulated material represented 1.6% of the total 


specimens include both extinet (Srhennrus gillt 
Merrilees, 1965, one individual) and extant 
(Macropus sp., one individual) kangaroos and an 
extant bandicoot (/yoodon sp., one individual), The 
discovery of the articulated partial skeleton of S. 
gilli is the first ever recovered and will be 
described elsewhere. The articulated bandicoot 
skeleton was encased within a mass of calcite. The 
arrangement of its bones was not consistent with 
an owl pellet. 


BREAKAGE PATTERNS 
Figure 5 shows the distribution of breakuge 


number of specimens recovered. Articulated patterns for the total fossil assemblage. 
TABLE 1, Characteristics of sedimentary units within the Cathedral Cave Fossil Chamber. 
Unit Clay CaCO! Sand Sediment colour Sand colour Grain 
No. % % % Munsell Munsell shape 
l x7 5 86.3 Yellowish red Reddish yellow SR-SA 
SYRS/& SYRO6/8 
2 6.5 7.1 86.4 Dark red Yellowish ted SR-SA 
2S5YRAS SYRS/8 
3 8.2 3 88.8 Reddish yellow Very pale brown SR-SA 
SYRO/S IOYR8/2 
4 8.9 2 SSS Very pale brown Very pale brown SR-SA 
LOYRS/4 OY R8/3 


Sand colour refers to the colour of dry sediment afler removal of the clays. 


Grain shape: SA = sub-angular; SR = sub-rounded. 


60 7 


50 + 


Frequency (%) 


Phi 


Fig. 3, Grain size distribution of all sediment units from the Cathedral Cave Fossil Chamber. Alphabetical prefixes refer to 


excavation pitand the sediment unit number follows 


FOSSIL. VERTEBRATES FROM CATHEDRAL CAVE, NARACOORTE Ss 


a 


TABLE 2. Summary of U/Th dates on calcite deposits in Cathedral Cave Fossil Chamber: 


Code number Date (ka) Comments 
CC FC FS-4 95.2 41.3 Overlies Units 1, 2 and 3, Pit B. 
CC FC FS-2 159.2 42:2 Overlying all sedimentary units; provides minimum age of fauna; 
sediment influx ceases 
CC FC FS-3 279.2 + 7.2 Underlies fossil bearing sediments: gives maximum age for Units 1, 2 and 3 
CCFC StI 399 + 19 Provides absolute maximum age of fauna 


Dates and sample code numbers from Aylifte er a/. (1998) and Ayliffe (pers. comm. 1998). Code number abbreviations: 
CC, Cathedral Cave: PC, Fossil Chamber; FS, flowstone; St, stalactite. 


Pig. 4. An example of one of the articulated specimens im 
wry in the Cathedral Cave fossil depasit. This large 
macropodine vertebral columm (cervical to sacral 
vertebrae) including some pelvic elements, was retrieved 
intact, Seale bar = 10 em, 


Approximately half of all specimens collected (48%) 
were entire (i.e. no breakage); another 25% showed 
evidence of clean recent breakage during excavation 
and/or removal. The remainder showed breakage 
patterns of more ancient origin that included 
irregular perpendicular (22.5%), crenulated (2.3%), 
spiral fracture (1.4%) and compression (0.8%). The 
majority of compression fractured specimens were 
concentrated in the lower Unit 3. 


PREDATION AND SCAVENGING 

Bone damage caused by predation or scavenging 
(including surface markings such as puncture 
wounds and crenulated gnaw damage) was evident 
on 2.5% of specimens. Predation or scavenging 
damage was restricted to bones of kangaroos of the 
genus Macropus with an estimated body mass of less 
than 60 kg. None of the very large extinet marsupial 
species (e.g. Zygomaturus, Procoptodon) exhibited 
predator damage. 


SURFACE FEATURES 

Few specimens from the Cathedral Cave fossil 
assemblage showed evidence of burning. Burning is 
commonly recognised by the carbonisation of the 
bone collagen, discolouring the bone to black 
(charring) or producing a chalky white texture from 
prolonged exposure to high — temperatures 
(calcination) (Brain 1981), 

A few long bone fragments had a unitorm deep 
brown surface discoloration that in places penetrated 
into the cancellous core. Specimens from swamp 
sites such as Rocky River on Kangaroo Island show 
similar discoloration. 

No evidence of root etchings or abrasions was 
found on any specimen [rom the fossil sample. Some 
‘pseudo-abrasion” patterns were observed (i.e. 
abraded cancellous bone), but these were interpreted 
us preparation damage. 

Few specimens showed evidence of sub-aerial 
weathering. Figure 6 shows the [requency 
distribution of bone weathering with the vast 
majority of specimens categorised as weathering 
stage 0. 


Clif S.P. BROWN & R/T. WELLS 


SKELETAL FLEMENT ABUNDANCES values obtained are for phalanges (0.7%). carpals 
The skeletal element abundances for large (0.6%) and metacarpals (0.2%). Although the 

mammals fron each fossiliferous sedimentary unilin absolute numbers of vertebrae and ribs are the 

cach pit are presented in Table 3, On average, the — highest for most units, their relative abundances are 

relative ubundances (mean Ri) of skeletal elements — close to the mean. 

representing large mammals are between 4.3% and 

6.3%. The highest relative abundance values are for — SpecIES MNI 

mundibles (mean of 19.6%), femora (11.1%) and Tables 4 and 5 show the MNI values for each 

tibiae (12.4%). The lowest mean relative abundance — species idenufied from the Cathedral Caye fossil 


I.P » 500 - 
Fs 
o 400 
&Cren. 8 
Cc = 300 
m@ Comp. 
P 5 200 
2 
O Rec/unk — 100 
3 
mSpiral 9 = a HE 
o - N ia) a uw 
Weathering stage 
M1 None pp Ra 
Fig. 5, Distribution of the various breakage patterns Fig. 6. Frequency distribution of specimens displaying 
observed on the entire Cathedral Cave fossil sample (N = characteristics of the various weathering stages 
545), Abbreviations: LP. irregular perpendicular. Cren.. (following Behrensmeyer L978) from the Cathedral Cave 
crenulated, Comp,, compression, Rec./unk,. recent (post fossil assemblage. 


depositional) or unknown damage. 


TABLE 4. Large memmal skeletal element abundance (No. ) and relative abundance (Ri%) from both excavation pits ane all 
fossiliferons sedimentary units in the Cathedral Cave Fossil Chamber, Naracoorte. 


Pil B B A B A 

Unit 3 2 2 l | Mein 
Element No. Riv No. Riv No. Ri% No. = Rive No. Ri% Rie 
Skulls 4 12.1 3 75 | zi I 11.1 0 0.0 7.7 
Masillae 4 6.1 I 13.8 | 3.8 I 5.6 () O.0 5.9 
Mandihles 13 19.7 22 27.5 4 15.3 3 16.7 a 18.8 19.6 
Individual teeth uo OS 6 4 13 2:9 0 0.0 Ls 1.2 
Vertebrae 46 5.2 él 3.6 19 54 13 5.3 18 8.3 6.0 
Ribs 24 2.8 21 20 10 3.0 3 1.3 2 5.8 3.0) 
Seapulae 4 6.1 | 1.3 0 0.0) | 5.3 l 6.3 3.3 
Humeect 3 45 4 5.4) I 3.4 i} 5.5 1 6.3 5.0 
Radi 3 45 3 3.8 2 7 i) 0.0 3 18.8 7.0 
Uinae | 1.5 4 5.0 | 3.4 2 I. | 6.3 5 
Carpals 0 ().0 0 0.0 I 0.5 0 0.0 3 2.7 0.6 
Metacarpals 0 0.0 0 0.0 I OS 0) 0.0 0 0.0 2 
Pelvic elements 13 49 9 3.8 3 2.9 3 42 | 1.6 3.5 
Femora 16 24.2 9 13 2 77 0 0.0) a 12.5 IL] 
Trhiae Il lo7 6 75 | AN 5 27.8 ] 6.3 12.4 
Fibulac 2 3.0 I 1.3 i] 3.8 2 1h. | 6.3 5.1 
Tarsals 10 2.2 7 1.3 5 27 () 0.0 3 27 1.8 
Metatarsils 13 49 lt 5.0) 5 48 0 0.0 ! 1.6 33 
Phalanges |2 OF |2 (16 § 1.1 2 ().4 4 0.9 0.7 
Totals stay 196 7 37 60 


Mean Rift 6.3 54 4.4 5.0 5.6 


FOSSIL VERTEBRATES FROM CATHEDRAL CAVE. NARACOORTE 97 


assemblage. A total of 103 hirge mammal and LO7 
small vertebrate individuals is represented by the 
fossil collection. The most comnion species was the 
Eastern Grey Kangaroo, Macropus givanteus (Shaw, 
1789). In total, kangaroos of the sub-family 
Macropodinae are the most frequently represented 
species (49.5%), followed by those of the extinet 
Sthenurinae (37.9%). The most prevalent small 
mammal species were rodents. 


SPECIES ABUNDANCES 

Figure 7 shows the proportions of each Jarge 
mammal species within the entire fossil sample 
based on MNL values. Herbivores are represented by 
97.1% Of all large mammal fossils. Approximately 
hall (51.5%) of herbivores in the deposit are extant 
kangaroo species of the genera Meerapus and 
Wallabia, Macropus spp. dominate the grazing 
niche, while the extinct sthenurine kangaroos 
(37.4%) along with Wallabia bicalor (Desmarest, 
1804) make up the majority of browsing herbivores, 


2 


Large carnivores make up only 3% of the total fauna, 


Boby MASS DISTRIBUTION 

Figure 8 displays the body mass distribution for 
large mammals (>5 ky). Body mass estimates were 
obtained from Calaby (1995). Jones (1995), Lee & 
Ward (1989), Merchant (1995), Murray (1984, 1991). 
Poole (1995), Rounsevell & Mooney (1995). Wells 
(1995) and Wroe et al. (1999) using maximum male 
weights. The large mammal distribution shows a high 
frequency of individuals weighing between 5 and 20 
kg and between 40 and 60 ke. Very few very large 
individuals (>100 ke) are represented in the fossil 
deposit. 


MAMMAL HABITATS 

Table 6 shows the preferred or inferred habitats of 
all the maramal species represented in the Cathedral 
Cave fossil assemblage. The mijority of species 
inhabited an Open forest or woodland environment, 
Some species are known to occupy a wide range ol 
present day habitats and, consequently, are less 
informative. Zygomaturus trilabus (Macleay, 1858) 
has been suggested by Murray (1984) to have 


Tash 4. Minimo number of individuals (MNI) far large manimedls from all fossiliferous sedimentary units, Cathedral 


Cave, Naracoure. 


Pil A A B B BR 
Uni 1 2 I 2 3 
Thylacinus evnacephlin i 

‘Total Thyliaeinidae MNT ! 
Lasiarhinus latifrons | | 
Vormbatidae Indet. | | 
‘Vor Vorbatidae MNL ! | 2 
Mireropus uigenlens 2 4 4 a) | 
Macrapus rufogriyeus I 2 5 
Macrapus sp. Indet. | 3 | 0) 6 
Wallabia bicolor 2 
Tota! Macropodinag MNI 3 7 5 22 14 
Sthenurty galt | i 5 5 
Sthemuris brawitel | | z, 1 
Sthenuras accidentally 2 ] 
Siheouilas sp. tocet. ag a 2 5 fy 
Proceplodon wslial | | 
‘Poul Sthenurinue MNI | | Ps 15 4 
AVGINTEPUS Willies | ! | | I 
Tolal ZAygomaturinae MNI | | | | \ 
Mivlicolee camifer | | 
‘Total Phylacoleonidae MN] | | 
Toll No. extinet species } | a 4 8 
‘Total No. extant species 3 + 3 3 4 
Total No- species 6 8 A] iI 3 
Total MNT extinct species 4 5 3 Is 17 
Total MNI extant species j 8 ty ad 16 
Total MNI 8 13 ” 40 44 


NB. Sarcaphilys barrisi( is present in the assemblage bul was recovered during a previous excavation and the 


stratigraphic origin is unknown. 


O8 8S. PR. BROWN & R.T. WELLS 


inhabited wetlands, such as swamps or billabongs 
and its diet may haye included vegetation growing 
along the banks of water holes. 


Discussion 
Geology 
Cathedral Cave lies within the Naracoorte East Dune 
which contains a series of potential sediment sources lor 
cave fills, including Pleistocene beach-dune, estuarine- 


lagoonal and lacustrine facies, and Phocene marine and 
fluviolucastrine facies (Cook et al. 1977; Grimes 1994). 
These sandy facies, individually or in a combination, are 
a likely source for the Cathedral Cave sediments, 
although soil formation, leaching and/or mixing makes 
it difficult to establish firmly sediment provenance. 

The sedimentary umits within the Fossil Chamber 
appear to be continuous between excavation pits with the 
exception of Unit 3. No distinction between sedimentary 


TABLE 5. Minimum number of individuals (MNT) for small mammals, reptiles, amphibians and birds from all fossiliferaus 
sedimentary units, Cathedral Cave, Naracoorte. Indeterminate family or class individuals were not included in total 


extinci/extant species calculations. 


Pit 
Unit 


ee 


Smiinthopsis urine 
Amechinus flavipes 
Amlechinas sp. Indet. 
Phascogale calura 
Dasyurus viverrinus 
Dasyurns maculatus 
Total Dasyuridae MNI 
Perameles bougainville 
Perameles gunnii 
Perameles sp. Indet. 
Peramelidae Indet 

‘Total Peramelidae MNI 
Cercartetus Hanus 

Total Phalangeridae MNI 
Bertougia penicitlata 
Potorous platyaps 
Potorous tridaetylus 
Betiongia sp, Indet. 
Total Potoroidae MNT 
Mastacamys [uscus 
Pseudomyy australis 
Pseudomys shoriridul 
Notomys mitchell 
Pseudomys sp. 

Total Muridae MNI 
Aves Indet. 

Total Aves MNI 

Tiligua rugosa 

‘Potal Reptilia MNT 
Limmedynastes sp, Indet. 
Total Leptodactylidue MNI 
Total Noo extinct species 
Total No. extant species 
Toritl No, species 

Total MNI extinet species 
‘Total MINI extant species 
‘Total MINI 


_ 
lo 


or 
i> 
1 
few 


— 
Nm ee 
i) 


i 


RK MM Mh ww 


ta 
m5 
+t 
am 
wh 


- Ww 
iio. 
i 
4! 
™os 
ed 
hot 
vimeeey 
wn 


Brows, S PB 1998) A Geological and Palacontological 
Lyanination af the late Pleistocene Cathedral Cuve Fossil 
Accuimulinon, Naracoorte, South Australie BSc (Hons) Thesis, 
The Flinders Linversity of South Australia (unpub. | 


FOSSIL VERTEBRATES FROM CATHEDRAL CAVE, NARACOORTE 


oo 


o 
Ee 25 
c 2 
w 
nol] 
5 2 
3 le 
B 20 
~ 
215 
> 
~ 
wo 
> 10 
; 
(0) - oe 1— = f= fe b i. { a || - mm || a 
w& . a w * = a. : 
c Q = & o = un th ~ 
sf eg * 8S Fe fg 8 3 
a) “ 5 = So 
3S g 5 i ij 3 s § qi = 3 = 
= a 8 5 § § G =e cs oF e : 
= rs) = S ms] a oa a] 3 
. 5 a “ 5 ; G vi 8 . y 
~j a . = ov i] oO I Ni = 
oO od = KR 
= = ‘ 3 8 BY 
= 2) uy ia) 
K 
Species 
Fig. 7. Frequency of the number of individuals of the various large mammal species expressed as percentages of the total 


number of individuals from the Cathedral Cave fossil assemblige. 


60 7 
Ss 50 
& 
— 
40 + 
3 
c 30 
o 
= 20 
o 
— 
u 10 
o+ ee | 
co a = 8 &B SS S 
ont: 
Body Mass (kg) 
Fig. 8 Distribution of all large mammal species based on 


MNI values plotted by weight classes from the Cathedral 
Cave fossil assemblage. 


units could be made based upon the grain size distribution 
or gram shape. Calcium carbonate and clay content varied 
between the sedimentary units with litle similarity with 
the sediments of the region (Brown 1998!) The 
variation in calcium carbonate content of the cave 
sediments thay haye occurred following incorporation 
of cave limestone via fretting from the Fossil 


Chamber roof (Wells e¢ a/. 1984). Pleistocene beach 
dune facies are prevalent in the region but are not 
interpreted as the source for the Cathedral Cave 
sediments due to their very high amount of calcium 
carbonate content. Moriarty et al. (2000) suggested 
that the cave fills at Naracoorte were sourced from 
surface soils during periods with a wet climate regime 
with abundant vegetation (Le. interglavials, stadials 
und interstadials). However, Units 3 and 4 contain 
fittle or no carbonaceous material suggesting that 
surface soil development may not be significant 
during the deposition of these units and they may have 
originated during more arid periods where rainfall and 
vegetation cover were low. 

The speleothem dates provide a time trame and 
suggest environmental conditions under which 
sediment accumulated within the Fossil Chamber, 
The buried flowstone (CC FC FS-3) gives 4 
maximum age (279.2 + 7.2 ka) for sediment and 
fauna accumulation in Units 1, and 3 and a 
minimum age for the underlying Unit 4 sediments 

A U-series date from near the lower end of the 
buried stalactite (399 + 19 ka) provides a maximunt 
age for Unit 4, as burial of this speleothem had ta 
occur following its formation. 

The speleothem developed on the upper surface of 
the sediment cone, dated at 159.2 + 2.2 ka, provides 
uminimum date tor cessation of sediment deposition 


1a) 


8S. P. BROWN & R.T. WELLS 


TARLE 6. Preferred or inferred habitats of nueamnial species recovered from the Cathedral Cave fossil asseniblaye, 


Naracoorte, 


Species b Ss H QO, F. W R 
Antechinus flavipes x xX x x 
Sminuthapsis mirina xX x X x 
Dasyarus viverrinns x x x 

Dasyurus maculatus x x xX 

Sarcophitus harvistt x x Xx 

Thvlacinus eynocephalus X xX x 

Phascovale calura x 

fyoodon abesulus Xx X x 

Perameles bougainville xX x XxX 

Perameles genni x xX x 

Bertongia penicillata x xX Xx x 

Potorous platyops x x 

Pororaus tridactylus x x x 

Lasiorhinus latifrons x 

Macropus viganteus x x Xx 

Macrapus rifagriseus x x x 

Wallabia bicalor x x x Xx 

Srhenurus gilli x 

Sthenurus browne x 

Sthenurus occidentalis x 

Procuphodon goliah x x 

Cercarlens narns x x x 
Zygomatnris wilobus xX x 

Thylecoleo carnifex x x 

Masiacomys /uscus xX x x xX 
Pseudomys ansivalis x xX 

Pseudomys shortwridgs X 


Natomys mitelelli 


x 


Data obtained from Archer (1981), Bradley (1995), Christensen (1995), Edgar & Belcher (1995). Fox (1989, 1995), Friend 
& Burbidge (1995), Godsell (1995), Happold (1995), Heinsohn (1966), Jarman and Phillips (1989), Johnston (1995), 
Murray (1984), Seeback ev al. (1989), Tate (1947), Turner & Ward (1995), Walton (1988), Watts & Aslin (1981) and Wells 
(1995), F = Forbs. 8 = Savannah. H = Heath. O.F, = Open forest. W = Woodland and R = Rainforest, 


in the chamber, which probably occurred following 
blockage of the solution tube entry point. In other 
words. the entire Cathedral Cave fauna from Units t, 
2 and 3 dates between 279.2 + 7,2 and 159.2 + 2.2 ka 
corresponding with oxygen isotope stages 6, 7 and § 
(Shackleton & Opdyke 1973; Martinson ef al. 1987). 


laphenoriy 
ACCUMULATION MODE(S) 

The fossil evidence supports accumulation of 
amimals via a pitfall trap. The Jow number ol 
mammalian carnivores and the seurcity of carnivore 
tooth markings and gnaw damage (characterised by 
crenulated breakage patterns), suggest that the fossils 
were not accumulated by mammalian carnivores and 
the chamber was not used as a den or a lair (el 
Lundelius 1966: Sutcliffe 1970; Brain 1980: Haynes 
1980; Scott & Klein 1981: Cruz-Uribe & Klein 
1994: Skinner ef c/. 1998). Purthermore, the absence 


of root etching and the small number of burnt or sub- 
acnul weathered bones argue against a surtace 
accumulation where animal remains would be easily 
accessible to carnivores. The few specimens 
displaying characteristics of carnivore aclivily were 
most likely hydraulically transported jnto the cave 
from locations proximal to the entrance or resulted 
from an entrapped carnivore within the chamber. 

Although water transport of adimal remains into 
the Fossil Chamber may account for some post- 
mortem damage, the aforementioned evidence 
suggests that bone accumulation did not occur by 
this means. All evidence is consistent with a pitfall 
trap. 

The deep brown discoloration of some bones 
Seems to contrast with the paler colours typical of the 
deposit. The deep brown specimens are comparable 
to bones found in swamp deposits at Rocky River on 
Kangaroo Island (Wells ef a/. 1999) where the colour 


FOSSIL VERTEBRAPES PROM CATHEDRAL CAVE, NARACOORTE Wd 


has heen attributed to tanimin uptake or stining. 
Perhaps this type of surface colousiig present on 
same Cathedral Cave specimens madicates local 
ponding within the cave system. at a time when 
outside conditions of high vegetation cover increased 
the quantity af tannins inte the downward 
pereohuting ground waiter. 

The data obtained for the small vertebrate hauna 
(Table 5) suggest at least two modes of 
aveumuliion. “The low toll number of individuals 
and the small number of arboreal species are not 
Inconsistent wilh w pitfall trap. Arboreal species 
would be mote able to climb outolthe cave had tiey 
fallen Ge climbed in, However. the bivzher number of 
rodent idividtials fecoyvercd from Pit A, Unit 1 and 
Pu BR, bait 2 Sugeesis (hat an avian predator may 
have roasted within the cave or solutior# tube ur in an 
overhanging tree One nightinfer Fron the relatively 
small niiinber of individdals compared with other 
tatensive owl deposits al Naracoorte (MeDowell 
20007) that (his only oecurred for a short tine, 


AHCUL ALLY SPLCIMENS 

The presence of articulited: fossil speeimens 
Suyeests thi some vintmals entered (he cave intact. 
These were presumably live animals, teapped by the 
pial mechanism. which either died From the fall 
ts the Give and decomposed on the cone or 
survived the fall and were subsequently able to move 
boul within the chamber As the majority of 
wleulhited miuterial was revovered from the distid fin 
royains Wallin dhe Fossil Chumber. the hitter seenurio 
seems the more ikely, Observations by one of the 
duthors (RTW) ob contemporury accumulations 
suggest (hut following choaipmicnl starving: animals 
became thienmotisic and tended to seek Gut the 
aecurily Of walls and crevices and died there, Theos 
evidence Furtber supports the hypotheses thi a pitfall 
tmiechanisi wus the primary mole of accumulation 
for luge nkunmuls aod indicates thar the burial af 
some bones occurred rapidly before disarticubition 
Gould ecu, 

The Tigh of ariicululad material representing very 
large mammals (100 ke) (he, Pracoptecdian sp. am 
Avauiaiirisy sp) suwests thar either their reniains 
were Hransperted tate the caye ot the dimetee of the 
solution tobe aeted as at Shody tass sieve’. 
preventing the passive OF larger, intact animals. As 
Already disclosed above, the absence ot weathering, 
JA resrons. Fro AYUrAn Te Fraaspart aie wily extensive 


VE Tho VPP ROG Die Suet Hh rineepbalss cnt sea lipenic of 
Kotrvisen Conc, Worl Petijge Syotaccectie Cures Cormspry nn 
Wph Sei Keesha lest Or poster presen ay ihe 
Chaeniors Stidiee Meseiter Rowtorial aitilvab. of Austeation 
ULCET TAT ALG ee STE TTSS PE De aIMe ne tis tere 
Rel Pobre s TU0) bantu 


bone surtace disealorition suggests that the bones 
did noe aecumulate outsnle the eave. ‘The sive of 
indivitoals of these species miy have prevented their 
falling directly into the chamber below. The 
individuals would be tapped and then die within the 
solution tube. wath their remains gradually 
incorparaled info Lhe fossil deposit as the caresses 
decomposed. The low MNI values for these lure 
herbivore species suggest one or all three 
possibilities; that few became tapped, that they 
could readily extricate henmelwes, (hal Uneir 
numbers were low in the dmmedhuc vicinity of the 
pitfall. 


SKALBTAL ELBMENT AKWNDANCLS 

The skeleial element abundances tor eueh anit aed 
cxcavahion pit indicate thit the relubve umber ob 
skeletal elements recovered (RIG) is low, Accvepting 
the hypothesis oF a pittall mechanism capturing live 
wnimals a random, (he majority of hone in the 
deposit would result (rom decomposition of whele or 
Near Whole unimuls within the Fossil Chamber In 
this case, dn Mdividial’s entire skeleton should be 
represented within the (otal fossil deposi and se 
would give high RIG values were the entire deposit 
lo be sampled. The low observed relanive ahundinces 
are thus interpreted as an artelaiel ol sampling These 
values qse iideate dispersal oF elements following 
uccumulalion under transport regimes such as 
sediment miss Movement. water How and/or 
biogturbation. 


Palaconiolowy 

COMPARISCHS WELD VICTOR IG FOSSIL CAVE 

Vieloria Fossil Cave. located about 700 nm from 
Cathedral Cave. contains several fossil sites iat 
have yielded an army oF Middle Pleistocene fainits, 
(hough onty (he main Fossil Chamber hoy been 
tharouhty researched (Wells efal 1984: Moriarty et 
a. 2000). The Fossil Climber ussemblige is by tar 
the Lirgest andl richest in all the caves al Naracoorte 
(Wells ev ul. 1984), Possil-rich sediments appear te 
have been deposited prior io 274 ku and span muy 
thousands of years (Ay eal M99k. Moriarty ev 
ui) 2000), A pofall trap is sugpested ta be the primary 
inechumism responsible for the accumulation ot 
uniimils (Wells ere 1984). A majority oF the mare 
conmMmon jnegafaudal Species alse occurs within 
Cathedral Cave (es, Sthemeras gilt, S brownes 
Merribees, 167. 8. cecideniciliy Ghanert, (910 and 
Zyeomotias Utebus) The mayer luna dllerences 
heiween the lwo assemblages lie in the presenee of 
the rurer species in Vicreri Fossil Caye tee. 
Paulorchestes azeel Oneuiw, LATS NL eterveni 
Mitreus, 1862. 5. sudeklocks Wolls & Murniv. (97a § 
pales DeVis, S95, Stheniuas sp anv, 4, Helttey! 
Pridvaux & Wells, (908 and Pronvdeerlur corchas 


Wi 8, P BROWN & ROE WEILS 


Owen, I874) nd are likely to reflect differences 1 
simple size, 

Preliminary faunal samples have been obtained 
from diated deposits in Spring, Chamber and Grant 
Hall within Vieloria Fossil Cave (Moriarty eo ul 
HOO). Many species detected in the Cathedral Cave 
fossil assemblage were not recovered from Spring 
Chamber. This may be an artefact of the small 
sumple size from Spring Chamber. U-series dites 
obtained indicate that the Spring Chamber (auna is 
older than that ol Cathedral Cave (Aylitfe had, L998) 
Moriarty ef af, 2000), althouulr some overlap oecurs 
during the period 282-21) ku corresponding to the 
Upper py whion of the Cathedral Cave date range (274- 
154 ki. 

Grant Hall sediments appear lo have a higher fossil 
content thin those oF Spring Chamber and the fina 
is more like that of Cathedral Cave. The megatiuina 
represemed Closely resembles the Cathedral Cave 
fossil assemblage, Moriarky ef al (2000) suggested 
un awe beiween 140 and #O ka for sediment 
acculmulation in Grunt Hall tlowever. they ancue 
that (his muy not be the true age of the sediment 
Titus andl Haun die bo the potential for reworking of 
previously deposited sediments, 


PROXIMAL COMMIUALEY AND PALAROENVIRONMEN TA 
RECONSTRECTON, 

An understanding of the laphonomic rises wilhin 
Tlic fossil qasemblage gives more contidence bo uny 
mildcoenvironmental reconstucnon, but. as the 
fauna accumulated ducing a period of approximately 
120 ka, time-averaging way comproniise this 
Witerprenition 

ocun be voneluded that the huge manminal 
component of the proximal community consisted 
prunarily of herbivores, Of these, graying “lypes 
(Mucropuy spp were the inmost abundant, The 
bawsing herbivore fauna consisted’ oF the extinet 
sthenunine kangaroos. Wallabie  hiceler wind 
ZAvgomatnitns irilohuy, Vegetadon allowing this tbs 
of herbivores to coexist within ihe sume region 
would (neliide shrubs, trees and grasses. Ne open 
forest or woodland wilh a grassy Understorey ts the 
most likely environment. However raging 
kangaroos could aso. forage in nere open areas 
withit oa foresi, on the forest edge ur aajacent 
measshads. Murray (lO84) suggested tar tine 
Worpholugy all the Aveenieuruy nusahy may be an 
Ailaptution for browsing on reeds within shalluve 
walter, suggesting the possible presenes il werlanels 
OP SWANTIPYS. 

The presence ol Lasianiiiny falifrons (Dace, 
TH45) in the deposit appears Theapsistent silly Uips 
environment) reconstruction, Todi. (its species 
Inhabis semiarid lo arid savannial regions (Wells 
1945). However in the Mid-Pleistocene. this species 


mity have inhabited the outer cde of open forest ar 
woodlind, as even in historie tines its ninge 
extended inte higher ramnfall areas (Wood-Jaies 
1994). 

Most of the small animal fauna is indicative of sa 
(pen farest/woodland cnviranmnent. Ao smaller 
number vecurs today im osavaniah and heath 
vegetation suyvesting it inay have occurred in clase 
proximity ta Cathedral Cave during the Middle 
Pleistacene. 

Mediuit-sized mammniils sich us bandicnots 
(Perameles and Jsecden) need sufficient ground 
vover lor refuge and the presenee al low-lying serub 
is suggested. Avtechinny flavipes (Waterhouse. 
(838), Pheisecoxale calura (Gould, thd) and 
Cercarieluy nandy (Desmarest. DSTX) are arbor) 
species. This further supports the presence ol 
ubuidhimt trees vansistent with oan apen 
lorestiwoodlind. ‘The only arid species recovered 
from the fossil deposit was the Ploins Rat, 
Peeudomys australis (Gray, 1842) which Walton 
(1088) sSupgests prefer todiy rocky arid regions. 

The Ritts Suggests VER tITION STFUCLUTe star fo 
that of the region prior to European lund clearmg. 
Croft er ai (1999) indicate that the vegeuition 
community of the Naracoorte region was dominated 
by open eucalypt forest/woodlands with jtermitient 
tussock grasslunds and sedgelund prior Ws European 
derlement, The diversity in vegetation forthe region 
during the Middle Pleistocene as indicated by the 
Cathedral Cave fossil hunasuezesis thal ana focal 
seole the vepeluion may bave heen ecotunal. 


PALARUCLIMATE 

During the period bracketing the Cathedral Caye 
fossil assemblage (between (59 and 279 kad a ghia 
moxinim occurred at I70 ka ulowe wath an 
Hilerehickil Wokyeen jaotope sige Jey und warm 
iIntershailial perils centred on 240, 220 ind 195 ka 
(Aylitte er ak LOY: Winnerad er uk 1907), 
Martinson eral s)987) reported three radiahiriin 
Heh temperuure peaks for the Sourhern Ocean 
(RO V-120h durme the trae ol ruta) aeeunudanon 
Ab 240, 220) 199 kewl a inaxiinuin Lemiperiire 
ot ubeul U3 °C bieher that present umd lowest 
Lemp tures wenn 3 90 below present Che teh 
lemperutine peaks correlate will) Che Tnteesliebal oF 
wart Toterstudiateverts of tits period. Aylitle eb ad, 
C8) Sivees) speleathend deposition lollowed these 
[Hises corresponding te ostadials and evel 
mieradiih, Late bene wecimulatcd darting perils 
Gl speleaitiony lammation suesostitig thatthe majority 
Gt mth retitis were aveumiluted during the 
warner (nore lacials andl iiterstidtals. Accepting this 
fivpollieses cyables (he dates for the span during 
which the Ganmal remains were accuniubalirus fe be 
refined to the period 240 kv li PS ka, 


FOSSH. VERTEBRATES FROM CATHFDRAL CAVE NARACOORTE NS 


Aylifle ev af (1998) Tndicated that the regional 
hydrological balance at present is an analogue tor 
Miereluci conditions, ahd liking this into secount 
his conghuded that the majority of animal remains 
decuoulued during periods of local climate senile 
ta Unit af the present time. 


Acknowledgments 


We thank M2 MeDowell fur his help in surveying, 
(ipering, fossif collection and ideprification of rodent 
species. “Thinks Wo 1. Reed for ber mpul into the 
interpretation of the taphonamy wnd critical analysis 
OF an eurher dint. CG. Pridemix eontubuted to the 
excuvalion ind helped this study with critical input 
Wto sthenurihe species jdentification wat was much 
uppreetaled Thanks ware alse due to D. Meeirian for 
his helpful conmments of the manuseript, 


AL Numivoorte, the assistance af District Ranger B 
Clark aod all the staff at) Naracoorte Caves 
Conservation Purk, purliculurly S. Bourne and A, 
Hansford wats much appreciated, 

We iso thunk PL Daenke whoo prepired miny 
specimens to near perfection. Many thanks also go te 
(he Ureless helpers mn the held; D. Burtholomenuse, K. 
Brelt, 8. Dalgairns, L. Gaston, G. Gully. L-M. Hall 
POMayes and f Thanakhantry. We also wish to thunk 
the many students from Flinders University 
participating inthe Vertebrate Palaeontology | course 
who helped with the excavation. Thanks alsa to P. 
Miariuneliand LO Aylitfe who collected and dated the 
$pclaothem simples. RK. Moriarty s dssrsuince in site 
Heology and speloathen sampling is gratefully 
acknowledged, Finally, we wisi to thank A. Baynes 
for his constructive erificism of the manuseripl: bis 
helphul advice was much uppreckied 


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SPECIES OF RAILLIETINA FUHRMANN, 1920 
(CESTODA: DAVAINEIDAE) FROM THE EMU, 
DROMAIUS NOVAEHOLLANDIAE 


By MICHAEL G. O’CALLAGHAN*, MARGARET DAVIES* & ROSS H. ANDREWS* 


Summary 


O’Callaghan, M. G., Davies, M. & Andrews, R. H. (2000). Species of Raillietina 
Fuhrmann, 1920 (Cestoda: Davaineidae) from the emu, Dromaius novaehollandiae. 
Trans. R. Soc. S. Aust. 124(2), 105-116, 30 November, 2000. 

Four new species of Raillietina Fuhrmann, 1920 (Cestoda: Davaineidae) are described 
from the intestine of the emu, Dromaius novaehollandiae. Raillietina australis 
(Krabbe, 1869) Fuhrmann 1924 is redescribed from specimens collected in Australia. 
The new species differ from R. australis and from each other, in the size and number 
of rostellar hooks and in the dimensions of the cirrus sac. 

Key Words: Cestoda, emu, Raillietina, new species, Dromaius novaechollandiae. 


Transeetions af te Rayal Seetery af & Aaye (20001 12412). 105-116, 


SPECIES OF RAILLIETINA FUHRMANSN, 1920 (CESTODA; DAVAINEIDAE) 
FROM THE EMU, DROMAIUS NOVAEHOLLANDIAE 


by MicnaAriG. O'CALLAGHAN , MARGARET Davirs’ & Ross H. ANDREWS 


Summary 


O'Cautactan, MG. Davins. M, & Axbrews, RLU, (2000), Species of Reiietivg Pubrmnn, 1920 (Cestodic 
Duvaineidie) fromthe emu. Dromeains nevdchotlandiae, Trois. KR. Sac. S. Attst 124(2), 105-116. 30 November, 


MOO. 


Four new speeies ol Reifedina Baba, 1920 (Cestoda: Davaineiic) are deseribed Prom the Witestine of 
the emu, Droniiiis audchollemdiae, Raiiedha auserals Uxrabbe, 1869) Fuhrman (92+ is redeserihed fron 
speermens catleetcu in Austinalin The new species differ (am Ro castais cid Wom caeh eather ai the size anid 
nuinber of postellar hooks und inthe ciimenstons of the cierus sae, 


Kay Worns: Cestodia. emu. Raifledia, new species. Dronainy noveehellandiae. 


Introduction 


Vhe emu, Dromainn veveehelfanfiae (loathun, 
1790), is one of only two ratites (Struthioni formes: 
Dromaidae) infubling Australia. Restricted to 
mainhind Australia, emus are now farmed for meat, 
Ou, leather and cays, In L995 there were 650 licensed 
emu farms to Australia with a population of 71,000 
emus produce 7&.000 chicks annually (Mannion ef 
ai 1995), Reeently Clarke ep ef (1996), Tully & 
Shane (1996) and Shane (P95) reviewed the 
Infectious and parasitic diseases of firmed emus 
although they did nor include any ioformation on 
cestode parasites. 

Cestades were first recorded in emus in |869 when 
Krabbe published a deseriphion of faerie austretiy 
(Davainedae) recovered (rom the intestine of a 
captive cmu which hid died in the Copenhagen 
(Kjuerboeling’s) Zoological Garden in 1867. The 
emu had arrived from Australia [5 years-curher as at 
lully-prown bird (Krabbe [869), 

Later, Fuhrmiin (1909) deserihed another 
dhivaincnl species. Cofmgiie collin, from an emu in 
the Museum for Natural Sciences in Berlin. The 
wade locality for this specimen is not known becutise 
the geveraphic distribution of the host is reported cas 
custern Australia. No subsequent records of cestodes 
in the cru exist and the ientily of the cestodes 
infeeling these birds in Australia is unknown, Both 
of the described purasites oF emus are recognised as 
valid species (Schmidt 1986) although Krabbe’s 
(1869) desenption of whit is now Reaillietne 


 Beparincal ol Pavironnentel Biology, University of Adelaide 54 
S05 

} South Atetnitian Resguebs iid Developryent basil. GEO Hox 
807 Adglaide SA S00) Fo pil oewlhighvtn richer satiety, say 2a) ve 


australiy tucks & number of niorphometrie and 
deseriptive characters. 

Our study of farmed and wild emus fas resulted in 
the recognition of five species of cestode. All are 
assigned to Raillredina Fuhrmuann, 1920 (yensu Jones 
& Bray 1994) on the basis of the possession of two 
rows of numerous hammer-shaped Hooks, wni-hiteral 
venilal pores, i small cirrus sae. which does not 
reach or just crosses the osmoareeulatary canils and 
exe capsules containing several cays, One species is 
identified as Raillrerina auytraliy whilst the other 
four are undescribed. Here we redeseribe A, ausireles 
from the holotype and new material and deseribe the 
new species, 


Materials and Methods 


Cestodes were collected from farmed and wild 
emus in South Australia, Additional materiil was 
obtained from the Australian Helmipthological 
Collection of the South Australian Museum (ATO). 
Cestodes were relaxed in rap water. fixed tn 10% 
buffered Tormudin and stared tn 70% ethanol, Whole 
Mounts were  stuved with  Heidenhain’s 
haematoxylin, Celestine blue or Semiction’s 
iicelucarming, dehydrated in a grided series ol 
cthanol, cleared jn clove oil and mounted in Canada 
bulsumi, Seoleves were mowited in Debaure’s 
medium. Measurements are given in (he text. ini, 
us the range followed, in parentheses, by the mean 
and the number of observations. Deseriplions and 
ineusurements Gf cestodes ure based on the 
exUmuiuon oF up lo S00 specimens of cach species. 
Drawings were mide with the vid of a camera lucida 
attached to an Olympus BH ynicroseape. Type 
Speciniens have been deposited m the Austealian 
Helminth Collection (AHO) South Australijar 


16 M, G, OCALLAGITAN, M, DAVIES & RL TL ANDREWS 


Maiseunt. Adelaide (SAMA) and the British Museuin 
pNatird History). Londen (BMNH), 


Kailheving australis (Krabbe, 186% Fubrovani, (924 
(FIGS |-7) 


Tuenia australix Keabhe, 1869. Ko Danske 
Vidensk Selsk Ske Nalury. Og Math Atd. 8, 249- 
369, Figs 206-298. 

Duwiinea anstealix Ckrabbe. 
{&Y] 

Reansomia auatratiy (Xtabbe, US69) Fulbrmann, 
1920 

Korlania auaidlis, (Krabbe, 1869) Loper-Neyra, 
)OA| 


1869) Blunehurd, 


Holoaype: tn Zonlogisk Museum, Copenhagen. 
Dciennark 


Veraivypes: Radin South Austratia (Si) (33° Saf 8, 
ar 48" G). Cot, M. O'Callaghan. Llevii 1995, 
SAMA AHC 31376. BMNH 2000.3 17.1-10. 


her melerial cvanined. Werribee, Victoria (View), 
Coll Ko EL Varrigan, April T9ks, SAMA ATIC 
A341; Shelley River, Queensland, 2.7%, 1907, 
SAMA AMC 227) Kinehegia, New South Wales 
(NSW), Cold. Beveridge, 31th O74, SAMA ALIC 
hOO0G: Yunta, SA, Coll G, Ey ord, bin, 19ad, 
SAMA AHC TITS: North West, Western Australia 
Coll. Tbh Johnstum, SAMA ATIC $2043, 


Deseripiion 

Cestodes of moderitte sive, up to 50 in unrelaxed 
specimens and up ta ThO in relaxed specimens. 
Maximum width 2. Strobila containing 
upprodimmiltely [ESQ proglattides. Scolex (.416- 
OS68 (0.498, n=20) in diameter with eversible 
rostellom (Pigs 1,2). nirely everted at lixcd 
specimens, Rostellum 0. 200-0288 10,240. n=10) in 
dhumeter urmed with 280-362 (326) hatnmershaped 
hooks urranged an two rows. Lurger rostellur hooks 
0,021-0.030 (0.025, n=250) in length, simatler 
rostellar hooks O.016-0,023 (0.020, 9=250) (Fig. 4). 
Buse of rostellun armed with 16-20) rows small, 
rose-Lhorm-shaped aecessory spines (,Q02-0,004 in 
length, Suekers O.136-0.168 (Q.140. 7=30) in 
dhvoeteratined with eight diagonally ranged rows 
of hooks 0.005 40.011 in length (Pig. 4). 

Proglotiides craspedote. Mature proglottides wider 
thin long, 0100-0, 184 (O17 Dx C8000, 848 (0.823, 
n=10) (Pig. 5), Genital pores single, anilateral, 0.016 
in divmeter: genital ducis  pussing belween 
longitudinal oosmoregulatory — cunals. Dorsal 


osmoregukitory canal O.048 in maximum diameter. 


lying intern da smaller ventral osmoresulatory 
canal, 020 ip dimeter, Transverse osmorepulatory 


cunals connecting tell and right ventral canile at 
postefior Margin oF each prog lots. 

Genital atrium snvall, situated in anterior hall oe 
literal proglouis marein and surrounded by an 
accumulation of cells, Cirrus suc elongate 0,152- 
O.164 (0.158) x 0.0 16-0024 (0,020, H=10) (Pig, @) 
extending lo but not beyond dorsal osmoregulatory 
canal. Distal region of cirrus of greater internal 
dianteter than mid rerion. urinature net seen: 
proximal region forming small, spherical, internal 
seminal vesicle, O.016 in diameter, Civrus aac in 
hololype specimen 0.149 x 0.023, who with internal 
seminal vesicle (.016 in diameter. Cotled vas 
deferens pissing Lowards centre al proglottis where 
iL beceites convoluted, occasionally overlying 
seminal receplucle before passing posteriorly 
fowards ovary, Testes in poral and: aporal felds, 
huinber 4-7 (5) poral and V1-13 C11, = 10) uporal, 
bounded by lateral osimoregulatory canal. ‘Testes 
0,044-0,052 (0.048, n=10) in dhameten 

im Opening to wenital atrium posterior to circis 
sic (Fig, 7). Distal revion slightly enhirged 0.024- 
O32 (ON29) & O“£.070-0,016 (0,018, n=i0), Mil 
regron. narrow, comed O.00S5 th chameter, leading to 
seminal receptacle medially posterior lo vas 
deferens. 0.O84+-0.128 (0.122) % 0.024-0.082 (026, 
n=!0) and tying anterior and dorsal to poral lobe of 
ovary, Qyury distinctly bilobed, situated jn mud fine 
OF proglottis, Poral lobe 0.040-0.072 (0.049) 8 .040- 
0.060 (0,048. n=10). uporal lobe O.014-0.072 (0.060) 
8.0.03.2-0.072 (0.043, n=10) with 3-5 lobules mi) cach 
Johe, Vitellarium irregularly Jobulate, post evaciair, 
slightly aporal, occusionally dorsal to aport lobe ot 
ovary, (.060-0.080 (0.068) & 0.0360-0.044 (0.042. 
n=10). Uterine duet passing anteriorly to developiny 
Ierts. Gravidl proglotides extending transversely 
0.720% Q.350 with large osmoreyukitory canal up to 
O.120 in diameter Ege capsules irregularly ovoid 
O.108-0.132 x 0.08-0.104. Ege capsules 76-110 (88, 
H=10) per gravid proglotts contuining TO-+ Ci, 
hati cees. Terminal proglottides extendiny 
trunsversely, us wide as long O.S80-0.800 x O.600- 
O.880. Oneosphere 0.012 in ditmeter. ondospheral 
hooks 1.005-0,007 long. 


Has/ 
Dramatis wovachetlaredine 
(Suiutnomnformes, Dromwidae), 


(Latham. 1790) 


Lacation in lest 
Sinall dotestine, 
Remarks 
Krabbe ()469) onntted the dimensions of the 


scoles, rastellum and siekers in his description of &. 
dastratis and the strobida Was (nadequately 


SPECIES OF RAILLIETINA FROM THE EMU 107 


~. 
ZS, Wry, 
Fe, ty 
gs Ya — ih, 
g “al tg, 
: “iy 


y 
et 


Figs 1-7. Raillieting australis Krabbe. |. Scolex with retracted rostellum, 2. Scolex with fully everted rostellum. 3. Rostellar 
hooks 4. Sucker hooks. 5. Single mature proglottis. 6. Cirrus and distal vagina, 7. Female genitalia. Scale bars = 0,1 mm. 
1, 2.5-7; 0.01 mim, 3, 4. 


1 M G.OPCALLAGHAN, M. DAVIES & ROH. ANDREWS 


described. The material described above. based on 
(he chaminaiion of SO) cestodes, indieites that 
Krabbe’s (1869) measurements of the postellar hooks 
(12-14 mn) ure consistent with hook width but not 
hook Jengti although this canon be confined 
beaitise OF fhe ubsence Gf a scolex in the type 
inalerial examined, The frysments Ob type material 
obtained, however, do not differ [romp specimens 
cxumibed in this stdely and lave thus enabled the 
redoseription OF A. australis. 


Raillielina heveridgei spy. ov, 
(HGS 6-14) 


Holuivpe: Keith, SA (36° 06'S, lane 19" By, Coll 
M (Yr Cullaghan, 30.70.1999, SAMA SHC $28300, 


Paratypes: Lock SA (33° 34'S. 135) 4a By. Coll. 
M, O'Calhighan, 91.1996, SAMA. AHC S283], 
41377, BMNIE 2000.5, 17.1 1-30, 


Onier material evemined: Yuma. SAL Coll, Go bk. 
Ford. Tax lO8L. SAMA AHC ILLaAl, 521347; 
Werribee, Vie,, Coll, 1, Beveridge, 23.7, 1995, 
SAMA AHC 26698; Mundulla, SA, Coll, Dinning, 
February 1933. SAMA AHC 1187; Bairnsdale. Vie - 
Coll. | Beveridge, 5.x. 1904. SAMA AHC S27717, 
S277 18: Condobolin, NSW. Coll, Ryan, 27.1,1971, 
SAMA AHC 8179; Vie, Coll D, Turner J99d, 
SAMA AUC §26205; NSW. Coll, TO A, dohoston/t 
L. Baneroft. (914 SAMA AHC 520450, 8 204334: 
La ‘Trabe, Vie. Call 1 Beverilge. 24ov11Y72, 
SAMA AHC S20837: Biirnsdule, View Coll & 
Beveridge, Scolex only, S.viil9%4, SAMA AHC 
S27717, STITIS: Vie. 1994. SAMA AHO 826205, 


Description 

Larve cestode, tp to 160 long tn dareluxed 
specimens and up to 600 in relaxed spectinens; 
vravid ostrobiht contiining approximiutely 750 
seuimients, Strobilie wilh max, wilh 3,4 i relaxed 
spevimens. Scolex O.480-0.736 (0.604, n=25) wile 
i suckers (Pig, &). Retrueted rostelluns 0. 192-0, 25% 
(0.234 n=t0) din. with 304-412 (370, n=10) 
lammershaped Hooks i two vows, Larcer rostellar 


hooks Q.O16-,021 (0019. n=250) Tongs smaller 


rostellar Hooks (.0T4+-0,019 (0.016, n=250) lone 
(hips 91, Very small uceessory costellar spines 
approximately O.00)-0.002 th length only visible 
tinder high thagnifieation, Suckers circular Qn) 46- 
H.16X (U.150, f= TO) in ditmeter armed with 12-18 
rows hooklets O.004-0,.011 long (hr, 10) Neck 
variable, up to O.250 i length, Craleareous 
corpuscles present in posterior hallo scolex, 
Proglortides oraspedote. Mature proglouides wider 
than long, 1S34-1.942 (1.730) 8 .0,273-0,3949 (0.426, 
n=20) (Fig, 11). Genital pores smgle. unilateral, 


Litree. ventral. longitudinil osmorezulitory ean 
O.108 imax. dium. joined by transverse canal 
connecting deft and right biteral candls: ih posterior 
margin of each proglottis, Dorsal canals not seen, 
Genital anlize appear in approximately segment 
150. Male and female genital nature in 
proglottides 200 and 300 respectively: first eges 
appear in +80, 

Genital avium small sittuled in anterior hall of 
luleral proglottis margin, Cirrus sae 0.256.328 
(0.29%, el) x G.O80 extending te ventral 
osmoregitatory cunal (Pig. 12), Distal region of 
eirrus Hined with spines, of grewter internal diameter 
than sindous amid region; proximal regiai forms 
spherical internal seminal vesicle 0 060-0092 
(0.079) x 0.052-0.060 (0.056, n=O). not detectable 
in proglottides Of every cestode examined, Vas 
Uelerens greatly cored. extending anteriorly across 
midline oF caeh proglottis then returning posteriorly 
towards ovary. Testes distributed in paral and aporal 
fields within wea defined by ventral osmoregulatory 
cunals. puntber 5-8 (7. 9=30) poraland 12-18 (15. 
p=40) uporal. Testes sub-erreulan 0.060-0.100 
(O.08K) x OLO80-0-088 (CLO83. n=10) not overlying 
ovary On yatelarium, 

Viwing opening (o genie aritum posterior lo cirrus 
sae. Distal region wih thickened myusenhir wall 
O.088-0.120 (0.106. n=10). Mil region of vagina 
nanos, colled, leading medially, posterior to vas 
delerens fo seminal receptacle varying in length fom 
D.ARS-0 240 (0.100. n=20) in length. Lying anterior hy 
Teves and par) labe of ovary. Sperm dock pussine 
posteriorly from semtinul receptacle. Ovary hilohed. 
U.084-0, 132 (0,170) & 0.080-0,088 (O.082, n=O) 
with 4-6 lobules in each lobe (Mig. 13), Vitellarncm 
ovoid OI 20.136 (0.120) x U.080-0.100 (087. 
n=1)) Situated pustenor to oyurys Uterine duer 
passing anteriorly to developing uweris Gravil 
proglotiides wider thin long, 25-27 4 (4-05. 
Terininal proglotlides longer Chin wel LO « O09 
(Pig. 14). Grayvid proglotides containing 30-40 (35, 
n=10) egg capsules, O,168-0,200 (0,084) x 0144 
OA9G (Q16L). n=10) euvh containing 10-12 exes, 
O.040 m0) clameten Onevosphere 0.014-0.016 (0.0161 x 
WOLIMLOL6 (0138, n=l) Oneospheral hooks 
O,00420,006 (0,005, 4 =10). 


Hesr 
Dromaius nevaehiullanitiae 
(Strutiioniformes: Dramaidaet. 


(Latham. 1740) 


Law ition i dost 
Srnall intestine. 


Bivnnlegy 
This species pamed tor Dr l. Beveridge in 
revognidion of his outstanding eontribution ta aur 


SPECIES OF RAILLIETINA FROM THE EMU 10u 


3 Ee veh >) 
/ # My, 
3% ES 
oe 4 
oe ty, = 
i so EM 
a % 


\ 
Te 


Tilteagee 
{rug ie 


ee 
oe 


Figs 8-14. Raillietina beveridge’ sp. nov. 8. Scolex. 9. Rostellar hooks. 10. Sucker hooks. 11. Single mature proglottis. | 2. 
Cirrus and distal vagina. 14. Female genitalia, 14. Terminal gravid proglottides. Seale bars = 0.) mm, 8, 11-14; 0.01 mm, 
9.10, 


WW M.G CY CALLAGHAN, ML DAVIES & Re HE ANDREWS 


knowledge of the parasites of the Australian ender 
fume ad His wuidanee to the senior author 


Raillietina chiltoni sp, ov, 
(PIGS 15-22) 


Holotype Keith, SA (36° 06'S. 140° 18'E) SAMA 
AHC 528502, 

Paruiypey, Kersbrook, SA (4 47'S, La8" SUE), 
Coll. L. Beveridge. Liv. 1989. SAMA AHC 31378. 
BMNH 2000,5.17.31-40. 


Dexeriplion 

Cestodes up lo 0 in relaxed specimens. maximum 
witlth La. Strobilt eontain approximately 360 
progloutides. Scolex 0,545-0,832 (0.643, n=20) in 
diameter wilh cversible rosteluim, 0.436-0,480 
(383, n=10) in chameler, retracted in majority ol 
specimens (Pigs 15, 16). Rostellum armed with 302- 
378 (335, n=10) hammer shuped hooks i bya cows. 
Larger rostellar hooks 0.026-0,039 (0.042, n=250) 
i) length; sauidler rostelha hooks, 0.022-0034 
(0,027, n=250) in leneth (Pig. 17), Buse of restellum 
armed with rose-thorn-shaped accessory spines. 
(0003 in length, visible under high magnification 
only and iv specimens with fully everted rostelluin 
Suckers O.136e).200 (0171, 9=30) in diameter, 
armed with B-14 rows af hooks 0,005-0,013 long 
(Fig. 18). Neek variable in length. O.4-0.8 in relaxed 
Specimens, 

Proglottides: erispedote, Mathire  proglottides 
O.890- 1.400 x 0.0720. 7400 CFig, 19). Genital pores 
single. unilateral; venilil ducts passing between 
osmoregulatory cunts. Dorsal osmorezulatory eanatl 
extremely narrow, diam. 0.002, lying internal to 
vertal ostporeeuhitery cioal, 0.012 onax. dian, 
Transverse osmorepulitory canis connecting rh 
and Jef ventral canals at posterior margin of exw 
proglouis. Dorsul commissures vot seen. Genital 
anlige uppearing in proglottis 40 approximately; first 
mattire proglatis 160: first gravid proglotis 280. 

Genial atchim small situated i anterior hall of 
literal progloltis margin, Cirrus sae 0. L040, 1/2 
(O18) § 0.036-0.040) (0.034%, n=10), not reaching 
lou tudinal osmuregulatary carats (Poe. 20). Cities 
Un-eurmicd, distal region of greuter intemal diameter 
(han nid region; leading uncotled do iiennal seminal 
vesicle 0.015 (LO12-0.016) © 0.012 (0012-0014) 
Vas deferens grealy coiled, passing towards centre 
Ol proglottis, Testes distributed in two fateral groups, 
3-5 (4, n=20) poral and 7-11 (9, n= 20) aporal. Testes 
1).056-0.068% (0.062, n=20) in diameter; not overlying 
lemiule genital organs. 

Va Opening (6 SOT ta abriim posterior bo Girhuis 
suet distal region surrounded by cells, 0.032-0.036 
(0,035) & 0.012-0,0280 (O.0IS, n=l0), Mid region 
coed, often dikded with spenn, leading medially 


posterior lo vas deferens, greatly dilled and saccular 
anterioe to poral Jobe wf ovary (Pig. 2). Ovary bi- 
lobed, situated in progletiis midline. enlareniye tn 
CONBeCUTiVE Hattie proglottides, maximum size 
0.220 x O,080 in posterior miuture proglottides, 
Vitelhuium — similarty enlarging, — maxinnin 
dimensions O.184 x 0.080. simtated posterior uni 
distal to aporg! lobe of ovary. Sperm duet passing 
posteriorly between lobes of ovary. uterine duct 
passing anteriarly to developing uterus, Grayid 
prawlottides 1.200-1.700 x 0.200-U.440 (fig. 22) 
containing 32-50 (38, n=10) spherical eee capsules, 
O.146-0,184 x 0156-0, 192, with 4-17 (15, n=10) 
eess per cupsule, Oncosphere circular, (070,020 
indiameter, oucospheral hooks 0.006-0.008, 


Hest 
Dromaius noeyachatlandiae 
(Struthioniformes: Drorvatidie |, 


(Lathan, = T1790) 


Leceation 1h host 
Small intestine 


Erynnluey 

This species is named for Dr N. Chilton of the 
University of Melbourne for bis contribution to 
parasitology ih Australia, 


Raillietina dromains sp. nov. 
(VIGS 23-40) 


Halotype, Keith, SA, SAMA AHC S28403, 


Puritypes: Kingston, SA tae 14S. 14021" Ey, 
Coll M O'Callaghan. lO. l998. SAMA ALC 
S 28304, 828305, 31379. BMNE 2000.3_17.41-60, 


Other material examined: Wagpa, NSW, 7.9111 904, 
SAMA AHC 27716; Kinehega, NSW. Coll, 1. 
Beveridwe, S1ai.1974, SAMA ATIC lO00S5: 
Menindee, NSW, Coll 1 Beveridge, 1O.viii1977, 
SAMA AHC 11008, Pine Plains, Vie Coll 1. 
Beyeridee. Ty. 1971, SAMA AHC L051); 
Conuobolin. NSW. 27,1971. SAMA ATIC 9179: 
Wagga. NSW, Coll. | Beveridge 7.41 WO4, Senlex 
only. SAMA AHICS27716. 


Deseriprion 

Cestode Upto 45 long in ineelaxcd specimens qnd 
up to 200 in relaxed specimens. Gravid strobita 
contain 940) progloitides, In relaxed specimens. 
strobila with a musximum width of 2.12, Seolex 
O.AKX0-U.752 mm (0594 n=20) wide al suckers 
Rosiclum everted, O.436-0.448 (O397, 1=20) in 
dianteter (Fir, 23), with 124-156 (142, nel) 
hammer-shaped hooks in two rows. Larder, inner 
rostcllar hooks O,050-0,063 (0.056, t= Til) long, 


SPECIES OF RAILLIETINA FROM THE EMU] WW) 


é “; vie aos ‘ 
f ( Udideiaae HHO 
B 


Pigs 15-22. Rufllictina chilteni sp. nov. 15. Scolex with retracted rostellum. 16. Scolex with everted rostellum. 17. Rostellar 
hooks. T8. Sucker hooks. 19. Single mature proglottis, 20. Cirrus und distal vayinu. Jt. Female genitalia, 22. Gravidl 
proglottis. Scale bars =O mm, 15, 16.19 21, 22; 0.0) mm. 17. 78, 20. 


112 M.G. O'CALLAGHAN. M. DAVIES & R. H. ANDREWS 


ys 7 


Lp odao i 
ee 


Figs 23-30. Ririllictina dromiaius sp. nov, 23. Scoles. 24, Rostellar hooks. 25. Accessory rostellar spine. 26. Sucker hooks, 
27. Single mature proglotis. 28. Gravid proglottides. 29, Cirrus and distal vagina. 30, Female genitalia, Seale bars = 0) | 
mm, 23, 27, 28: 0.05 mm, 24-26: 0.05 mim, 29. 30. 


SPECIES OF RAIL TINA FROM THE EMt 414 


smatler outer books 0.043-0,054 (0.04¢8. n= 110) 
(Pig. 24), Base of rostellum armed with [5-19 (17, 
n=25) Uigonal rows oF pose-thorn shaped decessory 
spines O.008-0.0 100) (0.009, n=20) long: (Fig, 25), 
Suckers sub-circular 0.192-0,280 (0.234. n=20) x 
(108-0260 (0.231. n=20) armed with 6-12 rows oF 
hooklets varying in length fron O.008-0,020 (Pie. 
26), Neck 0. 160-0.400 long, Calcureous corpuscles 
present ju the neck und less frequently in posterior 
halal scotes, 

Privlottides craspedote. Mature proglouides wider 
than long, 0.722-1,050 (0.893, i=10) long < 0.205- 


QA70 (O.A0T n=lO) wide (Pig, 27), Chravid 
progvloniides O.920-0.9800 4  OL740-0.790, 8-10 
iweminal. urheshaped  proglottides 0.500-0.730 


(0.596) & O.430-0.600 (0472, n=10) (Bin. 28). 
Geniuil pores unikteral openings wit a tausculsr, 
phicale genital atrmm 0, 1 14-0.135 (0.123. n=10) wide 
x (.041-0.082 (0.052. n=10) (Pig. 29). extending 
from the mid-point into posterior half oF lateral 
proglowis mario, Lateral dorsal osmeoregulatory 
ganitls 0.024-0,.032 in diameler joined by transverse 
commissures. in posterion rewon ef proglorddes, 
Ventral osmorczulutory canals not seen. Hlongite 
cirrus sac. O,.246-0,97 1 (0.257) » O04 0053 (0044, 
N=10), extending anteriorly and towards but nol 
reaching lateral osnmmregulitory canal Distal region 
of cirrus narrow. remainder wide. un-coiled. Ves 
delerens coiled. voluminous. extending transversely 
in anterior margin of proglotties, Testes (O-PS. ip 
por! wod aporal yroups, 2-6 (4, n=15) poral und 4-}2 
(LO. n=15) aporal. 0041-0057 (0,048. n=15) 4 0.040- 
O.0S0, (040. n=t5). lying within lateral 
Osmorewulalary camails, 

Vain opening 10 senilal alll posterior to inate 
gent pore, Distal region of vagina enlarged. .040- 
0.050 (0.048. n=3) x 0,020-1.024 (0,022, m=). Mid 
revion smiuous. Jeading unteriorly and mediilly, 
vecasionally overlyilig estes, into a large seminal 
recephicle, O.088-0, 120 x 0,024-0.040, Lying anterior 
ty poral lohe of ovary: sperm duct passes posteriorly, 
lined with bristles. Ovary bipartite. each Tube of 
approsximatcly copeil size 0.090-0. 130 (0,106, n=10) 
* O.041-0.061 (0.050, n=10) (Fig. 30). Vitellariun 
medial, post ovariin, sub-eircular (074-0090 
(0.082) % 0.066-0,094 (0,08, n=10), Litenne duet 
passing antertorly (o developing ulerus, Ege capsules 
O<1S6 (ON36-0.190) x O14 (0080-01400, 
spheroidal, 12-18 (15. n=20) in ctich proglottis; 
vonniniig 15-22 (17. n=10) eggs. 0.045-0.05 | 
(0.049) § O.038-0.041 (0.038. n=5), Oncosphere oval 
OONT-O0LS (01S, nS) x O.O1EO.O16 CODLS, 
n=3). etnbryonie hogks 1005-0007 lon 


Flay! 
Dronains novachaltanidiac 
(Struthioni formes: Promaridae ) 


(Latham, 1) 740) 


Lacationt ae hot 
Small intestine, 


Eivaniosy 
This species ts muned ater Le Host, Deanteiies 
poveeloaltenadion 


Raillictina mitchellé sp, nov. 
(FIGS 31-48) 


Holotype: Keith, SA (3600078, 140° loo By SAMA 
AHC 528306, 


Paraivpes, Keath. SA. SAMA ALC S28307, 3140. 
BMNH 2000.5_17.61-65- 


Ofer Material examined: Yuna, SA. Coll Go E 
Ford. tix 981 SAMA AHC [118] 


Dexeriplion 

Cestodes up iy 120 long in relaxed specimens, 
Strobilu containing approximately | 120 proglomides. 
Seolex small 0.224-0, 340 (0,298, o=45) in diameter 
(Figs 31, 32), usually with eversible: rosteliun 
O.1O8-0.154 (O48, n=40) in divineter, Rostelum 
armed with 296-380 (316, n=20) harmmer-shaped 
hooks jo two rows. Larger iimer rostellar hooks 
O.009-0.012 (O.0TL n=70) longs Guten, sinadler 
rostellar books OLOOR-O.010 (0.009 7=70) lone (Fig. 
33), Surface of everted rostelurn, aertor to roster 
hooks, covered by minute accessory spines. 0.040- 
0.020 long. visible under high magnifieation onty, 
Suckers Q.055-0.088 (0.072. n=40) in diameter, 
armed with 4-6 rows of hooks 02004-0.010 long (rig. 
34). Neck absent, 

Proglottides craspedote, Mature proglouides wider 
Than long.0,A00-0,900 ((.822) & O21 K0-0.220 (0.204, 
n=l0) (Pig. 35). Crenital pores, stigle, unilateral. 
Genital ducts passing between osmoregulatory 
cunals. larger ventral osmoregulatory eanul. 0.020 in 
Mma, Unt. lying tternat to dorsal canal. 0.012 in 
mux. diam. Ventral canal joined hy. tranverse 
osmoregulatery canal ih pestertan marein al 
prowlottides: Transverse dorsal canal nor see 

Genitalantage fitst appearing in proglotides 400: 
S20. Mile and female vendatia mature tn 
progiottides 640-750. First gravid proudotides L000 
Will) 100-120 gravid) proglotides terminating with 
10-20 coimpuel proglottides becoming provessively 
longer than wide. 

Genital auridny Soiull, sitated in aptevir halt el 
lateral proglonis. (musi, Chris sac OTS 2-0 [76 
(161) a QhOST4LO44 (0,038) n=10) (Pig, a6) nl 
reaching vealed psmorceulutory eqn, Bistel peenin 
of cirrus Tined will) Spines, of greater titertal 
ameler (han siniais pa) nepian: proxsnul ceein 
forms spherical miceind serial \esiele OR is 


tl4 M. G. O CALLAGHAN, M, DAVIES & R. H. ANDREWS 


Figs 31-38. Raillietina mitchelli sp. noy. 31. Scolex with everted rostellum. 32, Scolex with retracted rostellum. 33. 
Rostellur hooks. 34. Sucker hooks. 35. Single mature proglotlis. 36. Cirrus and distal vagina. 47. Female gemttaha. 38, 
Terminal gravid proglottides, Scale bars = 0.1 mm, 31, 32. 35. 37, 38: 0.01 mm, 33, 34: 0,05 mm, 36. 


SPECIES OF RAILLHETINA FROM "THE EMU Hs 


(0.040) x 0.020-0.032 (0,026, n= 10). Vas deferens 
slightly coiled at midline of proglottis. Testes 0.048- 
0,060 (0.053, n=10) in diameter, dorsal to and 
overlying female genital glands. Testes 5-6 (5, n=2()) 
per proglottis, one frequently overlying vitellarium 
with additional testes, | poral and 3-4 aporal, 
Vagina opening to genital atrium posterior to cirrus 
sic, Distal region, dilated, 0,082 x 0.024-0.032. mid 
revion, narrow. straight, leads. medially posterior to 
vas deferens, terminating in fusiform seminal 
receptacle 0.124-0.152 (0.143) x 0.024-0.036 (0.028, 
n=10), Ovary bilobed (Fig. 37), Poral lobe 0,064- 
0.096 x O.F040.112. consisting of 1-3 transversely 
elongule lobules. Aporal lohe, 0.112-0,160 x 0.125- 
0.160 consisting of 3-4 Jobules. Vitellarium 
irregularly ovoid, 0,056-0.076 (0.070) x 0.040-0.056 
(0.049, n =10), Mehlis gland spherical, anterior to 
vilellarium O.024-0,032 (0,028, n=10) in diameter. 
Uterine duct passing anterior to vitellarian, 
terminating dorsal to ovary. Uterus absent. Gravid 
proglottides wider than long O.O84-1,120 (0,964) x 
0272-0360 (0.316. n=Ll0) containing 9-15 ez 
capsules Q.140-0.170 (0.150) x 0.100-0.150 (0130, 
n=!0) each with 12-18 (15, n=10) egus 0.041-0,049 
(0.045) x 0.035-0.045 (0.040, n=10). ‘Terminal 
scoments shrivelled (Pig. 38). Oncosphere 0.015- 


O<.Ol8 (017) x O.OIE-O.017 (0.016. n=lOis 
oncospheral hooks 0.004-0,006 long. 

Host 

Dromains  vovechollandiae (oatham, 1790) 


(Struthionitormes: Droniatidae), 


Location in hast 
Small intestine, 


Enyinalogy 

This species is named for Sir Mark Mitchell in 
acknowledgement of support of this project through 
the Sir Mark Mitchell Poundation. 


Comparison with other species 


OF the species of Reilieting with hosts tn the 
Struthioniformes, Ro dramas sp. nov. resembles 2. 
easuarii Yound in the New Guihean cassowary, 
Casuarius picticollfy in the size of the costelir hooks 
(Kothin 1923), However &, drone is smaller than 
RK, casuaril, bas fewer rostellar hooks (142 y, 250), 
fewer and smaller testes and there are fewer eees per 
cupsule. Poraniella appendiculate Vahemann, 1909 
Ueseribed from an unknown bost in New Guined is 
similar in sive tO Ro dramas with TAO) rostedfar 
hooks D.036-0,043 in leneth, However 2 
appendiculata Mis only one ex per capsule 
(diagnostic lor the genus Parodie a smaller 
Chrus sae and more testes than Ro eres 


Raillietina chiltoni sp. nov. resembles R. 
infrequens (Kotlin, 1923) in the size of the strobila. 
sclex und rostellar hooks, the huniber of rostellar 


cS 
xz 
-= 
=~ = 
ms 7 Ff 
ch fic 
ec We ra 
=< i: cS 
=3 -= AF 
2 ~~ 
DS mm > 
& oF 
=n orl 
.g Seu 
ea Sf es 
_= = 
a = > 
cost —™ 
='5 ™ Ww 
2 == 1S 
wists 3D 
ec 2 7 i> 
45 cr ere cl 
ae a Ta 50 
3 St Sone 
= os Ssoaic 
E Lm 
= +S 
= Ose 
- + 
,c oo = +t , 
ex ms int = 
=) oS cs 
= = 
2ayjoc Ss 
>t as 
v/s <4 
ae = 
= i 7 7 7 
Zl Ss 2¢ 9 
= el fom aa om 
= pend = SR 
ss eos Soma 
= ch, 
co oS 
: rr 
5 cit mom! a 
= rm ml ae S 
oi,os =0 
= a * q 
a os 
— os 
ome | or 
ai rr, 
= = 
5 i 
Ot o£ 2 
—t as re) 7 
os ri =i 
os <a tS 
+3 
4 
re ost 
ao sc = 
a an = 
2:0 o 
os = 
om se ot 
j meri om oa | 
= 27] pcg? Se 
= St) oc <= << 
5 S125 at 
a D>; a oa aa oe 
= =e a moe = 
= =s oo —-ocde 
= =c aa £2 
= = — Tas 
> = oso 
= = iad 
> ee 30 Sl 
<a cae ma = 
2 Sf So eS 
Ss zijcocs =< 
2 
= eo 
po == PA 
ad 1 =. 
=> x S 
= at = = 
“i 4A 5 
= eHoBes 
= ren pr— = 
= 1 Se 
3 ro DS 
= hat Ms 
— _ a= 
=. I a= 
Prt _ ae 
= = ee 
= = Ss 
4 5 £5 
a = E= 
” 9 —t to 
— sa 4a 


116 M. G. O°CALLAGHAN, M. DAVIES & R, H. ANDREWS 


hooks and testes. However, R. ciiltoni differs from 
R. infrequens in the size of the cirrus sac (0.108 x 
().038) compared with (0.180-0.200 x 0.060) in K. 
infrequens. Io addition, the cirrus of R. chiltont has 
no armature and the internal seminal vesicle is 
smaller (0.015 x 0.012 v. 0.054 long). Raillietina 
chiltonit has a larger rostellum (0.383) than R. 
infrequens (0.250) and has testes in distinctly aporal 
and poral groups that are never in the midline. 

In the struthioniformes, Cotugaia collint can be 
distinguished from Raillietinad species by the 
presence of two sets of bilateral genital organs. 


The species of Raillietina described here can be 
distinguished from all congeners in the 
Struthioniformes by the size and number of the 
rostellar hooks, size of the scolex and size of the 
cirrus sac (Table 1). 


Acknowledgments 
We wish to thank I. Beveridge for his advice and 
comments in the early stages of this study which was 
supported by a grant from the Sir Mark Mitchell 
Foundation. 


References 


Charke, EoD, Kibiy, E. J. & Prinuirs. S. N. (1996) 
Necropsy finding in ratites (70 cases), Agri-Practice U7, 
34-35. 

FUHRMANN. QO. (1909) Neue Davyaineiden, Centra/bl, 
Bakteriol. Parasitenk. J Abt 49, 94-|24, 

Jonrs. A. & Bray, R. A. (1994) Family Davatneidac 
Braun, 1900) pp. 407-441 /n “Keys to the cestode 
parasites of vertebrates” Khalil, L. F., Jones, A. & Bray, 
R.A, (Eds) CAB International, Wallingford, UK), 

KoTLAN, A. (1923) Avian cestodes from New Guinea. IT. 
Cestodes trom Casuariformes An. Trop. Med. aiid 
Parasitol, 17, 45-57. 

Kragee, H, (1869) Bidrag til Kundskab om Pulgenes 
Baendelorme. K. Danske Vidensk Selskab, Skrifter. 
Naturvidenskab. Og Math, Afdel. 8, 249-363, 


MANNION, P. F, Kent, P. B., BARRAM, K. M.. TrAprel, P. 
C, & Buran, G. W. (1995) Production and nutrition of 
emus pp. 23-30 Ja “Proceedings of Australian Poultry 
Society Symposium, 7”. 


Scumipr, G. D. (1986) “CRC handbook of tapeworm 
identification” (CRC press, Inc., Boca Raton, Florida, 
USA), 


SHank, S. M. (1998) Infectious diseases. and parasites of 
ratites, Ver. Clin. North Am, 14, 455-483. 


Tuuny, T. N. & SHANE, S. M. (1996) Husbandry practices 
as related to infectious and parasitic diseases of farmed 
ratites, Rev. sci, tech. Off. int. Epiz. 15, 73-89. 


A NEW BLOOD-FLUKE, CARDICOLA FORSTERI, (DIGENEA: 
SANGUINICOLIDAE) OF SOUTHERN BLUE-FIN TUNA 
(THUNNUS MACCOYID IN AQUACULTURE 


By THOMAS H. CRIBB*, MARTIN DAINTITH? & BARRY MUNDAY 


Summary 


Cribb, T. H., Daintith, M. & Munday, B. (2000). A new blood-fluke, Cardicola 
forster1 (Digenea: Sanguinicolidae) of southern blue-fin tuna (Thunnus maccoyii) in 
aquaculture. Trans. R. Soc. S. Aust. 124(2), 117-120, 30 November, 2000. 

Cardicola forsterl sp. nov. (Digenea: Sanguinicolidae) is described from the heart of 
captive southern blue-fin tuna, Thunnus maccoyii (Scombridae), from South 
Australia. The new species is distinguished from other species of Cardicola by its 
very extensive testis, the length of its oesophagus, the length of its gut caeca and the 
form of its ovary. Cardicola smithi appears to be associated with heart and gill 
lesions’. 


Dransnetions of He Raval Society af S. Aust (2000), 124(2). 117-120. 


A NEW BLOOD-FLUKE, CARDICOLA FORSTERI, (DIGENEA: SANGUINICOLIDAE) 


OF SOUTHERN BLUE-FIN TUNA (7: 


by Thomas H. Crisp’, 


HUNNUS MACCOYIH) INAQUACULTURE 


Marin Darintiri’ & BARRY MUNDAY! 


Summary 


Crips. T. H. Datsitin, Me & Muspay, By 


117-120, 30 November, 2000. 


1 ! (2000) A new blood-lluke, 
Sanguinieotidae) of southern bluelin tuna (Mins meccovil) io aquaculture. Tris. AL See. 


Cardicala forstert, (Digenea: 
NL AMtiyt. 124 (2). 


Cordicala forstert sp, noy, (Digenen: Sanguinicolidae) is deseribed trom the beart of captive southern blue-fin 


tuna. Fhiiis diicces® (Scombridae), trom South 


Australia, 


The hew species is distinguished fon other 


species of Cardicaly by us very extensive tests, the leneth of its oesophagus, (he length of its eul cueca und the 
form of its ovary, Candicola sate appears to be associated with heart and gill lesions!, 


Introduction 


The southern blue-fin tuna (Times merceayii) hiss 
been used for aquaculture in southern Australia since 
1902, 
juvenile fish and their subsequent fattening over a 
period oF 6-9 months. The tuna have been subject to 
remarkably few. diseases so far Here we report a new 
parasite, a sunguimicolid: blood-Muker the associated 
pathogenesis will be described elsewhere. 


Materials and Methods 


Trenmmtodes were collected from the hearts of 
lreshly-killed fish hosts and fixed by pipetting them 
into near boiling phosphite bullered saline followed 
by immediate preservation in LO% neutral buffered 
formalin. Whole-mounts Were stained witht Mayer's 
haematoxylin, cleared with methyl salicylate and 
mounted in Canada balsam. Specimens tor 
seclioning were embedded in parallin wax, stained 
with haematoxylin and eosin and mounted in 
DEPEX. The following abbreviations are used: 
AHIC, The Australian Hebnintholovical Collection at 
the South Australian Museum, Adeliide: OM, 
Queensland Museum, Brisbane, 


Department oF Microbiolowy and Munpsitotouy, The University of 
Queenshine Beishane Ohl 472 
Speneer Institute of PARE, 2 Limi St, Mort Lineolo SA F6tis 
School of Bromediea! Seienes, University of ‘Hosmaditit, Locked 
Bay {320 Launeeston Ths. 7250, 

'Comgurrr, So Bo 11999) Pistopaitiolowical chanies ine und irmnune 
mesponse bh Sothern bliin Wine COREY Hee vall infeese 
WHR Cadivole sp. (Olio!) SaAplinigolidde) Petiekirs thesis, 
Valvers ty ob Fasmiariia Lanipub. |, 


Vhe industry is based on the capture of 


Systematics 


Family 
Cardicala Short, 


Sanguinicolidae von Gratf, 1907 
1953 


Cardicala farstert sp, nov. 
(FIG, |) 
maceover 


Type est Seombridiae - Thin 


(Castlenau, 1872), 


Type locality: Ol Rabbit Island, South Australia, 
34° 36'S, 135° 59' E 


Other localities: Louth Island, South Australia, 34°" 
35° 8, 135° 57' BE. 


Site. heart, 


Material examined: V5 adults including 3 sets of 
histological sections from Rabbit Is. 1) trom Louth 


Is, 


Deposition af specimens: Holotype and 9 
paratypes (including 3 sets of sections) AHC 28331 
~ 28340) 5 paratypes QM G 21 8017-21. 


Description 

(Measurements in tin of 10 gravid adults (means 
1n parenthesis )) 

Body lanceolate. highly compressed dorso- 
ventrally, almost Mat ventrally and convex dorsally, 
2512-3688 (3228) x 608-928 (759), Tegumental 
spines restricted to distinel yentro-huteral rows (Pig. 
lac, d). Nerve commissure dorsal to oesophagus 
and just posterior to anterior end of body: main nerve 
hundles highly prominent in anterior halh of hody 
und: discernible aliost tu posterior end of body, 


Ibs 


T. H. CRIBB. M. DAINTITH & B. MUNDAY 


POP ag, ats 
Ss a 
PAP Pep, etiy, 

- 


Fe J OG ag Hop, 


ms 


HN 


i 


ms 


ventral yiew. C. Marginal spines, ventral 
= 500 mm A; 200) mm B; 50 mm C.D, 


~ LLCs, 


Vin. 1. Condicele farstert sp. noy. A. Adull, ventral view. B, Terminal genitalia. 
view, D. Marginal spines and lateral muscles in transverse section. Scale bi 
fernale pore, In — latecal nerve, mf — niusele libres, mg — Mehlis’ gland, mp — male pore, ms — 
imei spines, o — ovary, oe — oviduicul chamber, oe — oesophagus. sv ~ semunul vesicle. L — lestis, u 


Abbrevierenss Tp 
silellariun, val — vas deferens, vd — yielline duet, 


ANEW BLOOD PLURE PROM SOUTHERN BLUOE-FIN TUNA WW 


Moulh inconspicuous, opening ventro-subtermi nally. 
Ovsuphugus highly museuhiun straight ki6-1146 
(O14) long, 24,1-33.4 (31.59% body leneth, Cueca 
Heshaped. Siiotis: extending witeriol|y to 672-928 
(834) from unterior end of body; posterior cuecu 
usually of distinctly uneven Jength, exteading to 
S60-912 (777) frum posterior end of body. Testis 
sully indistinel and ditficul to discern, intra ay 
extmbcaveal extending from ovary posteriorly tp just 
behind nervous commissure anteriorly: penetrated 
by dorso-ventrally orientated muscle hres 
throughout Vas deferens brodd,  prontinent, 
ofan Ud ventrally to tests, rani: sintously 
posteriorly, dorsal to ovary and: ventral to uterus 
before cutering seminal vesiele. Cirrus-sac absert. 
Sentinal vesicle clongute, evenly curved, b16-235 
(174) 8 26-64 (45). Male genital pore sinistro-dorsal, 
close to hateral margin of body. Ovary trregularly 
lobed, penetrated by dorse-ventrally ortentited 
musele Hbres throughout, (38-321 (237) x 263-462 
(376), Oviducl originating posteriorly and passing 
posteriorly jmedidtely to expand inte ovidieal 
chamber ol viarnuble size containing either oveytes 
Lind perhaps ¢ygotes) or sperm: it filled with sperin. 
chamber may become relitively enormous — up to 
‘57 s tlh. Duet emerging from oviddeal Chamber 
joined by vitellineg duct (hen tlm adtera-medially 
vis) fermiqag paiwpe surrounded by prominent 
Melvin’ wand cells. Vitelline follicies ane dillase ane 
throughow! body from level of anterior margin al 
ovary (sometimes fiteral lo ovary ay well), dersal 
unl verteal ty testis, und as Cur anteriorly ay nervous 
commissure. Vilelline duct passes ventral to bests 
and avery, Llerusy Tiled with vos. wandine 
sinuously ta ovary and ther posteriorly to female 
genial pore, directly anterior to and well separated 
fram male pore. Egws very thinewalled and 
compressed against euch other [9-27 (23) . 11-16 
(14) Exeretory system ot observed. 


Payimlags 

‘The species is mamed for Mr Ron Parster South 
Australian ium firmer in recosmoon af lis 
contibuboa to the development of the enlightened 
Miunlagement Gb euptive Lond, 


Discussion 


The pew species shows close alfiity wilh the 
sens Cevedioola Short. 1953 and is here Wentilied as 


a new species in that genus, Caradivela os 
distinguishable Cram other wenera of faring 


Suneninicohddge by the combimition ol un A-stiaped 
ful a sitele Jartely jter-caceal testis. lack of a 
CUTUS Sc, postenvarian uferis and sepucite 
submiveinal genie: pores (Herbert ef af, 18944. The 
present species agrees will all hese characlers excep! 


that (he lestis ts both tnler- and extrascaecal, although 
one other species of Ceveticola, Co nivgilry Yanauts. 
1970, olsci fas ao purtly extri-cucenl (esas, Only 
species of Deontaevliy Linton, TIO and 
Pearsenellum Overstreet & Kae, 19K9 also have 
exird-caccu) testes, Species of Dewtucviiy Linton. 
O10) huve a testis comprising “longitudinally 
clongutedt wings” (Yamdguti 1870) which extend 
literal to the cacen ina form entirely different fram 
that seen in the present species. The distribution of 
the vitellariuin ih the sale species of Pearvenel/iiam, 2 
Carventimn Overstreet & Roe. 1989, is conparable ty 
(hat of the-presctit spectes, bejng both witerior to the 
cutcal brlyrcation wad daberal fo the posterior eaeci, 
but that genus ts distinct from the present species i 
possessing a cirras-Sac [Overstrcet de Kaie 199). 

Curndicola wis erected (Short 1959) lor Co lari 
Short, 1953 irom lwo species of Cynayerut 
(Sciaeniday). Subsequently, nine further species have 
been deseribed or combined with this genus (Smith. 
1997a.h) nanely OC ahi Yanuguu. 1970, C cardicale 
(Menter, 1947) Shoet, (53, 6. chaetodontys 
Yumagul, W870. C, ceripidiets Munter, 1354, C, 
wrandi« Lebedev & Mantaey, 968 (not mentioned by 
Smith, }097a.b), C. vaieiis Yarnweu, 70. C. 
whined Manter 1954, Co cegiinntnd Lebedev & 
Maines, (96% and C. bresrtievisty Knoll & Appate, 
192. Two of the species. €. gf and ©. conerneia 
have been reported trom tina amily Scombnidae. 
subfamily Thonninge). 

The present species is unrcdiately distinguished 
from all these species by the more extensive 
distribution of the testis which ts both anteriar to the 
cuecal bifurcation und well lateral to the posteriorly 
direeied cacga, In this stucly, however, we found the 
disiribulin Of the (estas excecdingly diteull to 
interpret and, though we find i convincing as a 
species-level churaeter, We cimelude That it is nat an 
ideal character for recognition of species in this 
genus. Fortunately, several other characters also serve 
ty distinguish this species, The leneth of the 
vesophigus, vecupying 29-34% af the body tenth 
serves to-distingnish it from Co adi inwhieh wis very 
short (approx. (So) and species tn whieh wis very 
long ©. curdieate (414) and Co lariet (0%), The 
reéhitively very Short postermr cuca: al OC) conerienner 
and the short divergent unteriurly directed cacea of C 
breyiliensts, CC. checetodeediy, Co miugilts und ©, 
whiten are distract Cronr the relatively lone poster 
cucen and the paralle) anteriorly directed cieca of the 
present species. The present species generally 
resembles CL cepredacis but hits ud rekiovely larger 
and irreeularly lobed rather than samouth ovary ind 
has relatively shorter anteriorly directed wal edewu. 
Hinally, ©. grandis rom ac malin OMakerrie sph ty a 
vouch larger worm (4.77.0 mim Jong compared: with 
2.5-3,7 mm tor the present species). Us general 


12) T. H. CRIBB, M, DAINTITH & B. MUNDAY 


organisation is similar to that of the present species 
except that the testis is described as a single muss 
immediately behind the caecal bifurcation. 
Overstreet & Kote (1989), Herbert ef al. (1994) 
and other authors have frequently referred to the 
presence of numerous dorso-yentrally orientated 
“ducts” or “structures” in sanguinicolids. These 
often pass through the gonads. Such structures are 
abundant in Cardicola forsteri and are here 
interpreted, as suggested in Herbert ey a/. (1994), as 


muscle fibres, This interpretation appears reasonable 
in terms of the appearance of these refringent 
structures and in terms of fttnction in trematodes 
where the requirement for flattening against the 
walls of blood vessels is clearly of great importance, 


Acknowledgments 


We thank D. Scott and T. Wright for assistance 
with the preparation of the specimens. 


References 


HERBERT, B, W., SHAHAROM-HARRISON, FL M. & 
OvnrsTREET, R. M. (1994) Description of a new blood- 
fluke, Cruoricola lates neg, nm. sp. (Digenea: 
Sanguinicolidae), from sea-bass Lates calcarifer (Bloch, 
1790) (Centropomidae). Syst. Parasit. 29, 51-60. 

OvERSTREET, Ro M. & Koi, M. (1989) Pearsonellim 
corventum, gen, el sp, noy, (Digenea; Sanguinicolidae), 
in serranid fishes from the Capricornia section of the 
Great Barrier Reef. Aust... Zool. 37, 71-79. 

Snorer, R. B, (1953) A new blood fluke, Cardicola laruein 
#, nm sp., (Aporocotylidae) from marine fishes. J. 
Parasitol, 39, 304-309. 


Smith, J. W. (19974) The blood flukes (Digenea: 
Sanguinicolidae and Spirorchidae) of cold-blooded 
vertebrates: Part 1, A review of the literature published 
since 1971, and bibliography. Helminth. Abs. 66, 255- 
294. 

(1997b) The blood flukes (Digenea; 
Sanguinicolidae and Spirorchidae) of cold-blooded 
vertebrates: Part 2, Appendix 1; Comprehensive parasite- 
host list: Appendix [: Comprehensive host-parasile list. 
Helminth, Abs. 66, 329, 

YAmAGutt, S, (1970) “Digenetic trematodes of Hawaiian 
fishes” (Keigaku Publishing, Tokyo), 


A NEW GALL MIDGE SPECIES (DIPTERA: CECIDOMYTIDAE) 
INFESTING FRUIT OF PUNTY BUSH, SENNA ARTEMISIOIDES 
(CAESALPINIACEAE) IN AUSTRALIA 


By PETER KOLESIK* & SAUL A. CUNNINGHAMf 


Summary 


Kolesik, P. & Cunningham, S. A. (2000) A new gall midge species (Diptera: 
Cecidomyiidae) infesting fruit of punty bush, Senna artemisioides (Caesalpiniaceae) 
in Australia. Trans. R. Soc. S. Aust. 124(2), 121-126, 30 November, 2000. 

A new species of gall midge, Contarinia sennicola Kolesik, is described from fruits of 
the punty bush, Senna artemisioides (DC.) Randell in south-eastern Australia. Yellow 
larvae of Contarinia sennicola live within fruit capsules of Senna artemisioides and 
prevent seed formation without causing superficial deformation. In 11 localities in 
New South Wales, all plants examined were infested by the new species, with the 
level of damaged fruits being between 10 and 90%. Despite the high frequency of 
infestation damage caused by the new species, it did not appear to limit substantially 
reproduction of the host plant, as indicated by the overall large seed production. 

Key Words: Gall midge, Cecidomyiidae, Contarinia sennicola, Senna artemisioides, 
punty bush, Australia. 


Tractors of the Reval Soctery af S. Aust (2000), (2402), 121 126, 


A NEW GALL MIDGE SPECIES (DIPTERA: CECIDOMYIHDAE) INFESTING FRUIT 
OF PUNTY BUSH, SENNA ARTEMISIOIDES (CAESALPINIACEAE) IN AUSTRALIA 


by PETER ROLESIK: & SAUL A. CUNNINGHAM 


Summary 


KOLESTK, Bad Cunminciian, SoA. (2000) A new gall midge species (Diptant Ceerdoniyiidae siifesting fait of 
punty bush. Seve ertemivieidey (Cacsulpiniaecac) i Australia. Tris. Ro See S, Aust (2442) 12 120. 10 


November, 2000), 


Anew species afgall midge, Comurinie seniicola Kolesih, is deseribed from fruits of the panty bush, Sere 
aremistoides (DCo) Randell in south-eastern Australi. Yellow larvae oF Comariode seni le live within trait 
vapsules OF See cmremisioides und prevent seed formation without causing supertickl deformation, Wy Ul 
localities Th New South Wales, ail plants examined Were Tifested hy the new species, With the level oF dimaged 
froits being behween Hagin 906. Despite Ihe high frequency of Tifestalion chimage cunscd By the Hew species, 
Hedi tot appear (7 Limit substantially reproduction of (he host plant, as indicated by the overall large seed 


productions 


Kiy Words: Gall ndge, Cecidomytidie. Connienid senmeos. Serer arrenistoides, punty bust. Austr 


Introduetion 


A new species of gall midge, Conterinia sennicole 
Kolesik, is deseribed from fruits of the puity bush, 
Senna untenisiaides (DC Randell in south-eastern 
Austrilia. The new gall midge species was found 
independently by SAC during a study of the effect oF 
habia fragmentation oy repreduction by plants in 
central New South Wales during 1997 and [9% and 
by PK in 1998 during a South Australian Museum 
veologicul survey in the Seotie Sanctuary, New 
South Wiles. The host plant, Semme artemisiondes 
(DC) Randell (Cuesulpiniuceac). commonly known 
us The punty bush, is an endemic species widespread 
throdgh the takind of mainkind Australia (Harden 
1990). 1Lis a variable spevies, with 10 subspecies and 
Aothosubspecies  fecagiised (Harden 1990). 
inchidime what wis curler consilered to be Caso ver 
eremuphila. Senna artemisivides is invasive in 
erized land in Western New South Wales 
(Cunmimghiam ef ad. 198)) und commenty oeeurs in 
distirbed arenas such us rowdsides. 


Materials and Methods 


Branches of Senna aremisioides tearing lruits 
infested’ wih larvae ef ihe new species were 
collected in the Scotia Sanetuary. New South Wales 


Dyparkiivnt Of Pveticntiuire, Viteuliane: aiid) Qunolory Wade 
(yoy. Phe Liniversity vit Adelaide PM) Clete Osnatindl SA 
S0tb Lill: Peter Roles @sle bride ed au 

CSIBO Lntonoligy. GPO Bos (700. Canberra ACT 2601 b-iitt 
Suh. GOTH TI HAs irene 


int November 1998. Branches were brought lo the 
laboratory and the frufts processed ta ane of two 
ways. A small number was dissected and the larvae 
preserved in 70% ethanol, A hirger number was cut 
open and the larvae transferred with enlomological 
forceps into rearing pots conning wet sand inte 
which they dag themselves. Pupition took place in 
the sand. Emerged udults together with pupal skins 
were preserved i 70% ethanol, Canada balsa 
mounts of lype specimens were prepared according 
to the technique oullined by Kolesik (1995a). The 
types are deposited in the South Australian Museum, 
Adelaide (SAMA) and the Australian National lseect 
Collection. Canberra (ANIC). Dried samples of 
infested plants are deposited inthe Stare Herbarium 
of South Australia, Adelaide (AD). Measurements 
refer (othe holotype ward paratypes. 

To determine the distribution of Contarinte 
senmicola 20 fruits were vollected from two plants al 
euch of LP sties (ce, 40 friits) i Deeember 1997 ane 
199M, Sites ranged from a large reserve (ie 
Nombiinie Nature Reserve >140,000 had ta narrow 
roudside strips of vegetation in central New South 
Wales (Table 1). ALL Hrimts were Opened and inspected 
for the presence of Contarinta semnmicela larvae, 
Because it is possible to overlook hiryue al they are 
presentin small numbers or when they wre young arid 
thus very small, (he Frequency of occurrence recorded 
here is likely to be a conservative estimate. 


Genus Comarinid RondaniL bso 
Contarinia Rondani. 1860; 289 
fype species: Tipila lati De Geer, 1776 by origial 
designation 


123 P. KOLESIK & S.A. CUNNINGHAM 


Tasue be dvesation af Senna artemisioides fruity by larvae of Contarinia semnicoli. 


Yeur Site Latitude Longitude Se Lruits with larvae 
(plant 1, 2) 
198 Stickpoole SF 34° S06" 145° 50.0 95,82 
19Os Roadside near Stuekpoole SR 33° 4h" 45° 51,2° 55.35 
1997 hoadside near Denny SP 34°01, 9" 145° 51,2! 45.45 
190% Pulletop NR 33" 58.1 146° Ua.o! 45.80) 
1907 Roadside near Pulletop NR 33° 56.2" 14 074! Wh 50 
1997 Nombinnie NR 3a 020 he 06.0 65, 65 
1907 Comipaita SE We Si" IQ" 244° 75,45 
1907 Roadside near Conupuira Sh V4 STR LNG 23N" 55, 20 
1907 Roadside near Taleeban 33° 53.3! l4o> 28." Ah, 25 
1997 Ciibhatta NR 33° 3X3" h46° 43.0" 35, 44 
1997 Roadside near Gubbatla Nie 43° 35,3" 146° 31.5" 45, 30) 


Sk = State Forest. NR = Nature Reserve 


Contarinia is a large, worldwide genus used as i 
catch-all category for the tribe Cevidomyiini. I 
includes species with long, tapered evipositors, 
bifilar male flagellomeres and terminal larval 


papillae consisting of two pairs of setose and one of 


asetose, stublike papillae. So far 12 species of this 
genus have been found that are native to Australia, 
with TL of them forming a natural group feeding on 
inflorescences and seed-heads Of grasses (Harris 
1979), The new gall midge together with Coutarinie 
bursariae trom fruits oF  Burseria spinosa 
(Pittosporaceae) (Kolesik }995h), are the only mon- 
grass feeding species of this genus known from 
Australia, 


Contarinia sennicola Kolesik sp. Woy. 
(MIGS |-9) 

Holarpe. S, Scoug Sanctuary, New South Wales, 
Australia (30°) 8, T4011 B) Tea 1998, P. 
Kolesik, reared front fruits of Seana antemlstardes 
(DC.) Randell, larvae collected 21,8). 1908, (SAMA, 
121480), 
Panuvpes; 2S a3 22. 3 pupal skins (SAMA, 
[2146 1-12)488), 2 4d. 2 FPL 2 pupal skins 
(ANIC), same data but emerged [3.xii, 1998) - 
17ai.1999; 3 fiarvue, (SAMA, I121489-121491), 2 
larvae (ANIC), collected with holotype, 
Ohler meterigk galls, collected with holotype, 
ADI07823, ADJO7824 (AD). 


Mele (Pigs |-4y 

Colour, Head yellow with eyes dark brows, 
unlenmic brown, thorax brown, ubdemen with 
selerotised parts grey und non-selerotised) purts 
yellow. 

Head: Postverteal peak present. Antenna: seape 
and pediee! as broad as Jong, Magellonmieres 12 in 
number, first dnd second fused: circumfilar loops 


reaching midlengih of next node. Pulpus four 
segmented, Eve facets rounded, close together. eye 
bridge 8 - 10 facets long. Labella large, triangular io 
frontal view, pointed tipically, each with 7 - 9 Jateral 
setae, rons with 4 - 6 setae per side, 

Thorax: Wing length 1.2 mm (10-13. 0 = 5), 
width 0.8 mm (0.4 0.5): vein C broken at junetare 
with Rs, Rs barely visible, in form of pigmented area, 
My, not visible; C. Rs, Cu pigmented. Chiws simple, 
curved at midlength, empodium as long us claws. 

Abdomen: Sclerites with a pair of anterior trichoid 
sensilla and sete more or less evenly distributed. 
Genitalia: gonocoxites cylindrical, setose, setulose: 
vonoslylus about sume width entire length, sparsely, 
evenly setose. with small setulose area at base, 
distally With strong looth: Cerci rectangular slightly 
broudened distally, Separated by shallow, wide 
incision, setose distally, setuloses lypoproct Meshy. 
bilobed, lobes round, euch with few setae upicully, 
setulose; aedeagus tapered distally. shorter than 
COU L, 


Female (Pigs 7.8) 

Head: Flagellomeres. with necks about '/) length 
nodes. Cireunifila appressed, consisting. of two 
irunsverse rings connected by two longitucinml 
bands. 

Thorax; Wing length LS mm (i4- 1.6, 1 = 5). 
width 0.6 nin (0.5 - 0.6). Colour and other characters 
us inmate, 


Pupa (Fig. 9) 

Colour: antennal horns, prothoracic spiracles, 
dorsal spines light) brown, renuining parts 
unpizmented. Length 8 mm (1.5 - 2.2, 1 = 5) 
Antenmal horns simul. angular seleroused. Cephalic 
papillie with long, robust setae, Two pairs of lower 
facial papillae, one of each setose and one asetose, A 


ANEW GALL MIDGE FROM SENNA ARTEMISIOIDES 


att) 
PecRirt ni a 


Figs 1-4. Male of Cantarine sennicala, |. Wing. 2, Last tarsomere with claw and empodium. 3. Sixth flagellomere, 4. 
Genitalia in dorsal view. Scule bars = 50 um (Fig, | = a. Pigs 2.4=b. Fig. 3 =e). 


pair of triplets of lateral facial papillae, one of each 
triplet with minute seta, two asetose. Prothoracic 
spirucle long. narrow. trachea ending al its apex. 
Intequment of abdominal segments covered with 
spiculac, slightly larger and denser dorsally. Second 
to eighth abdominal segments with sclerotised, 
simple dorsal spines, 


Last instar larva (Pigs 5, 0) 

Colour: yellow. Length 2.2 mm (2,0 - 2.4. n= 5). 
Integument smooth except several ventral transverse 
rows of spiculae on anterior half of abdominal and 
second and third thoracic segments. Head with 
postero-lateral apodemes as lone as head length 
Spatula with long shaft, narrow apical enlargement 


124 P. KOLESIK & S. A. CUNNINGHAM 


= 
ate 
ENSlenen F 


Figs 5-9. Contarinia sennicola. 5, 6 larva, 7, 8, female, 9 pupa. 5. Terminal segment in dorsal view. 6. Sternal spatula 
with adjacent papillae. 7. Sixth flagellomere. 8. End of ovipositor with cerci. 9. Anterior part in ventral view. Scale 
bars = 50 um (Figs 5, 6 = a, Fig. 7 = b, Fig. 8 = c, Fig. 9 = d). 


ANEW GALL MIDGE PROM SENNA ARTEMISIOIDES ag 


with small rounded Tohes divided hy shallow 
ineision. Busi papillae typical for superutbe (Gagne 
1989), terminal papillae: One pair stublike, three 
pairs With tinck setae, Anus ventral, 


Envmelouy 

The specific name isa combinatiin of Sena, te 
venene name of he host pluntamd “cok. Latin for 
iweller/inhabitant 


Fruit damage, bielogy and geographical distribution 

Larvae of the new species five inside fruit capsules 
of Senna uriemisieides wilhoul causing any apparent 
dleformution of the cupsule but reducing the number of 
seeds that develop. In tamsmnitied light, 5 - 50 larvae 
aan be eouyised feeding inside the capsule. Late 
instar larvae cree single or multiple openings m the 
vapsules and leave the fruits by junipiig up several 
centinetres. Pupalion takes place within the soil, The 
higloey and infestation symptoms of the new species 
are very simple to hase of its Australia congener 
Comarinia bursariis a species that infests Iruit 
capsules of Aunsearia spinesa Cay, (Pittosporaceuc) 
(Kolesik 1995h), The incidence of Coverire 
senicola larva in fruits cxzumined was very hah All 
(1 (he 22 plants sampled, in sites separated by as much 
i LOG ki had larvae in one or more fruits (Table | ). 


Remarks 


Coniariia sennicvele ditters morphologically from 
the other known Australian. non-grass leeding, 
congener C. bursariae in several characters. In C. 
senmicald. the male céret are broudened distilly, the 
female cercr have one long proximal sensory. seta 
each and the Jurval spatula bas rounded apical lobes 
dunce attrow. equally wide shalt, by. bursartae. (he 
Inwle cere) ure not browdened distally: the Temule 
cere) live (wo short proximal setae each and (ie 
larval spatula bas angular apreal lobes une at distally 
widened shall. 


The frequency of aborted and darmaged seeds in 
fruits of S. ateniyieidey occupied by C senivale 
lurvac suggests thatthe larvae might be responsible 
for reducing seed production i (his leaumtnous 
pliant, In some plant species, especialy in legumes. 
predispersal seed predation by insects is an 
nportint factor in low seed production (Auld 1983, 
YK: Culiinghain M87, 2000b). Cuopinghan 
(2000u) found high lovels of predispersal insect seed 
predation in S, arremiyaides daring a study of plan 
reproduction in labat drugnieats tn the areas 
considered 0) rhe present paper Courtine 
senmicalr was Tound at sites will) relatively lew 
fruiting shrubs as well us al those with abundant 
fruit production. Larvae were found i [ruts with 
lew seeds as well us in those with many Undamaped 
seeds, The pervasive presence of Consfarpre 
semnicnla, im spite of this helerogenous Truilins 
patiern. night indicate that it causes witespread 
seed loss. bub rs not key determinant of yariation 
in-seed production by S. antenistaides. Comarina 
seonicala may nevertheless phy a role as one ob the 
factors Tn the population dynamics of the plant 


Acknowledgments 


D. bE. Symons and M,C. O Leary of the State 
Herhariim of South Australia Adelitide. courteously 
identified the host plant species, We thamtk J Kalesib. 
Department oof florticullure, Vitreadture ane 
Oenology, Cniversity of Adelaide for preparing 
microscopic slides and ROL Gagne and DB. Colless 
for constructive eriiicism of the manuscript. RG. 
Simms, SA Musetum, and Bo Parsons, Scotia 
Sunetuary are thaoked for organising the ceologiedl 
survey in the Seotia Sanetiary, This) study owas 
funded by the Australian Researeh Counei (PR), at 
ARC fellowship (SAC) andl Macquarie University 
vrant (A, J, Beatie und SAC), 


References 


Avi TT DO (YRS) Seed predugan im mitive legumes of 
south-custen Australi. ai 2. Beal, 8. 367-376, 
CIY80) Varnitton in predispersal seed predagort 
drseveral Australian Vedco spp. Grav 47, 314 326. 
ClNalsebtaae GML MEL AM. WOR Minton, Bob ak& 
Lidone J) Th (Toe tp Pants of Western New South 
Wiles” (NSW Govt Printing Office, Sydney), 
CUSNINGHAM. So oA. (17) Predator control af seed 
produetion by qn forest understorey palin. Oikos 7 
8224). 
120000) Etfeets of habitat fmagmentation on the 
iyprodielive ceclogy at oar plant spevies in mathe 
wowdlafd, Cramer Brat 14, Th 768, 
”, (2000b) What determines the qitober of seed 
produced in ae flowerne event!) A case study of 
Culytpragyne ghiesbrechitane (Apecicene), Agst A Bas, 


48, 054-0105. 

Dr Gtk. C1776) “Memoires pout serie a Plastoire des 
fiscetes” (Pierre Husselbere. Stoekdotied, 

Givone. Rob (9889) The Plane Feeding Gall Midges vf 
North Ameren’ (Comell University Press, Pinca, New 
York}. 

(1904) “The Gall Midges of the Neotropics 
Region” (Cormell University Press, hava. New York. 
Harnen, G1 (19908 ~Vlory of New South Wales.” (NSW 

(oniversity Press, Kenstimtoi). 

Harris. KM. (1979) Descripmons wind host ranges of the 
sortie midge. Chania soreiioate (Coquillery 
(Diplern Cocnlomytidae). idl af eleven new species dl 
Connon reared from Graminie aid Cypirweue in 
Australia. didi cnt. Key. 69 1ot-1s2. 


126 P. KOLESIK & S. A. CUNNINGHAM 


KOoLesik, P. (1995a) A new species of Eocincticornia Felt RONDANI, C. (1860) Stirpis cecidomynarum. Genera revisa. 


(Diptera: Cecidomyiidae) on Eucalyptus fasciculosa in Nota undecima, pro dipterologia italica. Atti della 
South Australia. J. Aust. ent. Soc. 34, 147-152. Societa italiana di Scienze naturali. (1859-1860) 2, 286- 
(1995b) Contarinia bursariae, a new species of 294. 


Cecidomytidae (Diptera) infesting fruits of sweet 
bursaria, Bursaria spinosa (Pittosporaceae) in Australia. 
Trans. R. Soc. S. Aust. 119, 177-181. 


INTESTINAL HELMINTHS OF FIVE SPECIES OF SCINCID 
LIZARDS (SAURIA: SCINCIDAE) FROM WESTERN AUSTRALIA 


By STEPHEN R. GOLDBERG* & CHARLES R. BURSEYT 


Summary 


Goldberg, S. R. & Bursey, C. R. (2000) Intestinal helminths of five species of scincid 
lizards (Sauria: Scincidae) from Western Australia. Trans. R. Soc. S. Aust. (2000). 
124(2), 127-133, 30 November, 2000. 

Intestines of five species of scincid lizards, Ctenotus brooksi, C. pantherinus, Egernia 
depressa, E. inornata and E. striata from Western Australia were examined for 
helminths. One species of Cestoda, Oochoristica australiensis and eight species of 
Nematoda, Kreisiella chrysocampa, Maxvachonia chabaudi, Parapharyngodon 
kartana, Pharyngodon kartana, P. tiliquae, Wanaristrongylus ctenotl, W. 
papangawurpae and Abbreviata sp. (larvae) were found. Fifteen new host records 
were reported. 

Key Words: Cestoda, Nematoda, scincid lizards, Australia. 


Lreiitienions af Hie Ravel sactety ap S. Aan (2000), 124(2), 127-133. 


INTESTINAL HELMINTHS OF FIVE SPECIES OF SCINCID LIZARDS 
(SAURIA: SCINCIDAE) FROM WESTERN AUSTRALIA 


by STEPHEN R, GOLDBERG & CHARLES R. BURSEY 


Summary 


Cuinpeure. So RO & Borsey. CR. (2000) Intestinal helminths of five speetes of scimeid lizards (Sauria: 

Seincidie) fom Western Australia. Jiri. KR. Sec Anse (2000), 12462). 127-133. 30 November, 2000, 
Intestines of five species of seineid Taards. Coeiotin dvoks), C. paaiieriins, Exeriid depressd. bo arin 

and 7 sift Trom Western Austral were caumined for helminths, One spears of Cestodit, Qoeherstica 


iniveraliensos andl eight 


species ol Neimuloda, Aressielle chryyocampa, 


Mewvechonta cheba, 


Porupharviweden kartana, Phivyneodon kevin, Poiliquae, Waniristrone vii etenord, Wo pape viiypee anal 
APbreviate sp, daevae). were found, Piftecn new host records ire reported, 


Ry Worts: Cestoda, Nemateda, seme lizards. Australia. 


Introduction 


Scineidae 1s the dominant dizard fannity in Australia 
IL contuns some 373 species (Cogger 2000) which 
constitute wpproximately 57% of all lizard species in 
Australia (Greer 1989), Helminth records exist for 49 
species (Mawson 1972: Goldbers and Bursey 1905: 
Goldberg er al 1999: Pichelin ef al 1999). The 
purpose oF this paper is to report addifonal helminth 
records for Crenaty brooks! (Loveridge, 1933), ©. 
paiuhernuy (Peters, 1860). Meera inornata Rosen, 
1905, E se7ate Sternteld, 1919 and the first helminth 
reeords for A. depresse (Giinther, 1875). Patterns of 
helminidi infections in Australian skinks are examined 
md 19 new host records are added to the elicek list of 
Pichelin ev al. (1999), 

Crenoetus brooks’ habits sandy deserts of south- 
eastern Western Australia and adjacent desert areas 
of South Australia. the Northern Territory and parts 
a! Queensland und New South Wales. Crenauis 
panthertivs is widely distributed in south-western 
Western Australia, oorthern South Australia, the 
Northern Territory and western Queensland. Egernia 
depresse oceurs in centralewestern coushil regions oF 
Western Australia, Ereenta inernate ts widely 
distributed throuvh the southern hall of Western 
Australia fram Seuth Australie lo western 
Queensland, in western New South Wales and north- 
western Vieloria, Ergenta siviate is widely 
distibuled through (he interer of Western Australia 
ly south-western Northern “Permlory aind) north- 
western South Australia (Copger 2000), 


HG rentol Biohigy, Whi ier Cotes WIITEF CA 90608 OSA, 
Porntils euoldbo anh iiier outa 

Departinienl of Loology. Vetueylvini Stale University Shenunge 
Cpus Shara PA HA-TN E SAL Lomitl ixbh rte psieeelit 


Materials and Methods 


Ninety three preserved lizards were borrowed [rou 
the herpetolagy collection of the Natural History 
Museum of Los Angeles County (LACM) and 
examined for intestinal helminths, These specimens 
hid been collected between October L9G and 
October 1968 lor use in an ecological study (Piunka 
1972) and Were subsequently fixed ih formalin and 
preserved in alcohol, Because the ecological study 
ineluded stomach analysis. only somal and large 
intestines remained with the carcusses. Stamachs had 
been deposited in the Western Australian Museurn, 
Perth, Western Australia und carcasses in LACM, 
Numbers. of individuals, mean snout-venr length 
(SVL), muscum accession numbers und vollection 
sites Congitudes, lititides) foreach species dre given 
inthe Appendin, 

The small and large intestines body cavity and 
liver of cach livard were examined for helininths, 
Using ao dissecting microscope. Hach helminth wus 
placed ora glass slide ina drop of unditoted glycerol 
for study under a Compound microscope, Nematodes 
were identified Irom these preparations: (he cestode 
was stained with hematoxylin and mounted in 
balsam for identification. 


Results 


Graviel individuals af one species of Cestodis 
Oochorisiica ausiratiensiy Spasskin. 195) and seven 
species of Nemuloda, Arefsiella elrysecampa Jones. 
1985, Muaxvachontia chabaudi Miuwson, 1972, 
Yrapharvagodan kernine ohaston & Mawson 
O41). Pharvercodon kere lohoston & Miowsen 
Oe), Po otiftenee Baylin, WSO Werrristrencey lity 
crereti Jones, O87, Wo papauuenuipiae Jones. LYS? 


by 


lizards 


scineid 


ISCUSSION 


D 


olf Australian 


ecuions 


Inf 


occur on stomach walls (Jones 1995) and may also 
nematodes are summarized in Table 2. Infections 


be under-reported in Table |. 
by trematodes, cestodes and acanthocephalans are 


& C.R. BURSEY 


7 
ae 


+ 


tata 
in 


§. R. GOLDBERC 
, Mean intensity 


ctions by host 


ce 


f helminth inf 
ind 15 new host records are presented 


Table |. Because physalopterid nematodes normally 


containing larvae of Abhrey 


er-report Kreisiella chrysocampa. 
Cysts containing larvae of Abhreviata occasionally 


on oO 


a 


=) 


range, locat 


ible | may und 


& 


3 


habit the stomach (Anderson 1992), the values in 


sp. were also found. Prevalence 


were found, Cysts 


SD, 
species 


T 


P1OIII ISOY MANY =. 


aUTSAVUL ade T 

oT = - ~ = — _ — ~ = apdinmpsundpd snxs vos sLipuny, 
UNSAIUL adIeT 

_ * _ os = _ fT OFFR1 OF — Or fe NOUAL) SNINBUOMSLADUDMA, 
aepipydorgiydury 

JUNSAUT OsIR 7] 

CFVIF Che LLC VSS FOG BS - ee ae apnbi]iy Uoposuripye 
aUsaqul sare] 

o é = = = x é = — cl LOLFLL Sls = = - DUDLADY UOPOSUNAPY 
SUNSAIU! BHM] “AULISATUT [RUS 

tas 7 - ~ = - ~ TST VOTRE IL SEs = - - puppy uoposucipydnand 
arpiuoposudaryg 

DUST ASIN] “PUNNSAIT [CLUS 

Sls ~ v1 Bre IPNBGHYI DINOYIAXD]Y, 
ANPINIA9OWSOD 

OUNSAIUT ABA] “ALLSAYUL |PeLUS 

TI SOFEL OF DAM IONAAY.D BEPIISIAAY 
uimnauojtirad 

[RABISEA UT SISAD 

- - - = - = = = — Ith LtrF9E 0% = - Wl te (araaey) ds puomarqqy 
sepuaidopes<yg 

Ppoweuen 

AUTSAIUr |[PULS 

= = = = - = = Ol Ele - - Ee = - - NISUANDUSND DIUSLIONIOO 
aepreydasojdouy 

Rpolsay 


| 

| 

' 
aa) 

’ 
— 
Nd 
+ 
al 
i 


+ 
Ww 
foal 
ne 
nal 
rn 
a 
wy 
f 
Oo 
s 
a 
a 
2 
Cc 


Ta 
ina 
cl 

+1 
aa 
cl 
+ 


~ = — OE F9OFES le - 7 _ tl 


+ dsS¥* ot 4 aqs*¥*¥ % + dqdS*X % I -s+*¥ & + Gr+* © HQMRAa] 
TOLLS uypusou] assa.dap sTuliaynd Isyooug WUT aH 
piuesy pnuad DULAST smoualy smoualy sOoH 


‘Spapzyy piguias upijossny Jo saioads aay mods syiunujay 4of UEYDIO} Pun (4) ISUBA (CS FX) adisuaqu ual ‘(3,) aauayorarg “| IW], 


ards, 


cincid fi 


5 


T 
? 


. Reported nematodes of Australia 
i 


a 


LE 


TABL 


HELMINTHS OF AUSTRALIAN SCINCID LIZARDS 129 


Wanaristrongylus papangawurpae Jones, }987 vet a _ 
: 4 Bi o, 
Wanaristrongylus ctenoti Jones, 1987 -|{m--2- a al 


Veversia tuberculata (Linstow, 1904) 

Thelandros trachysauri Johnston & Mawson, 1947 
Spinicauda australicnsis Baylis, 1930 
Skrjabinoplera goldmanae Mawson, 1970 
Pseudoriciaria dipsartis (rwin-Smith, 1922) 
Paeumonema tiliquae Johnston, 1912 
Physulopteroides filicuuda Jones, \985 = = aoa 
Pharyngodon tiliguae Baylis, 1930 

Pharvngodon kartana Johnston & Mawson, 1941 mi 
Pharyngedon hindlei Thapar, 1925 

Pharyngodon australis Johnston & Mawson, 1942 
Pharyngodon asterostoma Adamson, 1984 
Purapharyngodon kartana (Johnston & Mawson, 1941) Py La 
Parapharyngodon fitzroyt Jones, 1992 
Maxvachonia chabaudi Mawson, 1972 BTSs ™ ie m9 we. e 
Maavachonia brygooi Mawson, 1972 

Kreisiella lesueurii Jones, 1986 

Kreisiella chrysocampa Jones, 1985 - | rn — 


Johupearsonia egerniae (Johnston & Mawson, 1947) 


Hedruris longispicula Thomas, 1959 


Abbreviata antarctica (Linstow, 1899) 


Crenotus calurus Storr, 1969 


Crenonts dux Storr, 1969 


on] 
oo 
oy 


ay. 1838) 


a 


> 
=> 
3 
2 
E 
g 
wy 


D 
oS 
a 
= 
e 
E 
2 
a 
= 
2 
= 
x 
~) 


1839) 


th species 
Crenotus brooksi (Loveridge, 1933) 


‘co 

— So 
fp eo 
oO — 
oo - 
nm wn 
Z a 
LP 

a, 
a) oe 
3 =| 
5 = 
fata] = 
= vm 
& s 
x = 
~d S 
= 

= —— 
= med 
<= = 
S = 
= S 
S E 
S = 
# G 


Cryptoblepharus plagiocephalus 


(Cocteau, 1836) 
Crenotus ariadnae Storr. 1969 


Crenotus australis (Gr 
Crenotis labillardiert (Dumeéril 


& Bibron, 


Crenotus quattuordecimlineatus 


Clenots grandis Storr. 1969 
(Sternfeld, 1919) 


Crenotus helenae Storr, 1969 
*Crenotus regius Storr, 1YTI 


Ctenotus piankae Stor 


4Ctenotus leonhar 


Host 


S. R. GOLDBERG & C. R. BURSEY 


130 


(A661 } 7p fear UAL AN ul past] yOu, 

“STOI WedeyL 17 *7661 SUL “H “LF6| 

UOSMEJA] 2 UOISUYOF “QOS UOSMLPY 2 [ABU :suodal saIyI “OT *TPHT UOSMETAL FW UOISUYOS “G] 2186] 24g “COST UOSIRag 3 aUAyUETTeY “/p6] UOSMeEyA] IP uOIsUYOL *¢ [6] jUlIG 
‘suodas moj "ST S166 ‘AUAIUET[EY “L1 7OL6L SAP ‘OL 76S6T “SRUIOUL “C] 2O86] “ASNOYLOOPY 7 UENO “FL 716] UOSMEY! ‘89G] LOSMPAAL 2 [adUY :suodar OMI "ET ULSG] SOUTH 
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HELMINTHS OF AUSTRALIAN SCENCID LIZARDS Is} 


fisted in Piehelin ered (1999). Additional records for 
scunmad fivards ure given in Goldberg & Bursey 
(1995) and Goldberg eral, (1999), lachiding the data 
Irom Uns paper, helminth records now exist lor SO 
species of Australia shinks. 16% (50/313) of the 
Australian scineid taunts. Mean hamber of helminth 
species per skink species wis 28 + 2,1 SD, range | 
12 helminth species. ffliguer sefreurdey hal the 
niealest helminth diversity (12 spectess. 1S different 
skink species ane reported to harbour a single 
Hielnimh species. 

OW the Trematoda that infect Adstralian lizards, 
Toradisteunune cructer UNicoll Ite) has been 
reported trom the scincids, Aeniersis pero, 
Lerista bougetivilliy, Tilique seinestdes vod 
Trachydoyanrns rigoxus, ax Well as a pygepadid, & 
gekhonid. ane a varanid, Mesoeaehii aiicraun 
Nicoll (414 Tron Titigne setrcordey, Micraphealtis 
sp. front Vruchvidesiiciy rigs. an umdentitied 
Wematude from Lert dengetvillii, and 
cmidentified dienmeoeliids Hom an agimid (Pichelin 
chal, (999), Meyeeoelivm meroon lias also been 
reported from amphibians collected in Queensland 
(Nicoll (914), Species oF Microplate are parasites 
ol dreshwiter lishes, although experimental 
infections have been established in aiaphibians. 
replies and inammals CYsomiauty 195), 

Vive spevies of Cestoda have been reported trom 
Australian scincid lizards, mimely, Cwinedretdenia 
dlisonde (Sehmidl LS) fram Meniergiv peronn 
and Lerista bemgainedlii, Co hickmant (omnes 1985) 
rom Lanwproplialiy deliccia (De Vis, 1888). L 
wiuictenod  (Dumeri and Bibron = | S34), 
Neroroxemeny dneceay’ (Lucas und brost, 1894), 
Saprossnens chatlengert, and 8. jaustelints 
(O' Shauelinessy, 1874). Gechariiow anvtrafiensis 
SpusshiL, 195) fram Trechvdesanras rugesus, 0. 
tracliysanrr (MacCallum. L921) from 72 rteweyiny und 
OC. veiioletd Mickman, 1954 from Egerniid whi 
(Pichelin ep al F999) Me Jones (1987) reported 
Ovindrelaente allivonie lo gecuralso ina gekkonel, 
MacCullum (192)) described Veena treachysaurt 
from specimens diseovered fn the intestine of u 
speemmen ot Pracivdosaures tugosey chat bad died 
mete New York Zoological Garden. Buen (1997) 
moved Foire tO Cochevisted, Johnston 
(1932) reported Go trrcdisatini in To bipasis 
Spusskil (1451) believed thar substantial differences 
existed between thie specimens deseribed by 
MurCallum (1921) aod Johnsen ch32) and 
established Quehary tice ausiraliensy foe Johastan’s 
specunens. A mujer difference herween 0. 
caistredionsis and OF trachy seed 1s the anumgenient 
OF the testes; CL aastrilieais. has om cluster, O 
Prneiysetere Tas bwo, Vhe speeimen bond Eyeriie 
depreava exiibited one cluster al testes, Unidentified 
speeies OF Oorhoristia taye heen reported from 


Heimerets powniti and Lerivte hoagennviltic (Angel 
& Mawson Look) 

Cystieanths of Acaithocephate fave been reported 
from Austrifan sete laards Splaercehine 
riences rotiidocupoaiy Jolinston and Deland. 
1920 from farlapens quevii (umeril & Bibrou. 
L839), Meonievgls deeesiensis (Cuvier, (829) and 
Lamiapheliy gurchenai. collectad in New Soulte 
Witles, ain) uimidentified eyshacanths from Heniergis 
pero) collected in Seuth Australie (Piehelin ee al 
1999), 

The pentistome Mullienelta senmeoides Ali, Raley 
& Sell 1984 wis described trom Tilfguer seineides 
collected in Sourh Australia (Al Led a L884) aoe haa 
been reporied lrom a gekkonid (Bursey & Goldberg 
1999), Pentustomids were not listedin Piehelin ep al 
(hogy). 

Neimitlodes reported from Australian semen 
livards ace listed in Table.2, Nor included in Table 2 
wre reports OF unidenilied species Of Sknjahinelasia 
from. Crehatis sclomibyredie collected in South 
Australian (Goldberg d& Bursey 1995), reports ol 
Porapharvigedan  karnine and Shijahornlan 
leristee from sospecies of Lerista (= Rhodorta) tron 
South Australia (Mawson l97)) and reports of 
pharyngodonid on physulepterid larvae (tones 142, 
1995, Goldbere ef al 14999: this paper) An 
unidentified species af SAnedtnelecia was also 
reported froma pekkonid lized fron) South Australia 
(Angel & Mawson 1968: Mawson (971). Males of 
this Speeres of Sérjahinelacipy haye yel lo be foun: 
thus no species of Skvfrbirelacia has been reported 
from Australian hosts, 

Unidentified specimens of Pharynvodunidie were 
reported from Cryatoblepliuras plastoceplalis by 
Janes (1995) whieh vould belong te any one wl the 
nine uxvurid species fisred in Table 2. More 
UM cull fo ussess ure reports (ones 1992, 1905: 
Goldberg er af. 1Y99) of encysted Larvae denulied 
us APbreviata sp.  Physaloplera sp, oor 
physaloplterid furyae. Seventeen species of 
Abbrevinit and twospecies of Skejebineptera occu 
in, Austrulinn reptiles (Ruker 1987; udully ab 
species of Phyyulepierd are net known as purasites 
oF Aastralian repules but seven species are know 
fron Austeilian mmmals, five fram miirsupials 
dnd wwe from nulive rodents (Norman & Bevertha 
1999), Physaloprercidl larvae are widely distributed 
in Australia and fave heen reported from the 
sein divans. Criyvedlepharis plagreceplialts, 
Crénotus Calais, ©. dit ©. wrandis, ©. lrelenteae, C. 
pauntertias, t — Guatturrdeemilineats, 
sohiwnhiurehli, Boerne feat, Fo stefan, 
Eufompras quoy aid Leresta ayrelferd (riselves, 
P8817 cy awell as fron again, gekkenid aml 
yuranic ligirds umd several species oF squkes ones 
1995) Studies on diet trav stewie chub sau 


\32 SR GOLDBERG & CRO BURSEY 


livwds and the feral cat, Melis eaty L., L758. feed 
on skinks Jones & Coman 198d; Shine (986, Tames 
vlul 1992), Because these liryae are cneysted and ih 
relatively bigh prevalences, the shinks may Serve as 
purdtenic hosts. 

OF the nemutide species harboured by Australi 
seine lizards (Table 2), Medius longispicula; 
Johapearsonia vgerniae, Pharyngodon aylerostana, 
BP oaustealis, Fo tinder Po riliqnite,  Preninonenta 
tiliqnee, Spinieanuidea  eustratiensis. Thelendras 
veichysaurt and Veversia nibercularad are known only 
from shinks, Abprevidia antaretica ts know Har 
seineids, agamids, varanids and snokes, Mreivielly 
chirysocempa, K lesueurii, Paraphiryngenton first 
are known from setncids and agannds. Miceeornia 
bryvgool is khawn from seingids, agamids and ao 
varanid, WA chabaudi is Known trom seeds. a 
gekkonid. a yaranid and a snake. Porapliuryngedon 
kartand occtrs in seticids. agamids and wehkKonids. 


Pharvagedon — kartana amd — Warturiscrornes lity 
pupengewurpae oecur in scincids and gekkonids. 
Physalopreroides: filicauda ts Known from scineids, 
iugumids, gekkonids and varanids, Preudorionlarte 
disparilis ooewrs in scincids, amphibians und 
nanmas, Skrjabinoepierd celedmenee is known fram 
seincids, ugamids, a pekkonid and varanids and 
Wanuristroneylus elenolt is Know from seifeids, an 
agamid, a gekkonid and a varanid (Owen & 
Moorhouse 1980: Piehelin eral. 1999), 

Helminthological studies on udditionul species are 
feeded before the helminth diversity of Australian 
skinks is Known, 


Acknowledgments 
We thank R. L. Bezy (Natural History Museum of 


Los Angeles County) for permission to exariipe 
specimens, 


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Appendix 


Seineid lizards borrowed from Natural History Museum of 
Low Ativeles County (LACM) with longitude and lite 
af collection sites and helminths deposited in the tS 
Nitional Parasite Colleetion (USNPC), 

Clenotiy broaksi (N= 25. Mean SSVI = 42 jam + 3 SO. 
range = 47-48 min) callected 1967 Western Australia (WA), 
LACM (55525, 26° 32° S, 1257 58! Ej, (55529-55541. 
128) S10" bi), (55558. 55500-55501. 55565-55564, 55566, 
SSIN). SSSTADSSTA. SHSTK, SSSR4, 28" OR’ SH, (2R° 55! 
[), (55585, 28" WO" S, 123" 56" EE): Northern Territory, 
LACM (55544. 23° 14 8. 120° Sa° Fy (55544, 55548, 
SA5S0-SSAS1, SS5S3. 23" 13’ S, 129° 54" BL. USNPC 
SY250 Abbrevian: sp, (4rd stage larval USNPC 89247 
Kreisiella chrysocumpas USNPC 89248 Mavvaelonia 
whahaid: CSNPC BY249 Wena rinedeatignhin cretotl, 
Crema pautherins US = 20, Mew SVL = 42 mint 3 SD. 
runge = 37-18 mm) collected N67. WA, LACM (45986. 
SSUN8. 28 OFS] WY 05) Ey. (55991, 56000-56007, 
SHO FP BAS, 125° 50" Ey, (56032. 36025, 28" 08'S. 
PPA S55) BSR, SONU SO043, IR! OBS, 129" 50°), 
(56046, JA" VES. 121 13? 1), (S6083-S005+4., 56058-56059, 
SOU, 2OP PPS. 121 DOU). CSNPC $9256 Abbreviate sp, 
(AU Stage harwils LISNSPO ROIS Krvlvielle mlarvseeemipet 


USNPC $0252) Warvuehonia chatty USNPC SU255 
Pourapharyugedon karlina; IESNPC ROI54 Pharyawoday 
karte USNPC $9255 Wiristinevliny erenar, 

Aeeriia depresse (N= 8 Mea SVE = 8b mm + 7 5D. 
minge 81-101 mim) collected 968, WA, LACM (S040 
S404, 28 27S. 114 05" EY (SGAOIAOIT A, SO4TK, 27° 
OS'S. 119 37! fs). USNPC $9257 OQachorivticu auyira 
liensis; USNPC $9258 Pharvngenon diquae. 

feernia inorneta ON = 19, Mean SVI = Ja inm +4 8D, 
rubies GOe8O nin) collected 1966-1968. WA, LACM 
(AO434. 56436, 56438, 56440, 56442-56443, 2H 27! 5, 
119° OS" BE), (50447, 56450-50452, 28° 08'S. }24° 55) Ep 
(40458, 56463-50404, 56466, 56472. 560474, 50477-50479 
TRe 30' 8. 125" Si RY) USNPE 89259 Kreisietla 
clryyvecumpas USNPC 89260) Maxvuchonia chabaudi 
USNPC 89261 Phapyngodon tilqnie; USNPC 89262 
Wand ristrang iis ppangwiipae. 

Meer sila (NS = 21, Mean SVL = 95 nin #8 SP. range 
T8103 mov collected 1967, WA. LACM (56513-56517, 
S0521-S6525, 56530-50531. 50533. 50545-50537. 50534 
2B) ASS. (22° Sw 1), (56541, 50545, 75° 28'S, ADs! 
BE). (50046. 50548. 28 IS So 12 So" by, USPC Sulne 
Paraplaryisedian Katinas USNPO B94 Marvel 
Lilies CSNPC R265 Wererisironey lis cleat 


RE-EVALUATION OF THE DISTRIBUTION OF GEOCRINIA 
LAEVIS (ANURA: LEPTODACTYLIDAE) IN SOUTH AUSTRALIA 


By STEVEN J. WALKER*} & PETER M. GOONAN* 


Summary 


Walker, S. J. & Goonan, P. M. (2000) Re-evaluation of the distribution of Geocrinia 
laevis (Anura: Leptodactylidae) in South Australia. Trans. R. Soc. S. Aust. 124(2), 
135-139, 30 November, 2000. 

A survey of the known range of the Smooth Frog, Geocrinia laevis (Giinther, 1864) in 
South Australia was undertaken to determine the current distribution and abundance 
of this species. A total of 58 locations was visited throughout the South East and G. 
laevis was collected or heard calling at 13 sites within or near the Reedy Creek / 
Dismal Swamp drainage system. Despite very few reports of this species in recent 
years it is locally abundant and under no obvious threat of decline. 

Key Words: Geocrinia laevis, distribution, frogs, South Australia, frog census, status, 
conservation. 


Transactions of the Royal Society of S. Aust. (2000), 124(2), 135-139. 


RE-EVALUATION OF THE DISTRIBUTION OF GEOCRINIA LAEVIS 
(ANURA: LEPTODACTYLIDAE) IN SOUTH AUSTRALIA 


by STEVEN J. WALKER’ & PETER M. GOONAN* 


Summary 


Wacker, S. J. & GOONAN, P.M. 


(2000) Re-evaluation of the distribution of Geocrinia laevis (Anura: 


Leptodactylidae) in South Australia. Trans. R. Soc, S. Aust, 124(2), 135-139, 30 November, 2000. 

A survey of the known range of the Smooth Frog, Geocrinia laevis (Giinther, 1864) in South Australia was 
undertaken to determine the current distribution and abundance of this species. A total of 58 locations was 
visited throughout the South East and G. /aevis was collected or heard calling at 13 sites within or near the 
Reedy Creek / Dismal Swamp drainage system. Despite very few reports of this species in recent years it is 


locally abundant and under no obvious threat of decline 


Key Worbs: Geocrinia laevis, distribution, frogs, South Australia. frog census, status, conservation. 


Introduction 


There have been few comprehensive studies to 
document the distribution of the frogs of South 
Australia. Brook (1984) produced an atlas of the 
known distribution of the frog fauna of SA by 
condensing published and unpublished data from 
various sources. Other published studies have 
generally been focused on unusual range extensions 
and first records in the State (Tyler 1971; Bird & 
Tyler 1990; Johnston 1990), Overviews and species 
lists for the State are given in Tyler (1977, 1978, 
1994, 1997), 

Since 1994 the South Australian Environment 
Protection Agency has conducted an annual frog 
census in September (November in the first year, 
September thereafter) involving the public making 
tape recordings of the frogs calling from waterways 
throughout South Australia. This work has 
highlighted the distribution and a measure of the 
seasonal abundance of frogs, mostly from the more 
southern parts of SA (Goonan er al. 1997, 1998; 
Walker ef al. 1999), Some species are poorly 
represented or have not been recorded through the 
method being applied by the census, including 
Geocrinia laevis (Giinther, 1864) which had not 
been recorded (Goonan ef al. 1997,1998; Walker et 
al. 1999). Geocrinia laevis is mainly an autumn- 
winter breeder, calling only infrequently during the 
period in which the frog census has been carried 
out. 


Environment Protection Agency, GPO Box 2607 Adelaide SA 
S001 

Department of Environmental Biology, University of Adelaide 
Adelaide SA 5005. 


Fig. |. Geocrinia laevis from Canunda Conservation Park 
(SVL = 33 mm). 


Geocrinia laevis is a medium sized frog (22 — 35 
mm snout vent length) with short limbs and smooth 
skin (Fig. 1) that may be easily confused with Crinia 
signifera Girard, 1853 or members of the genus 
Pseudophryne Fitzinger, 1843 (Barker et al. 1995), 
Distinguishing characteristics include pale pink 
patches underneath the legs, in the groin and 
sometimes in the axillae (Woodruff & Tyler 1968; 
Tyler 1978; Barker e7 al. 1995), 

Like Pseudophryne, G, laevis does not breed in 
water. Males call from the ground in moist leaf 
litter and amongst grass. The advertisement call is 
a long slowly pulsed rattling or creaking sound. 
the first note often being the longest - “cre-e-e-e- 
e-e-e-e-ek —cre-e-e-e-ek = cre-e-ek = cre-e-ek” 
(Woodruff & Tyler 1968; Barker et al. 1995), 
Geocrinia laevis lays large, unpigmented eggs in 
loose, clongated masses attached to moist 
terrestrial vegetation. Major development occurs 
inside the egg capsule and following flooding 
tadpoles hatch in the water, with complete 


136 S.J. WALKER & P.M. GOONAN 


development taking about six months (Tyler 1994; 
Barker et al. 1995). The habitat of G. laevis is 
reported as being leaf litter in dry Eucalyptiy or 
pine forests subject to temporary flooding (Tyler 
1978; Barker et al. 1995), 

Geocrinia laevis was first reported in South 
Australia from a specimen (South Australian 
Museum, Adelaide (SAMA) R8118) collected near 
Mt Burr in 1966 (Woodruff & ‘Tyler 1968). Before 
this it had been found in Tasmania, King Island, the 
Grampians and in South West Victoria from 
Dartmoor to Pt Campbell (Woodruff & Tyler 1968: 


Beck 1975). Beck (1975) surveyed the South East of 


South Australia between 1968 and 1974 and found 


that G. laevis was confined to the Reedy Creek and 
Dismal Swamp drainage system in the lower South 
East. Since then, there have been no major reports of 
this species, 

With the major and continual modifications to the 
drainage system in the South East of South Australia 
it seemed pertinent to determine the current status of 
G. laevis in the region. As G. laevis may inhabit 
areas. which are vulnerable to agricultural 
development and because there is no detailed 
knowledge of its current distribution it is possible 
that any future development may — impact 
significantly upon populations of this species. The 
purpose of this study was to determine the current 


« Sites with Geocrinia 


Sites without Geocrinia 


SAMA Sites 


| 
i] 
PENOLA 
et 
19 20 | 
MILLICENT ‘80° * 
u el? 
22 921). 
® eo! 
53 | 
a? 
1301 
MT GAMBIER 
a 
\ 
Ts j 
a 
Sd ng | 
a en 2) | 
LA~g 
“ae - | 
ee ea a a 
PT MACDONNELL 


Fig. 2. Surveyed distribution of Geocrinia laevis in the South East of South Australia. Sites from SA Museum records are 


included for reference. 


CROCKINIA LALVIN TN SOUTILE AUSTRALIA IX? 


Wistibubion and status of G. leevis in’ South 
Australia. 


Materials and Methods 


Existing data sources (published and unpublished) 
including SAMA. records und NP&WS: regional 
surveys Foulkes |998) Were examined. 
Locations were subsequently superimposed an 
floristic vegetation maps of the South Bast inorder to 
predict possible suitable habitat for Go luents. B. 
Grige from Forestry SA alse provided taps of 
Forestry land und suggested jtreas where frogs might 
occur 

Surveys were undertaken during Mareh, Jane and 
August 1999. A total of 58 locations was Visited (Piss. 
2), (helading LO Sites based on SAMA records and 
fwo sites from the NP&WS survey The revorded 
coordinutes for some of the SAMA sites appeared to 
be imprecise or inaccurate because the sites did not 
have suitable habitat for G, feevey: in these cases, 
sites wilh suilable habitats which were nearby the 
recorded coordinales, were sampled instead. Each 
site with @ fwevis present was visited only once. 
with the exception of some sites visited in’ March 
which were revisited in Jtine and SAMA sites which 
did not huve G, daeviv calling in June; these sites 
were sampled again in August. Sites visited un 
MulGiple oeeasfons did not have G. Jeeviy calling 
during subsequent visits. 

Calling G. faeviy males were sought by ear or by 
use of a directional microphone attached to a Sony 
DAT reeorder, Where possible any calling miles 
were located, usally by triangulation, and captured, 
The call of Crinia sfenifera is quite variable and can 
sometimes sound very similar to the callbol Go faeviv 
or Peendopliryne sp. Therefore. any culls which 


(see 


could not be identified immediately were recordeal 
for later exanination, 

In addition, searches were carried oul at each site 
This involved looking under logs, leaf litter. stones, 
ind umongst! vegetation, fora minimum of one hour, 
during the day or early evening. Any frogs found 
were collected und pliced in large colon or pliasuc 
bags Tor later examination. A number of frogs was 
eolleected when they were seen on wel rouds at night, 
but no G. lgeviv were found at these ties, Frogs 
were released on sile al the conclusion OF collecting 
und identifieation. 

Numerous plant samples were also collected for 
liter tdentifigation to determine the common 
composition Of Mora assogiated wilh the sites al 
which G, laevis were found. 


Results 


Geecrinia laevis was present at 12 sites within the 
Reedy Creek / Dismal Swamp drainage areal and 
also From asite in the Canundit National Park (‘Table 
1). ft was not found inthe Pt MacDonnell urea where 
it has heen bisted in SAMA peeords, A total of six 
faeviy was collected (lwo from “The Marshes” 
wetland. two from Mt Burr, one from “Honan’s 
Serub” and one from Canunda National Park). The 
presence of calling males permifted a positive 
identification of the species at these and other 
locations (Table |). Analysis of the recordings of 
unidentified cally using & computer biased 
spectrograph (Specht 1998) identified only one other 
site (site 17) where G. leeviy gecurred. All other 
recordings were confirmed its being ©. slenifera. 

Sinec the Beck survey a small umber of GL laevis 
hus been collected in South Austrabia, some reported 


Taner J. Swammary af sites where Geocnimia laevis were detected. 


Site Site Name Species Present Northing Kusting 
11 Hlonai’s Seruh | GL.CS. LD. LE SAIS1T1 467855 
12 (Bogey Piekl) 20 kar S of Ruliungddoo Gh. LE 5825430) 4A 7020 
14 Honus Serub 2 GL, CS, LE S825057 4OSSES 
\4 Honais Serub 3 GL S82560) 460067 
1 Brookshy’s Late (it Lake Leake) GL. CS. Le SRARYOY 462002 
IN ML Burr Forest 1 GL, CS. LE S541 164 ASTSAN 
I) Mt Burr Forest 2 (rie Quarry) Gl SRADYI 459437 
i] Rowdside (ie Mt Burry GL, LE Sk4A2629 461544 
4 The Marshes | GL. LE S835O207 4SOINT 
22 The Marshes 2 GL. CS. LB S857195 4578S 
si Roadside 21 Mingbool) GL_CS 5834331 492 )58 
Al Canunda CP 2 Gl S833737 433837 
aa The Marshes 3 GL 5Ra4d 445 450415 


Northings ind Gastings as an Australian Map Grid, Zone 54, 


(GL = Geacrinia laevis. CS = Crime stanifera. LD = Lannedvadsies dinkerili, LE = Liraria awinen. 


14k S.4, WALKER & POM, COONAN 


tothe SAMLA (M, Hutchinson pers, comer, 1999) and 
others tothe SA Frog and Tadpole Study Group C&, 
Baskert pers. contm, 1999). Tneluded in the SAMA 
necords are (WoO sites to the west of Pt MacDonnell 
Nee We cenist, Ohe location (SAMA record listed is 
“Blanche Buy”) yas wcoustl shrubland / sedgeland 
insand duges which seemed to bean unlikely habrlal 
lor G. laevis. The closest location. just ialand: from 
ihe sand dunes, which may bave heen suitable 
habitat for frogs didi have Limmodvrates: peront 
(Dumerd and Bibrom (4h) and €. sfeaifera, pul 
there was he mdication albany G. laevis A number nl 
Sites Sampled around the other southern location 
(Section 346 Hundred of Kongorong”) also yielded 
ne sign of Ch feevis, There was nothing obvious to 
Suupest that There had been any significant land use 
chanees an the urea sige the lrogs in the SAMA 
were dolleated there i 1883. "Phe preduminunt land 
ase appeared to be ery iioof Tivestoek Wall mast al 
thre find eleureet Of prin) veneration, 


Discussion 


Cre riaia treyis was Cound at 23 silos te the Seveth 
Buse or South ANustralia dining this study, Apart tron 
ihe Site in Cunudda Natianal Park all of the siles 
were Wathin the Reedy Creek / Disniul Swanip 
drajmawe sirew. TAs correspaids to the disiibution 
revonked hy Beek (1975) with the wddilton of the 
Minwhaw! site further to the Gast 

Beek (IY75) speculated that the site at Cununda 
wi probably the fesult of “eres or larvae washed 
Jown one oF ihe mine made drains wineh etess the 
wea between (he Millicent Hills and the coast” Ll 
svenis mure Whely however that the populion at 
Cunwida National Park is a reliet ob a previous 
Uistiibution (hil eovercdl ynueh ool the South Bast 
north OF Mt Gambier, Prior to the pleavirise selicnie 
ithe South East. whieh Fest began uround T&6? 
mueh al the Upper South Bast OF Sour Austrutia 
experio¢need periods ol severe Tlooding ond 
Pmupdation (South Rast Dramage Boutrd (980), with 
ry having permunenh or peut 
periment waters. The Water movement in the 
Millicent area tended fo he direcked North West 
towards Kingston SE. or South West lowprds Like 
Bonney (ie. ta the cdireetion af what is. iw 
Comunda National Park), 

Geocriant laevis were found in depressed clearings 
subycet to iiuineaton dt Ure edges GF rittive Forests ot 
pine plantations (Fig, 3), ahthough ane site wis ad 
bogey farm paddock (site 12), "Phis se was \acited 
only a few hundred metres: (rom uw nearby forested 
area. Geo rinid laevis was also Four al sites. 17, 20 
ind At in vlearings near forested ureas ulongside 
min) cad, 

The vlauwrnws ustiully Comprised reeds, wiisses anu 


lovalities 


Tis, 2. Waring i Mt Burr Forest: typmeut hahitay 
Geocria laevis othe Southy Gost al South Austeatis 


soddes, With the oecisiondl shrub and: herhuccaus 
plan, The mayor plants collected fram the siles were 
the nohhy elubrush (/vealepia nodosa (Roubs), bs 1), 
sea rush (diicus Arausy7? blochal, 1845), and vanable 
sword-sédue (Lepidusperma literade Ro Br, TAU. 
Other plants commnooly seen inelided the buttercup 
(Remirculiy spy Linn. Spry tees (Byeyelaregy iis 
spotnescens (BR. Be) Viekery, 1952) and otherassirled 
orusses. ALJ number ol fallen branches ainkh artes 
tinber Tomy logging alse provided Iabital onder 
which frows could sheller, 

The dead and dying, reeds, sedves and: arasses 
formed a dense mat which retained moisture yin 
provided w network of refuges m which Gr. fev ind 
other frogs coulel Hide, ASG resuTl, Tf Avuus abies 
impossible locate the frogs, even wher 
Wiangulation suggested they were only a lew 
centimelres fram the collectors. At intensive seureh 
(hrouwh the undergrowth dnd under Gillen troaber 
produced tithe nmore success, His Quite posible: Hat 
non-calliog individuals niuy have been presen, bul 
not deteeted, al same sites 

The locations where G. fvewis can now be found 
are a areas whith previously had) permanent 
swirnps und Wetlands, inching the Cununda site. 
and would howe formed a continuous or teddy 
conlinuous expanse of witer during fhe qvet montis 
(South Eastern Drainage Board 1980), Even though 
min-tnide drains were ureoted to jnerease surhice 
low to the Lake Bouney aren. to clrain land far 
ugrivuliunal development and to allow expanded 
seUWlement in the region. this aren always had a high 
ravotall and nattiral drainage features that probably 
crablod populadons 1 colonise the Cannnidi 
loculion prior lo driuniwe aeayites, 

Although €) daeniy has a cestrieted distribution, ihe 
nnijority of locations identified had more thin SO 
males culling. The species is st Tound in the ane 
where th wis reported ta 1974 and consequently does 


GEOCRINIA LAEVIS UN SOUTEL AUSTRALIA io 


nolappear to be under aby obvious threat of decline 
in the region. Both “The Marshes” wetland area und 
“Honan’s Scrub” are large native Forest Reserves 
With the same statis as Conservation Parks, and 
therefore ure not likely to be planted or disturbed (B- 
Grigy, pers. comm. 1999), The sites within Mt Burr 
Forest ure located ti dnused areas that are unsuitable 
for planting due to llooding (B. Grigg pers. eon, 
1999), [Lis possible that these sites may he planted at 
the next rotation, in approximately 25 years, but only 
if flooding could be exeluded. 

Following the survey recorded above the EPA ran 
acensus of the frogs calling from South Australian 
walerways in September 1999, Geoeriiid laevis was 
recorded from “Honan’s Serub™ and “Crouches” 
within the Dismal Swamp / Reedy Creek area: 
“Crouches” was not included in the present study. 
Fewer than ten calling G. faeviy were recorded from 
these locations (Walker et ad, tenpub,), 


Acknowledgments 


We are greatly indebted to M. Bradbury and 
particularly B. Smith (University of Adelaide) for 
their considerable enthusiasm aid assistanee it the 
field trips. M. Hulchinson provided the SAMA 
records and §. Carruthers (Phinning SA) produced 
the floristic vegetation maps. B. Grigg (Porestry SA) 
was especially helpful in ullowing aecess to Porestry 
land and sharing information about the region, D, 
Rovers (University of Adeluide) kindly gave us the 
use of his DAT recorder and helped with the analysis 
of calls Of computer, D, Gooding (EPA) helped in 
the preparation of the distribution map. This project 
wis funded by a grant from the Wildlife 
Conservation Fund and both support and facilities 
were provided by the Enyironment Protection 
Avency, 


References 


Bakkir. b, Grice, G. 0. & Pyibe, Moh (i995) A Field 
Guide To Australian Vrogs” (Surrey Beully & Sons, 
Chipping Norton), 

Beck, RG. (1975) Factors alfeeting the distribution of the 
leptadaetylid frog Geeertig laevis i the south-east of 
Sou Atistralia, Tras. Ro Sac. S. Anse, 98. [ARS 1-47. 

Bikp, Pw Pyine Mob (1990) First South Australian 
revord of {he Tossortal leptodietylid) Tog bends 
(pervert: Gray. bal U4, 223-224. 

Brook, Ac J ClORE) Athis af Frogs of South Austratia” 
(Dept Aoglogy, University of Melbourne, Publ. No, 4). 
POUL KES, 1 C199%) Inhabitants of the Sauth Bast swamps : 
wiant ymogdies (oorare marsuphals. Parks cd Wildlife. 

Winter 1998, 4, 

Cups, BoM. Goon, BD. A. & Hint. BoM. (997) 
“PROG CENSUS 1995 and 1906, AO Report on 
Community Monitoring of Water Quality and Hibiutt 
Condition in South Austratie using Frogs as tdicators.” 
Environment Protection Authority, South Australian 
Department for Pavironment and Natural Resoureys, 
Adelaide. 

HILL, BM. & WaLker. S.J. (1998) “FROG 
CENSUS [997A Report on Community Monitoring 
of Water Quality and Tlvbjtal Condition in Soudh 
Austalia using Frogs as Indicators.” Environment 
Prolvction Agency, South Austeflian Departinent for 
havironment Vlerituge and Aboriginal Attairs, 
Adelarde, 


Jonsston, GR. (1990) Cyelanane matad andl Noten spe 
Additions to the frog fauna of South Australia. Tray. R. 
Sew. So Must. V4, 229, 

SOUTH EASTERN DRAINAGE BOARD (1980) “Bavironmental 
Impact Study on the Effeerof Drainage in the South Bast 
of South Ausiaha” (South Eastern Draimve Board, 
Adelaide), 

SpreHt, Ro (1998) Avisalt SAS-Lab Pra Version a.5b 
(Computer Programme) 

Pronk. MOL CY 71) Discovery in the Eygrurd Riaiges ata 
species of leplodaetylid frog new te the Gina of South 
Australia. Trans. R, See, 8. Angst 95, 205-217. 

(1977) “Frogs of South Australia” (2nd Tdi 


(South Australian Museu, Adelaide). 


a (1978) “Amphibians of South Australia 
(Government Primer, South Australia), 
(1994) “Australian Frogs: A Nutural Pistary 
(Reed Books, NSW). 
(1997) “The Action Phin for Australian Frogs” 
(Wildlife Australia, Endangered Species Progra). 
Wansen, Sod. Aint. Be M, & Goosat, PM. (1999) 
“FROG CENSUS J998, A Report on Community 
Monitoring of Water Quality ind Pabitit Condition in 
South Australia using Progs as Indicators.” Environment 
Protection Agency. South Austtalian Departnent tor 
Laviionment Heritage and Aboriginal Athans. Adelaide 
Woon, DOS. & TYLeR, M.L C19 68) Additions to the frog 
fins of South Austria. Ree 8. Avst Mus, 15, 705-709, 


REDEFINITION OF THE AUSTRALIAN FROG 
LIMNODYNASTES DEPRESSUS TYLER (MYOBATRACHIDAE: 
LIMNODYNASTINAE) 


By MARGARET DAVIES* & THOMAS C. BURTONT 


Summary 


Davies, M. & Burton, T. C. (2000) Redefinition of the Australian frog Limnodynastes 
depressus Tyler (Myobatrachidae: Limnodynastinae) Trans. R. Soc. S. Aust. 124(2), 
141-150, 30 November, 2000. 

Limnodynastes depressus has been known from the holotype since 1976. Examination 
of newly collected and reidentified material indicates that the species can be separated 
readily from morphologically similar congeners L. fletcheri and L. tasmaniensis by 
the loss of a phalanx on the thumb, the absence of a preorbital process on the pars 
facialis of the maxilla and differences in the musculature of the jaw, pectoral girdle 
and first finger. These include the origin of m. depressor mandibulae from the dorsal 
fascia and the tympanic ring but with no fibres originating from the otic ramus, 
exposure of the anterior margin of the m. coracoradialis, and pennate insertion of mm. 
lumbricalis indicis brevis and flexor teres indicis on the palmar surface of the 
metacarpal. 

Key Words: Limnodynastes depressus, Anura, frog, osteology, morphology, 
musculature. 


Tranwections of the Royal Suciety of S, Awa (2000), (2402), PT 050 


REDEFINITION OF THE AUSTRALIAN FROG LIMNODYNASTES DEPRESSUS 
TYLER (MYOBATRACHIDAE; LIMNODYNASTINAE) 


by MARGARET DAVIS 


& THOMAS 


C. BURTON’ 


Summary 


Davis, M, & Bertow. PoC 


(2000) Redetinition of the Austealiin frog Rindedviestes depressus Tyler 


(Myobulruchidie, Linnodyoustiage) Hes, & See, 8. Awst, 124 (2). Pal- 150, 30 Nayember. 2000, 

Linnedynustes depressity bas been known from the holotype since 1976, Examination of newly collected and 
teidennificd material indicates that the speeies can he separated readihy front morphologically similar congeners 
L, fleicheriund L. faymeienyiy by (he loss of u phalanx on the thumb. the absence of a preorbital process on the 
pars fachilis of the maxilla and differences in the musenhiture of the jaw. pectoral girdle und first finger. These 
mclide the orwin of ia depressor mandibutae from the dorsal fascia and the lyrnpadie ring bul with tia fibres 
originating from the olie ramus. exposure of (he anterior margin of the m. coracoradiutis, und pennate insertion 
ob mn, luinbricalis indicis brevis and flesor teres indicts on the palmar surtace of the metacurpal. 


Kray Wotths: Lineedvniites depress. Anura. frog. osteology. norphalogy, muscular, 


Introduction 


Linnodynastes depressus Tyler, 1976 was 
described from a single specimen collected in 1972 
bya Western Austuliag Museum survey party from 
near the Argyle Homestead in the Kimberiey 
Division of Western Australia prior to tts inundation 
hy the Ord River, No further speeimens were found 
until 1998, when Lo Morris recorded frogs in: the 
Keep River National Park that have been identified 
on the basis of cull and developmental biology as 
conspecific with Linnodynusres depressis (Tylor & 
Davies 2000: Morris & Tyler unpub; Watson, Tyler 
& Merris unpub.) 

We have examined some of the Keep River 
specimens as well us material colleeted eartier but 
nol positively tdentificd as 4. depressty, and have 
identitied morphological Teutures that can be used to 
separite the species Tom similar congeners such its 
Lo taymraniensiy Giinther and L, fletcher’ Boulenger, 
We have also cxumined lurther material from the 


Northern Territory to chirify the entity of records of 


Lo rasmaniensis trom WA and the NT. Dlere we 
redescribe (he species incorporating osleologiedl aid 
inyologicul dula unavailable for the ormvinul 
description. 


Materials and Methods 


Specimens examined ure housed in the Queens: 


Dept ol Eavironmental Biclowy, Gniveesily oF Adelaide, SA S0bS 
Division ot Biolowidl Scienees. Li Prohe University Bendigo. 
HO) Wars POO neti Vies 3552. 


lind Museum (QM), the Western Australian 
Museum (WAM), the South Australian Museum 
(SAMA), the Museums and Are Galleries of the 
Northern Territory (NTM) und the University of 
Adelaide Osteological Collection (UAZ). The 
inuscles of the throat, jaw, pectoral girdle. hand, 
leg and foot were dissected with the aid ol a Wild 
MY — dissecting micrascape. — Ostealogicul 
preparations were made aller the method of 
Dingerkus & Uhler (1977). Measurements (in mit) 
were liken using dial calipers reading to 0.05 min 
and follow Tyler (1968). Mlustrations were made 
using a Wild M9 dissecting microscope with 
atluched camera Jucidit. 


Material examined 

QM J55565-71, Keep River, NT. SAMA R23862- 
4, Newry Station, NT, WAM R S8s33 Kununurra, 
WA. NTM RISTO. RISTI2. RIS122, KI3125, 
RI3127. Keep River NPL RES60X, RES623, RI48K 1. 
R24912. R24V9T3. Bradshaw Station, NT R24332. 
Auvergne Station. NT. R24467. Fish River, of! Daly 
River, NT UAZ Bo23, Newry Station, NT, UAZ 
B2l645, B2647 Cockatoo Lagoon. Keep River NP, 
UAZ B2649-50, offspring of mating of aduils fron 
Cockatoo Lagoon. Keep River NPL 


Results 


Lunnadynastes depressus Tyler, 1976 
(FIGS 1-12) 
Jinnodvnastes depressus Tyler, My d, (1976) Ree, 
West Aust Mus, +. p45. 


42 M, DAVIES & TC, BURTON 


Holotype 
WAM R 44896, adult male, 


Definition 

A muderute-sized speeies (males 30 - 44 mm S-V, 
females 34 - 37 aim S-V) characterised by an 
clongale thuinb comprising one phaluna, prominent 
upper eyelid julling oufward as a shell, outer 
metatarsal tubercle absent. moderately long 
vomerine teeth, breeding females with well- 
developed Hoger Manges, frontoparictal fontanelle 
widely exposed, maxillary process of nasals absent, 
preorbital process Of pars facialis of maxilla absent, 
narrow wnexpanded alue of parusphenoid, origin ol 
i, depressor mandibulie from the dorsal fascia and 
the tympanic ring but with no libres originating from 
Lhe Oe ranvus. exposure of the anterior margin of the 
mM. Cormeorudialis, pennate insertion of mim, 
Jumbricalis indicts brevis and Mexor teres indicis on 
the palmar surfice of the ietacaurpal 


Pig.) Male Liniudvnestes depen (elite Capps 43 
win S Vo. Bran Newry Station NT 


Morphology 

The external morphology of this species shows 
little variability und conforms with the type 
description (Mig. 1). The palmar tubercle at the base 
of the first divit was extremely laree jo the material 
examined, notindicaed inthe iWustration of the bype 
(Tyler 1976). Breeding females have fhinges on the 
first two lingers. 

Measurements, in im. are as follows (means 
followed by ranges in parentheses): S-V - mules 24.6 
- de 1. females 34.5 - 36,6; TLIS-V 0.42 (0,39-0.47); 
HL/HW 117 (0b - 1.25); HE/S-¥ 0.36 (0,32 - 
0.38) E-N/IN O86 (0,73 - 1.00), 


Osteology (Description ron WAZ B2645) 

Skull moderately ossiticd. Sphenethimoi poorly 
ossified. not in bony contact with nasuls (Pig, 2A). 
extending anteriorly between yvomerine teeth and 
posteriorly about V/s length of orbit in ventral view, 
Prootie and execeiptital ieompletely uscd, Crista 
parolica short and stocky, widely separated laterally 
with poorly expanded ote mamus of squamosal, 
Vrontoparietal  fontanelle widely exposed 
Frontoparictad poorly ossifted, anterior extremilies 
‘Ay dength of orbit, Orbital edges of frontoparietal 
straizht, then angled slightly posterolaterally. 
Anterior margins of frontoparietil fontanelle Formed 
by sphenethmoid about '/s anteriorly along length of 
orbit, Posterior margin undefined beeause of lack of 
medial ossification of execcipitals, Nasals well 
ossified, perforated centrally, Creseentie wmteriorly, 
Masillary process absent, Nasals notin contact with 
pars facialis OF maxilla, Palatines moderately slender 
medially, underlying dentiverous processes of 
vomers; widely expanded literally, not reaching 
palatal shell of maxilla posterolaterally (Fig. 2B). 


Hig. 2. Memale Lianedvnistes depresses (UNA 26045), A, Dorsal and B. Venttal views of the skull, Seale ba > 5.0 mm 


REDEFINTTION OF LIMNODEN ANTES DEPRESSELS i 


Parusphenoid moderuely robust. extending about 
Yi length oof orbit, Alae slender, not expanded 
laterally, Plerywoid robust, Antenoar ramus long, in 
bony conluct with palatal shel ol maxilla, Plerygoid 
process absent Medial ramus moderately lon, 
SUBUCUITte, Hot overlying alae of parasphengid 
Posteniod nus slender Junenon al the three nani, 
extremely robust, 

Quadritojugal slender and entire, Squamosal 
inoderately robust with modemely long, shender 
zyeonalie minus aid short slightly expanded: ope 
VMS, 

Masilhu and premesilia dentate. Pars daeiahs of 
Maxilh) moderately deep: preorbital process absent 
Alury processes of premasillae proud, slihtly 
bifurculed unl directed posterodorsally, Palatine 
processes sbort. now im medial contact. Preryeoid 
process Of palatil shell absent. Vomers reduced 
anteriorly and medially with moderately long 
dentigerous processes, Columelht bony and sige 
in Shape. 

Pecumal girdle areiferal und robust. Slender 
Girtilavinous omesternunt with stalked. knobhed 
anterior extension, Xiphisternum broad, not bifid. 
Sternum cartilaginous, Clavicles slender strongty 
curved und widely separdled medially. Coracoids 
robust, widely sepurnited medially, Breapnate scapulit 
robust, Suprascapula aboot '/ ossifted. Moderately 
well-developed anteroprosimal crest on humerus, 

Curpus of five elements. Single sesumoids ul 
Juncuion of meticarpals and proximal phalanges of 
(ats Joh unl al junchons of adjacent phalanges. mn 
digits 4 ind 4, Medial flange-on meneu'pal 1 absent. 


Phatiogestl formula 1, 2, 3, 3) Fiest metacarpal 
vlongare (Mig, 3C). 
Seven procaclous. non-imbrieate presacral 


vertebrae. Vertebrae Land I fused. Relative wrdilis 
of transverse processes I=[V>SDstl>V>Vl>Vil> 
VILL, Sacral diipophyses poorly expanded (Fig, 3B), 
iy estending to uboul fhe centre of the sacri 
Ulapuphyses, Urostyle crest upproainmately “i dength 
OF Uirasty le. 

Hial crest absenl. Dorsal proniinenee prominent, 
Hromticiling. Dorsal protuberqnee posterolateral 
(Trig, 4A), 

Three larsul elements. Prehatlis narrow, (Pi. +A). 

Hyaid plate wider than Jung: posterior processes 
inoderitely Slender Anterior processes expanded, 
Aueromcdial provesses of anienor hvale well 
developed. brow. Pasterioy cornitih ossified (Fie, 
43) 


Verio 
Material esaminedl 

(IM 155568 11. OM ISSSTI LF). BOB. (etD. 
B2649 (subadully. B2650° (subsdully, B26ds (4, 
NTM RI40)3 62 4. 


The alae of the parasphenord are mire clonpate in 
both NTM R24912 ahd AZ BO23, UAZ B623 also 
has extremely broud palatines with extensive anterior 
espunsions about half way along their lenvth. A 
sigle nurrawer extersfon an the left palauine is 
present in. NTM R249(2- The hyord on AZ 8623 is 
extensively caleifiod (Pig. 34. The fronloparietals are 
more extensively ossified poslerolaterally in UA 
62649 und the palatines are more extensively 
expanded laterally in QM 155568. 


Myolowy 

The om. depressor mandibulue arises from the 
dorsal faseia and the tympanic ring but no fibres 
originale from the ole ramus, Phe anterior miarein ol 
the m. corucoradialis ts exposed rather than being 
completely bidden by the m. supraearacoideos and 
m. episternohdmeratis (Pig. 9), The min. linibriealis 
indicis brewis and Mexar teres indicis insert pennately 
on the palinar surfice oF the inetacarpal (Mig, 1B) 


Comparison with olher species 


Linmodvnasies depressus is morphologieally 
similar to L, fasmenioasts and L. flethers. W differs, 
from both of these species in the loss ofa pbalwnx en 
the lirst Jinger. although i shares the externally 
elongated thumb of 1. flercherd (Pig. ©), Prom 2, 
Helehert, Le depressuy is Turther distineuished by a 
more widely exposed lrontopurietal fonranelle, 
absenee ofa maxillary process on the fatsils cud bry 
iS narrow unespanded alae of the purasphenoid (Fig. 
Th From £. retwicfensin, Lo -depressuy ts Turthes’ 
distinguished by its narrow unexpanded alae of the 
paraspbenoid. by it more extensive ossification of 
the prootie and by its shullosver pars facilis of the 
nail (Fig Sy 

Myologioully, Lo depressus resembles closely, 
Hetcheri und Le tasmuanensiy in the miesculature ol 
the throat, Jeg and foor Consistent dittersnces oecus 
inthe musculature of the jaw, pectoral girdle und fipst 
finger. 

Inf. fletchert and Lh. lenmmetrtensin, Une a. 
Jepresser mandibulite originales trom three stless (he 
dorsal fascia. the olig ramus of the squamosal and (he 
posterior margin of the tympanie rings in’ LZ, 
depressay The miserGon on the otic rumius is licking. 

The pectoral) tiuseufature as Sinilar te all three 
species, except thal the mm coraceradialis is 
completely bidder by the in, supracaswedeuds ahd 
ne cepestermohumerais om Le fletcherr and Lb. 
wasniaists, Whereas in de elepressin, the anterior 
margin of the m. coracoradialis ts exposed (Pig. Y). 

Ty most frogs. meludime Lo fleroherd and b- 
fasmieuieiyts. the mn. lumbriealis indies brevis dnd 
flexor teres indicis maert Go the bash plait ol 
digit WoeF the hand (Pig, HOAY Ti Lo depeescis, 


Ida M, DAVIES & T. C. BURTON 


Fig. 3. Female Linvodynasies depressus (UAZ B2645). A. Lateral view of the pelvis. B. Dorsal view of the vertebral 
column. C. Dorsul view of right hand. Scale bars = 1.0 mm A, C; 5.0 mm B. 


REDEFINITION OF LIMNODYNASTES DEPRESSUS {45 


Fig. 4. Pemale Linnodynastes depressus (UAZ B2645). A. Dorsal view of the left foot. B. Ventral view of the hyoid. Scale 


bars = 1.0 mm, 


Pig. 3. Male Linmedynasies depressuy (UAZ B23). 
Ventral wew of the hyoid, Heavy stippling indicates 
calcifivation. Seale bar = 5,0 mm. 


insertion is pennately on the palmar surface of the 
metacarpal (Fig. 1OB) 
The larvae of L. depressus are similar to those of L, 


Hletcheri in general morphology (Davies 1992; Tyler 


& Davies 2000), although those of L. depressus are 
often much longer. However, they differ greatly from 
those of L. fasmaniensis in their pigmentation and 
the structure of the oral disc. Features of the 
chondrocranium also separate the species (Tyler & 
Davies 2000), 


Other material 


Prior to the collection of the material from Keep 
River NP. a species identified as Linttodviivtes 
fusmaniensis was collected at Kununnurra and was 
considered to be an introduction (Martin & Tyler 
1978). Further material collected at Newry Station, 
NT near the Keep River, was also alribuled to £ 
tasmantensis (Watson ef al. 1995). This latter 
material exhibited abnormalives of the fingers und 
loes (Figs If. 12) and the clongate thumb was 
considered to be another abnormality. A high level of 


146 M. DAVIES & T. C, BURTON 


Fig. 6, Female Linnodynastes tasmaniensis (UAZ AI461), A. Dorsal view of right hand. Female L, /letcheri (UAZ A1733). 
B, Dorsal view of right hand. Scale bars = 1-0 mm A; 5.0 mm B. 


iy. 7, Female Limnodvnastes flercheri (UAZ A1733). A. Dorsal and B.. Ventral views of the skull, Scale bar = 3,0 mm. 


REDEFINITION OF LIMNODYNASTES DEPRESSUS 


\47 


Mie. O Indes (first) digit of right han of Lininoedviastes 
depresses palmar surface with lendosuperhiciulis severe 
did cemoeed, PY son flexor teres: LB = ay luinbriedlis 
brevis: OPP = ni opponensindicis, US = severed tendon 
of tendosuperficialis, 


hig. 10. Ventral view of right side of pectoral girdles of A 
Linnedynastes fasmatienyix ant B. 1, depiressius, CR 
Mm vorgcoidialiss SC =m supravoracoideus. 


148 M. DAVIES & T. C. BURTON 


Fig, IL. Male Limnodvnastey depressuy (OAZ B623). Dorsal views of bones of A. left and B, aight hands. Ventral views of 
C teft and De right hands, Scale bar = 5.0 mm, 


REDEFINITION OF LIMNODYNASTES DEPRESSUS {49 


Fig. 12. Male Limnmodynasies depressus (JAZ, BO23). Dorsal views of bones of A. left and B. right feet. Scale bar = 5.0 mim. 


150) M. DAVIES & T, C, BURTON 


abnormality was detected in other frogs collected af 
this site (Watson er a/. 1995), Parker (1940) recorded 
disarticulation in the terminal phalanges of 1. peronii 
(Duméri] & Bibron) (which usually shows a similar 
reduction in the phalanges of the thumb as in’ L. 
depressus), similar to that recorded in the Newry 
Station specimens. Parker’s material, however, had 
two terminal phalanges. He commented that the bone 
may penetrate the skin, This was not apparent in the L. 
depressus specimens. Limnodynastes peronit dilters 
substantially in morphology from L. depressus. 

One of us, TCB, has dissected the material from 
Kununnurra and has identified the elongate thumb 
with associated musculature found in the Keep River 


material, Together with the abnormal material from 
Newry Station, we attribute all of this material to 
Linmodynastes depressus, We have further examined 
material collected from other localities in’ the 
Northern Territory and attribute all these specimens 
to L. depressus. 


Acknowledgments 


We thank P. Horner (NT Museums and Art 
Gallery), P. Couper, (Qld Museum) and K. Aplin, 
(WA Museum) for the opportunity lo examine 
material in their care and S. Walker for assistance 
with the figures, 


References 


Davits, M. (1992) Developmental biology — of 
Linmodynastes ierraereginae and L. fletchert (Anura: 
Leplodactylidae; Myobatrachinac). Trans. R. Soe. 8. 
Aust. 116, (17-122. 

Dincerkus, G. & Unter, L. D. (1977) Enzyme clearing of 
Alcian Blue stained whole small vertebrates for 
demonstration of cartilage, Stain Technol. 52, 229-231. 

Martin, A. A, & TyLer, M, J. (1978) The introduction into 
Western Australia of the frog Limnodynastes 
faxmarniensis Giinther, Aust, Zool. 19, 321-325, 

Parker, H.W. (1940) The Australasian lrogs of the family 
Leptodactylidae. Newit, Zool, 42, 1-106. 


Tyter, M, J. (1968) Papuan hylid frogs of the genus Ayla. 
Zool. Verhandl. (Leiden) 96, 1-203. 

(1976) A new genus and two new species of 
leptodactylid frogs from Western Australia. Rec. Wess. 
Aust. Mus, 4, 45-52. 

& Davirs, M. (2000) Developmental biglogy 
and Jaryval morphology of the frog Linmnodynestes 
depressus Tyler (Myobatrachidae: Limnodynastinae), 
Trans. R, Soc, S. Aust. 124, 169-175, 

Warson, G. F.. Davies, M. & Tybur, M. J. (1995) 
Observations on temporary waters in North-western 
Australia. Hydrobivlogia 299, 53-73. 


NEW RECORDS OF THE CESTODE GENUS 
PSEUDOTOBOTHRIUM (TRYPANORHYNCHA: 
OTOBOTHRIIDAE) FROM AUSTRALIAN FISHES 


By I. BEVERIDGE*, R. A. CAMPBELL? & M. K. JONES# 


Summary 


Beveridge, I., Campbell, R. A. & Jones, M. K. (2000) New records of the cestode 
genus Pseudotobothrium (Trypanorhyncha: Otobothriidae) from Australian fishes. 
Trans. R. Soc. S. Aust. 124(2), 151-162, 30 November, 2000. 

Pseudotobothrium dipsacum (Linton, 1897) 1s reported from the Australian region for 
the first time from various species of teleost fishes and is redescribed and compared 
with specimens from other parts of the world. Pseudotobothrium arii (Bilgees & 
Sharkaut, 1976) comb. nov. is redescribed based on specimens from the catfish Arius 
graeffii Kner & Steindacher, 1866 from Queensland and is assigned provisionally to 
the genus Pseudotobothrium. The relationships of Pseudotobothrium with other 
otobothriid genera are discussed. The analysis of anatomical features presented 
indicates the validity of the genus Pseudotobothrium and that it possesses an atypical 
heteroacanthous armature consistent with its position within the family Otobothriidae 
Dollfus, 1942. The family Pseudotobothriidae Palm, 1995 is therefore considered a 
synonym of Otobothriidae. 

Key Words: Cestodes, fishes, new host records, Pseudotobothrium, Otobothrium 
dipsacum, Otobothrium arii, Otobothriidae. 


Transacsions af ie Raval Sov tery of 8. Aust (2000), 12442), 151-162. 


NEW RECORDS OF THE CESTODE GENUS PSEUDOTOBOTHRIUM 
(TRYPANORHYNCHA; OTOBOTHRIDAE) FROM AUSTRALIAN FISHES 


by 1. Beverinar. R.A, Campenni, & M. KK. Joses’ 


Summary 


ievewtoor. d, Campari, RAL & Joss. MLK. (2000) New revords of the cestode genus Myeadoreboririin 
Chiypamorhyneha; Olobothriidve) from Australian fishes, Tras A See S$. Must P24) 151-1020 40 November, 


200), 


Peeudorobotirin dipsacuen (Linton, (897) is reported from the Australian region for the Hirst ime tron 
yavious species of teleost fishes and is redeseribed and compared with specimens Tram other parts ofthe workd, 
Pseveotobothriwn avil (Balgees & Sharkaut, (976) comb, nov. os eedeseribed based on specimens from the 
catfish Arta eraeffit Kner & Steindacher, 1866 from Queensland and is assrened provisionally tm the genus 
Prenlotobathrinm. The relationships of Preudofoboplietane with other otobathriid genera are discussed. The 
dnalysis Of anatonneal features presented indicates the validity of the genus Pyeuderohoriridee and that it 
possesses din atypied! hetcrowwanthous armature consent with (ls position walhin the finily Orobothriidiae 
Dollfus, 442. The Himily Pseudotobutiriidae Palm, LYS ts therefore considered a synonya of Olathe tie. 


Kiev Worps: Cestodes, fishes. new host records. Pyeadetaborinin. Orebatiriane dipruciin, Oreborliriian 


ari, Olobothriidie. 


Tntroduction 


Cestodes (tapeworms) oof the order 
‘Trypanorhyncha Diesing, 1863) occurring in 
Australian fishes are sull telatively poorly known 
since large numbers of potenbal host species as well 
as peographic regions around the continent romain lo 
be oxamincd. Amongst the most poorly investigated 
funilics of this cestode order is the Otobothriidae 
Dolllus, 1942. currently represented in this region 
only by Otehothetum ningiliv Uiseock, 1954 
(Hiscock 1954) and Poeedlanecisuan caryephayliuu 
(Diesing, 1850) (Beveridge & Campbell 1996), 

In the present paper, the oecurrence of the genus 
Pyendotobothriin Dollis, 1942, based on its only 
known species, / dépsaenin (Linton, 1897), ts 
reported for the first time in fishes from northern 
Australia Onhothrinn aril Bilqees & Shaukat. 1976 
alse reported front teleost fishes rom Queensland 
and is placed within the genus Myenderobatiriun. 
Becuuse the definition of the genus und deseripuons 
of both species included in this report are incomplete 
(Beveridge er ul 1999), redescriplions based on 
Australian speemmens as well as other specimens 
available for cxumination in museum collections ae 


Dypariiient i Veterinary Sciaee, The Universiyy ot Metbourne 
Parkville Vig WB 

Depultnent of inlay. Hiversity of Mussiehosetls Durtioutl, 
Noth Dudmouth 20705 USA 

Conine for Microscopy & Mideeaatvsis the University al 
Quevnshind St Luci Qh 472 


provided. Inaddition, the taxonomic relittionships of 
Preuddobothrivn witty the family Otobothriidae 
aire reassesace, 


Materials and Methods 


Cestodes collected by the authors were placed in 
Lap water to induce evagination of the tentacles and 
were (hen fixed swith 10% formaldehyde or 70% 
cthanol. Specimens were subsequently stitined with 
Celestine blue. dehydrated in an ethanol: series, 
cleared in methy) salicylaie and mounted in Canada 
balsam. ‘Tentacles were detached from strobilae 
usthg a scalpel and were then mounted in glycerine 


jelly Jor examination of the tentacular armature, 


Drawings were made usmy a BH-2 Olympus 
microscope with Nomarski interference optics, fitted 
with a drawing (ube, Measurements are presented in 
micrometres unless otherwise stilted as the range 
followed by the mean and number of specimens 
measured in parentheses, 

Specimens in the collections of the South 
Australian Muscum, Adelade (SAMA), the 
Queenshiid Museum, Brishine (QM), the British 
Museum (Natural History), London (BMNIL). the 
Muséum national a Histoire naturelle, Pars 
(MNUIN) and the United States National Parasite 
Collection, Washington (USNPC) were examined, 

A complete synonymy for Jt dipsnoune wats 
provided by Dollfus (1942), Consequently, the 
references cited here are those in whieh novel host ar 
scouraphic records are listed including those ened by 


I. BEVERIDGHE .R, A, CAMPBELL & M. K. JONES 


{52 

Bates (1990). Records such as those of Linton 

(1914), Pinther (1934), Joyeux & Baer (1936) and 

Southwell (1913, 1930) which merely repeat 

previously published accounts have been excluded. 
Host nomenchiture follows Paxton ef al, (1989), 

Robins er af. (1991) and Allen (1997). 


Pseudotobortrinn Dolllus, 1942 
Type species > P. dipsacum (Linton, 1897) 


Pyeudotobothrium dipsacum (Linton. 1897) 
(FIGS 1-9) 


Orobothrium dipsacum Linton, 1897, pp. 806-807, 
pl. 64, Migs 1-5 (Pametomus salratrix (Linnaeus, 
1766), Atlantic, North America, Massachusetts): 
Linton, 1901. pp. 412, 451 (Pomatomus sattatrix); 
Linton, 1905, pp. 329, 331. 375 (Centropristes 
stridta (Linnaeus. 1758), Atlantic. North America, 


North Carolina) : Southwell, 1912, pp. 270. 278. figs 
19-21 (identified as QO. insigne) (Bpinephelus 
undulosus (Quoy & Gaimard, 1824) (=Serrannus 
undulosus), Parastronateus niger (Bloch, 1795) 
(=Stamateus niger), Diagramma crassispinun 
Rtippell, 1838, Balistes spp.. Sri Lanka (illustrations 
are not of O. dipsacum); Linton, 1994, p, 2, 53 


Figs bea. Prendoioborhrinn dipsacun (Linton, 1897). | 
Scolex, 2. Bothridial pit, lateral yiew. 3. Tentacular bulb. 
showing oriia of retractor musele. Seale bars = 0,1 mim. 


CESTODES FROM FISUES \S5 


(Ahiferus sehoepfi (Walbaum. 1792) 9 Massachusetts: Treehinotus falcarts (Linnaeus, 
(=Ceratcanthus schoepfi). Xiphias  gladius 1758) (=Mycteroperca falcata), Atlantic, North 
(Linnaeus, 1758), Atlantic, North America, America. Florida); Southwell, 1924. p, 489 


Figs 46, Pseudotobothriun dipsacwn (Linton, 1897), basal and metubusul tentacular arovature. 4. Antibothr: ne surface ol 
tentacle showing origins of ascending hook rows wilh slight space between files Fund [in metabasal region. 5. External 
surface of tentacle showing ascending rows of hooks from left to right and band of hooklets on bothricicl (right) side of 
tentacle, 6, Bothridial surface of tentacle showing band of hooklets in centre wih prominent space on either side of the 
band. Scale burs = 0.01 mm. 


154 I. BEVERIDGE , R, A. CAMPBELL & M. K. JONES 


(Epinephelus undulosuy, Diagramna crassispinum, 
Sri Lanka); Southwell, 1929. pp. 291-292, 311, fig. 
47 (Epinephelus didulosus, Diagramma 
crassispinum, Sufflamen fraenatus (Latreitle, 1804) 
(=Balistes mitis), Litjanus dodecacanthus (Bleeker. 
1853); Lethrinus ornatus Valenciennes, 1830, Sri 
Lanka, p. 311, Abalistes stellatus (Lacépéde, 1798) 
(=Balistes srellatiy), Lethrinus ornatus, 
Parasiromateuy niger); Yamaguti, 1952, pp, 69-70, 
fig. 105 (Chelidenichthys kamu (Cuvier, 1829), 
Japan). Palm et al, 1994, pp. 153, 156, 159, fig. 4 
(Cynoxulossus senegalensis — (Kaup, 1858), 
Petrocephalus bane (Lacépede, L803), Gulf of 
Guinea), 

Orahativium  (Pseudotobothrium) —dipsacum: 
Dolltus, 1942, pp. 253-255, fig. 157 (Polynenuts 


guadrifiliy Cuvier & Valenciennes, 1829) East 
= \ 
AO 
Ls | 


Atlantic, Africa (Congo))(identification uncertain); 
Cruz-Reyes, 1973, 25-29, figs 1-5 (Balistes polvlepiy 
Steindachner, 1876, Pacific Ocean, Mexico). 

Pseudotobothrium dipsacnim. Ward, 1954, p. 255, 
fig. & (Sphyraena barracuda (Walbaum, 1792), 
Mian, USA): Palm, 1995, pp. 102-103, 154-162 
(Haemulon plumieri (Lacépéde, 1801), Brazil); Palm 
et al.. 1997b, pp. 71, 72, 75 (Pseudupeneiy 
maculatus (Bloch, 1793), Haemulon plunieri, 
Atlanuc, Brazil), 1997b, p. 84, figs le, Id. 


Types: Holotype (larval) from Pomatomus saltatrix 
(Linnacus, 1766) (USNPC 4794), 


Material examined: 
From Australia: 
from Abalistes stellatus (Lacépede. 1798); 5 


Figs 7-9. Profiles of hooks from the tentacular armature of Pyeudetabothrium dipsacum (Linton, 1897), 7. Metabasal region, 
trom left, first hook in row, hook in middle of row, hook near ead of row, terminal hook of row, outer hooklet (rom band. 
Inner hooklet from band. 8. Hooks of rows 10-15 from the base, from left, first hook of row 10, first hook of row I5, 
hook in middle of row 15, hook at end of row 15, outer hooklet of band, inner hooklet of band. 9. Hooks fron base of 
tentacle, from left, lobed hook on antibothridial surface, outer hook of band, inner hook of band, Scale bar = 0.01 mm. 


CESTODES (ROM PISTLES 155 


specimens. Heron Island, Qld (QM G21 7928-32), 

front Cuphalopholix cvdanostigne (kubl & vin 
Hlassult, }$28)° | specimen, Hergn tshand. Qld (QM 
C2 14950): 

fron Epinepheluy suilliiy (Valenciennes, 1828): 7 
specimens, Cape Clovelind, Qld (SAMA 31342): 

from Usfiopharus platypleruy (Shaw & Nodder, 
1792): 4 specimens, Cape Bowling Green, Qld (QM 
C2165, ITY), 

Irom) Meheiva medica (Cuvier, 1832): 2 specimens: 
Cape Bowling Green, Qld (QM C2) 2166, 212800), 

from Makatra moa Gordan de Snyder, 01); 2 
specimens. Cape Moreton, Qld (QM G212785, 
212798); 

fron Nase vlemningi® (Valenciennes, '835) 1 
specimen, Heron tslund, Qt (QM G21 2900), 

from Pyendecawe dientex (Bloch & Schneider, 
TSUL) 2 spediniens, Heron Island. Qld (QM G2t4904_ 
217930): 

from Plectaponres leapeardus (ateépale, 1802); 2 
specimens, Heron Island. Old (QM Gi2|4962), 

from Plectrapomus maedatis (Bloch 1790}: | 
speemmen, Eleron Psland, Qld (OM G206964); 

from Ritnecanthas rectaiiwitatis. (Blow & 
Sehuvider, FROM: 3specimens, Heron Ista, Qld (QM 
C2179OIAD) 

ron) Gynmevirdd uniedlor (Ruppel, LRaG) 2 
specimens, Clerke Reet, WA (SAMA 31343). 

From Indi Oceans 

trom Lanpiiy pibbus (Borsskal, (775); L specimen. 
Hulile, Malutives (QM GL 10508). 

froin Past Africa: 

lrom Cephaleyholis sonmerar (Walonciennes, 1528), 
| speeitnen. Zanzibar (BIMNE 196 (.6.26 12065); 

lant Epitephelus matabarions (Bloch & Schneier, 
PHOT), S specimens. Zanzibar (BMNEL }961.6,26.1 20- 
Ay; 

trom. Epinephatus tanvina Forsskal, V77S. 3 
specimens. Zanzibar (BMNE [961,6.26,1 20-5): 

from Aprneplielity chlorostigma (Valenciennes, 


PRI); 4 specimens, Persian Gull t&8MNEH 
1992, 7 13bTS), 

Bron Sei Lanka (Ceylon; 

from Sufflemen freeralis (Latreitle. 1804) 


(=Balhstes miss: | spectnen (BMNE 1997 |) 15 33- 
Ty 

from Abafivtes sfedlaniy (Lieeptde. 
specimens (EMANI L9O7Z 11S. 39 7); 


ITON): 


from Lellitiuas enmnetin Valenciguites. (830: 2 
specimens (BMNEL 1977, 1.15, 31-2) 
Fron Levhvinis spa 7 specimen CBMNIT 


L977 ALAS. 38-62): 

from Epinephelis tndilasis (Quoy & Gaiimard, 
R24) (SSerriniy nidilosus): 3 specimens (USNPC 
081A), 

From Saath Amerca: 

from Heenan planet (Lavépede, TsOL) | 


specimen, Brazil Gin personal colleetion of HL Palin). 

From North America: 

from Hoamaianiy saltatris (Linnaeus. (760): type; 

fron Myc rerapercn plonas Jordin & Swath Ta84: 8 
xpeciinens, New York Aquurliim (USNPC 35777): 

trom Haemulan purrs (Desopsrest, |8I3) (= 
Neomeenis pare: 2 specimens, New York Aquarittin 
(USNPC 35780, 35781); 

trom Canthidennix sufflamen (Mitehill, TSiSi: 2 
specimens (UISNPC 35782); 

from Cuntiderms macialus (Bloch, 1780) | 
specimen, New York Aquanumn (USNPC 35852); 

trom Qc yuris clirysreay (Bloch, 1791): | speemmen. 
New York Aquarium (LISWPE 35783): 

from Harpe rufa Linnaeus, 1754: | specimen, New 
York Aquarium (USNPC 35850): 

fram Scorpacnd plunteri Blovh. 1789: | specimen 
(USNPO 35851); 

from Diarape analis Cuvier & Valeneienies. }840 
(=Neomaenis analis): | specimen, New York 
Aqnarium (USNPC 45653: 

from Laclmolainas meaninus (Waulbauim. 1792). 1 
specimen. New York Aquariti (USNPC 36028). 


Desevaprens 

Meastirements from Australian Specimens. Scolex 
3.3-5,0 (4.2, 9=10) ti long maximum wielth at 
postertar extremity (.83- 2 C122) ne LO) nibs pars 
buthridnilis 1.05-140 (1 38, n=10) nm, 2. bothricive 
with thick mucins posterior margin of ouch 
bolhiidido with 2 prominent fossetes: pars 
vaginalis virible in length, depending upon sttte ol 
gontraction of specimen. 1.683,40 (2.58, b=10) min, 
prominent longitudinal museles belween tentacular 
sheaths; bulbs elongate. 119-170 (142. n=) mm 
fon. 0.20-0,30 (225, r= 10) tim wide, lengthswidth 
iitio 4.10-7,35 (5.92, n=), postertor exwenmities oF 
bulbs direeted laterally, terminating at postera-hiteral 
margin of velam: prebulbur organ absent: retraetor 
muscle onginating from anterior hall of titernal wall 
of bulb; no gland cells present within bulbs; scales 
ratio (pborpyagipbulb = Ps T.2a 1.10); scutes 
eruspedote. length of velum 0,26-0,50 (0.39) n=O) 
mm lon. pytidiiin O40-0.65 (0,54. n= 10) nid tong 

Tenucles of relauvely uniform diameter for most of 
Joneth. without busal swelling. slightly narrower ab 
bane. SO-150(720, H= 10) in diameter at base, }00-L60 
(140, n= 10) in diameter wi metabasal regions 1450- 
2910 (1940. n=10) long when Milly extended. rapering 
at up. sith apprax. 125 rows of hooks. Ariatune 
haiaraucanthous. bereromorpbouss hooks hollow. ne 
distinct basal armiuture. Llooks. arranped in aseending 
half-rows) yows heginnine on sAtibothridial surkice 
terminating on bothridid surfaces slhr spave 
belween hook Gles Danid | or dntibothridial surfer, 
most prominent between rows 6 ahd 20. number ab 
hooks changes wong tentacle: 10 rows [ron base, 1 


156 !. BEVERIDGE , R. A. CAMPBELL & M. K. JONES 


Figs 10-13. Pseudetobothrium arti (Bilqees & Shaukat, 1976) comb. nov. 10, Scolex showing lateral extension of posterior 
region of pars bulbosa. 11, Bothridium showing bothridial pits (p) at posterior extremity of bothridium. 12, Tentacular 
bulb, showing origin of retractor muscle. 13. Basal and metabasal armature, bothridial surface, showing interlocking files 
of hooks at base and origins of ascending hook rows of metabasal region. Scale bars = 10-12, 0.1 mm; 13, 0.0L mm. 


CESTODES FROM FISHES (57 


we NM 
nt \| 


Wye Lf 

CNL) 

ih, We 
{ 


| 
Ht 


Fivs 14-18. Pseudotobothriun arii (Bilqces & Shaukat, 1976) comb, nov. Tentacular armature. I4. Metabasal region, 
bothridial surface, showing ongins of hook rows. 15. Metabasal region, external surface, showing ascending rows of 
hooks on left and band of hooklets on right. 16. Metabasal region, antibothridial surface, showing central band of hooklets 
with flanking file of hooklets on each side. 17, Basal region, external surface, bothridial uspect with interlocking large 
hooks on right hand side, bill-hooks on antibothridial surface on left hand side. |8. Basal region, oblique view of 
antibothridial surface showing bill-hooks on antibothridial surface of base and origin of band of hooklets in metabusal 
region Seale bar = 0.01 mm, 


}5e | BEVERIDOE, ROA CAMEBRIE & MOK. JONES 


linoks per prineipal row; 20 raws frond base, 14 hooks 
per rows, 25 rows from base, 40 hooks per pow; 
increase iy Number OF hooks occurs al origins of rows 
on ualibothridial surface: number of hooks per row 
diminishes towards tp ol tenticles LO rows: from 
base. 30 hooks per row, 120 rows trom buse, 20 hooks 
per how. Hooks change slightly in shape along 
tentucle. AL base, hooks on antihothridial surface 
wneinile, blade tips hilbous, buses broad, hooks 20-33 
(28. 9=10) long. base 12-20 (18) n=O): on bothridial 
surface oF Lenicle hook tips sharp, bases elongates 
frat rows 5 lo 10, figoks 1) i small, uneinale, 21-33 
(26, n=I0) lone. base 21-25 (23. n=I)). blade 
becoming much longer than huse alone pow, larwest 
hooks ol rows elongate, faleate to triungular, 50-62 
(56. n= 10) long, base 13-2001 7, = 100; Front rows FO- 
LS, hooks 11) diminish in sive, remain uneinte: 
hooks 70779. SCR") clongaite. flee to spinilorm; Tram 
row TQ distully. hooks 1) te 20 (20) siall, 
Linen’ With namow base and sharply rebated hook 
Hip, 13-19 (15, n=10) long, base 6-9 (7, n=10), closely 
packed; hooks on internal and exterimal surtiees 
elongate. spinifonn. longest hooks of cach raw 52-67 


(60, n=l) long. hase 14-30 (18, n=LOy Pinal file of 


hooks Of each principal row 45-50 (44, n=10) lone, 
hase $-17 (14, 0=10), On bothridial surface, distil to 
row 4, Space presen! between finul hook al principal 
row ind eentral bund OF LO files Gl smaller hook lets 
At base of tentacle. all hooklets of band uneinate, with 


clongale hase; i Metabasal revion, houklets of ouler 


Hlep with short bases, 32-36 (35, n=10) long. buse &- 
LH (TO, n= 10): Hooklets ef inner files retain elongate 
buses. 26-44 135. n=10) long, hase 12-22 (16, n=10), 
Band ob haoklets with | row of hooklets tor each 
prime hook rows cerungement of band of haoklers 
Hot perfectly restile Adult unknown, 


Pscudotaborhiiiin ar 
(Tilqecs & Shaukat, 1976) comb, nov. 
(PIGS 1-bs) 


Oebotirime ari Bilqees & Shank, 1976. pp 
TTY 124, lig, | vedo serranis Day. 1877, Karachi), 

Wes > School ol Parasiialogy, Department of 
Analogy. Civiversity of Karachi (Mor exwniied), 


Malernthexcanmed: 

Front Artis servers Dayy US77 > Pakistan oc | 
specimen. Rirucht ident by Dr FM. Bilyees, 
BMNT 19S4L5 Ps. 14. 

Prom Aries geese? Kier & Sterndschner 1866: 
Austiline 2 specimens, Brisbane, Queensland, coll, 
M. K. Jones, Rax_1997 (SAMA 2827)), 23.4.1997 
(SAMA 28266) 


Deveripnine 
Measurements from Austealkin speciinens Scoles 


3.4, 35. 4.0 rom long, maximum width in purs 
vaginitlis O15, 0,16. 0.22 mii. at posterior extremity 
0.94 (I, 1.23 mins pars bothridialis O45, 019, 
H.55 niin, 2 bothridia with distnet miurgins, posterior 
murgin of each bothridivm wilh 2 small bur 
prominent fossettes: pars yugsialis: variable in 
lengih. depending upon siile of contruction at 
specinien, 2.74, 3.11, 3.50 min; bulbs elongate, 0,50- 
O70 (0.57, (=5) tim long, 0.15-0.22 (18. n=) mm 
wide. lenpihwidth atin 2.73-3.68 (3.26, n=5). bulbs 
directed almost daterally, terminating ab poster 
lateral inarsin Of vellum; prebulbur ore whsent 
relniclor muscle originating dy anterior extremity ol 
internal wall OF bulb; no whand cells presenr within 
bulbs: seoles craspedote. leagedr of yelum Q.15 nin 
scoles nitio (phogevagephulh) = 16.241 1a. 

Tenticeles with distinet basal swelling, 50, 60 in 
digmeter at base, 40. 40 in diameter in metabasal 
resion, Armature helgraacanthous. heturomorpheus: 
hooks hollow: distinct basal armature. Basal 
armatare with hooks Ll) an bothridial surface 
groully enlarged, bases clase together such {hat 
blades interdigitale, 40-70 (57, n=10) long, base 2U- 
A126, n=5), Hooks on posterior part of internal und 
exrermal surfaces of basal swelling arranged ine 1 
rows. Houks spiniform with narrow biases, [9-27 (22, 
N=10) long, base b-8 (6, n=10): anterior part af basal 
swelling on both internal and external surfaces 
devoid of hooks, External surkive of basal swelling 
with row of billshiuped hooks, 25-32 (28) w=lop 
Jong. base 5-8 (6, n=10) Metubasal aimature; hooks 
urringed ih adscendine hallrows; rows begin on 
bothridial surface, terminate on antibotlridiat 
surface: 6 hooks per principal row, very slight space 
between hook Jiles | and 1 on bethridial surface, 
diminishing in width posteriorly and disappearing al 
level oF base. Hooks 115) large, umeinte. 2839 (30. 
n=5) long, base 1826 (21. nea). Hooks 22") 
uncinite, somuler, 19-30 (25, n=lOy long, hase Tet-20 
(17. b= NO), howks 3(37) reel, Uneinate with alarply 
recuipved tipsy und narriw bases, 17-32 (22, w=10) 
long, Wase 5-8 (7, n=lO) hooks 4145) ty @(@)) 
spiniform with very narrow biases and recurved lips. 
20-31 (25, n=10) long. base 4-1 (6, m=10). On 
anlibothvidial surfaee, space present between ful 
lok of principal row (666")) and central band af 
hovklets. Outer file of hooklets spinieun, 17-40 (23 
n=10) long, bise 4-6 (5, n=l). separated from 
central band Of 4 files of spiniform hooklers [8-30 
(23, n=10) long, base 4-6 (5, n=lOh single maw vl 
hooklets per principal row, Adult vakroawe 


Discussion 
Pseudotehoihrinn dipsacuin has beet desc bed 


by severul authors Linton (h897) gave a briel 
deseription of the species bul did not describe: the 


CESTODES PROM EISHES (Su 


virmatire in deka, apart from. nating the closely 
spaced rows oF hooks ane. (he merease i number of 
hooks per row along the tenkicl The redeseripion 
provided hy Cruz-Reyes (1973) was ere detailed 
hut the description of the arnmlure was limited bo one 
suiface Of the tentacle. that on whieh the hook rows 
COTTIER 

‘alm oor al. (1994) provided seuining cleeien 
Wicroweaphs of the aarmatire. while Palm (1985) 
provided a bret summary of the morphological 
lealures of P dipyee di and a re-interpreuition al its 
armaiure, He (1995, 1997a) proposed that the 
urimalure was of the typical heteroweimibous type 
with the hook cows beginmnig on the internal surface 
ald terminating on the extermal surtuce of the 
lenlacle, without extn rows OF hooks On the fatter 
surface, ‘The armature tag previously been 
considercd ta be of the atypical heterowcanthous type 
in Whieh additional hooks wre usvally present on the 
caleral surface of the tentacle (Dallas 1942). On 
The basis of this resinterpretanon, Palm (195) 
ereated a new family, Pseudotobothriduc. for the 
species, although le cited it subsequently as 
’pecudétobothriidae Ward, 19547 (Palm L997b. p. 
75), The current re-description af the species 
provides. morphological evidenee whieh contradiets 
Pulni’s (1945. 1997a) hypotheses, Firstly, the hook 
rows in Po dipyacue begin oo the antibothridia! 
surface OF the tentacle und end on the bethriial 
stuirfaee, a feature which has not been recorded 
previously In this species, In most irypanorhynchs. 
the priacipal rows begia on the internal surlitee and 
ferminate on the extemal surface (Dollfis 1942; 
Campbell & Beveridge }994). Fxeeptions to this 
patiern are (he veners Aulorelia Euzer & Rudujkovic. 
JO8Y and Prochristiuretia Dolltus. 1946, sith rows 
besinming on the bothridnd surface of the lenticle 
(Campbell & Beverulye 1994. Beveridge & Jones 
2000), Palm ef af (1994) and Palos (1098. figs 155. 
157) mnislakenty identified the antibothridial surflice 
as dhe ester surface and the bothridtalsuplice as 
the internal surface in deseripuons of the armature, 
Palm subsequently (1997b, fies led) identiled the 
fuoltyriatial Suface as the extemal surlaee, 

‘The second sigmficant feature, noted here for the 
fies! time in this species, is the spave between the 
pomipal rows ab hooks and: (hase eccupying the 
ventee OF he barhridial surfuce of the tentacks. This 
apace iS clearky visible in various views of the 
lentiivle bul niav be obseured when detuched 
Lonlacles are jrinipulated in wlycerine (rig. 5), 
compressing supe Hooks avainst the surface of the 
endieley builds fig lo Le spaces observed, there are 
siunificunt differences in Shupe bebyeen the haoks at 
the ends of the pridcipal vows ind those in the eenue 
ofahe boleh suree of the jentiade (Pies 7. 5): 
For these coeusous. (he cite precarion dyanced here ts 


that the principal rows terminile as They approach 
ihe bothridial surfiiee and thar the hooklers on the 
bothridial surface form a “band”, wath cach row tn 
the bund conesponding to a principal hook row. This 
iiterprediiion conforms with the mite 
observations Of Linton (1897) and Dollfus (1942) 
that there are ne “extra” tows of hooks on the 
bothridial surluwe. The distinct “band” is a 
chargeterisiie of the pokverlowcuntlions trypunar- 
hynehs in the system at Dolltus ()942) or the 
otohothrioids in the syste ol Campbell 
Beveridge (1994) and is a feature af tbe relate! 
otobothriid genus Poeeianeisioine Dolttis. (929, 00 
whieh there are (hme sows ol hooklets fot euch 
principal row (Beveridge & Campbell 1996), 

Consequently. Po dipsacua is an plobothvierd 
Irypanorhyneh on the basis ef is armature aod ts 
alhed with Poeeilineistuin im whe family 
Otobolhriidae, An tdditionil fimily. Pseudi- 
wobothriidue, is nol} reqired and We theretore plitce 
Palms (1995) family asa synonyin of Olobothiridie: 

There are. nonetheless several unique features in 
the amare OF Po dipweetwn. Ve dramidtic inercase 
in-numbers of hooks per principal row along the 
tenhiwle hus been reported Jom he other 
irypanorhynch cestode, The number ot hooks i (he 
pliingipal row is usually constant of diminishes 
distully along the length of the tentacle. In addition, 
is unusual for the final oy penuliimate hook ef the 
principal row to be the longest. Ustully, (he Lirgest 
hook is abihe commencement of er i the muddle ol 
the prineipal row. Thos there are adequate reasons 
for Tiaintuning 2 dipxdewi us an independent 
species. Comparison of Australian specimens: with 
imuterad in museum collections stugeests that # 
dipsdcuit is a costropolinu species which ts 
inorphologicully unitorm throughout its geographic 
rupee, The materiil examined from Austratia und 
other regions considerably expands the host range of 
the species, 

Dollis. (1942) sub-divided Wie wens Oveahotlrnien 
ial) }wo subpeperd and erected the sub-genbs 
Pre lombotiring (with lype species OL afjscdteinn) 
primarily on (ie basis of aimature The sub-genus 
Grohethrinn was characterised us having extra 
hooks or heel rows on the external surface of the 
tentacle. while in sevdeteberdrrian, (he numbers of 
houk rows were the same on beth surfaces ol (he 
renacle = Tolltus (1942) alse noted hac in 
Pseudoiobothrivan the bilbs were clongite. while in 
Oteboifiriium they were pererally short Yamiyity 
(1959) also subdivided Ololwelie sia) into siib-genern, 
basing the division primarily on the lengils of the 
bulbs ahd desigmaling ) (asmeivrus We iype species 
OY the sub-ocnus Pseidarebotiivias. apparently 
overlooking the duxontoanie deesiai af Dallles 
(2942), Schiniedt (MURO) dik nat uceopte Yonnawun’s 


160) L BEVERIDOE. ROA CAMPBELL & MOK TONES 


(1959) subdivision, while Campbell & Beveridge 
(1994) ueeepled the sub-genera bat. following 
Yarmagut (1059). errancously ene QO. finsawi as 
The type species of the Sub-genus Peenederehotletunr. 
Sehimidt (1986) dil hot recognise the two suh- 
Benen, 

Crie-Reyes (1974) redeseribed OQ. e/pyudeium. and 
vonsidered Orehoiiviin invivne Lintan. (905 as its 
synonyin for reasons which were nol exphined, 
Consequently, the hast list for ft dipyacun given by 
Malm (1995), which follows Ceuz-Reyes (1973). is a 
vompositg, As Go iivigne is here regarded as a vail 
species bused on te redeseription by Hildreth & 
Lumsden (1985), the valid’ records oF G2 dipaacune 
une therelore (Hose Tisled in the synonyiny. Dollfis 
(1942) reported us possible specimens of ©. 
dipydcuin, pleracercd eollected front Pelviesiuy 
quadrifilis and Brachvdemetas duritus (Valenciennes, 
ISAL) (=Ofoperce anette) From Pointe Padrom: 
Congo (=Zaire), The iMestrations of the specinetis 
(Dollfus 1942. fig. 157) suggest that bey belay tv 
un undescribed species relited (6 OL fisjune, based 
On specimens Mai the sure geographical region in 
Dollis’ collection (MNEIN) There ane nu 
specimens ol Pidgin in the Dolllus eallection in 
MNHN aind therefore Dollis? (1942) recurs have 
not heen included in the eurrent Wst of hests of 2 
lipsuteuni, 

Bilqees & Shaukat (1976) described Q. ari based 
wr pleragerci from fhe muscuhiture oF ae catfish. 
Arnos serratny, Cant Pakistan. They distiniuished it 
ron congeners primarily on the basis of qumber of 
Files OF hooks and the bulendly diverent bulbs. They 
did not, howeven give w detailed deseription of is 
armature. The three Adstraliu) specimens examined, 
also [rom a cutlish, Weis oraepfii, are identical to 0, 
wen based on Comparisons wilh a specimen in 
KMNE from the type host jind locality, identified by 
Dr Bilyees. Both the seolex shape and the armature 
are highly distinctive. The pairs viueinalis os 
extremely long and slender and the bulbs extend 
hitcedy. as an OQ. penetrans Linton, 1907, 02, 
poplrikoes Dolllus. 1969 and O duds? Shields, LORS, 
Palin (1995) comsidered Q, fvirist toy hea synonyny of 
© peretcans, Gtahothrinn penetransx has a basal 
swelling and distinelive basil arnraiure as does 2, 
onli, However while the miefibasal armature ol ©. 
PeETUUS Consists Ol principal rows af seven hooks 
WIth interealary hooks beeween each vow (Palin 
P9US: Palm ev al 1993), there ave six hooks jr the 
pricipell row pry. cari followed byw file of hooks 
CCNY) separaded trom {he previons tile by a distinet 
spuce (Pig. 16), In the eentre of the anribothiidial 
surbive ofthe fentiele is a band at hooklets. four lies 
wide. The metihasal arimatuee of QO. anit lifters 
thenctove from every species GF Otibetiaitin i 
WHICH Tis feature las been adequately desenbed (0. 


penetraus, OQ. peplirikes, Qo tsigie, GO oinnin 
Subhuprudha, 1955) brit resembles that of the genera 
Meqderwholtrian ail Poecilaneisreim. both of 
Which have bands oF hooklets on the external surlaice 
of the reniaele (Palm 1995: Beveridge & Campbell 
1996), Otebarhriiin avit ditters Trout Po dipseewe an 
possessine d distinctive basa swelling and armature, 
as well as licking wn increase in the number af hooks 
per principal row long the tentacle and having the 
rows Of hooks beginning on the bothridial sdeface ol 
the tentacle rather thaw the wntibothridial surtice as 
InP dipsaewn. UW differs tram Peeeilneisrrun in 
having only one row of haoklets per principal row 
conmpaed with three in Poecihuie isu (Beveridge 
& Cuimpbell 1996), 

Olobothriaint avitalse resembles Puerimacenihien 
owen, -desertbed by Palm (1995) Tron ain 
Unidentified) cartish from Papua New Gained Me 
seolex ol PF oewent uppers lo differ in having an 
estromely thick tegument, being muel broader iy he 
pars vaginalis (O75 mim ta Rowen’, O17 nim in @ 
arity and taying Jonger bulbs (1 nim in 2 eweni, 
O37 mm in @. ae). The basal urmatire of the twee 
species is rennrkubly similar They ditler fowever, 
in that while 2 ower? tas a chametle composed of 
iMangular Pooks on the external surtice ol its 
lenlwele, Gani has a hand of hooklets. 

While irmay be possible to advice dirguments tor 
the erevuion Of a new venns to aecommodale 2, ari, 
the existence OF (wo mmonotypie genera wilhin the 
Otobottiidie, Prevdaebedieiin and Paveilaneiseun, 
avs Well os uncertiinties convening celationships willl 
the Monotype Lonus Paeeiocuciwnthyo, leads is to 
WdUpLa conservative approach ip allocuting OC. arif ta 
the pets Pyeudalobolirivn Vhe definition of the 
vermis Must now include species with aud without a 
basal armature am with the hook rows originale ei 
either the bothridial or antibothridigh supfaces. ob Me 
lenacle. The key feature which distinguishes 
Pyendoloboiiviin Tron Gieberhrinn remains the 
chariwer qdentified by Dollfus (142), namely that 
(he number oF rows af books an the extemal surlace 
of the lentqele i he Sauineais ce the imernal surtiee 
In Pywedebotiriine whereas there ure tare paws 
oo othe extemal surface in Otebuaplrpen 
Prendotuboririin and Poeeilateisirum boi ditter 
from Otoborli(it id possess a bund of booklets 
ov the tentacle rather Wan a single raw on! beoks 
Hiterpoliied helween the  intereahery rows. 
Pyediobothrian differs trom Pacotaneisivan i 
Possessing ane cow OF band hooks per principal raw 
compared with three aaws of band hooks ty 
Poecilauenvtruin. AML theee pene differ fren 
Poectoacaithia whieh possesses-a chuinette in ihe 
meuhisal fesion, Dewi Of fhe Sirabilia as well vs 
the adelition of ney speejes to these genera muy 
facihihite (he ela ficatiog of theit relationships which 


CESTODES VU ROM FISHES Gl 


remain abscure at he present dine. 

The venus Prendofobetirium, as deseribed bere, 
includes Iwo distinctive species of cestodes, 
ilerinediate between Olohothriin and 
Poocilancistram, Womay ultimately meed te be 
dismembered, bul indicates a hitherto unsuspected 
devree of diversity within the Otobouiriidae, 
sigeestingy that additional novel species await 
discovery, A revised generic diignosis ts given 
below, 


Pyeudoohothriun Dollus, 1942 
Definition 
Olobothriidae Dollfis. 1942. Scolesx. craspedote, 
Two bothridia with pin of fossettes on posterior 
mirein. Pars vaginalis elongate, Bulbs elongate: 


refractor muscle originating from anterior park ol 


bulb: prebulbar organ ubsent. Pars. postbulhosa 
absent, Armiture Neteroacinthous, heteromorphous. 
fiooks hollow. Distinctive Digi! armature present or 
absent. Elook rows bevin on bothridial or 


anubothridial surfaces of tentacles, terminate on 
opposite surface of tentacle; space present betwee 
principal rows and band of hooklets on bothridial or 
antibothridial surface ot tentacle; band regularly 
arranged, with same number of rows as. prineipul 
rows. 

‘Type species! 2 dipsacine (Linton, 1897), 

Other species: Po arii (Bilyees & Shaukat. 1976) 
comb. nov. 


Acknowledgments 

The authors wish to thunk LL. R.G. Cannon, D1, 
Gibson and R, J, Liehtentels lor access to specimens 
held in their collections, B.G. Robertson, Fe Spoure, 
5.C. Barker, R. A. Bray, J) HE Cribb, Re. G, Lester 
and TL W. Palin for collceting. specimens examined 
duribe this study. and M, Gomon, The Muscum of 
Victor. for advice on fish nomenchiture. Financial 
support was provided by the Austealian Biologicul 
Resources Study and the Australian Reseaeh 
Council 


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belonis fish from Philippine waters, fin 2 Mearasirad, 23, 

TAWTSS, 

, Oiueneete. A, ad Miike, He Clee) 
Viypanerhyachic fishes of conunereil inshore fishes al 
the West Afric cost. lyn Li Res, 7, 133-16. 

PAW ON, JOR. Hoban. DL. ALLEN. GR Masi ds 
(WYO) Pisces. Petromyvontidi fe Curangidae, 
“Zyologiedl Catilowue ot Abstali’. Vol 7 i Xustaliian 
Government Publishing Service. Cunherta), 

Prva. T) (1994) Broclisticke zur Renpinis der 
Kisselhandwariner, Zool dalicb. Mmen 58. 1-20, 


162 I. BEVERIDGE , R. A. CAMPBELL & M. K. JONES 


Ropsins, C. R., BatLey, R. M., Bonn, C. E., BRooker, J. R., 
LacuNer, E. A., Lea, R. N. & ScorTr, W. B. (1991) 
“Common and scientific names of fishes from the United 
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(1913) On some Indian Cestoda. Rec. Ind. Mus. 
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(1924) Notes on some tetrarhynchid parasites 
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(1929) A monograph on cestodes of the order 
Trypanorhyncha from Ceylon and India. Part [. Spol. 
Zeyland. 15, 169-312. 


(1930) Cestoda. “The Fauna of British India 
including Ceylon and Burma” Vol. 1 (Taylor & Francis, 
London). 


Ward, H. L. (1954) Parasites of marine fishes of the Miami 
region. Bull. Mar. Sci. Gulf Caribb. 4, 244-261. 


YAMAGUTI, S. (1952) Studies on the helminth fauna of 
Japan. Part 49, Cestodes of Fishes. Acta Med. Okayama 
8, 1-78. 


(1959) “Systema Helminthum, The Cestodes of 
Vertebrates, Vol. 2” (Wiley Interscience, New York). 


DORYLAIMUS BAYLYI SP. NOV. (DORYLAIMIDAE, 
DORYLAIMIDA) A NEMATODE COLLECTED FROM 
SEDIMENT IN A FRESHWATER ROCK-HOLE IN THE 
NORTHERN TERRITORY 


By WARWICK L. NICHOLAS* & M. HODDAT 


Summary 


Nicholas, W. L. & Hodda, M. (2000) Dorylaimus baylyi sp nov. (Dorylaimidae, 
Dorylaimida) a nematode collected from sediment in a freshwater rock-hole in the 
Northern Territory. Trans. R. Soc. S. Aust. 124(2), 163-168, 30 November, 2000. 

A new species of Dorylaimus is described from the sediment of a freshwater rock- 
hole in the Northern Territory, Australia. It is distinguished from other species by a 
combination of characters: the cuticle has about 30 longitudinal ridges in the mid 
region of the body, the odontostyle varies from 43 to 46 wm in length with an aperture 
covering 43 to 46% of its length and is about ten times as long as it is in diameter, the 
male tail is short and rounded and the female tail is conoid, terminating in a short 
flagellum, the spicules are 55-6lum long, and there are 22-25 supplements in a 
contiguous row. 

Key Words: Dorylaimus, freshwater, nematode, rock-hole, taxonomy. 


Hranwations ofthe Reval Sector of & Aust (2000), 12412) 163-168 


DORYLAIMUS BAYLYI SP, NOV, (DORYLATMIDAE, DORYLAIMIDA) A NEMATODE 
COLLECTED FROM SEDIMENT IN A FRESHWATER ROCK-HOLE IN THE 
NORTHERN TERRITORY 


hy Warwick L. NICHOLAS & M., Hoppa! 


Summary 


Nienonas, WoL, & Hoppa, M, (2000) Dervafue bavi sp, nov. (Boryliimidae. Dorylaimidiy a nematode 
collected Tronn sediment yea Peshwater rock-hole iy the Northern Territory. Pre. A Sane. 4S. Aus 12402), 103 


LOR. 30 Noyvenber 2000, 


Anew species af Dorviamias is deseribed from the sediment of a freshwater roek-hole mt the Northern 
fenilory. Australia. Mois distiizuished front other species by g combination of chanieters: the cuticle has about 
40 fongitudinal eidves inthe mid region of the body, (he odontostyle varies fron 43 (od pom in tength ayith ain 
dpernire Covering 43 ta 46%) of its length andl is about ten times ais long as (os dn dineten the mate tail bs shore 
um rounded and the female tinh is conor, lerininating ia short Thigellun the spicules are 55-61 p6m long. anil 


there ure 22) 25 supplements ia contiguous row, 


Biy Wokhs Depylarns, Trestiwater neniatode, rock-hale, hisonomy. 


Introdaction 


Nemidodes of tlie genus Doeryleiniiy Dujardin 
[S45 are amon the most commonly occurring 


freshwater nematodes and are obvious because of 


their large size. The genus has been little studied on 
Australia, Late list century Cobb described D. la tus 
Cobb (89) trom grass and PD. spiralis Cobb 1s¥3 
from carrots near Sydney and 2 mimes Cobb (893. 
D. subyimitiy Cobb 1893, D. pusiliny Cobb 1893 and 
D. pevfectas Cobb (893 (rom sugar cane in northern 
NSW. At the time. the concept of the genus wits 
much broader than it is now, Derylainus being the 
only genus i what is now regarded us the 
superlamily Dorylaimoidea. Cobb's descriptions are 
insufficient to phice these Goa even to genus, All 
were described from females, only the first and last 
named were ilustrated dnd po type specimens were 
designated. 

The USDA Nematode Collection contains three 
species labelled = Darvas — inenalyyteva, 
Dorylatnas aiser and Derylainies perfects 
collected from soil ander wheat at Nhill, Victoria and 
donated by Thorne in September 1963. Doryleimus 
monohystera was Jater transferred lo the genus 
Hennes and Do aniver to kadarvlatnus (Thorne 
197+) and the specimens of 1. perfects appear to be 
more correctly pluced in the genus Mesederyainus. 

Bishop (1974) observed that nematodes of the 


Diao Botkin ail Adeloigy, Adataliie National biniwersity, 
Caner ACT W200 
CSO) Lintormetoey CieO) Hay T700 Canberl ACT 2007 - 


genus Dervilaimus were common in teniporary 
freshwater pools near Sydhey bat published tig 
descriptions and kept nu voucher specimens, Hoddit 
eho (in press) collected aquatic nematodes 
extensively throughout southeastern Australia and 
confirmed that in that environment members ol the 
menus dre often present. 

This paper presents u description of a new 
Porylahnus collected by |. Bayly from a rock-hole 
(gamma) at Warambi Hill in The Northern Territory 
in 199], Subsequent sampling of the type locality by 
Dr Bayly yielded further dorylaims bul no additional 
specimens of the new species, 

Type and Voucher specimens are deposited in the 
National Nematode Collection (ANIC) at the CSIRO 
Division of Entomology, Canberra ACT, 


Materials and Methods 


Speemmens collected wah a O.1S min mesh tet 
were fixed in 70% alcohol, Por processing they were 
washed in water and tramisferred to S aqueaus 
glycerol, The water was slowly evaporated and the 
specimens were (ranslerred to anhydrous glycerol in 
which they were mounted for microscopy with cover 
slips supported by ghiss beads of the appropriate 
size. Measurements were made from camera eid 
drawings. All measurements were along the curved 
median line 

One specimen wae Washed in water and post-fixed 
In aqueous OsO., washed again in water und freeze 
dred. The specimen was mounted on a metal stub, 
couled will) vold/palludium and esamined and 
photographed in the scanning electron microscope. 


104 W.1.. NICHOLAS & M. HODDA 


Dorylaimus baylyi sp. ney. 
(FIGS |-14) 


Holotype: &, ANIC 81-340, |. Bayly. Warumbi 
Hill. near Papunya, NT, 5.v.98. 

Paratypes: 6 dd. ANIC 81-341 to 346.6 2 8, 
ANIC 81-347 to 351, L Bayly, Warumbi Hill, near 
Papunya. NT. 5.v.98. 


Measurements: Table 1. 


Description of Holotype male (Pigs |-5) 

Body large. slender. cylindrical. Tapered cervical 
region, six rounded slightly offset lips. Tail short, 
rounded. Cuticle very linely annulated in ceryical 
region (below resolution of the light microscope, 
but visible with SEM). with 28-32 longitudinal 
ridges al mid body. Amphidial foyea stirrup-shuped, 
aperture a longitudinal slit’ just behind lips, 
Odontostyle straight, strongly built, 10 x diameter 
or 2.3 x width of lips. aperture 45% of length, guide 


Figs IS. Dory lads bervlyi sp, nov. male. 1, Entire holoype. 2, Head with odontostyle filly protruded. 3, Pharyngeal 
region. 4 Junction of two testes with vas deferens, 5. Posterior of bady and copulittory organs, 


A NEW SPECIES OF DORYLAIMUS 


ring double. Odontophore slightly curved, shghtly 
longer than odontostyle. Pharynx cylindrical. 
muscular throughout is lJength. narrow at 
odonlophore junction, expanded half way along its 
length, nerve ring 25% of pharynx Jength from head 
end, with dorsal pharynyeal gland adjacent to 
expansion, Cardia triangular with length greater 
thin diameter, enclosed by anterior intestine. Gland 
cells between pharynx and intestine present, 
Intestine slightly sinuous to level of anterior testis. 
compressed by gonads, a relatively wide straight 
tube from level oF posterior testis to prerectum. 
Prerectum short, straight, tubular, set off from 
imtestine by sphincter muscle, terminating in 
narrow, cuticle-lned rectum. Diorchic. testes nat 
reflexed, mature spermatozoa filiform, i clusters, 
vas delerens a straght tube, on lett side of intestine 
near anterior end and ventral to intestine a 
posterior end, eyarculitory duct nol distinct from vas 
deferens. Prominent oblique copulatory muscles 
from anterior ta posterior of prerectum, Spicules 


TAhh dL. Measerenieits(pa) ef Dorylainius baylyl spate, 


169 


identical, dorylaimoid, ventrally arcuate. with 
capilulum and lateral euiding pieces (cruria) 
Supplements, adunal pair, then gap. then row of 22 
contiguous supplements, 


Purutype miles 

Sumilar to holotype. but numbers in row of 
supplements differ from 22-25, Number ol 
longitudinal cuticle ridges very difficult to count but 
probably in range 28-32. 


Paratype females (Pigs 6-14) 

Females resemble males in most characters, upirt 
from reproductive system and tail, whieh is conoid 
ending in short flagellum. Females didelphie ind 
amphidelphic with reflexed ovaries. Scanning 
electron micrographs of one additional female (Figs 
11-14) show that ridges present in the mid body region 
cease on tail and cervical region. Very line 
annulations. below resolution of hight microscope, 
evident in cervical region and vulya is small oval pore. 


‘Type Male Holotype Male/Para n=6 Female/Para n=6 

Mean Range Sb Mean Range sD 
Length 3748 3872 3345-4313 382 40-4 3425-4352 
Max. width 70 7S 62-82 74 76 66-83 
Width ait lips 19 is 16-21 (9 14 18-19 
Odontostyle length 44 45 43-46 [2 45 44-46 z 
Odlontostyle aperture 19 19 17-22 2.0) 1% 17-22 A 
Odantophore length S7 6] 55-05 37 60) 39-66 Wa 
Head to aimphid opening 5 6.1 3.9- 11) 23 7 3.9-9,5 24 
fled ta guide ring 26 26 23-28 [.8 24 25-35 4.9 
Hedd to nerve ring 190 | 175-204 {ho 197 IS-210 7 
Head to pharyngeal expansion 423 305 343-450) 4h 306 342-423 28 
Head fo end of pharyis 846 stu TOB-BOO 30 N77 840-887 7 
Width at cardi 70 55 1-71 dab 7 O81 63 
Head totip of anterior gonid) LESS 1302 LL28-1531 1S9 VALS | {28-1572 104 
Head to vulvar - - - - STI | 012-2257 403 
Hevel fo Vis deferens isl4 {72 125 S28 7 : 
Head Wo Lip of posterior gonad 2-470 2320 [S28 -20R2 309 2384 I7s1 2518 7 
Prereetum feneu ace 190 (30-250 44. 24) IU SS 46 
Reet length TY 72 GU NI M2 64 SUNG Mi 
Head to anus ATS AB29 Aa +4347 a77 3807 3753-442 3400 
Tull length 4d) 43 38-58 SY 337 190-295 4o 
Width at anus 45 45 4+)-49 3.2 36 W--f2 >| 
Spicule length 57 Sty 55-1 24 
Number of supplements inraw 22 a4 22-28 Jb 
Arius Lo supplement row 74 Ha TULL T \b 
Length ob supplement tow 71 OF FLSA 33 
De Man's il 54 55 16-06 ho | 1-0! / 
De Man's ob i 5.0) 44-5, | 3 it 3.-5.1 4 
De Nian's a O4 | TIO 79 VF 15-24) y 
DeMun’s u (14 It O.8-14 0.2 ih HAO TA) 
De Man's VG& - At 4 iad S4 


166 W_L. NICHOLAS & M. HODDA 


Differential diagnosis 

Darylaimus baylyt sp. noy, differs from all other 
species of the genus in spicule length, number of 
longitudinal ridges in cuticle. length of tail in adult 
females and lack of papillae near vulva. Dorylaimus 
baviyi sp, nov. differs from the closest species (D. 
siddiqii Anmad & Jairajpurt 1982) in haying a longer 
odontostyle (44-46 pm ef. 35-36 um in D. siddigit), 


a shorter til in adults of both sexes (De Man's c = 
15-2) ef. 14 in D. siddigii tor adult females and 77- 
99 cf, 53-64 in D. siddigii in adult males) and having 
fewer ventromedian supplements (22-25 cl, 31-34). 
Dorylaimus baylyi sp. noy, is also similar to D. 
deaconi Botha & Heyns (1991). Both have very fine 
annulations in the cervical region anterior to the full 
development of the longitudinal ridges but D. baylyi 


Figs 6-10, Derylaimus buylyi sp.nov. feniale paratype. 6. Head und cervical region. 7.Cardia. §. Entire female. 9. Tail. 


10, Vulval region. 


_ 


. 
— 


Ll 
_ 


— 


i 
| 
i 
i 
| 
i 
i 
i 


r 


ee 


ae 
oe 


Figs ()-I4. Scanning electron micrographs of female Doryladney baylyi sp. noy. 11. Head and cervical region, 12. Mid 
region of body showing cuticular ridges. 13.Vulya. 14+. Tail, Seale bars = 50 pin, 


168 WL. NICHOLAS & M. HODDA 


sp. nov. has fewer longitudinal ridges (28-32 ef, 33 in 
D. deaconi). a longer odantophore (56-66 Lim cf. 43- 
53 um ), much shorter spicules (55-61 pm cf. 71-86 
um), fewer supplements (22-25 cf. 35-42) and the 
vulva pore-like rather than a Jongitudimal slit. 
Dorylaimus baylyi sp. voy, differs from the very 
widespread D. stagnalis in the rao of length to 
diameter of the odontostyle (10 cf. 6.7-7.3 in D. 
stagnalis), the odontostyle being shorter (43-46 Lim 
cf, 47-51 um), the odontostyle aperture being 
relatively longer (0.37-0.43 of the total lengih ef. 
0.33), having fewer supplements (22-25 cf, 30-40), 
having much shorter spicules (S5-61 pm el, LOO-110 
win) and in having filiform spermatozoa (ovoid in 2. 
viagnalis) (Abebe & Coomans 1992; Mulvey & 
Anderson 1979). 


Type locality and habitat 
Freshwater rock-hole (gamma). 


Distribution 
Known only from Warumbr Hill, 3 km trom 


Papunya in The Northern Territory (23°15' S, 
131°54' E), Collected by lL. Bayly 5,v.98. 


Etymology 
In gratitude to Dr lan Bayly for the specimens, we 
named the new species after him, 


Remarks 


Baermann extraction of mud samples trom a later 
collection at the same rock-pool produced 
Mesodorylaimus retundolabiatus Basson & Heyns 
(1974) and Heterocephalohys sp. (Cephalobidae). 
These specimens are also deposited in the ANIC 
Nematode collection. M. rotundolabiatus as ANIC 
81-352. 


Acknowledgments 
We thank the Electron Microscope Unit of The 


Australian National University for the use of the 
Scanning Electron Microscope. 


References 


AbeBe, BE, & Coomans, A, (1992) Aquatic nenratodes from 


Ethiopia IX. One new and three known species of 


Dorylaimidae. Monouchius trancatus Bastian, 1865, and 
Diploscaprer coronanis (Cobb, 1893) Cobb, 1913 
(Nematoda). Hydrebjvlogia 353, 121-| 48. 


Agprassy, |, (1988) The superfamily Dorylatmoides 
(Nematoda) - a review of the Family Dorylaimidac, 
Opuse. Zool, Budapest 23, 3-03, 


Brsnov, J. A. (1974) The fauna of temporary rain pools in 
Eusiern New South Wales, Hydrobiologia 44, 319-323, 


Bora, A. & bleyns, J. (1991) Dorylaimoidea (Nematoda) 
from rivers in Kruger National Park. Koedoe 34, 1-24. 


Corg, N, A, (1891) Onyx and Dipeltiss New nematode 
genera, With a note on Dorvlainuts, Proc. Linn. Soc, NSW 
61,143-158. 

(1893) Nematode worms found attacking 
sugarcane, Agric. Gaz, NSW 4, 808-833. 

Muivry, ROH. & Anbrerson, R. V. (1979) Benthic species 
of Derylatmus Dujardin 845) (Nematodi: 
Dorylaimidae) and = Areridorvidimus on, gen. 
(Arctidorylaimidae no Jam.) from the Mackenzie and 
Porcupine river systems. North West Territories. Canada. 
Can. J. Zool, 37, TAS-755. 

Hoppa, M., Stewart-Rreep, E., DALLwitz, M_ J, Parse, T. 
& ZuRCHER, E. Z. (in press) “Freshwater nematodes of 
southeastern Australia” (CSIRO Publishing. Melbourne), 


DEVELOPMENTAL BIOLOGY AND LARVAL MORPHOLOGY 
OF THE FROG LIMNODYNASTES DEPRESSUS TYLER 
(MYOBATRACHIDAE: LIMNODYNASTINAE) 


By MICHAEL J. TYLER* & MARGARET DAVIES* 


Summary 


Tyler, M. J. & Davies, M. (2000) Developmental biology and larval morphology of 
the frog Limnodynastes depressus Tyler (Myobatrachidae: Limnodynastinae). Trans. 
R. Soc. S. Aust. 124(2), 169-175, 30 November, 2000. 

The larval morphology, early developmental biology and chondrocranium of 
Limnodynastes depressus are described and compared with the morphologically 
similar L. tasmaniensis and L. fletcher1. Advanced tadpoles reach a total length of 80 
mm or more and the mouth disc has three upper (one divided) and three lower (one 
divided) rows of labial teeth. Larvae resemble those of Limnodynastes fletcheri but 
are generally larger. Spawn is laid as a foamy mass in water-filled depressions in the 
ground. The chondocranium differs from that of L. tasmaniensis and L. fletcheri in 
the attachment of the processus ascendens of the arcus subocularis with the prootic. 
Key Words: Limnodynastes depressus, developmental biology, tadpoles, spawn, 
chondocranium, frog, Myobatrachidae. 


Transcctions of Me Baya Soerety af §. Mast (2000), 124(2), 169-175, 


DEVELOPMEN 


PAL BIOLOGY AND LARVAL MORPHOLOGY OF THE FROG 


LIMNODYNASTES DEPRESSUS TYLER (MYOBATRACHIDAE: 
LIMNODYNASTINAE) 


by Michiaun J. Tyke’ & MARGARET DAVIES” 


Summary 


‘Tytbe Mo & Davies. NE (2000) Developmental biology und larval morphology of the frog Linniwelysiiytes 
depressus Tyler (Myobatrachidie: Linmodynastinae), Trans R.See. S$. Ader, 124(2), 109-175, 30 Noveniber, 2000, 

Nhe larval morphology. curly developmental biowmey and chondrocraniun of Liincdviestes depresses ute 
described (ind Compared with the morphologically similiur Lo fasmeniensis and L. fletchert, Advanced Giulpoles 
cach ay total length ef SO mm or more and the mouth dise has three upper (one divided) and (hree lower (one 
divideu) rows of labial teeth. Larvae resemble those of Limavdyistes fletolerr butare generally larger, Spuwh 
is Hath us at founiy mass in water-filled depressions in the ground. The chondrocraniuny differs Pron thar olf. 
hisdiiensiy ond Le. fletcher? in the attachment Of the processis dseendens af the dreus subocularis with the 


poole. 


Kiy Worps: Linmedviases depress, developmentil biology, fulpoles. spawn. chomdrocriniun, [Hos 


Meobatchichie. 


Introduction 


Limnodynastey comprises three recognisubie 
lincuges (Tyler er af 1979: Mahony & Robinson 
1986: Roberts & Maxson 1986), one of which is a 
eroup OF saecalled miwsh frogs: that includes 7. 
depressuy Tyler. 1976, This species hats previously 
been known only fron the holotype but recent field 
and laboritory observations (Davies & Burton 2000; 
Morris & Tyler unpub., Watson, ‘Tyler & Morris 
Wipub,) will contuibute substantially to knowledge of 
(his frog, 

Spawn clumps and early developmental stages 
have been observed in the field and capiuve 
gpecimens have bred in the laboratory. Here we 
desenibe (he developmental biology of the species 
and include morphological dati oon the 
vhondrocranivin of the Jarval stages. 


Materials and Methods 


Hhitial Observations Of exe deposition sites und 
breedipy bebaviout were made at Cockatou Lagoon 
in the Keep River National Park, Northern Territory 
(15° 59" 8, 129° 03" FE) from 10-12 February 1908. 
barly developmental sizes were collected and 
preserved in Tyler's (1962) preservative. A captive 
pair Of Limmodyvnastes depressus collected at Kocp 
River spawned in the University of Adelaide 
Zoology Department Aquarium Room overnight 


Depa ciental Pavinoiicital aratogy, University of Adelide SA 
SOS 


VWAinAOOS and larvae Were reared te 
metamorphosis. The room was maintained at 30°C 
+)". Larvae were reared in shallow gliass aquaria (25 
x 25.4 8 cm) in verated, dechlorinated tap water and 
were provided with u diet of boiled, organically 
grown, lettuce leaves supplemented hy SERA 
voldfish Tlakes. Representative larval shiges were 
preserved in Tyler's (iid Stages follow Gosner 
(1960), Chondrocranial preparations were made alter 
the method of Dingerkus & Uhler (1977), Drawings 
were made with the aid of a camera lucida attached 
toa Wild M9 dissecting microscope, Measurements 
were made with dial calipers reading Co 0.05 min and 
with an eyepiece micrometer, Chonidlroeranial 
deseriptions follow Hits & Richards (1999), 
Temperitiire measurements in the field were taken 
wilh a Digttron D2060) digital lemperatare probe 
within the spawn vlump toward the base and in the 
vuleror mad beside the elunp. 


Results 
Field observanions 


Haurial 

Cockatoo Lagoon is an elongate billabong whose 
southern dimit ts within 3 km of the Victoria 
Highway connecting Kununurra. WA with 
Katherine in the NT. The billabong is approximately 
150 a wide for its entire length and extends for 
approxmmitely 1 Km ina northeasterly direction, The 
most detailed map readily availible is Keep, sheet 
4766 (12100000) of the National Topographic Map 
Series, 


(7th MAI TYLER & MADAVIES 


ay 
: 
ee 
‘ Js 
— 
| 
mA 
+ a 
Hip 1} Spawn clump at Ainedvaisres depressus ina 


depressing dudpleent to Cockioa Lagoon, Keep River 
NPONT 


Acthe ume af the visit tr 1998, there had been 
Hiile run ane the billabong wats reduced ta a series 
of jsoluted pools. The Observations reported here 
were winlertiaken mm the area adjacent to the Keep 
River National Park Office unc) Titerpretation 
Contre and the extreme seathern portion of 
Cockatoo | (oor extending for 400 m south of the 
Centre, 


SPAWN 

Seventeen clumps of spawn were hicuted al two 
areas Within the study site, on the periphery of a 
Shallow pool in depressions created by Brahini 
eallle (Pig. 1) and amongst dense patches of the 
perenmal grass Pyendoraphiy spinescens CR. Bry 
Vic. Details of representative clumps and site 
temperatures ave provided in Tables | and 2. Tyler 
(1994) reports spawn of Lituria rubella (Gray) i 
Water temperature of 42° Cand tadpoles of various 
species in water temperatures ranging Hom 35-45" C 
in other areas in northern Australia. 

Newly hatched lurvae were al stage 24/24, OF 25 
larvae examined, Hihimentous gills were present only 
on the Jett hand side in-all but woe which bad gills 
suill present on both sides. Adhesive glands were 
lighuly pigmented, The mouth wats perforated and (he 
beak wis keratinised. AL two divs post hatehie. 
larvac Were al stage 24 and stuge 25. AL stage 25, 
labial tooth rows remuamed unkeratinised, 


Lahordiory observations 
A spawn chimp was. laid overnight between 371i 
LOOX and Ai MY98. The cee mass wus a Tounry 


Pig. 204, Literal and B. dorsal views of Lamnodvniaves depresstys tadpoles Ste 26. Seule bar = 5 nin. 


LIMNODYNASTES DEPRESSUS DEVELOPMENT 171 


TABLE |. Dimensions (mm) of Spawn clumps of Limnodynastes depressus and deposition cavities at Cockatoo Lagoon, 
Keep River National Park. 


Reference Spawn clump Deposition cavity Cavity water depth 
A 50. x 45 160 x 100 45 
B 75 x 50 180 x 120 35 
Cc 60 x 60 105 x 80 20 
D 90 x 80 100 


TABLE 2. Temperatures (°C) in spawn chumps ef Limnodynastes depressus and the surrounding water or mud at Cockatow 
Lauvoon, Keep River National Park. 


Ident, Date Time Temp. in clump Temp. of adjacent water Temp, of adjacent mud 
| 10,11, 1998 1200 37,7 34.9 

| 114i. 1998 LOS0 31.1 30.5 
| 11th. 1998 1355 39.6 33.2 
| 11.41.1998 1600 36.3 35.9 
| hii (998 1928 28.8 315 
2 10.11, 1998 1210 39.5 35.0 

2 10.41.1998 1905 30.5 31.6 

3 10.11.1998 1300 39.8 37.9 

3 10,11. 1998 1900 29.4 30.5 

3 11.11, 1998 1100 27.7 28.2 

4 10.11. 1998 1215 375 36.3 

+ 10.11.1998 1930 26.0 30,4 

4 11.11, 1998 Ws 28.9 28.8 

5 10.41.1998 1220) 36.1 35.6 

5 10.11. 1998 1940) 28.6 30).7 

5 11.1, 1998 1130 31.8 30.2 
5 11.11, 1998 1400 37.0 34.3 
5 Li, 1998 lots 36.6 35,3 
5 Hi, 1998 1940 29.6 31,7 
6 1041 L998 1955 28.7 32.1 
6 111.1998 1140 32.6 31.1 
6 11.1, 1998 1350 AL4 34.3 
6 114i. 1998 1555 39.3 37.0 
6 11. 1998 1925 27.7 307 
7 10.11.1998 2005 29.7 31.6 
7 Tan 1998 1045 31.7 30.3 30.4 
7 11.11.1998 1355 38.2 37.7 35,5 
7 11 it.) 998 1600 35.4 36.2 36.5 
7 111.1998 1930 27,1 30.9 
8 2010 28.8 313 

8 (725 28.2 28.4 
8 1100 31,1 31.4 30.3 
8 1400 37,4 38.9 34.4 
8 1605 36,5 36.4 
8 11.1. 1998 1935 27.1 31.4 
9 10.41.1998 2015 31.6 
9 11.11.1998 0730 29.3 
9 11.41, 1998 1100 30.3 
9 11.11,1998 1610 33.8 


9 11.11, 1998 1935 30.4 


172 M1. TYLER & M. DAVIES 


clump which did not collapse until well after 
hatehing. An egg count could not be obtained as 
some of the lurvae had hatched quickly and not all 
could be rescued from the holding tank, The 
first larva reached stage 42 on 20.vi. 1998 and 
metumorphosis was campleted at stigze 46° by 
22.vi. 1998, LEO days afier spawning, 

Pigmentation of larvae was sparse io early stiee 
30, but by stige 32, larvae were well pizinented, 

A larva at stage 26 is shown in Fig, 2. The body is 
ovoid and widest behind the eyes. The snout is 
evenly rounded in dorsal view and slightly truncated 
laterally, The nares are more dorsal thin hiteral, not 
elevated and Opening unterolaterally. The spiracle is 
sinistral, relatively short and visible from above. I ix 
attached to the body wall along its medial edge with 
the diameter of its orifice being slightly less than the 
diameter of the tube. Tapering of the spiracle is 
mioonal, The anal tube is median and opens along 
the ventral edge of the ventral tail fin. The tail fins 
are well arched. the dorsal fin commencing in the 
posterior ia ol the body, The dorsal fin is deepest 
uboul '/) along its length and the ventral fin about °/\ 
long its length. At its terminus, the tail is pointed 
but not sharply acuminate. The tail museulature is 
moderately thick and tapers to a point posteriorly. 
Blotchy chocolate pigmentition on a cream 
bockground is Jocwed on the tail musculature with 
hittle melanism on the fins. The body is covered with 
bloiwhes of pigment over a poorly pigmented 
backeround, Neuromast cells of the literal lines are 
well differentiated (Fig. 3). 


The mouth is anterior. The oral dise is surrounded 
laterally and posteriorly with a double row of 
papillae that are pigmented at their bases and in the 
centre, Pupillae are absent anteromedially. There are 
three upper and three lower rows of labial teeth, The 
first upper and second lower rows ure divided (Fig, 
4). 

Measurements of larvae are provided in Table 3, 
Body length at metamorphosis ranged trom 25.0- 
41.9 mm in four individuals. 


Chendracranium 

The chondroeranium of a stage 36 larva is 
ilastrated in Fig. 5. The eartiligo labialis superior is 
composed Of a pars alaris (lateral wing) and wu pars 
corporis (frontal bar), The = pars alaris is 
synchondrotically connected proximally with the 
pars corporis. The articulation of the upper jaw with 
the cornua trabeculae is mainly by the pars. alaris. 
Viewed ventrally, the partes of the cartilago labialis 
superior form a simple arch. The cornua trabeculae 
ure the anterior extensions of the trabeculae cranii of 
the cranial Moor. These moderiely-broad diverging 
bars have a literal process near their proximal base 
to which the ligamentum quiadntoethmoidale is 
ullached, 

The cartilazo meckeli is sigmoid and transversely 
oriented, The medial part of ils anterior Face 
articulates with the cartilage labialis interior, The 
cartilago labiales inferiores form a U-shaped arch in 
ventral view. They are connected medially by a non- 
vhondritied symphysis. 


Hin 4. Dorsal view of Stage 34 tadpole of Limnedyoestes depresyys showin neuromust cells, Scale har = 3 nim. 


HIMNODYNASTES PEPRESSUS DEVELOPMENT Vi 


bxnie a. Measurements of cdpoles ef Lininoeyniistes 
Jepresats capressedd i nine ces rea) cet Peds 


Staae Body length Total length No, 
My 100 31.7 3 
46-114 35.7-45, 

7 1205 STAN Et 
VOO-T4T 397-444 

D4 12495 3H.14 4 
1-17) 321-51) 

nal) ais VS 3 
1] 2-li4 HNO-FK 

at J24 VS 2 
Wha 313K 8 

4) hls 3K. | ] 

me 12.5 FUR ye) 
1W129-13.4 35.9-44.7 

aD) (4.75 wis 4 
We TS.0 34 6-507 

a5 1k. 574 2 
168-210 S006. 

Ab 16.05 51,12 6 
12. 20,7 Ay (rely dob 

47 17,39 55,13 t 
P2718 AR HOS 

4s 19.35 O44 q 
(8.4 LYS 620) 64.6 

) 2h) b4o | 

4) 22S7 74h 3 
TI ZA, | 72570,2 

+1 21.0 SIU) | 

43 15.) 


rhe een rithm 
eer 


its lara 


aoe we SOMO ay 


Pie b Oral dise of Stage 26 ladpale of Jamimodynistes 
depressay, Seale bar = 1 mm 


view oof the ehomdreeruniin et 


Dorsul 
Linmedvvnresres depresyus (Stige 36), Seale bar = Sam, 


Fiz. 3, 


The rool of the britiease is wcomplete and 
formed by the fectui synouewnm and the tien 
ethruoidale, The cattilive orbitalis is high. The basis 
erumtias perforated hy the paired foramina carole 
and oeculomotora 

The processus museularis quadrath rises from the 
latera) margin of the palutoquadrate and lerminmates 
with a bluntly founded apex, The processus 
ethmoidalis is shorter than wide und (he processus 
pscudopteryguideus ts clongate und prominent. The 
areus subocularis is attached antenorly fo the 
neurocranivM by the commissur, quadratocranialis 
anterior, The larval processus basalis between the 
palitoquadrate und the otie capsule is prominent The 
processus uscendens oF the arcus subocularis is not 
ovellain by the otic capsule, The maximunr width oF 
the neurocranivo and pialaloquadrate is at (he level 
Of the processus asecuderns, 


Comparison with other species 


Linmaidvnesies depressus is morpholowealky ost 
similar to 2. davmenientsiy Gunther and £. fletchert 
Boulenger (Davies & Burton 2000), Hence here we 
conling comparisons oF the karval and chondroeranial 
morphology of 1. depresviy lo those lwo species. 
Larvae of L. depresses are very similar those of /. 


fletcher? (Davies 1992) withough advanced tadpoles 


oF 1. depressay reach w total length of 80 mim or more 
whilst those Of L. fleteheri are recorded as reaching 
69 fim total length at stage 40 (Davies 1992). ‘Phe 


\7| MJ, TYLER & M, DAVIES 


Vin, O, Dorsal views of the chondrocrimiy of A, Loneedyristes tasmmotenses Stage 32, Bok Merehore sme 44, Seale burs 
i Ee £ 


= wun 


oral dise. extemal morphology and pigmentation ace 
identical, Larvae of Lh. daymeiensis usually are very 
heavily pigmented (virtually black), although some 
larvae in western New South Wales approach the 
pigmentation of 4, flereheri and Lb, depressus (M. 
Anstis pers. eomm, 2000). The oral dise of /. 
HISPeasix has more tooth rews than do those of 
depressuy and Lo fletcher? (five upper and three 
lower) (Martin 1965; Martin & Littlejohn (982), 
The mayor difference in the ehondrocrania lies in 
the relationship of the processus ascendens of the 
arcus subocularis with the otic capsule (Pig. 6). In 
both 1. flereheri and L. rasmanieasis, the otic capsule 
overlies the processus ascendens. In addition. the 


Jurval processus basulis is very poorly developed in 
both these species. 


Acknowledgements 


MJT thanks the Parks and Wildlife Commission of 
the Northern Territory for providing scientific 
collecting permits and permission to work in the 
Keep River National Park. A. Phillips and D. Bryce 
are thanked for field assistance und to Morris. ts 
thanked for advice on the existence ol 
Limnodynestes depressus at Keep River, MD thanks 
5. Walker for assistance in the final preparation of 
the fizures. 


References 


DavIPsS, Me. (1992) Developmental biology at 
Linmodynastes tecraereginge wad Lb, fletchert (Anucu: 
Leploditetylidaes Myobatrachiniw), Trans. Ro Soe. 8. 
Atest, 1L6. LE7-122, 

& Burros, PC, (2000) Redetiniien of the 
Australian frog Jlinedynastes depressus ‘Vyler 
(Mychatrachidue; Linmodyniastinued fhid, 124. 141- 
150. 

DInGRRKES. G. & UHDUR. LD. (1977) Kneyme clearing of 
Alelan Blue stained whole small vertebrates for 
demonstration of cartilage. Stain Teclnal, 52, 220.94), 

Gosnek, K.L. (1960) A simplified tuble for staging anuran 
embryos and larvae with mores on identification, 
Herpetalowica Va. 13-190 


HaAds, A, & Rienaros, SL (1999) Correhations of erential 
morphology, ecology, and evolution in’ Australian 
suctorial tadpoles of the genera Liraria and Nvetiiniystes 
(Amphibia: Anura; Hylidae: Pelodryndinael /. Ader 
23h, 109-141 

Manony, M, J. & Rowinson. CoS. (1986) Nueleolur 
orginiser region (NOR) locarion In karyotypes of 
Australian ground frogs (Panily Myobatrachidae ). 
Crenelion 8, (19-127 

Morin A, A. (1965) Tidpoles of the Melbourne area. View, 
Net, 82, (39-149, 

oo & Lerroous, Me od. (hQ82)) “Lasmanian 
Amphibians’, Fuuna of Tasmania Hundhook No 6 
University of Tasmania. Plobart, 


LIMNODYNASTES DEPRESSUS DEVELOPMENT 175 


Roperts, J. D. & Maxson, L. R. (1986) Phylogenetic (1994) ‘Australian Frogs a natural history’ 
relationships in the genus Limnodynastes (Anura: (Reed New Holland, Sydney). 
Myobatrachidae): a molecular perspective. Aust. J. Zool. 
34, 561-573. 


Ty_er, M. J. (1962) On the preservation of anuran tadpoles. 
Aust. J. Sci. 25, 222. , Martin, A. A. & Davies, M. (1979) Biolog 
(1976) A new genus and two new species of and systematics of a new limnodynastine genus 
leptodactylid frogs from Western Australia. Rec. West. (Anura:Leptodactylidae) from north-western Australia. 
Aust. Mus. 4, 45-52. Aust. J. Zool, 27, 135-150. 


FIRST RECORD OF THE SOUTHERN RIGHT WHALE DOLPHIN, 
LISSODELPHIS PERONII (LACEPEDE, 1804) (OQDONOCETI : 
DELPHINIDAE), FROM WATERS OFF SOUTH AUSTRALIA 


BRIEF COMMUNICATION 


Summary 


The southern right whale dolphin Lissodelphis peronii is a small pelagic dolphin that 
is rarely observed close to land. What little is known about its biology has been 
gathered from stranded specimens'. The species occurs only in Southern Hemisphere 
waters where it appears to be largely restricted to the region bounded by the Antarctic 
Front in the south and the Subtropical Front in the north’. It is unusual amongst 
dolphins occurring in the Australasian region in that it lacks a dorsal fin’. Lissodelphis 
peronii appears actively to avoid ships and this and its unobtrusive behaviour when 
not alarmed may result in the species being under-recorded. 


Tinesactions vf the Koval Society of S. Aust. (2000), 1242), 177 178. 


BRIEF COMMUNICATION 


FIRST RECORD OF THE SOUTHERN RIGHT WHALE DOLPHIN, LISSODELPHIS PERONIL 
(LACEPEDE, 1804) (ODONOCETI; DELPHINIDAE), FROM WATERS OFF SOUTH AUSTRALIA 


The southern mht whale dolphin Lissedelphrs, pect is 
aamall pelagic dolphin that israrely observed close (o lane, 
What litte 15 known about its biology has been gathered 
trom Stranded speermuens'!. The species occurs only tn 
Southern Hemisphere waters where itappears to be largely 
restricted ta the region bounded by the Antarctic Front in 
the south and the Subtropical Front in the north) Tt ip 
unusual amongst dolphins occurring in the Australisian 
region im that it lacks @ dorsal Fin’. Livselelphin pereneet 
appears actively wy avoic ships ane this and ils unoblrusive 
behaviour when nol alarmed may resull iptie species being 
under recorded, 

Whilst conducting surveys of scubirds from the bridge of 
the CSIRO research vessel “Pranklin® during a yoyawe that 
included five days cruising Ih Australian territorial waters 
off South Australia, groups of whales and dolphins were 
sighted on several pegasons, On J] Auszust 1998. when the 
ship was 92 uautecal miles south of Cape Gantheaume, 
Kangaroo Island, South Austealia (97° 32" 10" 8, 137" 27! 
40" Ej and proceeding in a noyth-easterly direction a berd 
of small dolphins was observed surfacing 200 me Loi the 
ship. On the hasis of un absenee of any dorsal fin und the 
diriking combimuion of white ventral surfaces and a largely 
black dorsum the dolphins were idennfied qs 1. prevent. 

At the lime af the sighting (11.05 L107 a.m. Aust. 
CST) viewing conditions were good. Willa southerly wind 
of fess than five knots, the sea surface was vlassy smooth, 
wilh only ashght swell (~2 mm). Air terriperattire was 140°C, 
burometric pressure was 1020.3 hp and steady and the 
conditions were cloudy but bright. Water temperature at (be 
sea surface was [2,87 °C und sulinily 34.98 ppm. The 
dolphins were observed 44 nautical miles south oi the 
continent shelf in deep (4904 mi) pelugic waters. just 


south of the Subtroprestl Front. Slightly warmer waiter 
(13,5-1425 °C) was encountered later in the same day only 
several nautical miles to the north of this sighting. At lirst 
my {itlention was drawn fo a atea of small splashes ol the 
surface of an otherwise culm sea. By the use of 10% S50 
binoeulurs small dark backed dolphins were idendtied as 
the cause of this disturbance, The dolphins were travelling 
slowly (2-5 knots! uway from the vessel, When rising te 
blow they broke the surfare gently, exposing only the very 
lop of their dorsal surfices but (he euiplete ubsence of i 
dorsal fin, Wiis immediately ohvions. The dolphins were 
travelling as a cumpaet proup all Treading Gn the sane 
direction und it wis diflicul to assess their number This 
behaviour continued for approsifiately: 30 sec Nefore the 
dolphins abruptly changed directian placmy them on 
course heading across the bow of the stip, At ihe same time 
they besun porpoising clear af (he surface After travelling 
less than 30 m (he herd abropuy changed direetion again 
retuming to theiy initia! course heading away fron the 
yessel. The dolphins were now more easily counted and 1 
estimated the herd comprised 20 individuals. No young 
Were seen They continued lo move away Irom the ship on 
it fauly diree| course and wher fast seen, approximately tc 
rin dater, were sull travelling at u sustained speed ani 
porpoising clearor the waiter 

As cach animal leapt clear of the water, zoo views OF it 
were obtyined. The following composite description wis 
nvide from figld notes taken atthe Gine of the sighting. The 
dolphins were small and sicnder and were abour wwe metres 
in length. When porporsing their bodies appesred is 
proportionately elongated, although this feature was 
presumubly enhanced by the ubsence of dersul fins. Bach 
individual had a striking but simple pred pattern tha 


Taue l. Sightings and strandings of \.issodelphis peronii in the Australian region. 


Date Locabou Position Comments Source 
(1 Jan. 1802 OF southern Tasmania c. 44'S, lypé Specimen Peron (1807) in J 
MIP E 
7 dan, (424 ONSE Austealia eases havpooried u 
Pre [S84 Tusrania is specimen 7 
Aug 1968 co 100 miles SW of Ausrratia 4 sighling of six tncispduals 4 
Ot, 197s Cloudy Bay, Bruny Island, Tas. 43° 25'S. Stranding (Tas. Mus, | 
47° 1S E specimen # AL3014 
Aug. 1970 mid Great Aust, Bight (WA or SAd a siehting of 50 individuals 4 
14 Sept LUXS ON South-west Cape, Tus 41°41 8. siting af 25 1 W. Rades 
145° 40 B individuals pers, comm, [909 
Id Feb. 1986 South of WA 46° 03-8, sighting af ¢. S00 4 
126° 32 E individuals 
Sept. VOR6 Bendalong, NSW ay" 57S, apparent strand S 
1a) B2E (see LOX) 
29 Sept. 1995 Chinamuns Bay. Mana Island, Te. 42° 40° S, stranding R.M, Warneke 
48° 02 E pers. comin. }9YY 
1! Aug, 1998 92 miles olf Cape Gantheaume 37" 33 S. sighting af 20 Thesentudy 


Kangaroo Island SA 


13" 27 


individuals 


(78 


uppeared consistent ueross the herd. White venual surfaces 
extended dorsally to the rostrum and face. The upper 
surface of (he Mlippers, and the remaining dorsal surfaces 
were black. The border where black and white met was 
sharply defined. and curved from the middle of melon 
down (he sides of the head (o a portnt at, or just ahove, the 
flipper; it then curved upwards slightly before continuing 
along the flanks and tail stock as a Fairly straight line. 

To the best of my Knowledge this sighting 1s the first 
documented occurrence of L. perani in waters off South 
Australia and the 11! for waters around Australia (Table |), 
Previous records have included 4 strandings and/or 
specimens anid 6 sightings, with most being from waters 
south of Tasmanian, There are no records from Vietora 
amd just one or two from waters off Western Australia, 
However, one of these sighlings may have ovcurred in 
waters off South Australia us the record was simply noted 
as “in the middle of the Great Australian Bight’. The most 
northerly record of L. peranit in Australian waters is of a 
ateaided animal at Bendulong in NSW (35° SY, However, 
this record should be treated as uneonfirmed as enquiries 
by the author failed to locate any specimen, photographs, 
nites or first hand kiowledge of this individual. 

The complete absence of strandings along the southern 
eoasr pt maand Australia and the scarcity of records 
elsewhere in Australian waters: may be attibutable to the 
apecices’ preference for pelagic waters. The small size of L. 


Raker, A. N. (1981) The southern right whale dolphin 
Lissodelphis peroni (Lacépede) in Australasian waters. 
Nat. Mus. NZ Ree. 2, 17-34 


Gaskin, D. E. (1968) Distribution of (he Delphinidae 
(Cetuwea) in relation to sea surface temperatures off eastern 
and southern New Zealand. NZ J. Mar. Res. 2, 527-534. 


Newcomer, M, W., Jelferson, T. A. & Brownell, R. L. 
(1996) Lissadelphis peronii. Mamin, Species 531, 1-5. 


* Morzer Bruyns. W. F, J. (1971) “Field Guide of Whales 
and Dolphins (CA, Mees, Amsterdam), 


peroni’ further reduces the Jikelihuod of dead animals 
being wushed ushore as scavengers ure likely to consume 
such small carcasses before they are able to drift to the 
coast from pelagic waters. Although the species has been 
known to ride on the bow wive of vessels on dccasions, 
this behaviour is apparently uncommon’ An apparent 
avoidance of vessels, as noted in the obseryation 
documented here, hus been reported by other observers’. 
This avoidance behaviour, combined with the species 
smull size, and invonspicuous nature when nor alarmed. 
may result in individuals or small herds being overlooked, 
Indeed the mdividuals observed off South Australia ave 
unlikely to have been sighted had sea surface conditions 
heen more typical and white caps of any size been present. 
Combined with the knowledge that competent field 
observers have, until quite recently, had few opportanities 
to systematically or routinely visit deep pelagic waters of 
the cool temperate one of the Australian region the status 
of L. peronié in the region remains unclear. 

Thanks are due to N, Cheshire, Captain of the “RV 
Franklin’ and CSIRO for the opportunity to take part in the 
vovupe. R, M. Wameke kindly assisted with references and 
information on unpublished records while D. Eades readily 
provided details of his L, perenii sighting. bam grateful to 
M. Clarke and J. Ewen for helpful comments on a dratt of 
this note and to C, Kemper and Ro M. Warmeke who 
reviewed (he manuscript. 


* Liewellyn, L.,. Ellis, M.. Martin, J. & Ferguson, A, 
(1994) “Atlas of New South Wales Wildlife: Marine 
Maminals and Reptiles’ (NSW National Parks and 
Wildlife Service, Hurstville), 

" Fraser, F.C. (1955) The southern right whale dolphin, 
Lissodelphis peronit (Lacépede): External characters aud 
distribution. Bull. Br. Mus, Zool, 2. 341-346. 

‘Guiler, E.R. (1978) Whale strandings in Tasmania since 
1945 with nofes on some seal reports, Pap. Proc. R. Soc 
Tas, 112, (89-213, 

" MePhail, D. M. (1987) Cetatcea: Southern Ocean. Mar, 
Obs, 57. 15. 


ROHAN H. CLARKE, Department of Zoology, La Trobe University Bundoora Vic. 3083, E-miuil 


R.Clarke @7 oo, latrobe.edu.au 


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