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VOL. 121, PARTS 1 &2 
30 MAY, 1997 

Transactions of the 

Royal Society of South 



Smales, L. R. A revision of the Echinonematinae (Nematoda:Seuratidae) from 

bandicoots (Marsupialia: Peramelidae) ------ 1 

Zeidler, W. A new species of freshwater amphipod, Austrochiltonia dalhousiensis 

sp. nov., (Crustacea: Amphipoda: Hyalellidae) from Dalhousie 
Springs, South Australia --------- 29 

Bird, A. F. & McClure, S. G. Composition of the stylets of the tardigrade, Macrobiotus 

cf. pseudohufelandi ----- ----43 

Bird, A. F. & McClure, S. G. Studies of the eggs of Macrobiotus cf. pseudohufelandi 

(Tardigrada) from wheat fields in South Australia - - - - 51 

Kolesik, P. Two new species of Asphondylia (Diptera: Cecidomyiidae) from 

Halosarcia spp. (Chenopodiaceae) in South Australia 59 

Jago, J. B., Tian-Rui, L., Davidson, G., Stevens, B. P. J. & Bentley, C. A Late Early 

Cambrian trilobite faunule from the Gnalta Group, Mt Wright, NSW 67 

Brief Communications: 

Jago, J. B. & Haines, P. W. Poorly preserved trilobites and brachiopods from the 

Kanmantoo Group, Fleurieu Peninsula - 75 

Kemper, C, Dutton, J., Foster, B. & McGuire, R. Sightings and strandings of the 
pygmy right whale Caperea marginata near Port Lincoln, South 
Australia and a review of other Australasian sightings - - - 79 






VOL. 121, PART 1 


CONTENTS, VOL. 121, 1997 

PARTS I & 2, 30 MAY, 1997 

Smales, L. R. A revision of the Echinonematinae (Nematoda: Seuratidae) from 

bandicoots (Marsupialia: Peramelidae) ------ 1 

Zeidler, W. A new species of freshwater amphipod, Austrochiltonia dalhousiensis 

sp. nov., (Crustacea: Amphipoda: Hyalellida) from Dalhousie 
Springs, South Australia ------ 29 

Bird, A. F. & McClure, S. G. Composition of the stylets of the tardigrade, Macrobiotus 

cf. pseudohufekindi ----- _ _ _ _ 43 

Bird, A. F. & McClure, S. G. Studies of the eggs of Macrobiotus cf. pseitdohufelandi 

(Tardigrada) from wheat fields in South Australia - - - 51 

Kolesik, P. Two new species df Asphondylia (Diptera: Cecidomyiidae) from 

Hatosarcia spp. (Chenopodiaceae) in South Australia 59 

Jago, J. B., Tian-Rui, L., Davidson, G., Stevens, B. P. J. & Bentley, C. A Late Early 

Cambrian trilobite faunule from the Gnalta Group, Mt Wright, NSW 67 

Brief Communications: 

Jago, J. B. & Haines, P. W. Poorly preserved trilobites and brachiopods from the 

Kanmantoo Group, Fleurieu Peninsula - - - - 75 

Kemper, C., Dutton, J., Foster, B. & McGuire, R. Sightings and strandings of the 
pygmy right whale Caperea marginata near Port Lincoln, South 
Australia and a review of other Australasian sightings 79 

PARTS 3 & 4, 28 NOVEMBER, 1997 

Bourman, R. P., Martinaitis, P., Prescott, J. R. & Belperio, A. P. The age of the 

Pooraka formation and its implications, with some preliminary 
results from luminescence dating - - - - 83 

Vaucher, C. & Beveridge, I. New species of Potorolepts Spasskii (Cestoda : 
Hymenolepididae) parasitic in dasyurid marsupials from New 
Guinea ----- - 95 

Littlejohn, M. J. & Wright, J. R. Structure of the acoustic signals of Crinia glauerti 
(Anura : Myobatrachidae) from south-western Australia, and 
comparison with those of C. stgnifera from South Australia - - 103 

Watson, G. F. & Gerhard t, H. C. The breeding biology and advertisement call of 

Lttoria splendida Tyler. Davies & Martin - - - - - 119 

Barnett, E. J., Harvey, N., Belperio, A. P. & Bourman, R. P. Sea-level indicators from 
a Holocene. tide-dominated coastal succession, Port Pine, South 
Australia - - - - - - - - 125 

Gullan, P. J., Cranston, P. S, & Cook, L. G« The response of gall-inducing scale insects 
(Hcmiptera : Eriococcidae: Aplomorpha Rubsaamcn) to the fire 
history of mallee eucalypts in Danggali Conservation Park, South 
Australia -- - - ------ 137 

Olsen, A. M. An intensive monitoring study of two wetlands of the River Murray 

in South Australia; physico-chemical parameters and cyanobacteria 
concentrations. - 147 

Kolesik, P., Whittemore, R. & Stace, H. M. Asphondylia anthocercidis, a new species 
of Cecidomyiidac (Diptcra) inducing fruit galls on Anllwcercis 

it tto re a (Solanaceae) in Western Australia - - - - [57 

Brief Communications: 

Wallman, J. F. First record of the Oriental Latrine fly, Chrysomya megacephala 

(Fabricius) (Diptera : Calliphoridae), from South Australia - - 163 

Smales, I... R. The status of Cyclostrongylus mediaannulatus Johnston & Mawson, 

1940 - - ------ 165 

hisirl to Trcnisuclujin of tin Ro\a! Smit-fr r-f South Auxtniim Vol, J2I, parrs 3 & 4,28 November. I9V7 




By Lesley R. Smales* 


Smales, L. R. (1997) A revision of the Echinonematinae (Nematoda; Seuratidae) from 
bandicoots (Marsupialia:Peramelidae). Trans. R. Soc. S. Aust. 121(1), 1-27, 30 May, 

The name Echinonema being preoccupied the genus here designated Linstowinema 
(nom. nov.) is redescribed. The type species L. cinctum comb. nov. is synonymous 
with E. meridionalis (sic) Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 1980 
but not with E. cinctum sensu Inglis, 1967, sensu Chabaud, Seureau, Beveridge, Bain 
& Durette-Desset, 1980. L. warringtoni sp. nov. is established for E. cinctum sensu 
Yorke & Maplestone, 1926 and sensu Chabaud, Seureau, Beveridge, Bain & Durette- 
Desset, 1980. Linstowinema inglisi comb. nov. is synonymous with E. cinctum sensu 
Inglis, 1967 and E. inglisi sensu Chabaud, Seureau, Beveridge, Bain & Durette- 
Desset, 1980 and is redescribed and four new species L. latens sp. nov., L, 
tasmaniense sp. nov., L. maplestonei sp. nov. and L. peramelis sp. nov. are described. 
Key Words: Linstowinema, Nematoda, Echinonematinae, Isoodon, Perameles, 
bandicoots, Marsupialia. 

TrmMtViimt tff tfir ftflVtV* W». m ■.,/ \ l/xvl [Mtttti 121(1). I 27 


hy Lrsi i a R. Smaij K* 


S\iaiis |. K H'J*»7> A revision oi the Miinonenwiiinac iNctiuuo<_lu Seunitidaei trom bandicoots 
iMviiMlpialiuilVrainctiditc) Trews. U- Soc tL'Amh 121(1), I £?. JO Mty l l »7. 

t Ik niuiK- /-i hittonttint being preoccupied the genus here designated Uthumfiwnitt mom. iiov.i is redescrihed, 
The type species/., ( an imu comb, nov. is synonymous with £ rffWwftftftllfr iw. j Cltaband. Seme;ui- llcveiidgC- 
bttio & Durelte Desset, 1080 but not with A, aw;ow wnsn Iny.lis. |9fi7, WW ClwKmd. SewetHi. Ucveridgc, 
Hum A f Hiielte -DesseL 1 9HQ» /-. WMTitijilOtti sp. huy is c-MjjhliNhc*i (in £. t }nt htm sc/isu Yorke i; \laplcsioue. 
I02h and w/fw< (. liiibaud. Scuteau. BflvcrtdgB. Hum <k Dureiie f) # * >n* >. / if^ttiwinatia in«hsi comb- nov, 
is synonymous vviih /.. tfutium wfrftti Inglis. 100? alul A. l>i&j«J m pui t rtuhjud. Sciireau. BeccndgC- Bain & 
Oiirtlk'-fJessel. 10X0 tnul js retlescrthed and lour new speeics /._ (uh-tts Np. nnv., L tas/tuhiictiM sp Hoc, £, 
nmplVsUmoi sp nov. and /.. f.u^mtm'lis .sp. nov, arc (tescfibeti All .sewn species can Ik differentiated as follows; 
/. nmiiiHL U- IK iows til hodj hooks willi undulating cdA^v. oesophagus icnniinuing at ihe kwel ot the Klh- I llh 
io\\ /_ Uiitrtn^lntti, 0-1 3 rows of body honks Willi undulating edges, oesophagus H-innnolinL! al rhe level of [lie 
u ih I 3Ut row: /., itwfisf, 10-U rows of body hooks with undulaihtg edges, ocsopha£U% terminating ai die level 
ol ihe Hlli-^lli row: L kifetn, 0- II oiw> ot bod> hooks with undulating ed£es, oesophagus terminating 
posteriorly lo the I llh row. /_. U^mwiit'tw. H 15 pbws hook- without nndukiline edges, oesophagus 
lefiuiimting ui the level of the Hih-IOlh row; L inapU'sicftci ]2r1i| lows of body hooks without undulating edgev 
oesophagus terminating posteriorly to Mlh low ol hooks;,/,, pcuwulis- 1113 m\\s ».i hodv Imoks wilhoni 
undulating edges, oesophagus lenniiialmj; u! lesvi (*l'the 12th row A key to the species i^ pivcii. t.msiontnetihi 
t nulum oectli's Itl f^Hith'/rtfhtsuhjs, f tnth'H>nm\, l J craiHvlt\ nusufa and P. yjimtii I., iuviisi and /.. tti.uttttllii ] H:,< 
only in / •t}'rs>il'<\. /_., /dK'f/vonlv in / w>'rti>nn<\, f MtTTftlgit/tti in /, nun n>it<us, I tihrsnfn\ t I tmnifiiy and /! 
'insula, L. imwk>t<»u i to /. mu> • wants and /' mwttl and i., pt'KtWtrlk oily '» /• hsitytifiHtirik 

Khi Wmhhs 1 ZiniMif7'('»ic. N-dPTWlOfla, f I iiion^utaiinae. IwtidftH Pl'ftfMth f, bandicoots Maisupialia. 


When l-Htsluw H896dt) dcscrilx-d a llioruv headed 
iKMoalodc ueeumnt' in a bandicoot host, he called U 
Ho{)((Kcphahis iinclus. He subsequenil) found that 
Hophh-rj)httlu^ was preoccupied so he renamed rhe 
L*ettus lnhintmetmt kiler in lite same year (Liusiow 
IXMXh) h-fiirifwt'iiitt bus continued to be used lor Ihe 
genus nniil the present. Echinom'inu also is 
preoccupied, however, having been used previously 
for it yc-iuis of spoiiex-- hy Curler in IJffitj I HOVf 
propose the name Lnistowincnm tumt. nov 

The handieooi host tin^mally was identified as 
^entweli's obcsnlu\, Linslow I IS°Sa) described and 
figured a nematode with seventeen rows of hooks on 
the euiicukir dilation o[ the oesophagal region ol the 
body and oesophagus terminating on a level with rhe 
0th row ot hooks. Yorke & Muplestone tl*/2^) 
described a nematode with \%-\3 rows ol hocly 
hooks and a relatively short oesophagus which they 
idem 1 1 led as "A. ciiuutnt", They neither yuve 
measurements nor provided a figure lo show die 
relationship between the posterior end o( the 
oesophagus and (he rows of body hooks of their 

U'IMii.hri.! mi lliuln.n,' ■•■tm-'il On, . n-.l,iml I pi * v i i-ny 

Huckiwtuimn™ OW VfOl 

speemiens. Yorke & Maplesttnie t l*J2f>) uSMgned lire 
type and. ar that lime (inly species. /.. cinciuin 
tLtustow. i 8VM ). to the spimrid family Rictulariidae 
Later Johnslon & Mavvson (193V)) reported L 
vim-ium from ;t naiisc cat, l)(i\\mtt\ \'ivcniittt,\. near 
Sydney and re -evaluated the available mlormation on 
the host species They concluded thai 1 instows 
original description was hased on malenal eolleeled 
from Isoadon nbewitlus (Shaw. 17071 from the Upper 
Bmneii River in Queensland while Yorke & 
Maplcstuik\ redeseTiption was baseil on material 
from o bantlicoot. possibly /. nutcwurus (Could. 
\SA2). eolleeled in the vicinity of Townsvillc. 
Johnston j& Ytawson (l'-->30) further conclutled thai 
bitili Linslow (189Sal ami Yorke St Maplestone 
i \ l )2b) were desenhing material from the same host. 
namely, f. twicmutvs Since /. obestthts docs not 
occur in Ihe Burnett River region (Braiihwnile 1055) 
bur /. maeroitrus does, (Gordon iQScfy, ihis would 
seem to be a reasonable conclusion. 

The first confirmed record tif t in\t<n\iitvtii<i ftdttl 
the bandicoot genus Penuucles Ocofiroy. 1X03 is by 
Johnston &. Mavvson 1 1040) From P tuisutti Geoliroy. 
1 804 cidleeted near Sydney. These authors noted 
ddtereuces in the- male tail Of the specimens ihev 
e\atnme'U, Irom earlier descriptions by Linsiovs 
(ISOSu) and Yorke & Mapleslone (l l '26j but, 
nevertheless, assigned the specimens lo L (itiL-intn, 

2 l.k sviAii-s 

Ol Hit* differences noted by Johnston & Mawson 
(19401 1 lie number ol papillae on the male luil is 
pmtJlcrtMrtic j^ neither they. Linstow iisysa.i atii 
Yorkc & Maplexrone ( Wlh) described the number 
und placement Of papillae found i»u other specimen-. 
til linsunvtrufiut The expansion of the male body 
nrtmrul (lie cloaca! region desenbed ami figured by 
Johnston & Mawson il9-Hh is also n significant 
miucUmv. Although not mentioned by Linstow 
(lK^Sa) Yorkc & Maplestone | |02<jj G\ Chabaud e/ 
iii. (I98Q) in then descriptions of L wuciwn. it is 
comparable Ija the male tail desenbed by Chabaud (ft 
at, |1#8QJ Fob EtfifftWtfftti sp, fs/cMxcurnng ui R 
mtstnu horn an unknown loealily. 

No further work was done on the genus unul inglts 
tl^hl) re-examined the relationships p| the 
Mtperhnnilv Seuraloidcu. He redescfilvd L. < jblftfelfl 
from materia] collected trom /• ahatittlta near Pcrlh 
and placed the sole genus within a new suhraimly 
the F.ehinonemalinae. located wuhin ihe seuratoid 
i.n.iily Sehneiderneinalidae on Ihe basU of the form 
ol the mouth opening, distribution ol cephalic 
papillae, long spicules and short guhernaculrun 
Subsequently, the affinities wf ihe genus were 
clarified by Quenlin I 1970) and the EehiuonematHi.u 
included within the $etnutid;ic 

Chabaud e7 ui I NKOf re-examined all the available 
malerial. established two new genera Smuti htnrt 
Chabaud. Seureau. Beveridge. Bain ."C: Duicih: 
Dew^l, KJ.H0 i.nd /ne/< < /"'"' Chabaud, Seute.ui 
Beveridge. Bain & Dureltc-Dc^sel, I l >*0 tor wornis 
from dasyund marsupials and redefined fofrmvnctftti 
ItiWW IJiwewitiwna) and /:. tttu'titm (x/ct LinstuW. 
I S')S ntw Ingliv b)b,7_ In iheir description of L. 
i ut> iiint they noted difficulties in interpreting 
I mxlovv's original figure ol ISOSa, hut decided that 
an oesophagus t.8 mm long agreed with their 
dclmition ol' a "long" oesophagus, thai is, onc 
terminaling al the level ol the most posterior body 
hooks. Linstow"- (IS')S;o figure cloud j diows the 
oesophagus lernnuahug at the ievel ol the l )|fi ol" 17 
mws ol' body hooks. Chabaud ci tit. tTroUJ also 
described three new species, namely. /, tifiurmJM 
\\/i | man a dasyund. I- fttVSfdfttwDi tith i from / 
'.'/'<_ \uhl1 t.i\n\ F ifLjjtiXi tv/( i - E ii'fii'HMi x«'//.o< 
luglis, !*)(■» 7 also from /, f>hcutins, These .iiithors .*ko 
indicated that there were possibly additional species 
from the bandicoot genus fynumlcs but they had 
insulficienl material for detailed descriptions. 

The bandicoots (subfamily Perumehnuc i me 
rabbit -si^ed omnivorous marsuptals with long 
pointed heads and compact bodies. They forage by 
digging conical holes with their short forelhnbs and 
explore these holes with I heir pointed snouts 
(Gordon & Ihllberl 1^X9). fyatttwUs tutsttiu, the 
lone ii'i.sed bandici"*ol. is found along the east coast 
of Australia, from ramfoiest in the north ihmuy.u 

wetland antl dry wooUland to areas wilt) liltlc ground 
co\er in the south iSiuddao \ty5)i lis disiribniion 
overlaps Willi /. wacraitms. die northern brown 
bandicoot, winch i> found on the east coast, north of 
I he Hawkesbui) River and across the \orthein 
Territory to the north of Western Australia, in flrwi 
of low ground cover including grassland, woodland 
and open forest (Gordon 1 995 > To the souih, the 
distribution ol ft tia.sufu overlaps tlvat ol' /. ohoultt*. 
Ihe southern brown handico.ol. which is found across 
southern Australia 111 Western Australia. South 
Australia and Victoria, southern coastal New: South 
Wales and Tasmania and preteis -siudv muK with 
scrubby vegetation or low ground cover that are 
burnt out from time to rime (Biaithuuuc 1993). Flic 
eastern barred bandicoot. P. t>nwiir Gray. tK"^. now 
■ r a. i. .led to vvIk-ic lis disi.iboboii 
overlaps with that of /. fbe.siihts, and a tew rehcl 
colonies in southern western Victoria, jnefers open 
grassland, bur may also forage jn %cfub anil 
heathland (Seelvck !9M.S). The western barred 
bandicool l J btntwtinvilk- Qit"y & Gaiuvard, IS24. 
now eMsting only on Bemicr and Dorrc Islands 
Shark Bay. Western Australia was previously found 
actoss much of die southern half of Australia (Krie.nd 
& Btirhidgc 1WS), The only other bandic«»o| -.pecies 
still extant. /. auratus. the golden bandicoot, \v^\ 
survives only pn Barnoi and Middle Islands o\T Ihe 
coast of Western Australia, the north western 
Kimberley region i\\m\ sub- humid parts of die 
Northern Territory, having been previously recorded 
from u much wider range of habitats (McKen/ic . i 
al 19951 

In this study, all the available iveoi-its .mo mitten. d 
collected from peramelids. including matei (al 
dissected from hosts held in museum eollechons as 
Wfell a> live caught bandicoots, have been ex.amuicd. 
'Ibis I"ms provideil sufficienl material to rcas.sess. the 
lavonoiTiic characters available to use lot ,-r>ecies 
discnmmaiion, redesignate and redescribe (tic Ivpc 
^vcic- [ - 'tttrtitm (lanslow. 1-8^1 comb. nov. = /.. 
/>/< •ntJi(*tnrh\\ i\i(.) Chabaud, Seureau, Bc\cndgc. 
Bain tS: Durelte-Dessel, I9SU redesenbe and name 
/ WitrrWgWHl sp. nov. = /•_ (lucrum wttfll Yprlps A 
Maplestone l l )2<>; mjyM Chabaud. Seureau. 
Reveridgc, U.nn & Dttretie-Desset. I°xtl and t 
(Hytisi Chabaud, Seureau. Beveridge. Bain <V: 
|)uretle-I)csscl. l l »S2 and distinguish Ihe lout 
additional new species from bandicoois lhat art! 
described below. 

Materials and Methods 

M.itciMl and dissection o-.cind> tVoin _'H 
haudicoois were ti^iilitnid, Tho- malerial w;i- 
derived horn three sources, first ly. the eas|i«- 
intestinal tracts o( 5o bandicoois, collected belwefi 


• /. /tun nmnt.s 
Mitchell Plateau * 

i aims 

0\ A^Ioujn&iiiilt* 




Fig. I Present ;tnd lorntcr distributions til "Australian bandicoots (after Gordon &. I lulhert 1989). The symbols indicate the 
localities where bandicoots were collected between 1989 unit 1996. ImhhIihi ttunttn.\ t A Huiinmim. / oh{:\i<ht\ M 
l\tvmrlf\ hi)u^aimHU\ f\ mount, A. P. tutsn!«,A. 

1905 and 1988 and deposited in ciihcr the South 
Australian Museum (SAMA) or The Museum of 
Victoria (VM), were examined. These animals hud 
probably been fixed in 5-10% formalin before being 
stored in 70% ethanol. The nematodes dissected 
from these hosts were stored in 707/ ethanol 
Secondly. 79 animals were either collected as fresh 
road kills or trapped alive, m spring-loaded wire box 
traps bailed with peanut butter ur a peanut butter, 
honey and oats mixture, between 1989 and 1996. 
The trapped animals were killed by intraperitoneal 
inoculation of euthanasia solution pentobarbitone 
sodium (Nembutal ®). The digestive tract of each 
animal was examined under a dissecting microscope 
ami any nematodes found were washed in normal 
saline, fixed in glacial acetic acid in hut OfXoW 109^ 
formalin then stored In 70'V ethanol. All the 

available specimens o^ Linstmi'irwnut held in the 
Queensland Museum (QM). the Australian 
Helminlhologicul collection of the SAM A (AIIC). 
the Western Australian Museum (WAM). the CSIRO 
Division of Wildlife and Ecology (CSIRO) and The 
Natural History Museum* London (BM(NH)) were 
also examined. The preservation history of material 
from the former institutions is largely unknown but 
probably it was fixed in ethanol ur formalin. Material 
from the CSIRO collection was fixed in hoi [Q% 
formalin. All material is now stored in 7()9i ethanol. 
Specimens were examined from all (he extant 
bandicoot species (number of handicouts in 
parentheses* from 8J localities across Australia: 
Isoodon atimttis (5), /. innvrourus (8 1 1. /. ohcsithts 
(85), F J emmi'les hinti>a'nnilff (13). P, ftlitimi (o). P. 
ttu.Wita (14). baodicout, no species given (9). Host 

4 L K. SMAJ.hS 

TABU '. f)Mirfhttlii)H ttf hittulinu/t sfuuvs f )Ytti)fHi2tfji>l klllsloswru'liU \j>ttn\ h\ State Oi TtUWOty. When- Ml/ \ft\ijn 
loiiilttv has hc<ut Jf/Wfl fit 'he tthmthtn ttCttTd tin- loatd'W ft ftvFMf ffV Au.stntfin, 

<\}>l>rc\tLtitt>n\ r\ Australia, \\A \\c\iini AuMmfui. ,\7, Northern Tvnifinv. AV\, AV/l/rf? Australia ithltultin; 
h,it!i>>:m«> htmul on,} FraifUin f\t«it,l t <J. Oui:>>!\l,imh NSW \'rw S.uilh Wtitex, M \'t<fon<i; 7, Tosnioitht. 






hi'i'Jcu unrt<rn\ 

i nunnmrus 

i tibixttltis 

Peromely hoiii'tiiuviih' 

P. ujmnii 

P. tniMtta 









tlistribmions and the locations of the 79 animals 
collected since IsMOure given in big. I. Details ol all 
the localities where speeiinens were collected are 
lisleil In the descriptions of species given below- 
Latitudes and longitudes are provided Car all 
localities that are listed in the Australian Gamecr 
I lie locution, by slate, of all hosts species examined, 
is given in Table I 

Specimens were examined after dealing in 
lactophenol ar becehwood creosote. Measurements 
wctr made with the aid of an ocular micrometer or 
tlrawing lube and map measurer. Measurements ;ne 
given in p.m. as a innge from 10 specimens followed 
by the mean in parentheses, unless otherwise slated. 
Ml (he 'tew materia! htf* been deposited in the AH( 

Comments on Uixoimiiiic characters 

Prior io I4K0 only tine species was recogni/.cd in 
the genus l.tttshnvutcttia i.e. l.vhittoncmu iitictttnt. 
Then Chabaud ci nl. i 1980.) described four species, 
three I ion i petamclid hosts and one from the 
daryvund. D<ts\um,s hufhaams Gould, 1842, The 
species occurring in the dasyurid was distinguished 
fioin ihc others by having the lust row of cephalic 
huoks longer than the second. The three species from 
bulKllOOOlH WCPC (JiffCfCfltiBttHi fl*>W £flch other on 
the basis o\ (he arrangement ol hOOkti and spinas mi 
the body, the relative .sizes and positions ol papillae 
on the eloncal region, the relationship beiween the 
ventral spines ami pre-ckmuil papillae, the exu-tu [if 
small culicnlai bosses surrounding the cloaca, tin* 
number of papillae on the tail of the male and the 
length of the oesophagus relative lo the hooks on the 
(Ulfltcd part of the anterior body associated 
with the region ol the oesophagus, 

Although Chabaud tl til N9H0) recorded all the 
sensory organs on the male tail lip as pairs of papillae 

Inghs ( l L >67) had noted pairs of papillae and a pan ol 
phasmids. Thus lnglis (l%7) reported .3 pairs ol 
papillae and a pair of phasmids on the lip ol the lad 
of Eihttanutnu tyictUtn i sit ) and Chahaud t j t ul 
ll l J80i repotted 4 pans of papdlae for the same 
species. Spicule morphology is uniform across die 
genus, differing only in dual length and proportion Of 
body length. 

Chabaud ct ul. (t£80) described the oesophagus as 
either "long", extending It) the level of the last row ol 
the hody hooks or •'short", terminating within the 
dilated eutieular region. Detailed examination ol 
speeimens for ihis study has shown that, allhough the 
termination of the oesophagus relative to the 
■an rounding hooks is consistent within each species, 
the actual length ol the oesophagus (Mil its 
relationship to the body hooks may be difficult lo 
determine. Specimens that are not completely 
straightened prior lo or during fixation ate 
problematic. It is difficult to tell the extent to which 
the oesophagus may have contracted into the neck 
legion and the cuticle sunounding the oesophagus 
may also be contracted. I urlhennorc. (he range ol 
lengths of the oesophagus within populations o\' a 
species can also be quite variable, depending ou the 
mas and maturity of the worms being measured. 

feoth spines ami hooks originate in the cuticle, but 
spines in this context, are delhicJ as being small lo 
liny and rootless, whereas hooks consist ol a Ihoiu 
and a root anchoring the thorn within the cuticle, The 
smallest honks m.i\ be only slightly largei than the 
largest spines, The leluthc lengths of the three rows 
of hooks on the cephalic bulb is a consistent 
character, but the dimensions of the hooks may vary 
markedly hetween individuals of the same species 

from the fr&toc individual hosi, Therefore Iwofc 

dimensions are not useful lot discriminating between 
species. The dimensions oflhe cephalic bulb arc also 


unreliable because d! the potential variation caused 
by llie extern o) relaxation of the specimens pnoi to 
fixation and lite method of fixaliou. Olher consistent 
characters al lite Sp6c'«8 level, however, ate the 
number ol rows ol hooks on Ihc diluted anterior 
and llie morphology of these books. In some species 
I he roots i)[ llie books bave undulating lateral edges 
giving them a Hilly" appearance: in others, the 
ctlyes ol Ihc hooks arc more or less plain. 

Key to the species of /JtiKtrnvinema 

I, Kt oiw ol cephalic FlOftkS longer lhan 2nd row: 

parasiles of dasyurids ,.. LtdfttQtubl 

2nd row id cephalic hooks longer than Is! row 
parasites ijf bandicoots , . (2t 

j, Lind> imnks without uroJuifliiaa ajgw - -tf) 

Body liooks wilh undulating edge^ ((>) 

s Oesophagus iciminaies posterior lo hooks on 
tmiuilar (JilftliOtl of oesophageal region: male 
wilh 4 pans Hi papdlae, I pairol phuMiiuk uil lull 

up - M» terminates at ui anierioi l" poslciior 
row of body hooks on cuiicular dilalion or' 
oesophageal icgion; nude with 3 pairs of 
papillae. I pair ol pbasnnds on tail lip j5l 

I Male wilh K-IO. lemalc Willi 10-12 body books.. 

nude wilh (y pairs eloaeal papillae all same si/e. 
boil) spines extend along 'Mt'i of dorsal surface 
lerminaie at level rtf most anlerior pan of lateral 
pre-cloacal papillae I HI \enlral suilace..A, ftfftm 
Male with 1 1 -l\ female with 12-13 body hooks: 
mule with n pairs <>( cJottflfll papillae. I pair 
lateral ud -eloaeal papillae larger lhan other 5 
pairs; body spines extend along 13% of dorsal 
surface lerminatc markedly anterior lo eloaeal 
papillae on wntml Mirface J. nfctfp/t'WWW 

5. Male with ala-like expansions o\' body 
surrounding cloaca: Willi 6 pans of cluneal 
papillae- 3 pairs of lateral eloaeal papillae larger 
lhan 3 pains of ventral eloucal papillae spicule 
length about 'A) of body length: female with lail 

longerlhan 7<w ' • '-• t<i*inutiit'ti\e 

Male wilhoul ulu like expansions of body, with 7 
pairs eloaeal papillae all same size, spicule 
length •iboui '/m of body length; female wilh tail 
shorter (ban 500 / ficmincfis 

ft, Male wilti |3-16. female wilh 14- IK rows <\i 
body books; male wilh ala-like expansions id 
body surrounding cloaca, body spines lerminaie 
at level of mosl anterior pall of lateral pre-cloaeal 
papillae on ventral surlacc A. virwitmt 

Male with 'M2. lema-e ft Ufa | I -Id n>ws of bodv 
hooks; male withoul ala-like expansions Ol bod) 
saiioundtng cloaca, body spines terminate 
markedly anierioi lo eloaeal papillae on ventral 

surface , - - -O) 

7 Male with oesophagus shorter lhan 1570. Icmalc 
wilh oesophagus shortet than 1850. male with 6 
pairs of eloaeal papillae, pair of lateral ad-eloacal 
papdlae larger (ban other 3 pairs, 3 pairs of 
papillae, I pair ol phasmids on tail lip. spicule 
length about ^15 of hody length: female with tail 

shortet lhan 940 /.. inxhsi 

M;ilc with oesophagus longer than 1570. female 
with oesophagus longer than I860: male with 6 
pairs ol eloaeal papillae all same si/e. I pans of 
papillae. 1 pan ^i phasmids o\) tail rip. spicule 
length aboui '/:n of body length; female wilh lail 
Imvjei than osu , / iuaryfygW/ 


Older Ascaiidida 

Supei laiuily Seuialoidca 

family Seuruiidae 

Subfamily l:chinouemaiinac 

GenUS Unitow'nwmn iiom. uov. 

7r//e \{>c<lt\: LWUffllMflKKlto Wt&lJ mstow, IXVM 

comb, uov. 

Synonyms; NftpIwepIurbJS i (tu tn\ I inslow, IS U -S. 

I iiummi nut iinrhiioiuihs ( habaud. SfeureUU. 

Ht.-\eriilge. Bain & DttreUe Dessel. I9S0 nte 

t : .chiiu>nt'fti(i Mtntitm tfemtl VOfkti &t Maplestoiii-. 

\92fK Inglis. l l )f->7; t'habaud. Seureau, Bain & 

nureiie Desset. l u K0(in part ). 

iMisUtwiiwma tnoin, no\.i 

Gctiitii' ifiitxfio,\i\ 

Anterior end wilh cephalic bulb hearing 3 rov.s o\ 
I4^l(. files of large httoks. Vhnith opening triangular 
in ouiline, wilhoul lips or hp-like structures, with 4 
pairs double cephalic papillae. I pairamphtds. mnci 
circle of sense organs on cdpe of mouth (see Inytis, 
l%7. Figs 6 7). Neck with -I I tows of very liny 
spines. S-1S rows oi' 14-16 hooks surround an 
anierioi culicular dilation, associated with 
oirsophageal regjftn. Body wilh numerous rows ol 
spines, number iA liU-s i^' spines increasing 
progressively towards mid body, decreasing lowards 
posterior, continuing lo caudal tip of female, 
icrnunatmg at about Vi ol" length dorsally. anlerior lo 
cloaca senirally on male. Short oesophagus simple, 
club shaped, surrounded b> nerve ring at level of 
cephalic bulb. Deirids simple, eomcal. at level pf 1st 
row oi cervical hooks. Spicules long, equal, 
identical; small gubernaculum present. Vulva ut mid- 



region of body; monoclelphic ovejector directed 
anteriorly. Parasites of Australian dasyurid and 
perarnelid marsupials. 

Linstowinema cincfti/n (Linstow. 189S) comb. nov. 
(FIGS 2 14) 

Syjionyuis: Hoplocephalns ductus Linstow, 1898a: 
pp. 469-471, Figs 3-11. /•.'< hinotienia meridionatis 
Chabaud, Seureau, Beveridge. Bain & Durelte-Desset, 
1080: pp. 436-438, Figs 4, 5A, D; Spratt, Beveridge & 
Walter. 1991: p. 26. Eclutumetna cincta Linstow, 
1898b: p. 672; Johnson & Mawson, 1940: pp. 473 
474. Fig 25: nee Yorke & Maplestone. 1926; nee 
Inglis, 1967: nee Chabaud, Scurcau, Beveridge, Bain 
& Durette-DeSSet, 1980. F.chinoneina cincta 
Maekerras, Maekerras & Sandars. 1953: p. 62. 
Lchinonemu sp. 2. Chabaud. Seureau, Beveridge. 
Bain & Durelte-Desset. 1980: p. 438. Fig. 5C R 
Sprat L Beveridgfi & Walter. 1991 : pp. 23. 24 (in part). 

Material e.uttmned 

From Isootlon obestdtis New South Wales: IV. 
fragment Lismore, <28 L 49'S. I53 a I6'E). April, 
1965, AHC 4413. 46 6, Timbilliea State Forest, 
(37° 19'S, I49 l! 43'E), 15.xii.197S, CSIRO N733; 
[■4cJ'd\ 192*9, Sidlings Swamp North, Timbilliea 
State Forest, <37" I7'S. 149° 45'E). I9.vii.l994. 
I7.iii.l994, 20.vii4994, 23.iii.1994, CSIRO N42I3, 
N4074, N4228, N4075; 85<J<3, 88V V, Sidlings 
Swamp South, Timbilliea State Forest, (37° J8'S, 
149 n 45'F), I4.vii.1994, 2().vii. 1994. CSIRO N4212, 

From Victoria; 2dd, 142 2, Melbourne, (37° 
47'S. \W 59'E). 9.ix.l991, AHC 30292; 29 V. 
Monash Llniversity, (37° 47'S. 136° 59'E), no date, 
AHC 30293, 30294; 26 6, 2VV. Gorge Forest 
Road, (38" 2l\S, 141" 36'B), Sept. 1962. AHC 
30296; 26 6, II 9 2, 7 fragments, no locality, no 
date, AHC 30295, 30298. 

From South Australia: 1 6, 8 V 2, 6 anterior ends. 
Waitpinga, (32° 36'S, I38 Q 32'E), no date, AHC 
4460; 36 6 79 V. Scott Creek, (35° 04'S, 138° 
42'E), 5.x. 1 992. AHC 3029 1 ; 36 6, 3 9 9, 
Myponga, (35° 23'S, 138° 28'E), May 1966, AHC 
4446; Kangaroo Island, South Australia: 1^.292, 
Cape Willoughby, (35° SI'S 138° 08'B) 13.x. 1990, 
AHC 30301; 29 2, Seal Bay. (36° OO'S, 137° 20'E), 
4x1987, AHC 30304; 26 6, 22 2, Binowie, {37° 

47'S, 136° 59'E), 5.viii. 1987, AHC 30303. 

From Perameles gtmnii Tasmania: 66 6 , 14 2 2,4 
fragments, Grove, (42" 59'S, 147° 07'E), AHC 
30025; 36 6, 49 2, Kingston, (42° 59'S, 147° 
I8'E), 1 l.vii.1992, AHC 30056. 30057, 30058. 

From Perameles nasuta: Queensland: 2 9 V , 
Wongabel State Forest, (17° 20'S, 145° 30'E) 
9.vii.l982, CSIRO N 1753; 1 9. Mt Nebo Road, (27" 
23'S, 152° 47'E) ILviii.1993. AHC 30316. New 
South Wales: \6„ Epping, (33° 46'S, 151° 05'E). 
Il.viii.1993. AHC 30316; \6 Epping, (33 G 46'S. 
151" WE), I4.vli.1933, QM GL 12048; 36 6, 
29 V 1 , Sydney, (33 (l 50'S, I5L J I5'E). no collection 
data, AHC 1820; IV, Nadgee Stale Forest. (37" 
26'S. 149°54'E), 13. ii. 1978. CSIRO N493. 


Cephalic bulb with 3 rows of 14 (male) or 16 
(female) files of hooks. 2nd row longest. 3rd row 
shortest (Fig. 3): neck with 5-9 rows of tiny spines; 
cuticular dilation Of oesophageal region bearing I 3- 
IS rows oi' 14 (male) (Fig. 2) or 16 (female) files of 
body hooks; 1st and last rows smallest, 4tlv7lh rows 
largest; roots o[' hooks with undulaling edges (Figs 
10, 13): remainder of body with Up to 36 (male) or 54 
(female) small spines at each annulation, over whole 
body of female; extending over -"L of dorsal surface, 
terminating about 400 anterior to cloaca, almost 
reaching level of anterior pair ol caudal papillae, on 
ventral surface of male body (Fig. 7). Oesophagus 
simple, club shaped, terminating about level with 8th 
- I 1th row of hooks, within the anterior cuticular 
dilation; '/* to '/m body length (Fig. 2). Nerve ring 
surrounding oesophagus within cephalic bulb; 
secretory-excretory pore in neck; deirids conical, at 
level of 1st row of body hooks. 

Male: Length 12-22 (14.8) mm, width 460-730 
(600). Cephalic bulb 260-490 (395) long by 325-420 
(380) wide; cephalic hooks 1st row 145-225 (170), 
2nd row 170-235 (200), 3rd row 104-145 (130) long. 
Oesophagus 1445-2040 (1790) long, cuticular 
dilation bearing 13-16 rows body hooks. Deirids 
520-630 (590), nerve ring 320-420 (375), secretory- 
excretory pore 500-530 (550) (n=3) from anterior 
end. Spicules equal, similar, without alae, 935-1 150 
(1035) long, about 7u body length. Gubernaculum 
short, simple, sublriangular 60-63 (n=5) long (Fig. 9). 
Nine pairs caudal papillae; 3 pairs ventral and 
immediately pre-, ad- and post-cloaeal respectively. 

Figs 2-14. Linstowinema cinctitm (Linslow. 1898). 2. Anterior end, optical section (lateral view). 3. Cephalic end (lateral 
view). 4. Cephalic end {en face view). 5. Cephalic end male, optical section at level of first row of hooks (en face view). 
6. Female tail tip (lateral view). 7. Male posterior body spines (ventral view). 8. Vagina (lateral view). 
9. Ciuhernaculuin (lateral view). 10, Body hooks (lateral view). 1 1. Male tail (ventral view). 12. Male tail (lateral view). 
13. Body hooks (lateral view). 14. Female tail (lateral view). Scale bars = 500 gm 2: 100 urn 3. 4. 5: 50 pin 6. X. I I, 12; 
25 urn 7, 9, 10, 13: 250 urn 14. 



I pair lateral ad-doacal. 2 pairs lateral pic-t loacal; 
all same tf/ja, 3 pans pupil lye « pair phasmids well 
posterior to cloaca, near lail rip il 7 ig I II Cloucal 
region with small euticular bosses: a hi- like 
e.vpaitsion of body anterior and posterior to cloaca 
ilaw, I 1 ), Tail ?MMMJ i305Mnn.L< (Fifr 12k 

K-male; Lettglh lfr-22 (?0) nnn. width 8WMIB? 
iXXOi. Cephalic buib 455-58Bt3KQ Um» hy 4!EM0U 
t'SOO) wide: cephalic hooks l>i row 1X0-235 i2IO). 
'..d cow 22(iObS (24>>_ ,Vd WW MO- INI) i, LS5» long.. 
I 'esophagus 17X5 2125 1 1 4001 K»ng. euheulur 
dilutiou beaiiuy 14 IS rows body honks, Deinds 
h<. /so (fti% oervi ring 4411 52fl-i47fty, frceneiars. 
i. ioi\ pore 520, 5S5 fjj=3l from anterior tod 
Vulva 7*140-10370 (tffttfjl bom anlcnorend (Fig. X|, 
Vupna nbriui 300 long (td=IJ Tail 985-H2J* 114)50) 
lone 14). Lggs ovoid 45 5 t | IKi Idug by 5fr 

51 (4?) vwidfl 

Vv/x Auw 
Pcnmulcs tiosi.tii vi.olliov 1801 

l\{>, in, ,///m 
Sydn-'V, AnMiaha 

Sin. ill intestine 

Is {u v/jr/ iftKit 
NcolypcAdIC 1X20 


The present location ol [he material ilftotfihed b> 

l.msiow as Ilofihn cfihahks then renamed lt)ttW1 
iitniii is unknown. Ytirktf & Mapleslone tl92ot and 
Chabaud ct at, ( I9X0| neither gflVC fl foCIKfan forth*: 
lype material nor indicated whether Ihev had 
(jxaitiirted u I mstow was working in (lottmgcn in 
IX ( >X bui neither C hinnmina nor tlt'pkh (flu/ins is 
listed under specimens held in the 7-nolngishtf* 
Museum det IIinnbolL I'mveisital. Berlin Mi is 
Museum docs, however, list holdings til other type 
s|h\ miens from linslow The specimen^ are not held 
in the parasite collections t die HM <NH). tlH 
liilcrnalional Institute of Parasitology. St Alhatts. or 
tin- I S National Museum Parasite Collection in 

The type host was given by l.msiow (lS9Sa) as 
Pemmtft's nhi\nfus. one of two bandicoot species 
collected In Richard Scmun. These species were 
identified by Rorncr (1001), using Hie catalogue ol 
Thomas ( 1888) in the British Museum, as P'mmeUw 
nhrsuhis, from tturnclt River and P. wtn mm from 
Cooktown. with measurements being given far /' 
itnttrittv. Pemnteles inaavnt is formally listed as a 
svimnvm of hnmiiin huh rt/Hrn\ (Mahoney St Kide 

l°XX) but R <>hrsninv \* not- This i^ surprisine 
because the luxonoiMie stains of P. *>fr<\t(in\ was 
discussed by Maekerras ik Mackerras (l%0> who 
indicated thai PcmutcU-.s ohesJtltf -■ Diclrfphts 
i'ht'\n/ii - htunfiHi ohexuhts, t>ut thai smee die 
northern lulut of disti'ihitluMi of fjtootfotl vt>t?$ulti$ is 
near Sydney. bandie(U'ls recorded a> /. ol,w\uhis ffDlW 
Oueenslaud should be referred !o as /. mm nmnts, 
the species occurring from north Queensland 10 
nuiihern New South Wales 

Vnrke & Mapleshme 1 192(>) list ihC i>|K lnist fij 
PinuHth', i)ht'\ttlu Johnston & Mawson < I **3V* > 
decided that since Vorke & VlaptesUHie had dniW n tut 
original figure their ntalerial came from low/nssilk- 
ill itiMthvrii Queensland and Mated that I l]Vfr)ty 
nialerial from I\oni/i'ii ohesuhts earnc trom f pper 
(iinnra River also in Queensland On th-it basis, 
Chabaude/w/ ( IW0> concluded that the host ol bodi 
ihc I mslnw material and the Nurke & M;. pi. Minn 
iriaterial was htnuhm ma< nmmw the northern blown 
and lliTl / nhcMilii^ the souihern brown bandieout 

Thtsd^es iuit. however, explain wh> Komci ( I *>0 1 i 
m Ins identification ol ihc bandicoots collected b;. 
s'nrioii IJ^tS diem as two -.eparate spcues ratln t th-i.i 
as Perurnelcs watrtim now IwvdDH itmcivitrus. The 
DllalDgUC of Thumas (IX8X) however docs [ittl / 
iKixutn. so the 1. pper Hurneit River bandicoots weie 
l^l-.ibly /. tlUHSNtlUM 

Chabaud </ ,il, (l l >Xih noted IhuL I.X mm. Ihe 
ineiisureinnit given fni Ihc length of die oesophagus 
by Lmsiow (Ixr'Xai w;issundai to measurements ol 
specimens examined by ihcm that had been collected 
Iomm I tiNu munis ffoin northern Nt-w Suiiih Wak-s 
duomih lo northern (Queensland, In particular. |hc*C 
speciniLits had the oesophagus Urrmiiuiine at abom 
the level ol the posterior end ol the euticuliu dilution. 
j eh.iraeiei ihev tlescnbed as a ' lonu oesophagus". 
Yorkc lV Ma| | I l Po) dul noi nidh.ate |hC 
length of the oesophagus relative lo (he lows of body 
hooks on the cubcular dilation but they described I 2 
or 13 'circles" of 14 Ln lb rows of hooks. Two 
spepililCjlS <ol|r-Lted by Nicoll in 1915. currently 
held in the BMiNIl). whieh nuild possibly be the 
(ualcfial described by Vorke & Maplcsione 1,192(0 
.uid n-desL i iliL-d below also have the "long 
oesophagus" deseribed b> Cluibaud rf (u, i 10X0) 
The specimens described by l.inslovv ilS^Xa) are 
drawn mk\ labelled as having 17 "circles" of hook.s 
with ihc orMiphagus lermiiialing at the level of the 
L )lh row of hooks. These characters are eonsiMcut. 
not with /:. < iihrnm \t't)\t< Chabaiul ft ul i, l ( )X0) but 
with specimens described by Chabaud tf nL t I9X0J 
as £ nwridiontids. occurring in the southern brown 
handicool fxi><uh>n t>hi j stiln\ collected in Soulh 

In a re-e\amunuuHi ol" the nniiertal desi i ihed hv 
Chabaud tet aL (1980) as E nwritiiontili\. together 


Willi specimens collected liom long-nosed and 
barred bandicoots lor Lhis study, il was found Ihai al! 
specimens hud 14-18 ttjwa ot" (tody hooks and rlic 
oesophagus lei minuted al the level of the BthT Ith 
row of hooks. Measurements i»t the oesophagus 
ranged from 1445 lo 2040 lor males .md 1785 to 
2125 for females;, also consislem with fhe 
measurements glvfiifl by Linsiow t 189M 

Johnston & Maw son i K>40| described three 
females and tWO males obtained Jrom the intestine ul 
Ihc long nosed bandicoot /! tutstnu collected in 
Sydney and attributed diese to /:. iiruttun. I hey 
dc.sctibed and figured differences m the male tail. 
nameh <in expansion gf Ibe body surrounding "be 
cloaca, similar lo. bul not transparent, *i$ arc caudal 
alae Cnahntid tf ai ( 1 ^SO) figured the posterior end 
ol a male, which they designated FfhiiHttu'tHa sp,2 
I'rom /' titisHti/ registered in ihe Al IC* as IK20. They 
commented that Iheir specimens were comparable 
wnh those desciibed by Johnston & Mawson (1910), 
N'eitlier group described die anterior ends pf ihe 
woods tbey examined. The only material registered 
in the SAM'\ which might be the original Johnston 
& Maw son *j>eeuut:Ms is AHC 1X20. The 
moipholog\ of the anicriot ends ol" these worms, fc 
IS body hooks, ibe oesophagus icrmmatmg level 
with ilic °ah I til* row ol hooks, is consistent with fi 
trh •niti<uuili.\ \rii.\n t b.ibaud. Seureau, l>cveridge, 
Bal|V& L»metlc-l>essel ( [V{®. 

Chabaud r) til M l >S0i did nol comment on ihe 
pie.senee ci| absence ol an expansion to the cloaca on 
tlion specimens but coudast A. tn<>ritli<>inttts with 
CiMMJWtfSP ) (s'.v rhab.iud rt at I WO p, Cw, 
he. >) .slating Iltul /.t hinem -ntu sp. 2, was 
comparable with 1he lohnsion lV M.iwsoit (l l >40j 

A re examination ol all I he available -.pecimrm. ol 

/ tttft ittit'ttttitt \ wV| and fifh>tu>intm' s\x 2 |*/t \ has 
laded 10 show an> significant dil Terences between 
tfu m Such diligences as do ckisI can be attributed 
to ihe tact that Chabaud cl tfi, i K)J(U| were dealing 
v.ilb a mixed infection ot' two species, namely L 
* t/h twn ,\\h\ /.. \vm rin^tnui (described below } 
n\'i iiMin- m ihe ift||cClC4 , " ,,M VVulp.. j 
and Ylyponga. Tftfc male I. ctftvtWfo ill tlffe 
population were al Ihc small end ol the si/c range 
and oesophageal length varies with worm length and 

The description by I m-.iow | I NOHa | .if // r,*tr'«\ 
is congruent with Ihe revised description 0| matciial 
designated /:. tywndivtmlis by Chabaud l.7 <(/. ( l l >Htt|. 
Kit dr^tnpiion by Yttrke & Maplestoue 119201 ol 
xjKtuiu-iis lunn /. Mtniititnt.s and identified as t 
tiiiciii. i-s coiie/iicni with E ctnviwtt scusu t'habaud. 
Seureau. Bcveridije, Bain & DureUe-Dessei, [ l KS0 
ii<<< I in-stviw, rs^K, The re I ore T I designate a specimen 
tuitu AHC IN20 .ia the itL-olvpr <«1'/,. | nhhim 

l.instcwint'/nn vim turn il.mshws, 1898) most 
closel\ resembles /. mi>/i\i (redescribed bekivvi in 
thai the oesophagus is relatively short in relation to 
the number of rows o( hooks, ending within ihe 
euticular dilation- The males ol' both spei. u-\ h.iu- 3 
pairs of caudal papillae and spicules 7 j - / - bt>dy 
length- f.itisiiniinfitni rhwturft can be distinguished 
iVom L ingHXi m Slaving 13 16 unalei and 14- IS 
( female > body hoolcs compared vviih 10-12 and 1214 
in /,. tn^iisi. Although the ocsoplumus i^ relatively 
shorl it is, however. Longer than to /- ilffilfxu beiny /, 
brid\ length in male L. * rractutn eoiupaied with 7 n 
biidy length in /.. hlftuxi. The paii ol lateral ad- 
Jo.ical papillae \b no lai^ei than the »nhei pairs oj 
ventral papillae in L linciwn but is larger in L 
iitt*IiM (see Inglis l%7 ptgj SJ). Ihe ala-like 
expansion ttf the body surrounding the cloaca of L 
tint tarn is not found on /., Wftlhi \$&$ Inglis l u 67 
l r ig. 9J. Hie body spines ol '"/.. rimittm only covet 7j 
ol the dorsal body surrace but ctnet "7 ., of die dorsal 
body surlVc of /. inyhsi- Bg£S of/, cmrdttn dilfet 
Iroin those of all other species in that they are ovoid 
iMiherthan almost sphericaL 

I he material from /. ofextifitx from South Australia 
described by Cliabaud el al. [WHO] as L 
inc>ri(ii(mtiJi.\ w j as found lo be a mixed infection ol L. 
cittvwm. (females with up to 18 rows of body hooks^ 
oesophaeus irrminaune ai uboiil ihc jcvd of the 9th- 
1 1 Ih row ) and /-. mtnwghwt (letnales vulh up lo 13 
rows of body hooks, the oesophagus terminatine 
abnul the level oi the 1 0th -Hih row I described 
below A comparison of die measurements given by 
( luh.Hid ft o/. ( l°KU) ami ihe specimens cvmmu-d 
lor Ibis Mudy reveal thai the irtales (bey mejsured 
were smnllet in sire, the oesophagus, spicules and 
lail were shorter than in die specimen> examined rot 
this study The females, however, were within the 
s.nni- >i/l laitucas foi \\\]-, siudv and the compaiutiw- 
measurements arc more cOnsistaoli 

The material ihssetted (torn the southern brown 
bamhcooi collected al Scott Creek included some 
females up to 3b mm lone. These were larger than 
til© spe.i -nivih ftmii S.u.lli Ausli.tlta slodicd l>\ 
t Itabijlul // ttf, \ lOKOt, tfjp h> :7onn li>ii.C) Inun 

easicm baaed bandicoots from Tasmania, (up lo 
I2iq.rn long), and from lona-noscd bandicoots, from 
Tasmania. (30-32 mm loner OtJiet vatiaiions 
olisetved between female specimens collected in 
diffefetii busts atid lo*.ii|uic^ hi the present study and 
itiose ot Chabaud vl al, \\9WSt) included the 
oesophagus longest in southern brown bandicools 
from Secilt tYeek < J Sri3 23St)), and shoriest in 
southern buFtfd bandieoois fhltri Mvponya an J 
Wailpinga 1 1 l()0i; the tail longest in eastern brown 
bandicoots Ovm Tasmania (MSO-H22) and shoriest 
in southern brown bandicoots from S*oit ( 'ovk 
0SS4-*>Not. alut the \ulva sbelnly *\n»r pOSl^floi 

m I- k. SMALES 

(I().l)30-I4.45()| in soudiern hruwn handiconts from 
So tit Crfiefc than in eastern barred bandicoots tftirii 
Tasmania (7140-10,370) of southern brown 
bandicoots from Wailpingn G0.K00). The eegfi Irom 
iht* Scull Ciook specimens were smaller lhan those 
from Wniipinya or Tasmania, being 33 by 36, 32 by 
45, and 43 by 4S respectively. differences 
could be either variations between populations 
within (he species, or tfifl resulr of contraction during 

l,ttt\rt))\.hwn)ti <ini'(ttni, originally desenhed as 
occurring in /* ttkcHUh (\t) tVuni Queensland is' now 
recorded a> also occurring in /, o/'< ,w/u,v. /' nasitlu 
and /'. iitinti'ti, The geographical range now includes 
Victoria. New Souih Wales. Soiuh Australia 
including Kangaroo Island, and Tasmania as well as 
norlhetn and southern Queensland. The record ol' f 
aiwnmt in A ifbe&uPuti. from l.ismore, New South 
Wales. AUG 4413 is n dubious host record because 
llv southern brown bandicoot is noi Tiuind nearly as 
lai north us I ismore tBrailhwaile I995i. In .all 
probability the host was /. itiiicnmrus. 

l.insttminema warriiigtoui sp no\- 

iFIgs 15 24» 

Swuifivins: l:chinotn j itm ttmla senxt Yoikc & 
Map^Stqnc 192$: pp. 347-348 net Lnisiow, IS9K; 
//r< In^rlis. I%7:.\mw* Mundav & Green. 1972. p. 10 
tin pari); \i j nsu Ghabaud Seutcau, Bcveridew Bam iV 
Iliitellc-Dcsscl. |9,X(): p. 435-43f> (in part); ^n\u 
Spratl. Bevendge & Waller. I<ty1 p. 25 (in pail I 
lyitiit'uwinu cunwtn vmof Johnston & Mawson. 
WS3: p 33 

Material cuwnitctt 

From lxntf8i>f) tnirtttus. I A '. 1 v . no collection data. 

I mm iKrimton Wttrivttno Queensland Id, I Y- 
Hiilline.sKme. H9 OVS. 140 M'R). 24...IM15. HM 
iNUi |$5fl 12. 0.165 16(3: VIossinan to Oaiutrec 
Koad- <10 1VS. US |0'h), 2a. O'Oi.AHG JB275: 
22] J. 2ifo>y£. Vjossmau. GO JS'S* '42' LVU 1^58, 5ii.|95K, iiiiJVSH llviii 195*. 
Ij.n I95K. 20 li.l^SK. 9.n 195S. QM Gl 14351 
(,l 14301, GII43n3. Gil 13M.GU4565. CiLJ«<tfi 
(4.1 tOO. (.1.14372, lU 1 437.VGU4377. Gl 14^1 
GU4W; 2.;.'. V . . ( "aims u> Mossman Road 
do 55% 145 46T-). 1m.PW1.AMC 3027-1. I M, 
Gillies Highway near Lake Bamiie. (| / I5\S. I4. s 
SR'El 79 ( v109l. AHG 30270: frtf S I I 'i\ 
Athcnm. «)7 lo S. 145 29'Ui_ 2.V\.IOX2- GSIRO 
NI6IO; s Y Y. I I £ v. Vunyahurra to Atherton Road. 
(17 Irt'S. 145 3S'l-.i. 29.x 1991. AUG ^027N: 7 Y M7 24'S. 145 SS'HJ, 25Jx.W57, 
QMGI.M30S, IXI ';,', ISO- -:.Garad-eeG7 29'S 
I4fi OO'fclt. 20ii.0)so. qm GL 12653, GL.43&7; 

8c 3. 79 '2, Milhw Millaa to Inmslail Road. (17 
^\ , S. 14? 37'Hl 3 i.*. 19*1, AHC 3027b: 7f>o '. 
13V; 'V. Inmslail, (17 32'S. 146 Ol'l:). l(>.vi.lV50. 
5.x.l l J53. 20.vii,) l J5rS, 27.eii4^5ri. twiiJ'150. 
I9.vil957. IS.ii.l90t), AHC 452S : QM GU4357. 
I 4.14300. GLI4370. (iI4437S, GLI4379. GLI43K?. 
7 ; 5 , 3 5 1 V Athenon to Ravenshoe Roiid. ( 1 7 Ib'S. 
145 2947K 3l\i09l, AUG 30277: h: , -1 ? S. 
lugham. <l# u 39'S. 146 KTE), 31.x. Iwl. AUG 
302SO; 2J 5,6V ,\ Palmcrston Highway, 29,v.l9f>9. 
QM QL 14360: 7 V f J . Hlkm south of Proserpine. (20 
|4'S, 152 35T4 \\K 30266: J t J . Roekhampion to 
Ycppoon Road <L*> OS'S, 150 44I'». Id.viii Tvh#0 
3_vii 1992. AHG 30271. 30267; J$S, Ycppoon L0 
l.niu Park Road. (23' 10'S. 150 46'R), I'&ijlIW 
AHG 30270: 4-3^. 15 V v. Roekhaiviptoii to t ; mu 
Pail Road. {23' I.VS, 150' 30't) 17.iii 1992. AUG 
^0272: l_'. 2- - . Rockhamptou U. Keppel Sands 
Road. (23 20'S, 150 4ST.). IH.vii.1992. AHG 

3026K: 6°'.- Mt Glorious, (27 ( 2TS. 152 54'L4 
r/,v 1955. OMGI.I4359:69i ( ^ I2tr, . . Ash^rove 
(27 27'S. 153. 02'H), 27.i 1956. QM GI44347: 
120. ; -:. 20^4. Paddiu^ou, Btisbanc, 127' 2S'.S t 
155 OI'K),25.viii 1955. 21 \. 1955, I4,ix. 19.55. AUG 
4371. QM GI44350. GLI4371: 29 fl'fl , 55, , 
Brisbane, i 27 2H 'S. 153 01 'Hi. 2S.K4954; uM 
GLI4-UI.. (4.14380: 7<S<5, (K . M.^ill (27 1 
2"'S. 152 54'h). 12.x. 1967, GSIRG N157; 2.;". 
!8'S. Mi Nebu, (27 3.VS. \>2 57 T). I,\ 1954. 
23,vii. 1993. QM GL 14159. AHG 303l7_ 50S1S. 
$66, i^v . Bmokf.eld. (27 30'S. 152 55'h). 
20 v.| ^73. |9.vT967. AHG I9V,7. GSIRtJ N15I: 
I.G' ?, 21 v 1 , Rocklca Giossme. Uhx.V^. C2ilRO 
NI52: Nev\ South Wales: IV. no olhei data. AUG 
4462: No lucaJitV pven: 97 J h 171 V ;\ N ni i 
29.»\.I954, r5,x49.54. 28.i\.l954. 27.i.. I osr. 
t2.u..i95x, QM CLI4H5. Gl. 14346. G144^52. 
G|LM393i6l 14354. Gl .14355. tit 14362 

From fuunlnn ntw^mlus New Soulh Wales I 
Sidhnys Swamp Hoail 'limbillica Slate koivst. t^" 
I7'S. 149 45'L). GSIRO N42 
Victoria: I ! •:-:''. I0H x. mi collecilou doffi, AIK" 
44M. AHG30297:(v" r . A y Halls Gap H7 OS'S 
142 31 K). no dale. At l( 30297: Mtf. 3 f^OUW 
Reservoir, Grampians. (37 L i4'S. 147 X)T'.j. no dale. 
AHC 30299. S.uiih Austiabu. Vt '; . Mvpon^i, <^S 
Z3'S. l3>T 2S'P). HO date. AHG 44-16: : '.' , I 
anierittr end. Waitpintia. 1 35 36'S. MS 324-1. nil 
dale i\H< 4400. Kamjarou Island South Auslralia' 
97 V. I ;. V.vonne t^v. i 75 59"S, |G I3'l ) 
U.P'XS. AHG 30302: 16 i _ J . 39, _' . Puri»dana. (35 
47'S- 1-^7' 104... 3.tiTWXft. AUG 30307: 16: 
\V ' Hinmu\ic_ i.G if7% \ift So Hi 5.rihM9«7: 
AHG 3031)3. 

I loin h J rtuth-li< nusitw QueensJand. I > > ' 

Marina, (IT" 00'S U5" 1WV) 29 A 1991 AUG 



Figs 15-24, Lmshnvmctna wvtrift&totii sp. nov. 15. Anterior end, (luteral view), 16. Cephalic eml female, optical section at 
level o( first row of cephalic hooks (en face view). 17. Female, hand eul transverse seelion through body hooks on 
cuticular dilation. IS. Body hooks (dorsal view). \ l ). Body hooks (lateral vicwi, 20. Male tail (ventral view)- 21. Female 
tail (lateral view). 22. Vagina (lateral view). 23. Gubcrnuculum (ventral view). 24. Female tail lip (lateral view). Scale 
hars = 200 pm 15.21: 100 pm 16, 17. IS. 1 9. 20. 22. 24; 50 pin 23. 



Cephalic luilh with 3 ksivje ttf 14 [maltfj in lb 
tfcinale) tiles of llOOks (I -ig. 16\ Jhd row longest, 
3rd row shortest; net k wilh 3-7 rows liny spines; 
eniauka dilation of oesophageal region bearing 9 I 3 
rows t)P 14 (male) or lb tlemale) files of body books 
(Fifii 17), 1st and liisl rows smallest, 6lh - 8lh rows 
Unrest. fOOt* Of hooks With undulating edges (I rgfl 
IS, 19); remainder of body with up lo4N (male! or 54 
(female) small spines at each annulation. over whole 
body of female: extending t)\er '/$ of dorsal surface, 
terminating tthOUl 50(^ anierior 10 cloaca, not 
teaching level of most anterior pal] of Cauda] 
papillae, on ventral suiiaee of male body (fie, 20). 
Oesophagus. '/.»> - Vi*. budy length, simple club- 
shaped; lerminurine ul level ol U lh-I3th row o\~ body 
books (Fig. 15) neat posterior Clld of anterior 
eulieular dilation. Nerve ring surroundinc 
oesophagus within cepbalie bulb; seerclory- 
e.xeivtorv poie m neck: deiiids conical at Lovci 0t W 
tow ol body hooks. 

Male: Length 15-30 <'7.5) mm. width 4511-75(1 
I m.S). C ephalic bulb 270-340 (2W) long by 270- : * W 
(295) wide; eephahe hooks 1st row 110-170 (155), 
2nd tow I iSO- 1 'jo ( I 75). 3rd row 100- 1 30 1 1 10) tnng 
Oesophagus 1575-1925 (1743) long, emicular 
dilation healing JM I lows Of' body hCK>kft (JdriOfi 
4°-0-550 (530); scerclory - c:\ciclory pore 3o0 (n=l ); 
nerve ring 310 (11= I) from anterior end Spicule 
similar, equal, without akie. o l H)-IOOO (850) long 
$bCUIt '/"< body length duhernaeulum short, simple. 
SUb trianeiilar. 50-75 fu=5) lone (lie, 23), leu p.m ■. 
1 aintal papillae; 3 pairs ventral and immediately pre-, 
.ul and post t loatal rcspceli velv. I pair lateral tu] 
eloaeak 2 pans lateral pre eloucal. all same hllti 4 
puts pupil hu , pair phasmuls posterior 10 cloaca n^ai 
1 ; m I ijp [fr'tg, 20), ClOUUil region with small cuIicmI u 
blisses ala like expansions absenl Tail ^3(M 30 
1 IlKtj Ion-. 

1. inab! 1 tmgiii h 4i (3Ki mm. width mpam\ 

1M01. Cephalic bulb 350 VHi (370) long by «0 Mill 
(2fi0) wade; bearing 3 r->w>. Ill ho. ( k>, 1st tow 170- 
-!'"M l-SS), 2nd W MJMJOl l l >5) Jrd r»»w 120-150 
(HttJ long, Oesophagus IS70O500 C'2S(M lung, 
eulii ulai tJilaiiiHi bearing I 1-13 rows ol 'body hooks. 
Deirids 5X0-700 to70); m-< orm j, -rsueiurv pore 
370-100 iV-n, nerve .in- 300 3h0 (33ti] IHIMI 
in- nor eml. Vulva 1Js-I5.o (14,2) mm itom 
anterior end ibm ±2). lad 1000-NOO (1265) lot^j 

(Us* -'■ -•*' Vagina ftboin KK) long (n=U ' r- 

ahnost sphetteal M) 44(3Nl by AW? (40), 

This species is named aftct Wtiinugiou ftirke who 
with P. A Maplcslonc earned (Mil much ill ihe eatly 
w*m1 on this uenns. 

Type h'luliry 
Towusvillc. Queensland. AuMiiilui 

Type ki>$l 

hiwiinn ttiut'nhtrux (Gould I H I 1 1 

Small intestine 

/V/T A/?Ci /WfVJ 

Neotype BM <NM.) 1050. 12 b 


Tbe two worms. I .•:, I , BM (Nlh l<J50. 
I2.6.lb5-16b from I'ciwtuic) nhcsnld collected by 
NiloII in NI5 in Queensland .wc (he only specimens 
rcmsiercJ \\\ AlisLiahan or I lulled KlUgdotll patasite 
collections which could be the material desertbed by 
Yorke & MnplnMone (1 ( )2<0. On examination thesL- 
vsorms were louud to have the II (male) and 13 
(t'emale.) body hooks desctibed by Yotke &. 
Maplcsuau' (1926) for EchtttMMUA "'""" -'tul to 
have the "long" oesophagus ami all the other 
chaiaclers allributeil to /:', vhuiutu by Chabaud vt ,tl 
(l L )H0,i. I ihea'lore designate lite male specimen ol 
RM(NII) 1950 [2AJftJ Idd as the nc»»type. 

All the specimens WciUifiCd m lbi.S study as /. 
\\turiti',;hftii conformed to the dcsctiplion gi\cn by 
Thailand ri ul, (1080) of /. iimnmi- /ittvnu tthitht 
wturinxfoni sp, nov. diflcts bom all ot.hct species in 
the gemiS 111 llflVitlg lip h' u -l I rows ol hiX.ks (hiak ') 
01 IM3 (lemale) on the oesophageal cuticulai 
dilation ami a "long"' wesophagus, that is, the 
oesophagus tcimmaies ul Ml IKlil Ihe [foul IBW nt 
hotly hooks. Male L toWfibiRttwl can be 

li .im shed (TOm I ' /Wl tltlH b> [hfl i-oinhnialton of 

chaiacfcrs at ihe postcnoi eml. i.e., Ihe terminating 
staitial IhmIv spines Jo not extend to Ihe mosi 
anterior |Wlt orpapilkn- in /. wan m^Unti Imt .lo in / 
iitiitittti, the distance hehveen spurs and clOrtTd r, 
500 |_im uol 10ti as in /.. . iiufntn, ihe hniucd cucltt 
ot iiiii^ulai bosses riprrn^nding lllti cloaca m /. 
H tit rihvjtitii compared with 7 . rin* /(/'/'. I 
\vnrrin,i>t'rtii does not have an ala-like expansion 
sinnMiudiiiy Ihe Moaea IiliI i. < tiunnn does, ami four 
I'aus ol papillae not thtee \h\ Ihe tail lip The spu ulr . 
of J., wtitiviyjeni 'm hody lettetl' an* ihnitL'l lliilll 
those of /■ cinctum 7jj body leimth. The . ol /. 
wartni^htni are almost spherical but those of /,. 
1 tm tutu arc OVOid. 

Chabaud <'/ til, ( l ( >S0) in their description <»l /. 
litu nun {sit) indicated lhal they had studied 
numerous specimens from a ranee iy\ localities 
im indui'j Woolwcutga ^^\ Darwin in the Northern 

Temlory. A re examination ol the material fujm die 
Northern Territory has shown thai it represents a new 



species of Linstowinema, L. latens. described below. 
The measurements wid figures of specimens reported 
by Chabaud et ah (1980) however, are congruent 
with L. warringtoiU rather those oi' L. latens. 

Spratt et td. (1991) noted that there were no 
records of helminth parasites from /. aitrutus the 
golden bandicoot. The finding of L. warringtoni in 
one of five golden bandicoots dissected for this study 
is therefore the first record oi a helminth from this 

Material registered in the QM as GL14345, 14346, 
14352, 14353, 14354, 14355, 14362 was collected 
by Dr M. J, Mackerras. Therefore although no 
locality was given these specimens are probably 
from Queensland. 

The finding of L. warringtoni in /. obesnlus is also 
a new host record. The specimens from Waitpinga. 
Myponga. Timbillica State Forest and in two of the 
hosts from Kangaroo Island were found in mixed 
infections with L. ciintttm. The geographic 
distribution of L. wnrriftgtoni therefore has been 
shown, in this study, to extend from northern 
Queensland down the east coast of New South Wales 
to Victoria. South Australia and offshore to Kangaroo 
Island. A larger number oi I. obesnlus from the 
southern states needs to be examined before 
geographic distributions can be fully mapped. 
Populations of bandicoots in NSW, Vic. and SA, now 
have patchy distributions over a reduced range 
(Brailhwaite 1995) and attempts to collect additional 
specimens of /. obesnlus for parasitologies] 
examination have been unsuccessful to date. Further 
work on the southern geographic distribution of L 
Wttrrmgtoni will be problematic, as bandicoots 
become more difficult to collect. 

The specimens of L. wurringtoni found in a single 
/' uasntn suggest either a natural low prevalence of 
infection, or an occasional, incidental infection of 
this host. 

Linstomnema latens sp.nov. 
(FIGS 25-36) 

Synonym: Echhwneitta cinclnm sensu Chabaud, 
Seureau, Beveridge, Bain & Durettc-Desset, 19S0: 
pp. 435-436 (in part); sensu Spratt, Beveridge & 
WalLer, 1991: p. 25 (in part). 

Material examined 

Type material: Holotype o\ allotype V, from 
Isootlon maerounts, Walsh Point, (15° 08'S, 125° 
46'E). Mitchell Plateau, Western Australia, 
22.vii.19H2: AHC 30322. 30323, 43d 6, 65V V. 
paratypes AHC 13028, WAPvl 1 10-83, I 16-83. 

Other material: From fsoodon maeronrus Western 
Australia: i6. 4V V, Mt Hart, Kimberley Ranges, 

(16° 48'S. 124° 55'E), 22.x. 1993, 24.x. 1993. AHC 
30289. 30290. Northern Territory: 16-5 6. 18??, 
Darwin, (12° 27'S, 130° 50'E). no date, I9.vi4993. 28. vi. 1995. AHC 4703, 30287, CSIRO 
N4413, N4414; 16 c5, 79 9, near Byers Rd, Stuart 
Highway turnoff, ( 12° 30'S, 130" 50'E),, 
AHC 30284, 30285; 16 o\ 109 9, Adelaide River, 
Arnhem Highway, (12° 28'S, 131° 14'E>. 1993, AHC 30288: 13d 6, 12? 9, Bees Creek, 
off Stuart Highway, (12° 35'S. 131° ()4'E)., AHC 30286; H6 6. 89 9. Jabiluka, 
August 1979, AHC 6421; 39 9, Woolwonga, (12° 
45'S, 132° 39'E), 19. x. 1972, CSIRO N159. 
Queensland: 6 6 6. 1 1 9 9. Atherton ( 17" I5'S, 145 
29'E), 25.V.1928. CSIRO N1610: 1155. 149 9, 
14km north of Atherton (17° 15'S 145° 29'E), 
15.iii.1982, N1532; 1955. i 2 9 9 . Yungaburra to 
Atherton Rd. (17° 15'S 145° 3()'E), 29.X.1991. 
I.xi499l, AHC 30278. 30281; 1 5, Gillies Highway 
near Lake Barrine (17° 16'S. 145° 35'E), 29.X.1991, 
AHC 30279; 65 5. 89 9, Gillies Highway near 
Yungaburra, (17° 16'S, 145° 35'E), 29.x. 1991. AHC 
30283; 23 6 6. 116 9 9. Mareeba to Kuranda Rd 
(1.7 s 00'S, I45 D 26'E). 2. \i. 1991. AHC 30282. 


Cephalic bulb with 3 rows of 14 (male) of 16 
(female) files of hooks, 2nd row longest, 3rd row 
shortest (Figs 26, 27); neck with 2-9 rows tiny 
spines; cuticular dilation oi' oesophageal region 
bearing 14 (male) or 16 (female) files of body hooks, 
first and last rows very small. 6th - 7th rows largest: 
roots of hooks without undulating edges (Figs 29, 
30); remainder oi' body with up to 44 (male) or 60 
(female) small spines at each annulaiion, over whole 
body of female; extending over 7m of dorsal body 
surface, terminating about 300-400 anterior to 
cloaca, level with anterior pair of caudal papillae, on 
ventral surface of male body (Figs 32, 33). 
Oesophagus '/m - '/n (male) l f\s - '/'« (female) body 
length, simple, club-shaped, terminating posterior to 
the oesophageal cuticular dilation (Fig. 25). Nerve 
ring surrounding oesophagus within cephalic bulb, 
secretory-excretory pore in neck, deirids conical, at 
level of first row of body hooks, 

Male; (measurements oi specimens from Western 
Australia, followed by measurements oi' specimens 
from Northern Territory). Length 15-18 (16). 15-21 
(18.5) mm, width 425-625 (490). 515-715 (600). 
Cephalic bulb 280-350 (325) long by 280-380 (340) 
wide, 275-435 (350) long by 290-385 (340) wide; 
cephalic hooks, 1st row 150-180 (152). 105-165 
(140), 2nd row 160-190 (168), 125-195 (155), 3rd 
row 100-110 (108). 72 117 (94) long (Fig. 26). 
Oesophagus 1200-1825 (1555), 1325- !9 ( >0 (1715) 
long, cuticular dilation bearing 8-10 rows body 
hooks. Deirids 540-690 (61(f), 390-650 (530); 



figs 25-36. Linsiowinema fawns sp. nov. 25. Anterior end. (lateral view). 26- Cephalic bulb (lateral view). 27. Cephalic end 
female, opIicaJ section at level of first row of hooks (en face view). 2K. Vagina {hileral view). 29. Body hooks (lateral 
view), 30. Body hook (dorsal view). 3-1. Female tail (lateral view). 32. Male posterior body spines (ventral view). 33. 
Male lail (venlral view). 34. Female tail tip (lateral view). 35. Male tail (lateral view). 36. ("iuhenuiculum (ventral view). 
Stale bars = 200 pm 25, 26; 50 pm 27: 100 pm 28. 3 1 . 33, 35: 25 urn 29. 30. 32. 34. 36. 



sccieloiy-exueloly pore lipl RQtfn. 230-4°o (365); 
nerve ring 300 (u=l>. 22&-J&S (275) I'roin anterior 
cm! Spicules equal similar, without nlac. N30-lt»0 
tW?J. 8#)-l 100 i 10051 long, about Vi« Eft 7... bt"ty 
kivjih. ( uiheruaeulum short, .simple, suh triangular. 
Ol-dS (nrfJL 6f>-70 (u-fi) long (H& 3bl 10 pans 
caudal papillae; 3 pairs ventral and immediately pic . 
ad- ami post-cloac;il respectively. I p;iir lateral ad 
cloaeal 2 pairs lalerai pie-eioacal. all same \t/c. 4 
pairs papillae, pair phasmids, ala-like expansions 
absent posterior Id cloaca, near tail Up (Fig; 35)'. 
Cloaeal region with small cuiieular bosses- ala-like 

ejtjiaiiMQite absent (Ml i»wft (2M0j, 3&32G 

(.?05J lon^ CPip **JI 

I emale: I englh 3J)-3Hf3D), 25-41 (30) mm. vs.o.h 
420 750 ,vK)). (S^S (175 t*40). Cephalic blllli 30(1- 
3N0 <TSOi. 230-510 (AK5J IfinJ In SStf-SW (3K5». 
375-5?0 (I3(ii \\uie, Ikmiiivj } row* of hooks, 1*1 
row Ib0-10()<[75). 135 -IW 055), 2nd row 170 -NO 

i isoL N0-io5<i?o).3rdrow iuo- i :o oioi^o -I 35 

(IJIQ llW Oesophagus 1450-2075 <I7sH)1, 1585- 
2430 f|07Q) lomj: cntienliir dilation bearing 4-12 
h, K l> hooks,> 600-701) IbfiUJi 325-77lft4 , >5>: 
secrcloiy-escreloiy pore not seen. 2 i »5-5 t M) (420); 
nerve ring not seen, 205-105 (355) in=5j fiom 
anlcrior end (F%. 2M. ViflVa ".2 -14.7 (I I. Si. ».2 
17.2 ( I I 3>» mm from anlcnor ffrtii. Vagina about 1 15 
m-O). I'Vi ■ -. almost spherical 3G-60 (SO). 3342 ( J51 
by -33-W(34). Tail 7<->0-l 2I0 OW0). l >20 -IM5 | I IbO) 
lony (bie.s 3 1. 34). 


I he species name is deftved lioin the Laim IdT&M 
ni-'iHiinj' hidden, \jnils II wii« ruij KmiihI when Qtg 
material was first examined. 

/v/s lirfKl 
tsoixfott imuhHims ((innld. IJJ42) 

Mitchell I'lateun. Western Australia. Australia 

ffitf* (// //O.S7 

Small intestine 

Ixpc \ffci itttcii.s 

Molotvpe male. AIIC 3032 2. allotvpe female. AIIC 
10323. paraiypesAIIC H028 


/ htsieivuTi'ttitt Utu'ns sp. nov. resembles L. 
\uint/n;ii'tn. :ilso occurring in /. mm tVtlNl% in Iviny. 
ot similar si/e. Onales \>2\ mm in L iiUftn 
compared with I 5-20 mm lonj.' ill / \viirrini;((t/ii), 

having the oesophagus pf similar length ( I 575 1925 
in / Ittti'tis compared with I200-IX25 in / 
\\'jnvti!n»it males) and tour pairs of papillae on the 
male tail, Hie oesophagi's in /., tuiais. however. 
terminates posteriorly to the hooks surrounding the 
oesophageal legion whereas that o( I . \uttnni!h)ttt 
terminalcs at about the loci ol the dab - 13th row of 
hooks. /,. luf,n\ has K-10 (male) or l M 2 (Icmnle) 
i\»ws ol hooks. \oob' without undulating Cilfees, white 
/.. i\drriN\>ft>ni has l ' 1 1 unalel or 11-15 (feiriiile) 
raws oi hooks, with roois having undulating cilgcs 
on the dilated ciilietilai reymir The hody spines on 
the dorsnl surface tit nmle /. ItftfM e\iend further 
towards the posterior end, (aboul l, /iu v\ Ihe body). 
than on L WttwUtjprm'i Mhoiii >/s of the hody>- 
VVntrally the body spines c\icnd U) the same level us 
the mosl anterior |_nhr of papilhie on /,. /(//e^/v, 
but CJO nol on /.. Wfirf'tHffttittl TIW niale t;iil ffl lonen 
m /, \vumtvjj<>m (330-4 30) rlun in /, litnns (250- 
340) The vagina of / tuuws i\\S) is shoru>i than 
thai of./., wnmnvjoui (300). 

Specimens wltc found in /. ithtttom n\ from 
northern Western Ansiralia. the Northern 'lcnit"iv 
and northern Queensland. The popululion o\ 
northern brown bundicoots in Western Australia is 
isolated from lhal oi the Northern Terrnoiy and 
Queensland (Gordon l°3)5), bill measurements ot 
worms trout hosls i.euni'i.t^ in the Kiml.erh-y, 
Western Auslialia. are consistent with those from 
Darwin. Northern Territory- The only morphological 
difference observed between these populations was 
that ihe first row of cephalic hooks ol the Kimberlev 
specimens was almost as lony as ihe second row. 
I50-IS0. compared wiih l601V)o In males, but in ihe 
Darwin specimens the dilterciiee m lenyth lietween 
the two rows of hooks was more marked. 105-163. 
compared with I2n l ( J5 This difference is not 
considered lo be signil"icanl and as the Queensland 
worms were similar lo those from the Norilitin 
Tciriiory. the uuilerial from all three localities is 
considered to he conspecific. 

The northern brown handicoots collect ed \\\u*) 
iiorlliern Quet-nshiiiiJ vvt-re infecied uiih / , 
Mutt ihi*ttH)i. X hiNts. /.. lulrhs, 4 hosts. 01 liuth 
species. 3 hosts. Chahaml tt til, t I0S0I idcunricd alt 
the material they examined from I he Northern 
lc ntory and northern Queensland as I., (iminm. 
rio\v /.. n' ( ;r/7/;i,*/f'//i ll is not possible to determine 
from their paper which, if any, specimens from 
northern Queensland, presently lodged in the QM. 
S AM \ oi CSIKO collections, they examined. The 
specimens Ihey examined from Woolwonua and 
Darvsin in Ihe Northern Territory have been re- 
examined foi this study, and are all htlfiis, The 
Queensland material examined by Chabaud el til. 
(I0S0) could have been either /,. \v<nrim< t u>ni. /,, 
ltjf< , ti\, or both. 

Liustowiiwma inglisi iChabuud. Seuroau. 
Bcu-iidye. Bain & Durelle -Dcsset. 1980) 

comb, iiov 

(FIGS 37-47) 

Svnonyms; /u hiiti>nrtntt t ittctum \ensn [Rgfcfo, 1067: 
pp.122. L2Ki 131-133, lags B-JOi rchittonctnu ht$li$l 
Chtihaud, Seureau. Beveridgc. Btiffl & Duretie- 

Ocssei. IOSO: pp. 437-438: Spratl, Bcverulge & 
Waller. 1991; p 26- 

MtttfirM t'HtiitttiLil 

From isoorftvt nhr\nlu\ Western Australia: 4d ' : 
3 i -'. Murdoch. HI 57'& 115 51'FJ. 27.V.IVKI. 
AHC 800!; 40tfiJ, fW>9 *' ^>lh. (31 57\ 115 
51' Im Jan I90_\ y_j l09X 5iU003, I Mi A1IC 
W257. 3025X. TO25& 30200. 30262, 30263, 30264. 
30265; II ■ ! ■< . 44 ? S . Wattle Grove .32' 02'S. I 1(3 
(HJ'E), 5.V.I96& HM (NH» 1067. fi,l$-ti2& VVAM 25 
70, \il£t$\ IOV'1, Glen Koresl. I0.viii.60- AHC 
20760, MM (NH) ryff7, 473-52?? Wrf. lO'i " 
ho.csidalc. 15. til I07X. AHC K8K5; 40,/ J, 10 , : 
Jarrabdalc, (JIX 30'S. Ifi <)7'L», 2<Sa.I003, AHC 
102M: 4 J ■ '. Sv -t, Albany. (35 1 GO'S, 1 17 52'Ki, 
IRi«t(W4. I0.\i.|004. CSIUO N4242. N4243: 
Wrfrf, &ZS i . Maniimup. (35 I5'S. 116 WE). 
27 vi. 1003 lo.ivl 993, AHC 30255, 30256: Wo J ' , 
iSS'i j , no localitv given, 22a.I077. AHC NKKX 


Male: Leimth I I IN {15) mm. width 325-025 
1540). Cephalic bulb 2o6-325 (272* long by 247-350 
(309) wide; cephalic hooks, 1st row 140-1 NO i (551- 
2ml row 1 60-200 (]{H)), 3rd row 1 15-140 (125) lung 
(Figs 3, 7,i. Oesophagus 1 120-1565 (140(0 |0U£ 
cuheului dilation beanny 10M.2 rows body hooks. 
IX-irids 410-650 (520): scciciory-excretory [tone nol 
seen: nerve ring 312 <o-J) from anterior cud. 
Spicules similar, equal wilhoul alac 700-1200 (0S5) 
long, about 7,- body length. Guberriaculum short, 
sub- triangular. 50-5K(n=5) long il ; ie.45t. Nfine*pair3 
caudal papillae: 5 pairs ventral and immediately pre-. 
ad- and post cloucul. I Ugc pair lateral, ad-cloaeal. 
2 pairs laleral. pre-eloaeal; 3 pairs papillae, pair 
pliusmids well posterior m cloaca, near lad lip (l ; ig, 
47). Cloucal region with small culiculai bosses, ala- 
like extensions of body absent. Tail 150-350 (265) 
long (Fig. 46). 

Female: Leneth 15-30 (24) mm, widlh 500-0 <5 
(700). Cephalic bulb 312-357 (334i tpug by 29? 
(334) wide; cephalic hooks, 1st row 155-105 (18(1). 
2nd row 180-235(205). 3rd row 1 15-150 < 125) long. 
Oesophagus I385-IH3S (1655) long: euOeulat 
dilation bearing 12-1 I rows body hooky Den'uls 
4S5 570 (545): seeretoiy-e.ureiorv pore 3*.>0 (n=l) 
from anterior end; \mv\\: ring nol seen Vulva 9 K- 

10.2 mm (n=2) (Pig. 43) from anterioi end. Vagina 
about fell (n=I). Eggfc almost spherical 4tr4S (-1 I) 
|- t> Uv45 (40), Tail 6 UM>20 HM)) Innti (figs 42 44) 


Cephalic bulb with 3 rows of 14 (male) (Tigs 3«. 
30) oi 16 (lemale'i files ol' hooks 2nd n»w longest, 3rd 
n»w shortest; nccV With 5-11 rows liny spines 
uiticular dilation of oesophageal region bearing 10- 
14 rows. 14 (male) or 16 (female) files ot bpdj 
books. I si and last rows smallest. 7th - 0th rows 
hooks largest (Figs 3, 7); roots o\' hooks with 
undulaiine edges (biys 40. 41 1. Remainder ol body 
wilh up to 35 (male i. or 45 (female) small spines ul 
each annuiaiion, cvVcyf whole body of female, 
extending to 400 pm from tail tip on dorsal suilaee 
and about 30O-500 anterior to cloaca, nol extending 
10 anterior pair of caudal papillae, on ventral surlace 
Ol' male body ilig. 47), Oesophaeu^ '/ll (male) to 
l/l i (femjile) body lenyllr simple club-shaped. 
lerminatiuc level with Sth-Olh row of hotiks. Nerve 
ring surrounding oesophagus within cephalic bulb- 
seeretorv-evcrciory pore in neck- deirids conical, at 
level of Isi row of hod y hooks. 

h'('t j hits/ 
IxtHhli'ti clnsHhis (Shaw. 17971 

i'v/w loidltly 

Wattle Cirove. near Penh. Wesieiu Australia 

Site m host 
Small mtesiine 

7'v/'^ specimen 
NeolypeBM(NH) 1067 616 

Ui murks 

Materia! from /. obi,\nitt\ fomi Waitle Grove. 
collected on 5.V.I066 and identified by Inglis is 

FlgS 37 -47. hnMowhwnxi tytfUi (C'hah;unL .Scureau Bcvetiilge. Bain & Uurtilic-Dcssei, IVMO). 37. Ank'r.oi cud. (Imen.l 
-Vjpw), IK ( cptulie c-nd {Hi tint' view I. V). Cephalic eml male, optical section at level (*rfi«1 rdW (if books C*H fan \k\\ ) 
4(1, Body hooK (dOi^ul view). 41 Body htifii* (laleral vieu), J>2. Female l;iil ilaleral view). 13. Vauiiui (blend vicV/1. W 
(•finale tail lip (lateral V iew). 45. CJubeinaculuiu (venlnd view) 46. M.ile tail I lateral view I 47. Mills tail iveulnd \ .ev, i 
Scute burs - 200 uni 57. 42: 50 uni ^S, J9, 43, 44, KMl pTft 40. 41 . M>. 47: 25 urn 45. 


LB smai rs 

deposited m llic WAM and BM (Ml). This Uppfiarti 
to he the RfUteria] described iis /;. tiih imu by Inglfs 

( I*5ft7) I therefore designate n specimen from BM 
(Mil l%7. 1 <oo20 as the uc< My pe 

l_!ti\ii>\\>iiu-tmt tit^lisi (Chubaud. Seurcau. 
Hrvi.-inl'jc. Bam & DuretlL-Desset. |'->N<» can be 
distinguished r'rom /..'iti and L lawns in 
having ilk- oesophagus terminating within the 
anterior culieular dilation. lJnsu>\viiwma ///eV/w fifw 
10-12 (mule) and U 14 (Jemale) rows of body liookb 
rompared with 9-1 I -mm\ 11-13 rows lot /. 
wutritt^totu .m\ 8 II) and Ml? rows lot /. fcffw* 
The .spicules or L in^fisi (Vh body length) are 
relatively longer than lor /-. wvmu^fe/ti (•Ao hody 
Icilgth) and /. faiftrwtVi , body length), The male lail 
ol L /^e/'w'ako differs Iohii bOtliXf. wnniitylntn nn<t 

/ fo/ivM m being shorter tfi) %5fJ (365) eompuvd 

Willi 331M30 fmW and 250-341) t M)7) respectively 
and in having only 3 pairs of papillae rather Mum I 
Ihr letnale tail is also shorler (OI0-92O (7300 in 
f.. //^//.W compared Willi / 1 iooW>;e/o///and /.. hums 
i UMn-1900 * r2fi5li and 7'WA- 1615 (I070,i. 


The pair nl lateral papillae level with t|v Clofl&tl 
Opcrnitg is more prominent than the other pairs ol' 
pupil tap surroinulino .)„d anterior lo the dbttfti of / 

id^IIm In (hi-, respect, i tnvjist resembles / 
tihuoHtlsi hum dasyurid marsupials hut /.. in^h\t 
dillers horn L vihiumrtsi m havint: the 2\ui HOI flic 
1st onv of cephalic hooLs ihe longest. l.mMoutnctha 
ttii8Qfh$x\ ftCtiUh in V&syumx Imiitn uw\ horn die 
Northern lerii[i>r\ while L in\>li\i occurs in / 
>>ht'utht,\ honi southern Western Australia. 

/ hiui'witHtttu toijifhi can be distinguished 1'iom /, 
vim tutu, which also occurs in 7. 0/frfti/iM am! ha.-, the 
oesophagus terminating witlim the culicular dilahon 
ol ilu oesophageal region ill about the 9th row ol 
hooks, by the number ot mws uJ body hewks, 10-12 
(male) and 12-14 iTcmalc) compared wuh 13-10 and 
N-Ifi in /.- (inviHin, The differences between /.. 
i iiu ■•mat and A. m#fisi are disc lused m detail iniUei /. 

I 'he nie.tsuicmeiUs o| /,, Ui$Hti Irom this study are 
.oM-iucni unh ihose yiven by lujjlis t,l%7r Any 
dilTcienees belween the two sets ol mcasutomenls 
art- because lnj^bs < l%7> measured smaller worms. 
1 2')- 1 I 59 for the males compared with I I- IS in this 
study and 10.I-IX.4 lor die It-male*-. compared with 
IS-3JJ in this study. fhflbftUd et ul. < 1*980) described 

/_. irjtfiiAi males as having spines coverin-: vmiK -h ol 
the body dorsally. A vaielul c\ammalion ^\' 
specimens tor this study, however, has shown dial llic 
dorvd spines, although liny, extend alnng about ^l, : 
ul body, that is. luither towards the tail ftp than do 
Ihe venital spines. 

l.iiish>\vin<niii m\<li,\i has been lound only in ,' 
o/vWro I'lorn ihe south orWeslern Australia. 

hinxtowinema tasmaniensv spjiov 


Synonyms, hchintrnvma li/nimn wnsu \1unda> ..V 
Grten, \ K )12- p. in mi parti. /', hinoDvma ti$\m 
XHMU Spiatt. Hcveiidge is. Walfi-i. \W\, p. 20 (in 
pad). EehUumvMd *R.I C'habaud. Seure.iu. 
Beveridt;e, Bam & Durelle-Desset l ( )H0, p. 4% 
Spoilt. Beverid^c & Waller. 1901; p. 26. 

Mltlt'ttilf VMtlHIIh'tl 

7V/V ituiler'uti: Ifolotypr .;. alkWype .', Iiom 
ttoHiiw nhtMiin\. Kingston I 12 v>*s, 147" IXT-l. l|,vii.l»K)i. AIIC: 3lGB(l JM32I 
Paral.vpe.s 12 ' r , II v ".AIIC WI0, JWI | 

(>///( r nnnvritii t-rom \\iunhn <fhvsn(it,\ South 
Australia. Kan-aroo Island: 12 ftf^ 4 j 1 "» , Vooiuic 
Rtt) 135- ^)'S. 137 LVP), no date. I.\.l%8. AM<* 
44.SS. MIM)2: 13^'. 24^9.. Hundred of (loss,- 
Tune IW. Mir 3(1305: S^.:. f 7 V V . Karalta. | - 
M'S, I3|f TO^E-l, Sept, l'>.x\ :\nc 3030R; 13 - 
15 . T Parudaua. 1 35 47'S, 137* \<Y\\l till 1991. 
&A1 "9X0. MIC 3tr3(Hl, 3(1307; I'. I 1 - 1 . Seal Hav. 
m 00'S. 137 20'l:r 4.r!')K7; AHC 30304 
Tasmania; Ji .' 155) \ Beaconvtiehl. (II I2'S 
l-lo 4<T|-), loop aik j^jj; ,,:,.;_ Otagftrn 
'II 21'S, 140' 5,2'H. 1902, AIIC 3RWM I 1 
Hoba.t. (42 53'S, 147 HTR), 25 vi.I9H2. CSIUiJ 
NI674: 7 :. X , ', Co^ Kainie. |42 J ' 59'S, I i/ 
IS'b), IS .vn, 1992. AHC WHOft; S' rf, 13' ,' . 
MaojatL- tip. (43 1 02'S. 147 I0T;|. Ili.iii. to<>3. AHC 
303 I ?; I , , . Upper Dromedary- no Jalr ^ivcn. AHC 
4530; 12 : I'C ■ ■, "', no collecOon data. AIIC 


Cephalic bulh with 3 rows nf 14 (malei or 10 
(female Mh^.s 49. 50) files ol hooks. 2n.l row lon-jesl. 
ird row shorlcsl (Fig. 49); iu-.a. with 5 8 row,- Ouy 

tit's 1H-OI. Uitkttiwttimtl limuilikmi s, v no*. 4K. Ankn.*. cutl. UplSCtll seufkn (fctkCPal mcwl 19. CfephiTltC hulhOaieoU 
view t, 50. ( cptiatic end reniiite. optical section Ut level ffl liisi row ol hi mUio/ /;<,<■ viiwi. s| t-Vniali: Unl ('literal viou }, 
52. lyfyjc puskri-H Nuly s)Milfv isciiiMl vit-wi 5T rtoUy hooKs (lukml view! s4, tiiuly h.,,,^ view). Sfl Vagina 
diikoil mlu) so, Male tail ( \ icw i. 57. Curving nlmn (wiiiial viov,i *>*. Prinale lail tip (laioral \ ieu ). 59 MjiIc 
mil li|>(\enlnil view) 00. Mak uul lip <\eiitr;d viewr ftl Utllvrnacliliun lL.teo.1 vil-ui. Sr ;( K h..o ^ 500^1 4S s l 200 
Jan.!". oo, IOO M im.'>0.50hiu5: st. J4, s 5 . 5h- : 2S ,. m S7.5S. "9; 1 2 p.» 0! 




•vpotcs; eulieular dilation of oesophageal region 
bearing 13 15 rows fif 14 (mule) or 16 (female) Hies 
til body hooks Isl and last rows smallest. l >th MMh 
rows largest, mots of hooks vviihout undulating 
fijges (Tigs 3i- $41; tvmumdci of body with up (o 4*J 
<uuUj or 66 (female J small spines at each 
.tMuiil;iti"n, over whole hotly of female, extending 
itvtl ft tiF dorsal surface, terminating about 350 
-iim-i mi to cloaca, not reaching level of most tintprioi 
pan ol caudal papillae, on ventral surface of male 
bflljry (fig. 56). Oesophagus aboul V. '/ i body 
It'ftglh, supple, club-shaped, (eiiinuutmg at level a! 
Nlh-IUlh row of bad) hooks (Fig. 48) Nerve ring 
sin rounding oesophagus within icphalk bulb, 
seen lory-cvcu'iory pore in neck: cleinds conical. at 
li-\ci of I'-l tow of biyiv hooks. 

Male. Length II 14 H2.6) mm. width -155-715 
fffif I Cephalic bulb : 15 2 ( >5 (255) [ring by 270-3 l ><> 
(325) wide: bearing 3 rows ol hooks. 1st row 140- 
ls>S { 170), 2nd row~l75-225 (200>, 3rd row 105-131! 
(1201 lOflg (fiu 1°) Oesophagus I I0.v I5N0M ViO) 
long eulieular dilation hearing I2-I '< rows o| hods 
hooks; deirids £>5-4R0 (430). secretory-cwreiorv 
pore MHM40 i365i i.n=6i; nerve ring 240 300 (iSflt 
|Q=4] ftOltl anterior cud. Spicules similar, equal. 
widvnU alac 720-1030 (840.) long, about Vll botlj 
length. tjuberuacuhtm short, simple, sublnungulat, 
50-55 ia=.Vi Inng (Ffgsh 57. 61). Ten pairs Caudal 
papillae: 3 pans ventral and immediately pre .. ad 
.ind post-cloaeal respectively, 1 pair lateral ad 
i h'aial. 2 pans fUcinl pie cI«kic:iI (] rg "' f ')- 3 lateral 
pairs larger (fig 56); 3 pads papillae, pan phastnuT. 
posterior to cloaca, near lad tip tFig. 59J. Cloacal 
region with small eulieular bosses; ala-like 
expansion of body surrounding cloaca, Tail 235-325 
<2K5» longthg.b'o). 

female: Length 19-22 (21) mm, width 815-1155 
l%5). Cephalic bulb 1 95 -3 25 1 255 > Ions by 17(1-501 
(460) wide: bearing 3 rows of hooks, 1st row l l >0 
110 1305}, 2nd row 220 265 (245). 3rd row 125 ! 70 
»,1S0) long. Oesophagus I445»r99fl M6I.M Itijjg; 
culicukii dilution hearing 14-15 rows body hook- 
I Voids 355-450 (405): >ccretory-e\crelory pore 375 
tn^l ) from anterior ciul: nerve ring not seen. Vulvn 
5525-7600 (650O) .rr=3i lnrtg (Kg. 55/. Vagina 175 
in- It lone. Tail 7I5- C >35 (SI0) long (Tigs 51. 58). 
Cgg* ttlmOSt ^plKriod 36 4S (42) by 33-45 (3K). 


/ w.sttnvitn ttut f<L\t>iutut'ti\r sp. nov. resembles f.. 
iiighw and /,. cinttnin. also occurring in I. <>hr\nln\ 
and hav nig three pairs ol papillae on the male tad and 
the oesophagus terminating within the culiculat 
dilation of the oesophageal reeioi.. at about the Kth - 
I lib low of httoks. / histuwtHt »nt ki\uutt)itn\c can 
be dilfcreitlialed fp>m L. tnglhi in ba\uie 12-15 


(male) and 14-15 (female! rows of body hooks 
without undulating edges compaicd with 10-12 
(mule) and 12-M (female) rows ol body hooks with 
undulating edges and from /. vim Turn which has 1 3- 
16 (male) and 14 IX demalei iows '»l body hoi»ks 
with undulating edgus Unxtouint nm htxuitinittisr 
blither differs flout /., Utility in having all three patr- 
ol papillae lateral and anterior to the cloaca, largei 
than those surrounding the eloaut. ami Hi having the 
cloacal rcyion with ala-like extensions of the hlltlV 
Litisri'Wfin'md iijglfci has only ^\^ pair of l.ajii 
lateral ad- cloaial t^ajiillac aurl does imt ba^e the ala- 
like eMensions. Linsit>\\itt< tini Vtofltmh which does 
have ala-like extensions of the body, has ventral hody 
spines extending lo the level of the most anterior pair 
of lateral caudal papillae but L F i(t\t>t<>uic*i\r fJl)C* 
not.\tt>nirntntt n'/nlnm has all si\ pairs Cjl 
papillae lateral and anterior to the cloaca the saniv 
si/e but L tttwtuntcttse has die thiec lateral (»ans 
larger The gubernaLUlum i- more U-shaped in 
yclHl.i! view ol /, !u\>>i>t>net>\t th.u> in /. i'/i^mmh / 
iim UWit hemale L have a shorter lad 
TI5435 (XI0) than I i ntnntn 9S6 I 122 . 1050). The 
oesophagus is uhout *k body tenjjth in L. 
ftivnhtnii'nw males compared with '/x in L rim tutu 
and 1/11 iu L ittyitM I be vulva ts closo to the 
antetior end in L tits/nun ttti,s<\ abotil 0,6 mm 
compared with 1 .0 mm ill /„ itlfiifa and 1-2 mm in / 
chwtUTtl. The vagina of /.. utsnumiensc, 175. ts 
shorter (Iran in /.. t untittn 30(t and L itt^fisi _vS(f 

ChtibtUd t'( dL (l u ^«») figured the tail of $ maK 
specimen, registered as AIIC* 4530, fiom /, t>hi\nht\, 
Upper Dromedary. Tasmania. There is now only one 
damaged male specimen in die bottle available for 
comparison, but such characters a.*, can be -.een, jiuI 
tlk ilrawuigs of Cliahaud tf <tt, ( lUSUj (F,g. > l J . | 
p. 13S), are consistent with h. tit.stntttiu'ttsv, 

I iu\u>\\'tninut tasnmnu'nsc appears lo have :t 
^eoeraphic range which extends across Tasmania 
and Kaugarott IsIjikI. Three Imsts from Kangaroo 
Island were infected wilh two species ot 
Uttblnwtncnnt. one with L iuatmn .\\n\ i 
i(t.\t>ii(tu('ttsc, and two with /,. Witifin^cni and /.. 
tti\ntfttiirtt\c- This suggests past links between 
kangaroo Isluiul hosts, mainland Australian hosts 
and Tasmanian hosts. 


The species is named according to a label found in 
AHC IS20. "L. cutcnttn tns/nitnii'ti.sis", here 
determined to be /.. rititnwi* which had apparently 
been written by Chabaud ct a!, wlien pirparing their 
paper ol l l JH0. 

h'pt forulitv 
Kingston. Tasmania. Ausuulia 

type host 
Isoodon ohesultis (Shaw, 1797) 

Site in host 
Small intestine 

Tvpe specimens 

Holotype male, AHC 30320, all 
30321, paralypes AHC 30310 


Material examined 

otype female. AHC 

Liiistowinema peramelis sp. nov. 
(FIGS 62-66) 

Synonym: Echirumenut einetum senst 
Beveridee & Walters 1991: p. 22. 


lype material: Holotype 6 , AHC 30023, allotype 
?, AHC 30097, from Perameles bougainvillc. 

Other material: From Perameles boa\>ainv'dle 1 6 . 
I?. 1 anterior end, I posterior end. no collection 
data, AHC 30055. 30054. 30053: I 6\ I V. 3 anterior 
ends. 2 posterior ends, no collection data. AHC 
4522; 26 6. I?, captive. University Adelaide, no 
date, AHC 13928. 


Cephalic bulb with 3 rows of 15 (male) or 16 
(female) files of hooks. 2nd row longest, 3rd row 
much the smallest (Fig. 62): neck with about 6 rows 



Figs 62-66. Linstowiitenui pcrainctis sp. nov. fi2. Anterior end (lateral view). 63. Body hooks (lateral view), 64. Body 
hooks <dors;il view). 65. Female tail (lateral view). 66. Male tail (ventral view). Scale bars = 200 urn 62: 25 urn 63, 64: 
100 urn 65, 66, 



liny vpincs; cuticular dilation of oesophageal region 
bearing 10-12 rows, N Ouale) or 16 (female! files of 
body hooks, first and last rows smallest Blh-fitifl rows 
largest, roots uf honks wilhout undulating edges 
I tigs fi3. 64 f; remainder flr 1 body wilh a row o\' >.mall 
spines ill each cnricul;n .Uinulalinn ommbers uf 
spines nOt couidcd), over whole body ol leniale, 
LNlendnie over / ( dorsal surface, term innt tn;j aboul 
300 nnierinr 10 cloaca vcntrally. reaching level of 
rUUM anlerior pair ol caudal papillae, on ventral 
Mirtaee of male body |tfg T dh j OCSOphugUfi -ample 
club-shaped, ahi>ul '/n - '/q body length, terminating al 
1llh I2lh row 0( hooks (Fie. 62) Seuviorv- 
e\i letory pure, demds und nerve ring not seen 

Male: frt*?l Length 9, 9 nun, width 475. 475. 
(epii.ili. htlltl 23?, 525 l'>ne b\ 2U. 2<w wide, 
eephahe ho.4.;-, [*l io\\ |<23, I 2$ 2nd row I I 7 150. 
!nl row 75, \U lone OcwphttgUS 745, B7fl long, 
iiitieulai JiiaJUyi bearing II 12 rows body hooks. 
Spicules similar, equal, without abic. 050, 1050 |00g, 
..hoiil 7o boU-V IfthgHl (mheiuueulum short, simple. 
subhiauyuhu. I I pairs caudal papillae: J pair, 
ventral arid immediately pre-, ad- and port-* loacal 
li'speciively, I pan lateral ad-cloacak 3 pairs lateral 
pre cloaeal; 7 anterior pairs all nhoul same si/c* 3 
Paul's papillae, pair phasmids a ell pOfitOflui i" filoHA a 
ueai lad lip (Tie. 60), Cloaeal region wild small 
• •Mlit ular bosses ala like expansion absent lail 21JK 

Mil Fung. 

I runic: Oi-2) Length I I mm width 520, Cephalic 
hook* Isi ,ow< 2(.0, I 15. 2ftd row |4tt 2fr0, W row 
78, 140. Oesophagus | 190. 1250 lonp; cuticular 
dilation beating 10 12 rows body hooks, lail 400 
ItHIg (}t^ fifi. Vulva uoi Ktitfi. hygs moir oi less 
spherical 51-57 by 60-03 


Mlhougli there was only u small uiitnbei ol 
specimens and they were all tn poor condition, 
sieuihcunt differences beiweeii these specimens and 
other specie 1 , ol' LiitMttwitn-Dhi could ho found 
Although elements such as thr vulva uOflltl uol be 
seen, the worms appeared to he mature. ha\me 
lertili/ed eggs /// tin -n: Since P. htiUgiim\ l i}t? is now 
extinct on mainland Auslralia and fully protected on 
lhe islands hi Shalk Bay Western Auslralia. it is 
unlikely thai any more specimens will heeome 
available Accordingly, this somewhat incomplete 
description is presented stf the best possible ft] lhe 
specie -■ under the circumstances. I.insh^vhwntu 
p<r>int>t?\ sp, now most closely resembles /_ 
U'lJJ rinxiumi .\\\i\ lhe species |hal have been 
distinguished in the discussion ol / MiMtHgUmi, in 
i!i. n there arc no more than 1142 (male) and lo 12 
ilcotalet rows ot hodv hooks compared wilh l )-l I 
uu.tlt ■) and II M (female) and (lie oesophagus [\ 

retahvely lone. c\lcndiue lo the 1 Ith or I 2th row ot 
body hooks. The two species can be dillerentiaied b\ 
si/e. I tnsiowhwhitl Wdrriii'^ioni males, 15 20 mm 
and females. 2> 2-4 1 mm, aie much larger worms than 
/. (nr<att<hs. y and II mm, respectively. Lin 
\/tn\int'i/ui pvntmvlis has plain edge* on die roots of 
lis Imdy hooks hut L. w/ar+ingMm has undukanie 
edges. The numbers and uirangemcnl ol' papillae f,iri 
the posterior end ol the nrale are also dif'teivm / ith 
\htui>nnt<t iHnnmiis is the only species o\ 
.st<nvin<<tn<t wilh lout pairs of papillae lateral anil 
anlerior to lhe eloaca; all other species have three. 
The spicules ol /.. /VTi////W/v 1950-1050, Q*\ h. 4 . 
leuiHhj are relatively longer ihati those *>| /,. 
>i>t<ntit;l<>nt <<VMI I0"0. l/:o body len^lhi, Lino,* 
wmciini f>cr,i>ttfli\ has Ihree pails oi' papillae ileal the 
caudal tip hut L ivttmnxto/ii has bve. The lad oil 
pcnrmpHx '20). 240imalei. IhOileinaleo is shorier 
iImii (lull of /- ItHtWitlSWM (3 '0-420 aixl I0*10 
l*«H)). lheeggs«)i;. ( /'<'/ f ^/r//M.5|-5:' H WlWH 

larger than l hose ol L wvrriii\itotti \M>44 h> .M-4XJ, 

t-ii'sttnwwwa iwruinchs is similar tot. itt$l(si arid 
L ant turn- in havine three puJpH of caudal pupdkiL. 
It can be diO'erentiaied tfom thes:_- u\o fl?CCT i I 
haviiu 1 ;» ii'lativilv hnig oesophagus, letirniiiaue .ii 
lhe level ol lhe posterior rows ol hoof.-, on tin 
eutieular dilation of the oesophageal tegion. and nat 
within it. and m having up in 12 row, 1 - ol body books 
without undulatine edge,, compared woo / oivtt.u. 
Lip Ifl 14, Ulld I I "H'tUHl, up lo \>. iovvs ol bo4> 
hooks vsith undulahng edges 

f.tiiMnw invnta pcutmclis resembles L. lUMltttflieitXi 
in liavinj.- four pairs ol caudal papillae near the 
CilUdu] tip and hody Imoks withoui uinlukiling idees, 
hill ddt'cis I'lom /., fusntiittu -tiM' in ha\ mi' lhe 
oesophagus lernunating at lhe end ol lhe QLiiTmihll 
dilation and not within it. up to 12 tows ul btujy 
hooks rather than 1 5 and largcrepgs. 5 I -57 by nO-M. 
compared with 30-4.N by 34-45. 

Although lhe thiol un^ ol irphalie liouks is 
tLkiiively much smallei ihait the tlrst mid second 
onv, hi /., pvnmu-Hs than in oihei species of 
{jfiMoH'ifwtna. the cephalic hook sizes vary urc.idy 
between inilividuaK uttd so this may not he a 
consistent character. 

Ihc eight P- ht>ni>tjiitviil<_'. dissectetl tii pro\itle / 
penntu'lh for this study tilnve ot winch were 
Milreted), ^eie nil revistercU in lhe SAV1A m |^3d. 
but no other collection data were given in the 
museum register. The collection data lor AIK A 
and I302M are ec|ually sparse, the information lilt the 
kibels giving only the locality as possibly Soulh 
Auslralia and captive m lhe /oology Department i if 
the kaiiversity of Adelaide, rvaminaiion of (he AkK 
remrds indie ales that live additional huntflCWH.t'Wt$(r 
dissected loi helmuiths. Ivvit ok which wcte infected 
with / jh'ntmt-liK This suggests that the prevalence 


of in lection in p Inm^invitlv bV £. fn'iwnvits was 
about UNIt 

The specilic name ts taken from Lhc label of AHC 
45 22. (he mulci ial originally registered as 
i l i-hmoiwnni rim Uuti penmules, hen: determined to 
lie I fwrumrhs- 

ixfir lot tilify 
\ luknown, Australia 

lypi' ///as/ 
l J cnmwtcs bou^atmilic Quoy $£ Guimard, 1JS24 

Swp /'/ //ovt 
Small intestine 

'/v/w .sprcntu'ii* 

' Holotvpe male, AHC 3C0T123. allotype remale. fcltfc 

Lhistitwhumo mufftcMenti ftp tmv. 

Mdfrnttl <-\ttminrd 

/v/'< malrrutf: llolotype J* allotype »' . QoUl 
f'ri,htuic\ miMfia Dinner Creek. <I7 L 2ft S, 146 
OO'L), Queensland. I I v.60. AHC 30()o4. MHm. 
Kuaivpes4 r3 dr\ I t ' . 2 anterior ends. 1 ,? posterior 
cudQM 14363/L AHC K>763 

ft/A(? fttdttiittf: Loan I'mrnclc* tiit.wila 
Queensland; 2 V S 1 , no locality given. 5.i\.57, QM Gl. 

14457. I,:;.:;. 1 , i j fVagjrtont [musSfUlT 32 r s. 

146 Ot'K). I5.\ii.5 ( ). QM GL 14356: 2JJ f 3 ij 
southern Queensland no dale. AliC 1726 

I i'uiii lso<klott mu<n>uru\ Queensland; 2 Jl 
Mossuian, (16 2X'S, 142 " 23'K).»5,X, QM (31- 
14303/2: 2 *e3. M - Innisl'ail. (17 U'S, 146 
01 I:). 1726: ! J . BnSnaue. < 27 2S'S, 
151 01 4:>. no dak-, AHC |7J$;5<3 5.3?. 1 u.Her.o, 
end, I'addingto.i, (27 2S'S. i 5.V Ol'fchAug. FV55. 
A I IC 4371: from bandicoot, no collection data, 3 ( ' i , 
A IK- 10667. 

Drst ijptum 

Cephalic bulb with three rows of 14 uuale) ffjy 
60) t >r 16 (female) files of fijjgfi fiOQJtS-, 2nd row 

largest 3ul low smallest (pig, 6N): neek with 5-M rows 
ol liny spines: cutieular dilation ol oesophageal 
ivgion bearing 1 1-13 row sol' 14 ( male i or 16 (female) 
Hies of body hooks, first w\\A last 2 lows smallest. 7th 
- °th tows largesl: roots of hooks without undulations 
( bigs 72- 73). remainder of body with a row ol' n\\ to 
42 tniule) or 50 (female) small spines at each 
annulaliiin (l-iu. 7 1 ). over whole body of female. 

extending over 7- dorsal surlace. lerininaimg ahoui 
4(H) anterior to cloaca on ventral sutface Of male 
body. Oesophagus simple, club shaped aboul Vu-7 i 
body length terminal ing posterior to oesophageal 
eiilieular dilation (Tig. 67), Nerve ring surrounding 
oesophagus within cephalic bulb; secretory excretory 
pore in neck, deiritis conical, at level of Kt row ot 
body hooks. 

Male. Length 1:^23 (17.5) mm. width 475-560 
(550). Cephalic bulb 285350 (335) long by 260 310 
(200) wide; cephalic hooks Isl row 117-150 (130), 
2nd row 140 175 ( 160). 3rd row sMJ-MO (120) long. 
Oesophagus 1360 2210 ( |*0 u .l long, cutieular 
dilation hearing 1113 rows hods hooks. Deiruls I0> 
530 (475). neive ring 255 300 (2N5). secretory- 
c.xercloiy pore 325-405 t360) from anterior c\\<\ t 
Spicules equal, similar, w ilhoul alac l >20- 1 100 ( 1030) 
long, aboul 7 ft body length. I I pairs caudal papillae. 
3 pairs ventral and immediately pre-, ad- and posb 
cloaeul respeclivi'ly. I large pair laleml ad-cloaca), 2 
pairs lateral pre-cloaeuL 4 pans papillae, pair 
|ibasinids well posterior to cloaca, near tail lip fPiga 
74, 75), Cloacal rcpiou with small cuitcular bosses, 
ala-like expansions absent. Ciubernaculum short, 
simple, sub triangular in ventral \icw, 60(n=l) long 
<hg. 76). Tail 305-440 t?75) long tl : ig. IX). 

beniale, Length 1S-35 (28) mm, width 560-765 
(670). CcphaliL bulb 325 425 (362 1 l.xie by 274-435 
(310) wide: cephalic hooks 1st uav 150-176 (I60i. 
2nd row 170-225 1 1 «■>()». 3rd row 1 15 145 (125) louy, 
t>e>ophiigus 16(5-^740 (2272) |oug; cutieular 
'Jilaliou bearing 1 2-13 rows body hooks. Oeirids 405- 
W? (4 c )5t. nerve ring 260-390 (330), sccreioiy- 
o.cK'tory pore (n=2) 405. 470 from anterior end. 
Vulva not seen. Tail 565-970 (790i. gggi! noi 


(.ittshiwiiicnui ntapirsionri sp. nov. resembles / 
U'tirri)i\>titm"\n having up |o 13 towsuf boil} lii>oK^-. .; 
relalively hmg oesophagus and five pairs of cauJal 
papillae on the male, fl tliffers from L, wFlfflhffilNi in 
having 11-13 (male) and 12-13 (female) compaivd 
with 9 1 I (male) and I 1-13 ( female i rows of body 
lu-oks. The oesophagus ot i maplcsloiwi, extends 
beyond the cutieular dilution of the oesophageal 
region of the body vvheicas m /, luimnyloni it 
tcrmiuales yi uV level o\' the 9lli 13th rows of body 
hooks. The body hooks of L nniplt-sniHt-t are withoui 
undulating edges uml ihosc iA' L WQrritifiUJni aiv with 
undulating edges. Male body spines extend along Vi 
of the body dorsally and well above (he lateral caudal 
papillae venirally on /.. i)t<iplc\t<>n>'t but on L, 
^ittrmxtoitt thev extend aU»ng 'A ol the body dor.sally 
and almost to lhc lateral caudal papillae venirally. All 
6 pans o( pre-, post- and ad -cloacal papillae are the 





i 7 



















same si/f on /.. wQfrbtgtoiti bm f. tiKtpf&tfwi i»as 

larger lateral ad-tloaeal papillae. The spicule^ "I /. 
mapU\ionrt (l/i,, hotly length) arc relatively longer 
(Jiaii those ol Z_. wnmn^umi l '/_><> body length). The 
arrangement ( ( >i the papillae surrounding and anterior 
I'O I lie cloaca (it L mafth. sfenct lt\uSt closely 
resembles lliat lor /- itiylisi i.e., wilh Ihe lateral ad- 
eloaeal paii the largest and the ventral body spines not 
extending to the level of the caudal papillae. 
LtHslnwUti ota iH(ijih\tr/nit differs from /_. int> in 
having (he oesophagus extend posterior to the body 
hooks without undulating edges rather than an 
oesophagus which terminates within the utlieuluT 
dilation ai about Hie level of row.-, SO of die body 
hooks with undulating edges Male 1, itii>h\i have 
uiily three pairs ol papillae on the eaudal lip but /.. 
tn<if>!cs(<ntci have lour. Male U WNfflvMNMii o\\ 
averaec have longer (ails * ^05 440 iMy\ titan L 
i>i. : hst ilSO-.VSl) (2t>5»,i l Ihe body spines ol L 
ma/'h stottvi extend ftlOllg 7- of ihe hodv doi'sally 
compared with Ihe whole body ("At) on /.. 

I -insfownn-ntd /m//'/ev7o//W ean be distinguished 
tVtim A. hifi'iis. whieh also lta.s the oesophagus 
extending posterior 10 'he body hooks, body hooks 
with undulating edges and tout pairs of papillae on 
die eaudal lip. in having II 1.1 (male) anJ I- I 5 
Hemalei mws of hooks t:oril£itlt£d wuh X-10 (mule) 
and o 12 I female l rows ol" body hooks in /. hitetis. 
Male A. tnaj»tc\ioiu< have body spines cMendum 
alone V, of the body dotsally and well above die 
lateral eaudal papillae veutially compared with / 
hitats. whieh has body spines eMendme alone ' 'An 'I 
the bodv densallv Jnd level wilh the mosl anienoi 
lateral eaudal papillae vvtnTally. Ihe lateral a<l- 
eloaeal pair ot papillae is largest on /.. imtpU'.sioiu / 
compared widi all 1 pairs ol lateral eloaeal papillae 
heme the same sixe on /.. lultus feniaf- /. 
nuifilisttwt i have on average shorin tails iS65-O70\fi'\)'inct/m nh<i>U'Untw< ean be ditleroiitialod 
[him / run mm in having ||-M (male;. 12-13 
thaaak) bods hunks wirhoiu undulating edges 
eompaied walh 1316 (lualet I M.S ifcmalc) rip*ly 
hooks wilh undulating edges, a long oesophagus 
terminating posterior to ihe rows of body hoofs, noi 
a short oesophagus IciuunaluiL* a.\ Ihe level i)| ihe & 
I I rows of hooks. t.inMtwnutiid tfiaptt itomi hfl* lour 
P-ti's ol eaudal papillae at the tail lip. while I.. 
.iiiiimn has Itttec; ihe pair of lateral ad-ehuieal 

papillae is ihe largest on /. niupk'sfo/H'i but all six 
pan, (.1 iloacal papillae are the same si/e on / 
iinciitnt Unslin\itictn(i Htui>lt'slonci does not haw an 
ala like expansion surrounding the eloaea but /. 
ciih-hmi does; the vential body spines of /_,. vUwmm 
extend almost to the level of the most anietioi laicral 
pan ol pre-eloaeal papillae lull 1)1 L . >/M/'Av'"<"f they 
do not the female (ail of L. mupUsioiui |5(i5-V>70 
(7'JUji is shonet limn thai of /,. tinrm/ti ( v >Mf>! 1 22 

Tlu- itihn --.pities ni Lmsii.'Wunnui. I., /wnmu'li* 
ami /.. !<i\ithtnietiM\ wlueh have bndy hoofs without 
miilidaiiiig etlges have three pairs of eaudal papillae 
on the tail lip. These ean lie further differentiated 
I'ruin /_. hui{>ivsinm'i bv the number or rows of body 
hooks and Ihe rvlative lenejlis <»l the ^esophagus. 
lAtwtitwmciihi pcrami'li* has 10-12 rows ol bod> 
hooks with the oesophagus terminating ril the level u\~ 
ihe I2ih row: /.. tawttftoh'ttUP Has 134 5 mvvs ol hod> 
hooks wilh the oesophagus termiuaiing al ihe level ol 
Ihe *lh 10th row and L ffiaphfiWn£l lias I Mi rows 
\t\ body hooks with the oesophagus Lermuuiling 
posteriorly to die t3Ui low. UttMftwfticmti fn-nitttrh's 
has four pairs wf laleral ad- and pre eloaeal papillae, 
eompared with Ihree pairs on /_. iHtJpfaittotbtt ami 
relati ively shorter spicules (7- of body length I 
eompared with i'/i4, UtiMituinL'ina )uxUuniU*nw has 
three large pairs of lateral eloaeal papillae eompared 
with the one large ad-cloaeal pair ol /.. nuii'lcsiotur 

fieeaUse of the small nnniber o\' female speeimen> 
available, none was dissected, so allhoueh eggs vvete 
seen, it was diffieuh to determine which were mature 
and ihetvfoiv suitable for measuring- As t\ result, no 
ineasuremenis were tnade. 

Ltt)\i<i\i'fni-itui un.ipliKtnnt'i oeeurs in /* ihltyiW aue 
/, t]HH-r<>tint\ troui Queensland aue! New Suinh VVakv 
aldiough llrt evammalion of more hosts is require. 
\-)ciori: I ho lull extwii ni die gcngraphie range wl ihi> 
species ean be dciennined. hi two / rthinvtitn>.. I. 
>>hii>h\h>iti'i oeetined til mixed loieehotis with /.. 
*e f ./r/7//.t;/'"/'. 


The speLiL-s is named alier i\.V Maplestone. who 
together with VV. Yorke. eameil out pionceon;j woik 
on the nematodes ol Australian marsupials. 
hiluH"ticf>io nUfpltftitWi Wto IWCd on ;in undate.l 
niuseiim kan-l by (habaud c -7 at. I l*'N0) loi 
(JnhiiMiwmti speumeii'n Irom/. mm rt*utv\ thai were 
subsequently determined to he L. wtttnttytmt 

| <" 1)7 -■> / '/Mf»>u ith'nhi ttutf>tf'\ti>nii ^p. n<>v_ Wl'. XntLT-or end. i»|'Ik;iI stvlitni ( IdliJftil \ \£\\ I. OS < k Mill, i l.iH i.e 
vti-wi fi4. (Vpli;.ta i_nil male- opticul seetiitn ui W-\\i\ of In* a .ow <il tituik^ \tti Jm i 1 view r 70 renuk- lad il.iu ml v ti P | 
/I Mate, |M>Mi-iiiM luniy spines vit-w). 7^, \Uk\) lun«K (klkivil v aew '). I \ h">J> \»<>>k idilSSllJ VftfW) "4, Male Uul 
up * vii.iiil | hw. /-> Mak tail iveniial view) U>. * miIvouiluIuih ivenlial viewr 77. Iviuule Uiil tip i lateral view > 7K. 
\1 uk ■ t ui an.,'ii Mivvt. Se'iieHars-S(KtpnK>7: 2uU urn mK. 7i>. 50 um oQ. 75. 11. 25 \tW 71 72.73 7-1. 7(v. |4H)|iin 7S 



/V/n fey 

YY/T li>f(t!l(Y 

Dinnd : Creek (17' 2<VS i4(> <H)T) Queensland. 

>//e rfl rtcutf 

Small inlestine 



I.insh'Hinnttii svan'in^loni appears lo DC the 
dominant species of IjHStowiiittiut in eastern 
Australian bandicoots, heiny found from northern 
(JticensUmd through It) .Souih Australia, including 
Kangaroo Island. It occurs in all extant species of 
lUfWiltm as well as one individual ol /*. Wxttkt. 
indicating n low prevalence in this lalter hosl species, 
Although L. lawns was found in iTorffacTfl 
Ouccnsland populations of the northern brown 

bandicoot together wiflj I WiirvpigtMii, H was the 

only species occurring in norlhein brown bandicoots 
toiiii die Northern Territory And ihc norlh of WesLern 
Ausiialia. while L W/jj/tof was die only specie-, 
occurring in southern brown bandicoot* in die souih 
of Western Ausiialia. 

Only one species, L (nratticli.s. occurring in five ol 
|d pi hnii\>ufitvi!U' examined, has been found 
rvUusivcly in l J <-nwivlv\ spp. and three species / 
t<i\wat!i(>n\r, /.. Uncus and /.. inxlisi exclusively in 
lKnir<l<ni spp. / hitt'Hs occurs in /. huiiri>nn*s\ L and /., ttlfilhi 0CCU1 in t "/'< .w///a. Of 
I he cHher Ihree .species /., ttiuf)U>\tt>tu>i has a north- 
eastern disiribulion occurring in / nun ramus iind P 
mmnu I. uinnmt a metre soudi-easiern disiribulion 
occurring in /. -ihrsnlus. P. ?ut\ittit ami /' xntnut. 

further collections of material from south-eastern 
Ausiialia are needled before ai'iy hypothesis uu die 
distribution of species or tiasanvaitnta can be 
developed. Ii does appear thai bandicoots 1/ 
ahesulnw till Kangaroo Island, bur not Tasmania, 
may have been derived from Mock in which all three 
species </. wat i it)\>fi'))i. L ciinntnh aanl P 
fastaaaunsct were prevalent. I he emit inn my 
deli i menial cTlects id l-auopean sell ten iciil have 
resulted in a patchy distribution or" / "in'Mtfus ovei a 
reduced range (ttraiihwuite 19^5 1 This decline in the 
hosl population m;iv have affected the distribution 
and prevalence of species ol Unstomacma on the 
mainland The cm rem prevalence or The three speeies 
id'/ ,n\!»wwrttht in bandicoots on Kangaroo Island 

may reflect past prevalences of ihese species ii] 
bandieools on ihe mainland. 

The ability to trap bandicoots varies with Specie* 
(heir age. species and locality (Gordon & Hulbeit 
l ( )S9). Pcramcltw fU/Mttu is apparently more difficult 
to trap lhan species ol Isaadan (MenkhorM & 
Sccbeek I9A>5). This may be the reason forihe small 
number of Petfwte/iii compared with foyadtm 
collected in this study (see Table I t and in Ihe 
anionut of material deposited in museum collections 
from each hosl eenus A furihcr complicating faetoi 
IH ihe possihlo difference in prevalence ol infection 
with l.ntMouitiema between Ihe Ivvo. The iccords ol 
the S\\\A and QM indicate that v>0 P. ansuia have 
been examined far helminth parasites and. of these. 
only ci^hl were inleeled with /.. i/n//>lfMa/icL m\ 
with L ittHtitm and one with /. warrin^aan- 
Similarly, o| 51 / J yjmnii examined, otdy tour w-ete 
infected wilh L- iimutm and one w-ilh f 
WWWfnglctni. Ii is unlikely thai Linshn\iiic/na could 
have been overlooked duiine dissection as the worn in 
ate huui and cm be icndilv delccled in the small 
inlesline. These low prevalences ol inlcetion contract 
with the prevalences found for species ol /.u-iWc-i 
JissLcied in this study. Of 12 bandicoots examined 
S3 were infected wilh LinslowiHenni spp. 

A working hypnihe.-i\ would be that species of 
l.iust<n\tnvtmt are dominant in the helminth 
communities of fsmnJfW but not in those ui 
fr-ruHiclcs In some ufifiSfv spgpjes of bandicoot havi' 
oveilappinc gcogrjpljic ranges, / nun t\*mti\ and /' 
m/\Hlt> in the iu>rtb east, /. <>hcxii}U\ and I* thnttftt in 
the soulheasi and t obtst<fn.\ and ft xttnifii hi 
lasmania. Their habilal preferences w ilhm each 
^eoyrapbic region are different, and althotigli thev 
may not be in strict synipairy. opportunities ha 
incidcmal infection and host switching would c\isi. 
( Jhsrrvutions tiom ibis study su^^est that (pCtflCti Ol' 
tjn&UM\'fiie>wci may switch from /. tttu-Mihis tu V 
t>ntttui and r ntmtnr, front / hiwiilm to /' 
hi>t{umt)\Mf and perhaps front /' mtsithi to / 
mat -tintiv\. tins nuyln aceount for ilie otrunence \)i 
/ tnunwith'tna 111 Pevunu'lcs. 

Addiiioiuil collections of material fo>m ftfttftylti 
spp. actoss Australia and especially /• <>hc.\alu\ Irom 
south eastern Australia are needed to test tins 


Thanks are due lit the fallowing collectors who 
provided material for I his stud) ; l. Bevcndue. 
R. Beavis, T. IL Cribb, f. Dennis, M, Driessan. 
f. I : oIl->'. T- Friend. R. fuirner, R. Gosslinf. 
P. Haycock. P. Hayward. O. Iliekinan. & Mund;r v 
R. Nolan. R. Nonnan. D. Obemlorf. I.. Owens. 
(X Spratt, ihe late M .f Mackerra.s, P. Presidenle -aiiil 


tliC laic A. J. Beurup. Thanks arc due to C. Kemper of 
the South Australian Museum, J- M Dixon of the 
National Museum ol Viclonu and T. Friend of ihe 
Department ol' Conservation and Land Management. 


Western Australia for allowing me to dissect 
bandicoots held in the mammal collections and II. 
Smyth for the drawings. This work was supported by a 
grant from the Australian Biological Resources Study. 


Bi-.-Minw VlTla. H W- i W5l Southern Broun Bandicoot pp, 
176 177 //; Stratum. R. (F.d.l "The Mammals of 
AiiNtralia" (Reed Books. C'bulsvvood). 

CitrtBAi u-A. G„ Sii'kim . c. Bivi.Kini.i:. I.. Sain, o & 
hMUinr-Drssrr M.-C. il l JNl)> Sur les Nematodes 
lehinonemalinae. Att/t. AY//./,\//<>/ hum, rump. S$, 427- 

t*wn;k, Hi J. (ISSi) Supplemental y report tin specimens 
dredged up from the Ciull ol Manaar together with others 
from the sea in the vicinity of the Basse Rocks uud from 
Bass's Snails lespeelivcl}. presented to I he Liverpool 
Pice Museum !>> t'api II. Cuwric Warrcri. '*"" Mffjr, 

Pmi'M), I, A. & Bl'«Klix;i.. A. A (IVV5) Western Baiivd 

PunOlcoul pp. 178 ISO In Sicilian. K- (fd ) "The 

Mammals of Australia" (Reed Books. Chatswood). 
(ioKnuN. (i, tl'l l J5| Northern Bnmn Bandicoot pp, 174- 

175 flu,! 

_St Hi n 111 ki. A. II. (1589) Perumelidae pp. 6G3-62.4 

/// Walton. I). W, & Riehurdson. B. .1- (F.tls) "Fauna til 

Australia" Vol IB Mammalia (Australian Government 

Publishing Sei\ k e. Canberra), 
Inoiis, W, (i, (P)o7> The relationships of the nematode 

superlainilv Seuratoidea, ./. Helminth"!, 41. 1 15 136. 
Joiinsion. T. II. & Mavvsov P. M. i\VM)) Sundry 

nematodes rrorrt eastern Australian Marsupials, funis R. 

S.'u , S 63. 204-20* 

& ( PMO) New antl known nematodes ln>in 

Australian marsupials. Pmv, Linn. Sac NSW 65. 468- 

A: _ (l*Js2l Some nematodes li'oin 

an birds and mammals. Trans: R Soc. S Auxf. 75. 


LiNStovv, O. Von (IX'Wai Neniathelminthen Von I lei in 
Riehard Semon in Australien ^esammell f)nif.\rhr fried' 
fhtmru Cn'st'l/stft. fpfltt 8. 467*472. 

_ i |98*)h) Neinutlielminllien. von Hcrrn R. Semon in 
Australien yesanunelt, /ifflcy.isduw Central liftlti 
Leipzig 5. fY72. 

Mi kirs/ii N.I ., Morris. K-J>-A liu k\uv C, Elf 1995) 

Golden Bandicoot pp. 172-173 fa Strahan. R. (Kd.i The 

Mamrnuls ol Australia" (Reed Rooks. ClutKwood)- 
Mv "khKHAS, 1. M &i MvekUiKAS. M. J ( l l KiO»Tuvotiotiiy 

t)f the common short-nosed marsupial bandicoot of 

eastern Queensland. \tist. J. Sit, 23. 51-53 

_ & SaMWKs. TX F. ( ly.S^) Puiasilcs of the 

bandicoot. Pmc R.Soc Qld 63,61 fr3 
Maiioma .!, A. & Riur.. \V. L) L, ( H'SSl tViamelidae pp. 

3d 42 In Walton, [). \V. (Gd.) "Zoological Catalogue ol 

Aiislrali;i"* 5 Mammalia t Australian (iovemnienl 

Publisliiu^ Serviee. ll'anberra). 
MiNKiKmsT. R W. ik Si.ihick. J. H, i |995) Lon^-nosed 

Biuidteoot pp 77-7N in Seebeek, J. II. (fed.) 'Mammals 

of Vietoria Distrihution. ecology and eonseiT;irion" 

(Oxford l niversitv Press. Melbourne). 
MusDAN. II. L. A: tSukKN, K. Ci. ( 1**72 ) Parasites ol 

Tiismaniaii native and lenil fauna Pari II. Ileluiuiilie , 

Rtr. Queen Yh\ Mus.lA. 1-15. 
Qitniin. J. C. (W70) Sur le eyele evolmil de St annum 

cdJitnnhfnsv Desportes. 1947 el ses uniniU's usee eeus 

des Nematodes Subulurpi (Asearidia) el Rietulaifs 

(Spirurida). Ann. Pam.sitol, hum. camp, 45. bCr>62ft 
Komhk. [•. (19(11 ) Mononemalu und MarsupioJia. DtHtlsi ht* 

mi'tl-muunv (u'scUm ii, Jt'tnt, 8. 153-160. 
Sirmrk. I Ci. (tf?95) Hastern Barred Bandicoot pp. IS2- 

183 In Slrahan. R. ( "The Mammals ui Australia" 

(Reed Books. ChatsvvrMid). 
Si-kMi. D. M, Bi-vi:ktr>(a!. I. A Wai.ti a. 1:- I (Pi'H I \ 

catalogue of Australasian monotremes und maisupiaK 

and their recorded helminth parasites. AY< \, Aim, Mn\. 

Moiutiit sir. I. 1-105. 
Stoddakt. P- 1 I9M5* Pon«;-nosed bandicoot pp. IS4-IS5 hi 

Slralnm, R (Pdl "The Muitirnals ot Aus|i;dia" (Reed 

Books. Chatswood). 
Youm.. W. & MAri.hsmNf.. I> A, (1926) "The nematode 

parasites of vertebrates iChurchill. I.omlon). 





ByW. Zeidler* 


Zeidler, W. (1997) A new species of freshwater amphipod Austrochiltonia 
dalhousiensis sp. nov. (Crustacea: Amphipoda: Hyalellidae) from Dalhousie Springs, 
South Australia. Trans. R. Soc. S. Aust. 121(1), 29-42, 30 May, 1997. 
A freshwater amphipod Austrochiltonia dalhousiensis sp. nov. is described and 
illustrated. It is endemic to a few artesian springs amongst the Dalhousie Springs 
complex in the north of South Australia. Morphologically it is very similar to other 
species of Austrochiltonia found in mound springs near Lake Eyre South but 
preliminary electrophoretic analysis of allozymes supports the recognition of a 
distinct species. It most closely resembles A. australis (Sayce, 1901) in that uropod 3 
is two-articulate, but differs in a number of minor features, which collectively 
distinguish it from its congeners. 

Key Words: Austrochiltonia, dalhousiensis sp. nov., new species, amphipod, artesian 
springs, Australia, taxonomy. 




by W. ZfcinihK-- 


/.him in. W. ( IWi A new species of fivshw uier amphipod Austnn hthonui <kUiuni:,u'iisL\ -.p. nov. (Crustacea: 
Amphipuda: iivaieilidac) Iiuhi Dalhousic Spunk's. Si>t»lh Australia. Trans. R. $rt£. S. \tt\L J21( I h 29-42. M) 
May, \W7. 

A freshwater ampfupod AusmHhilu'iiui iinUn'n,\ivitst\ sj>. nov, is described and illustrated, h is endemic nj h 
lew artesian springs -juumgsi die Dalhousic Springs complex in (lie norili ul Snuili Australia- Morphologically 
ii is very similar 10 other fcpprftiS of AusinHlu(i<wUi found in mound springs near Lake Lyre South hui 
preliminary clcclrophoretic analysis Or nllo/ymes supports the recognition ul a di.slina species, ii mosi closely 
lexnnhtes A tut\lr<tliv (Sayec. 1901 ) in licit urnptKl ^ K Iwn-arlicutate, hul dilTers in \\ nnrnberol minor features, 
which collectively distinguish il from lis congeners, 


\nsttthhiftoniit lltilttatftii'WliH sp. nov. new species, arnphiptuL arie;sinn springs, Australia. 


Aiiiplupod species Of the genus Auttmihtlteniit are 
among Lbs iniDsl common crustaceans found in (he 
permanent freshwater* of southern Australia rtttHpra 
front New South Wales to Western Australia and 
including Tasmania. More recently Austnuhiliotna 
lias also been found in the inland waters of artesian 
springs in Soulh Ausiialia (Zcidler 1989) and ;il 
'Ldghasloif north-cast of Aramac, Queensland 
(personal coUcelum. May 1 988). 

When I re established die genus Anx!r<><-hiih>ni't 
(Xeidlcr 1 9KX) il was my intention to proceed with an 
Australian revision of I he genus beginning with (he 
description of species lotind in the mound springs 
near Lake Eyre South and aL Dalhousic Springs in 
northern Soiuh Australia. Since then I have examined 
a large number of specimens from wide-ranging 
habiiats in suulhein Ausiialia and have found (hem 
all to be very similar morphologically and difficult lo 
distinguish from (he only previously -described 
species. A ui/stnihs (Sayce. IvOI) mui A \'ithtt'tmis 
(Sayec, 1902). Williams ( I9d2| revised (he 
'•Aslcinaucs ul these two species based ^)\ type 
material and a range of specimens from New Soulh 
Wnles. Victoria, Tasmania and Kolinest Island. 
Western Australia and likewise found that, 
morphologically, specific differences are minimal 
However, a preliminary analysis of allo/ymes of 
specimens hum Hie Suiilh Australian muiind springs 
Ufilljtg electrophoresis, indicates that AusirocfulKwiu 
is mosi likely a very speciosc genus. Cii\ r en its 

: Smith Aiisluli.iti Museum. VhiIi IVii. iu- Ailchuilc S. Ausl 

potenlial enormity, the project was abandoned due to 
lack of resources 

J lie species found at Dalhousic Springs is most 
similar lo A, attstniti.v Sayec. 1901 in dial uropod 3 is 
two-articulate. It has a very restricled distribution, 
occurring al only three of ahotil 80 active springs in 
the region (Zcidler 1989), Two of Ihese springs are 
quite large, wilh large outflows of warm water 
o40 n O bul Ausimtiiittonhi is found only in the 
distant overflow where the waler is colder anil close 
to ambient temperature. However, one isokued 
specimen was collected from the edge of the pool of 
die main spring, which has a waler tetnperalure of 
about ^5"C The other spring is a small, relatively 
cold spring on the southern edge of the spring 
complex- In each case the animals were only found 
in the shallow edges of swamps or channels amongst 
the base of the sedge Cypents Itn'\'ii>aii4\ L., 1771 
and sometimes also the reed Hira^niitc.s anstratis 
(Cav., 1841). 

The restricted and isolaied dMnbuiinn u\ this 
species o\' Ausmn hihtm'uL makes il vulnerable In 
habitat disturbance even though Dalhousie Springs is 
within \Vit]ira National Park. The purpose ol this 
paper is 10 establish the taxon so ihai park managers 
and visitors can appreciate ils significance and 
potential vulnerability. 

Maleriiik and Methods 

The Dalhousic Springs complex (Pig. 1) consists 
of about 80 active springs all o\ which were sampled 
in !9X5 (Zeidler& Ponder 19X9) but Aii.sii'tnhilinnia 
was found in only three springs (Hg 2). The springs 
atC coded following Zcidler & Ponder I I9K9. Fig. 2). 



Animals were collected from amongst sedges and 
reeds With U small hand sieve or picked oW plain 
debris with forceps. A total of 424 specimens (230 
9 9, 174 cfcT. 2(1 juveniles) was collected and 

Physicochenheal data Tor the sites sampled arc 
limited but some measurements were made near the 
main source of the spring. These data are given in 

■AHLi! 5M<H||£ 


Northern Territory J Queensland 

[J iil-,. ,,■,,.- '-.r-r mq-:.« 1 




1 ,*,., ,, 

HI r. |> 


>Mnri !■■ 
South Australia 

': | 



M . ■ 








\ | 



Fig. I. Location of Dalhou>ie Springs, Sooth Australia, 
1 mm A'idler Wl. 

Table 1 and data on nearby springs are also available 
(Smith 1989); 

The new species was compared with the 
descriptions of AitximchHionia given by Williams 
(1962) and with specimens of A, aiistralis from 
Dandenong Creek. Victoria (SAM A C3872) 
identified by Williams and used in the study by 
Smith & Williams (1983). 

Material reported here is deposited in the South 
Australian Museum, Adelaide (SAM A) and the 
Australian Museum. Sydney (AM). All specimens 
are preserved in 75$ ethanol or 2 1 ^ 
formaldehyde/propy lene-glycol solution. Of the 
types, only the holotype and allotype have been 
dissected (partially), with appendages removed from 
the left hand side of the animal unless otherwise 
indicated. Dissected appendages are preserved with 
the carcass or, in the case of the holotype, the 
inouthparK uropods ami telson are mounted in poly- 
vinyl laetophenol on a microscope slide. 

Specimen length is measured along a lateral 
paraholic line drawn from the anterior extremity ^\' 
the head through the mid-line of the body lo the 
posterior limit of the telson using a pair o\' dividers 
and scale. 

The thoracic limbs arc referred to as gualhopod I 
and 2 followed by pereopods 3-7. Size comparisons 
of gnalhopods exclude the coxa and dactylus. and til 
the pereopods, the coxa, with articles beine 
measured along the mid-line. 

The following abbreviations are used in the text 
and figures. A J . A2 = first & second antenna; CI I . G2 
- first & second gnathopod: LL = lower lip: Md = 
mandible: Mxl, Mx2 = first & second maxilla; M\p 
= maxilliped: Q2-5 = oostegites from pereopods 2-5: 
P3-7 = pereopods 3-7: PI I = first pleopoda: 1 = 
telson; V 1-3 = uropods 1-3: UL =» upper lip: r = used 
as suffix to indicate that appendage was laken fmni 
right hand side of the annual. 


1. Temperature measuramrms ami physn <n hemn ahltiht i from Smith 198$) fM sp)WUSfrom whirl} amphifvuls WC'V 
iollettfii a! lime aj eotieetitm n:\rtptjoi Cat ■ data from i9&3 tiXftedilirm} 

Field Chemistry 














25 n C 



Oil - channel to main pool 




i loo 




Cal - main pool 








Cal - main discharge channel 








Cd2 SW edge of pool 








C\I2 - ai or near swamp 


I 1 

















\* «r. j 


*VS '■■ *t N 

• \i « 




^iWi life /*/^ /.; 


Dalhousie (ruinsj^ V\ 4 ^V 

, . V, will ' p*1 *i; ■ l ' l 1 .... v 

V-. I' ■ \ '"'' ;i ' k" ! Sat ■/« 

v/ f/ "apo 

,. .: >..-. ,..,, 

' V- ^°*dn a ; 

: > 


Fig. 2 Dulhonsie Springs complex showing springs (coded) from which Ausiroehiht'iiui tlttdwtisirnsix sp, nav. was col- 
lected- Collection Mies lor C;il and Cd2 are arrowed. Oilier major springs are shown as dots. Swamps from springs and 
«. reck beds arc stippled ( lijiht stippling indicates ephemeral stream beds and heavier stippling areas ol pet mniienf waiet). 


w /rim ,ER 


Uistraehilhviia delhousienm sp. DOV. 
(FIGS 3--S) 
flrttfrta fe/foiWw sp. Zc idler. 19S9- 83-X4. fig. J2 IB. 
I4WU IS.s 

/h'/eivfu- jT, TJjilhousie Springs SA. Irom amongst 
iceds mid sedges along eastern edge of swamp 
created by outflow from main spring (Cal), },$ km 
north of edge crfpld iiirsirip. 20 n 23'07" S 135 30'26" 
Is- k2MJ9H5, W. Zeidlcr & K,L. Gowlett-Holmev 
.SAMA C '5651 

Mfoiypc. O^CTtJUi V. SAM A {3652, Collected 
With holotype. 

7V/m/vyvx AM P1NS40. 10 9 £ . (U o"cf . same data 
!l,S rJoUjijpfr, SAMA CS6'53 ? 24 PQ (one 
ovi^eo'Lis), 17 Cfcf« same data as h<»h>t> |>tr. SAMA 
(ViM. 37 Q 9 (three ovigerous). II o"o\ same 
data .'s holotype exeepi 14 vi 1 985 SAMA C5655, 

25 9 v - IX o'ef. same data as hulolvpe bill I.I km 
fnrlher north. 26 22 26 S 135 30'26 V R. 

(tttnr material i\\utnhw<l; All I rem Dalhourdc 
Springs area [Rife. 2f. AM P4SS4I, ! Q idamaveU), 
spuug Cal . 26 2V0O" S 13^-29'5:V b. trom edge ol 
main p<>ot, W I, Ponder & D. Winn, 3.vU9N5. 
SAM A CSe56. 219 9 (three ovigerousj. 31 cf o\ 
spring Cd2, from edges of swamp formed by 
outflow, approximately 9 km N\V id source. 

26 '24'33*' S |35*2tf(5" H, W Zcidler & K, L 
Gow|eir-Holmcs r 6,v.,i9X5. SAMA CS&% 14 
V 9 , 21 cfc\ same data as previous M extvpi 

14 vi I9M5.AM P4.SX42, Iff 9 9 time pYfgcnjia.V -fi 

no', sprirtg Gbl. from edges of swamp. 26 11*12* 
s I «' 29'26" b. W. K Ponder A: D. Winn &V1.19&3 
SAMA C5&5& 90 3 9 ( thrttr ovjeemusi (1 I ; ■ I 
20 juveniles, same data as previous lol e\eepl 
collected \V1 Zcidler & K. b Gowleii-llolmes. 

/)rs, nation of thrftitypr molt (Figs 3-6) 

Length 3.X mm, I lead about as fi>fljj as deep, length 
almost equivalent LO first two pereouHes, Antenna I 
about 3\ length ol head; peduncular article I length 
I *i\ width, articles 2 and 3 subequal in kngllt about 

7>\ leuglh ol* article 1: flagellum slightly Icftger 
than t.5x peduncle, of nine articles with one ventral 
.irsihetasc at base flf each of last four article^. 
■\uicntiii 2 ahoul 7\ length ol Al with 
cluuacteristie eland cope al base; peduncular article 

1 slightly wider than long, article 2 width about 0.7 \ 
length. 2\ as lone as article i and 7s length of 
nitiele 3: flagellum slightly longer than peduncle, of 
ci^lit Liriielcs. 

I'ppcr lip slightly widei than lonp. apically 
rounded, be.aritit: numerous sh<*rt setae apically 
lower lip with vestigia! inner lobes; *iuter loK-s 
sLiOnxale with selosc distal aud inuct mat^nis. 

Mandibles vvtlhoul palp; left wilh incrsor of si\ 

teeth, laeina mobilis of live teeth, spine t'uw Of three 
feathered spines and iriturulivc molar; riehl with 
incisor of five teeth, laeina mobilis of three teeth, 
spine row of two leathered spines and iiiturauve 
molar with one lone feathered seta. 

Mjrxilltf I without palp, notched ai palp's itoniiul 
position: outer plate with nine combdike spines 
apically: inner plate very narrow with two feathea'd 
spines upiculJy. 

Manilla 2; outer plate about l.5\ length of inner 
plale, setal row restricted to apev. inncT plate with 
one laree seta medially about 0.4 from apex, selal 
row apicalK and medially, almost to large seta. 

Maxillipcd. inner plate large, .sub-reclangulai. 
reaching end oi' merus, maximum width about !M| 
length ol ouicr margin, widi ihrec apical spme leeih. 
the inner owe smaller, four plumose setae on innei 
margin and several apically: outer plate ovate 
reaching midway along inner marcin o( carpus. 
uboLit as wide n\ inner plate, apital margin With duee 
setae, inner margin with several setae for distal hull, 
palp large, 4-arttculale: merus proximally narrov,. 
sub-iriangul.'ir, outer margin about 2\ length of imtci 
margin wilh lw r o selac on inner distal angle: eai|)il.s 
slightly brt.mder lhan [otig. slightly expanded disUills 
distal luotlurds or inner margin \\i\b row of selac. 
two sciae on outeT distal ane'e and also ncni inner 
distal angle; propodus slightly narrower and shoiier 
tlutn caiptis, distal margin with scst-ral strong JCRIC! 
curved daclylus with sirone unguis. 

Coxal gills sausage shaped, present horn GZ u> l'6 

Gnmhtipod I: coxa shehdy longer than luaumuui 
width, proximal width about 0.7x distal width, 
anterior matgtn concave, postetior niiugm straieJit. 
dislai margin evenly rounded wilh several evenly 
spaced selae: carpus triangular with large 
posieroiiixnd lobe, sa ill* anterior margin abnosi 2\ 
length of posterior margin, maximum width ahoul 
I 5\ that of anterior margin, posterioi margin with 
elose set row of nine si out. pectinate spine-.. 
propodus sub rectangular, abotd l,4x length o\ 
earpus, slightly wider distally. widlh 0.6\ leugdi. 
posrerodislal corner with two stout spines on either 
side of dactvlu.s. cluster ol long setae on anterodislal 
fctirner, row of seven long setae medially, mixtuie ol 
long and shoO setuL- near distal margin: daciyhis 
slightly shorter than width ol proprnJus littmg ueally 
against palm. Gnathopt>d 2 length l.6x that of (.il: 
coxal gill length 2x width, little shortet than ca<iili 
eo\a rectangular, slightly longer than wide, ahoul 
QJifl length of t>asis. dislai margin evenly rounded 
with several eveniy-spaced setae, merus wilh rigtit- 
angled bend: carpus similar to Gl but without 
pectinate spines; propodus slightly shorter th.m 
basis, length anterior margin 1.5\ maunumi width - 
postciopjosiniul corner formme rounded lobe, palm 
oblu|ue wilh mmierous spines of varying lengths on 



Fig. 3, Auswuhilumni dalhousiensis sp. nov., holotype o\ Scale bars = 1,0mm (whole animal). 0.2 mm (A,U.T>. 



Fig, 4, Aiistiochihonia dalliousicttsis sp. nov.. hololype d\ Scale bar = 0.) mm. 



Fig. 5. Austrockiltonia daUiousiensis sp. nov., holoiype d\ Scale bar = 0.2 mm. 



Fie. 6. Atlstrochiltonia (hilhousiensis sp. nov,, hnlntype a\ Scale bar - 0.2 mm. 



either side of cutting edge followed hy shallow 
&n*)Vti lor tip of daclylus: daclylus claw-like, as- long 
as anlerior margin pf propodus. 

Pcteopod 3 with pail ol propodus and dactylus 
missing 00 right; eoxal gill length almost 2x width, 
about 0.75,x lengih of coxa; coxa like thai of G2 bill 
slightly larger, slightly shorter than basis; merus Sv 
as long as basis, anterodislai conier produced, cajpie- 

7^x length of merits. Pereopod 4 similar fen P3; 
shehtly longer iban G2: coxa with distinct 
posteroproxima) excavaLiiin. maximum width 
slightly more than length, slightly jongei lhaji basts. 
piopodus slightly longer lhan merus, daclylus sloul. 
length slightly less than().5x of propodus. Pereopod 
5 slightly longer dian P4. coxaJ gill length about 2x 
width, slightly: longer than basis; coxa width about 
l.5x thai of basts, anterior iobe slightly more than 
0,S\ length ot basi,>, posterior lobe about 0,S\ length 
u| basis: basis slightly longer Hum wide with typical 
expanded posterior margin and posterodistal lobe 
(cachme Xfy about midway of ischium: meius with 
posterodistal corner produced- length about 0,7s thai 
(fl' b.isis; carpus slightly sht>rter than merus: 
piopodns length L4x that of carpus, daclylus sloul. 
0,S\ lengih of propodus Pereopod 6 length I Vv that 

01 1*5: coval eill length about 2x width, about 15x 
lengih of hasis; coxa almost as wide as basis, anterior 
lube 0.3\ length ot basis, posterior lobe O.Xx lengih 
oi basis; remainmg articles like those of P5 hat basis 
wilh siraighi postcrupioximal shoulder and CtifpUS 
slightly longei ihau rnerus. Pereopod 7 longest, 
slightly exceeding Pf'i. like P6 but coxa semi-cm ular 
and lacking eoxal gill, widih I 4x length, about f>. Ix 
length ol basis: posterodistal lobe of basis more 
expanded icachiug to about midway of met us. 

Pleopods all nnmodilicd mot as in Citt'honkn 

Uropod I ahout L5x length of 172; peduncle with 
spme on inner and outer distal corner, three large and 
one smaller spine on dorsal outer margin, one small 
spme on inner margin: outer ramus slightly shorter 
than inner, length 0.7k (hat of peduncle, wilh twe 
median and three icrminal spines: inner ramus with 
CWO small and three larger terminal spines and ihree 
medially. Urnpod 2: peduncle wilh spine on inner 
and outer distal cornel and additional one on dorsal 
m.Kvin; inner ramus l_2x lengih ol outer ramus and 
[3% that ol peduncle, live large spines clustered 
terminally and three spaced evenly medially: inner 
ramus wiili 20 spines of vaiying si/es giadually 
elosci logelhet lowaids tip. Lropod 3 tWo-arlieuJaie. 
inatguially more than OJsx length ol Iclson: ramus 
0.5x length ol" peduncle wilh three long seiae an<l one 
shoo gctW terminally. 

Iclson entire, subiectajigulai. .slightly widei than 
long, distal maigin slightly concave wtih iwo small 
sciac ;u each corner 

Dcxoripiion of 'allotype female (Figs 7-0i 

Length 3.8 mm. ovigerous with 23 eggs in brood 
pouch, same as' male except for ihe following. 

Antenna 2, Oagcllum of seven articles. 

Gnathopod I, coxa relatively narrower and longer 
than for male, width disially <).8x length; poMenoi 
margin of carpus with close-set row of 10 pectinate 
spines; propodus relatively narrower than for male, 
slightly longer than carpus Gnalhopod 2 lengih L2\ 
lhat of Gl: eoxal gill relatively smallei. less than 
0.5* lepgth ol coxa' cov.i wilh posterior margin 
produced to point medially, maximum wtdlh N\ 
length, as long as basis; remaining articles like those 
ol Gl only relatively more slender Pereopod J 
lengih abuui I .3x that of G2; coxa similar in shape to 
that of G2. Pereopod 4 slightly shorter than P3; ccttft 
without distinct proximal excavation, almost as wide 
as long. Pereopod 5 only marginally longer than P4: 
coxa width about I.7x that of basis; merus. carpus 
and propodus relatively shorter (him for male 
Pereopod 6: basis with posterior margin rounded 
proximal !y.' merus, carpus and propodus relatively 
shoiter than lot male, Pereopod 7 stightlv shoiiei 
than Pf>: basis relatively narrower, and merus, carpus 
and propodus successively slightly ih.ui f6l 

Oosfegites on OXQg 5-5i Oil Witn vurlcrl margins 
and numeious small hooks, together lorinmj! light 
marsupium. First heart-shaped, length 1 .6\ 
maximum width, about ft, /x length of <J2: second 
liapc/oid. length almost 0.5x that of P3. maximum 
width almost 0.5x length; lliiid oval-shaped oi 
similar size to second; fourth sub-rectangular with 
obliijue distal margin, length anteriorly almost 0.5\ 
lhat of P5. maximum width almost equal to length ot 
posterior margin. 

Uropod I lengih I .(». tllSfl of U2. peduncle with 
five large and one small spme on oulcr maigin. mner 
margin sviih two small spines proxuualJy in addition 
to large, spine on disial corner; outer ramus as long as 
inner, length 0.8 x thai of peduncle, with two laig.- 
and two smaller spines lerminally and iwo medially , 
inner ramus wilh three large and iwo smaller spine* 
terminally and two medially- Uropod 2 pedunclc 
wiih two large spines toy outer margin; outer mmus 
slightly shorter than inner, length l.3x that ol 
peduncle, one huge and two smaller spines 
lerminally. ihree large spines medially, hmer ramus 
with iwo terminal spines, cluster of lour near tip and 
another two mcdiallV- 

Tclson with tfionp of ihree small setae at >;kI. 

t7\7//Wo, s "i 

i'ukrn ftotn ihe type htcaluy m rccogmlion ot ihc 
rcsiueicd dfstiibuuon ot the species. 



Fig. 7. Aiisfrochiltonia dalhousicnsis sp. nov., allotype 9 . Scale bars - 0.2 



Fig. H. Ausfrocfiillonia ihiUumsii'nsis sp. raw., ;illolype 9 . Ooslegite.s on P3-5 not illustrated. Scale bar = 0.2 mm, 



Fig. 9. Austwchiltonia dalhousiensis sp. nov. Oostegites from allotype 9 . G 1 -P4 from parutype 9 , 4.8 mm, from 
SAMA C5653. Scale bar = 0.2 mm. 




Apart from minot variations due \a size, paraiype 
and other material examined is very similar to either 
((tC ht »li»typc iir allotype. The maximum recorded 
si/e of nudes is 5.2 mm and lhat Of females 0t5 | TVH 
hut most specimens examined ate around 4 WW 
long.. Minor differences between specimens 
generally were noted us lollops. The numbci Of 
flagellar articles pTAl varies from eight lo ten with 
one small specimen having seven: A2 has" from six to 
nine flagellar articles hut must specimens have only 
seven or eiphl. The number ol" uesthetasex on A I is 
remarkably eonstaiil with only some latter 
specimens having an extra one (live). The number ol 
pCCtilKlte $C?4C on the CtifpUSOf I Wi«i <V<nn seven 
to nine iii males and right to ten in females (\imil. n iv 
lor G2 i'l tcniaJesh hi the allotype the COgOC of (72 
P4 ate ol an unusual shape, dillcnng bom males jjkJ 
tton ovigerous females (Fig, 9) in thai the posterior 
margin is produced |o u point medially and PI is 
without a proximal excavation. In the holotype the 
b:*sis of PG has y relatively straight posteroproximal 
shoulder bur in nearly all other specimens examined 
the posterior margin is evenly rounded. Pereopod 7 Ea 
usually longer dnin P6 bin in the allotype il is sligliilv 
<hdW, po'bahly beeuuse vif the relaltvelv shorter 
pcopodus which is normally longer than the carpus 
The spmaljon of I'l & 2 varies slightly with huger 
specimens having vine or (wo extra spines on the 
peduncle and rami- Uropod 3 is usually two 
artit ulate and only one specimen (a female bom 
i ViSVi h;ui U3 wilh one article and then only on the 
tightdvmd sule, Oostegites of females vary 
considerably ill si/e but are expanded, as illustrated 
(or the allotype, in ovicerous specimens 

The possibility lhat speciation may have occurred 
betvuen springs without any obvious morphological 
changes was considered and specimens lor allo/yme 
eleettophoretie analysis weiv collected ftom .ill three 
localities. A preliminary analysis ol ihis material 
tisiny tuelhods outlined by Richardson ei cii (I486) 
indicated fixed genetic differences of ]i) l \ or less 
(fill ?l loei). thus supporting the morphological 
evidence oi' one species with hllle variation, f. iivcu 
Ihese results, a more detailed analysts was 
considered unneecs.saiy. 


1 he new speeies described hetc closely jcsemblcs 
A. aiistrtjlis m thai 1)3 is I wo -articulate. Mowevet. .i 
uuiuber of minor features collectively teadtly 
distinguish il from Hun species aftd j|s only othet 
couecuer. A. \tfhlcuuis. The ntam distmgutshm;: 
lealiues ure as follow. Females icaeh a larger $Ufl 
I hi* 1 1 uiulcs and Ihe species is genet ally uo! as kugc .is 
A, uuslfuli\ bualcs up to 10.0 mm, females up to 8, \ 

inml or A. .\nf>U'ttitis (males up to 10,0 mm. females 
up to 0.4 mm i- Antenna I has fewer ueslhelascx (5-7 
111 A. iiLHiralis). Both antennae have fewer flagellar 
articles I At up lo 17. A2 Up W 11 in 4- iWMniit\) 
The coxae of A. itaihoitstMisis hip. now are relatively 
wider and the excavation on coxa 4 is uol as deep as 
in A tin\trtthx or A snhtt'twis. In both A. aitstrulis 
and A. stthtttiitts, the lateral margin oi' the- excavation 
ul u>xu 4 is at right-angles to rhe posleriot margin 
whereas in ,4. iLilhoushinis the angle of the coxal 
excavation is much greater than 90. In ovigeious 
females of A ttolhoush-ttsis the coxae of G2-P4 have 
the posieiioi margin produced to a point medially 
and coxa 4 is without a characteristic cuavaiion 
There are fewer pectinate spines on the cat pus of G \ 
(males* and Gl & 3 tltmniesi than in A. aibilftiih 
(usually »I0>. For Gl f males) and Gl & 2 
(females! the carpus is. sligluly shorter than the 
propodus whereas the reverse is true tor A tfHStfUlti 
bihei miuoi difletences between the new and oihe> 
species no doubt exist but wcte not evident in Hie 
present study, 

'\n.\mu hihon'w dtflhtmtiWMX is also similar lo 
l y hrr/mt< hilromti utut{>lnhahtm /etdler. 19V I. a 
phrcatic species which nlso has n limited distribution 
at Dalhousie Springs t/eidler P> l M). especially in 
thut cwigerous females of A. thtlluiuM'rnsis have co\a 
4 without an excavation, a feature characteristic ol /* 
«n<>phthiih/hi. Given the isolated habitat ol 
PHdhousie would suspect that these two 
species would have common ancestry. Howevei, 
since eleetrophoretic analysts has shown thai they 
differ at tboul SO^v of the 2 1 loei examined, lllis does 
not appear to be the case. 

The closest relatives geographically, apart from P 
anophtitttinhi. are species ol Atistrinhiltoua found if) 
the mound springs near Lake F.yre South. Although 
A tld!hi>u\'ifn\is is morphologically very similar to 
these species, eleetrophoretic analysis has shown lhat 
it differs from them at 73-SO r i of the ?l loci 
exununed. Clearly a moie detoileil morphological 
and geneltc study ot the yenus is iev|uncd |o 
determine relationships. 

A sinelc. damaged female of /t, tlaihoush ns'ts w as 
found in The pool t.l ^iimg Gal I AM IMSS4G I'hr- 
rCCOrd may he due lo contaminated collecting 
ec|iupmenl us the watci tempeialtnc at Ih-il hicahtv I* 
WCiinJ heshwatei amphiptuK prefer i 00 In w ; .iei'- 
(liaruatd & Barnard I^K.M, Il theicl<>ie seems 
unlikely (hat A. ttuthtmstcnsi\ occuis naturally m Ihe 
pool ol C'al but its pos.sible occuircncc al Hits 
locality watrants fuiihet imcsligatioil. 

The factois determining Ihe dishibulioii ol llus 
species are unknown, Its restricted de-tubulion ul 
Dalhousie Springs is ptu/lmj: as many uppaicul!) 
suitable habitats exist in which this species was not 
lound, Allhoueb re^tneled in its dishibulioii. iiu 
species ts tclulivcly abundat»t at all ol the gOllOCHotl 



Like P, (/nophfhiilmn, the presence of this species al 
Dalhousie Springs on the edge of the Simpson Desert 
suggests that it is a remnant of a once more widespread 
fauna during a time when central Australia was much 
welter than it is today (Kricg 1989). 


I am most grateful to Di W. F. Ponder (AM) For his 
assistance in organising the 1985 expedition to 
Dalhousie Springs. He is also thanked for his 

assistance in the field and for collecting specimens, 
as is Ms D. Winn (AM). Ms K. L. Gowletl-Hohnes 
(SAMA) assisted greatly with field work and the 
collection of specimens. Mr M. Adams Evolulionary 
Biology Unit SAMA conducted the electrophoretie 
analysis and his expertise is gratefully 
acknowledged. T also wish to thank Ms J. Thurmer 
(SAMA) for preparing bromides of the figures and 
Ms 1), Churches who typed the manuscript. 

This study was supported by funds from the South 
Australian Museum. 


B\kN.\Ki>. .1. L, & BAftNARD, C. M, (1983) "freshwater 
Amphipodu ol the World. I. Evolutionary Patlerns" 
(lla> field Associates. Ml Vernon. Virginia). 

Kkii « i (.. W. ( 19X9) Geology pp. 19-26 hi /eidler. W. & 
fonder. W. P. I Ed*] "Natural History of Dalhousie 
Springs" iSA Museum, Adelaide). 

RionkosoN, H. L &AVB&STOGK, P. K. & Adams. M. [1086) 
"Allo/yme Electrophoresis" (Academic Press, Sydney). 

SAYfi . O. A (I9()|) Description ol some new Victorian 
fresh-water Amphipoda. PrpC, tt. Soc. \ia. 13, 225-242 

(1902) Description of some new Victorian fresh- 
water Amphipmla, Na 2. (bid. 15, 47-58 

Smith, P. C. (1989) Hydrogeology pp. 27-» h, Zeidler. W. 
& Ponder. W. F. (Eds) "Natural History of Dalhousie 
Springs" ISA Museum. Adelaide). 

Sviiiu. M. J. & Williams. W, D. (1983) Peprodiiclion 
cycles m some freshwater amphipods in southern 
Australia. Mrm.Ausl.Mwii IS. 183-194. 

WILLIAMS, W. D, (1962) The Australian Ireshwaler 
amphipods 1, The genus Aiisirochiltonni (Crustacea, 
Amphipodu. Hyulclliduc). Altst.J. Mar. Fie.\h\y. Res. 13. 
1 89-2 1 ft. 

Zni>l.i R. W. ( 1988) A redescriplion of A/hn hifhmui capen 
sis (K. H. Barnard. 1916) with a review o! (he genera ol 
the family Ceinidae (Crustacea. Amphipodut. Attlh S 
Afr.Mtts^VH. 105-119. 

i 1989) Crustacea pp. 79-87 If) /culler. W. & Ponder. 

W. F. (Eds) "Natural History of Dalhousie Springs" tSA 
Museum. Adelaide). 

(1991) A new genus and species of phreatic ampfti- 

pod (ClTJfltaCea, Amplnpoda) belonging in the 'Chilloriiu' 
generic group, from Dalhousie Springs. South Australia . 
Trarit fi.Sm S. Aust. 115. 177-187. 

& PoMji k. W, P ( 19X0) Preface pp. \\-\\ hi /eidler. 
W. & Ponder, VV. K (Kds) "Natural History o\ Dalhousie 
Springs" iSA Museum. Adelaide), 




VOL. 121. PART 2 


By Alan F. Bird* & Stuart G. McChUREf 


Bird, A. R & McClure, S. G. (1997) Composition of the stylets of the tardigrade, 

Macrobiotus cf. Pseudohufelandi. Trans. R. Soc, S. Aust. 121(2), 43-50, 30 May, 


The chemical composition of the two stylets of tardigrades has received little attention 

in the past, probably because they are so labile. We have studied these structures in 

the tardigrade, Macrobiotus cf. Pseudohufelandi, and have shown, using 

histochemical techniques and energy dispersive X-ray analysis, that these structures 

are composed of calcium carbonate. 

The response of M. cf. pseudohufelandi to the environmental stimuli of anoxia, 

differences in pH and temperature change demonstrate that this tardigrade can 

reabsorb its stylets under stress and reform them when the stress is removed. The 

possible evolutionary links between tardigrades and molluscs that have darts 

composed of calcium carbonate are discussed. 

Key Words: Tardigrade, stylets, Macrobiotus cf. pseudohufelandi, SEM, energy 

dispersive X-ray analysis, histochemistry, salivary glands. 

tititt.utt twtts vftht AVw// N/», /to o/ V An\t, 1 19M7), 1210), 43- M), 


by K Bird* & Stuart G. McC 

I L'Kfc 


ItiKD, AT, & Mff.itiRi., S.lT (IO'J/1 Composition of ihe stylet* 0f the Ludimadi:, Ma<rnl>ii>hi\ el". 
fXwMiufehmtti. hvns. H. $<n. S.Aust. 121(2). 43-50, 30 VI uy, IW7, 

The chemical composition erf flic two Stylets of tardigiades has received little attention in Ihe p;isi. probably 
because they are so labile We have studied these structures in the tardigrade. \fa< tahf<ttu\ c\\ psauiithufvhtiuli, 
and have shown, using histoehemical techniques and energy dispersive X-ray analysts, thai these structures are 
composed ol calcium carbonate. 

The response ol M. cf. pst'tulalm/elandi to Ihe stimuli of anoxia, differences m pi I and 
lemperatmv change denioiisiiaic that this tardigrade tan reabsorb its stylets under stress and reform them when 
llic slress is removed. The possible evolutionary links between tardigrades and molluscs ihtU have darts 
composed of calcium carbonate arc discussed. 

Ktv W0fcr*3: Tardigrade, stylets. Macmhiiiftts cf, psctu/afmh.'ftrndi, -SIM. energy dispersive X-ray analysis. 
histochemistry; salivary ".hinds. 

m' "n'lim 

Materials and Methods 

Members of the Phylum laidigrudu iwuier heats) 
all feed using a pair of stylets whose composition is 
Mich that they dissolve am! break down on (he death 
of Ihe tardigrade and in some commonly -used lixa 
lives. Thus. Ihey arc frequently not included in the 
camera lueida drawings or photographs ul the bucco- 
pharyngeal apparatus commonly used for taxonoinie 
identification. The general composition, shape and 
purpose ol the twin stylets seems lo he similar 
throughout the phylum and so the chemical compo- 
sition of stylets ol one species probably holds for all 
Because ol their transient nature, dLssolving on death 
or fixation, tardigrade stylets have not been ana- 
lyzed. It has been suggcsLed that they are calcareous 
in nature (Kaestner l°oK: Kristensen I l )7n; Wenck 
|sH4. Kristensen unpub. both cued by Nielsen I l J%) 
ami that they resemble nematode stylets in form and 
function iKiggin 1%2 cited by Kinchin 19941 The 
sly lets of Mti<'t'(?hn>tus ef. pst-udahufolaudi are 
curved, sabre-shaped structures aboul 40 urn in 
length llfnl exhibit marked birefringence under polar 
i/ed light (Bird 19%), 

In this paper we examine (he dissolution of Ihe 
Mylets in various media under ihe pi.lari/ing micro- 
scope their staining |<ioperUes utuler bnght field 
microscopy and Iheir elemental composition using 
energy dispersive X-ruy analysis in the scanning 
eleciron microscope. We also examine the icforma- 
lion of the stylets after dissolution and discuss these 
structures in the light of tardigrade evolution. 

2 PUvhml RiMil MiUImmi.S Ait-a, '"or,: 

< MKri'l lind mil Wiii.-r, I'MH J ftm <Rwt>lUl S 4tKl- SUM* 

Locality, sad type and e.Wuctiaii 

The tardigrades were recovered from soil classified 
as a soloni/ed hrown earth from an experimental ploi 
on a farm at Avon. South Australia I Bird L$9o). Aflei 
thorough mixing, 50 g ahquots i)( ihe sod were 
placed in a misting apparatus and processed u.s pie- 
viously described (Bird 19%). The tardigrades so 
collected were picked out using, a dental No. 3 nerve 
broach. Water which had passed through the soil was 
collected I'rorn die misting apparatus anil filtered 
through a 0.2 (.un rnemhiane filler. The tardigrade* 
were placed in a shallow layer of this water in a sler- 
ilc fViii dish. Lender these conditions, tardigtades 
remained viable for a week oi more without feeding. 

Slylt'i dissahtluw 

The break down of Ihe slylels was observed, on 
slides with covershps sealed wilh nail varnish, in 
solutions with pH ranging from 1-8 for different 
lengths of lime trrom ihe start of the experiment lo 
several days) nud at temperatures ranging From Ii5 - 
Mi C Observations were made with polarized lighi 
and differential inlerfercncc contrast (Nomarskil 
optics using a Vanox Olympus AUBT research 

Stvlt! fi'/atwiuian 

Specimens were mounted in distilled water in 
sealed slides with pieces of No. 1 glass covers! ips 
achug as spacers lo prevent crushing. These spevi 
mens were kepi under observation, using Nomarski 
optics, until the stylets could no longer be observed 
(usually nhoiu 3 h). Ihe covershp was then gently 

H A. R I5IUD& s.(.i. Mrl'LURE 

removed under a dissecting microscope and the 
motionless tardigrade* pi, iced, together with sonic 
smull nematodes lo stimulate feeding activity, ill slifll* 
low distilled water in a Petri dish. After about 12 lithe 
tatdituades, which hud resumed activity, were placed 
m scaled slides and re-examined under Nomarski 

Optic S 

EtW&y lUspty WiV A' n/y tinalysi.s rfftbe N/v/e/.v tnnler 
ilu wanning I'lccfron mi* t<>s<<>pc (Si'Ml 

Because of the difficulties of ohluining intact 
stylets from tardigrade.*- on sealed slides or after ihe 
use of acid fixatives such as f'A 4:1, as used for the 
fkatinii of nematodes (Hooper l l )8G), a special Lech- 
niquc was adopted for these studies as follows. The 
! a rdi grades were transferred to a small drop of water 
which was then slumped with an excess of (/v 
gluteraldehvde in Sorenseifs buffer at pH 7..V The 
tardigradcs were then gently crushed under a cover- 
glib to permit rapid entry of the fixative without caus- 
ing undue damage to them, Alter 2 h in Ihe fixative 
the lardigrades were removed, washed lour times in 
0.2 pm membrane-filtered distilled water and placed 
in a small drop o\' the membrane-filtered distilled 
water on the surface of a polished carbon stub. The 
tardigrade* were manoeuvred close together and a 
second polished carbon stub was lowered on to the 
first so that the tardigradcs were eaughi between the 
polished carbon surfaces. They were left in this posi- 
tion for 24 h lo dry and then the uppermost stub was 
lifted directly upwards, thus pulling the rardigradc.s 
apart and exposing their inner organs, including the 
bueco-prnirvngcal region. This technique was also 
used on fresh, unfixed material. The stubs weie kept 
ni a sealed dry container until examined and pho- 
tographed in a Cambridge S 250 Mk } SI M opera- 
ted ai 20 kV using Ilford 120 roll film (FIM Plus). 

Energy dispersive X ray (EDX) analysis was by 
means ol a t ink IDX system attached in the SEM. 
EDX analysis of characteristic lines with energies 
less Ihan approximately OS keV is not possible with 
this detecLor because of the absorption ol the low 
energy X rays by the detector's beryllium window. 
Consequently, the direct identification of elements 
wilh atomic numbers less than that of sodium is noi 

ttiMinhennstry o] the salivaiy glands during .styfet 


The jtilhraquiuone dye alizarin red S , used as a 
stain for calcium, was made up as a 1% solution (50 
nig 5 ml ' ) either in a buffer solution at pit 4.0 or in 
distilled water ipil 5.0). The lardigrades were placed 
in a drop of the ali/arin red S solution and gently 

•- h 

■a i 

N : ?'^ 





Auirnh t'htimwl Cwnptuy 

Fig. I . Mead of living specimen of Macrohionts cl. pseiuio 
hutchtiuti immediately after being placed in distilled 
water on a sealed slide. Note lite two eye s[Htis in tnvw-t 
part of photograph and the two curved slylels lanuw 
lie;ids>. Nomarski optics. Scale h:ir = 10 pm 

\-\\i 2. Same specimen as in lag- I but photographed ptsl 
alter ihe stylets hint dissolved 166 mm later Arrow beads 
indicate Hie original location of the stylets. Nomarski 
optics. Scale bar - 1 (JlfT. 

tag. 3. The same as hg. 2 hut viewed under bright field 
optics. Arrow heads indicate the original location of ihe 
stylets. This photograph illustrates the buccopharyngeal 
structures used in Ihe taxonomy ot this genus. Scale har 
- 10 Mm. 

squashed, without causing gross damage to ihcni. by 
placing U glass eovn'shp over die drop and with- 
drawing Ihe stain solution from under Ihc eovectfip 
with pieces of filler paper. Ihc edges wf 'lie coverslip 
wlmc then sealed wilh nail varnish and the specimens 
observed and photographed under height field upiie>. 


DlSSolufl"}! <*f lift Shifts 

The lime taken for dissolution ot Ihc .slvlctsin dis- 
tilled water, when a slate ofunpxJM has been induced 
bj scaling the tardigrades under a coverslip on a 
slide, is temperature dependent Thus, at M\ t Mai 
-ayki dissolulion occurred within three houi* wh'-o.- 
as at l-V ( Ihe stylets were only partially dissolved 
aflci "ihis lime. 

The appearance of (he licad region o! M el. 
fj\( HffolmjchtmiL viewed under Nomurski optics, is 
h"\Mi prioi lo commencement o| stylet dissolution 
ai 25 ( dig. I ) and ailer its completion ihrce hours 
later (big. 2l Ihe head region is also shown alter 
thai- hours, photographed under normal bright field 
opt us (Fig s> 

The sequence of slaves in ihe dissolution Cd the 
stylets is illustrated most cleaily in a series of pbo 
lomicrographs taken u\kW( the pnlaiving micro- 
scope ( Fig 4 A-H ). Stylet dissolution is gradual o\er 
ihe fust two hours dig. 4 :\ Di but aeceleuues ovei 
Ihe Ihird hour (he. 4 I- Ht -o dial marke.l i ■!<an>:i- 
were detected at appro \ i male I \ IVnimuie intervals 
from l\vo hours onwards, 

Style.t dissolution occurred more rapidly ill acid 
solutions ihan in neutral or alkalutL solutions, h look 
place within 5\>9fl mill in 0.05M HCl and occurred 
immediately in IM IICI and so is similar |q CaC'O, 
in this rcspeei 

The hirefringcuee cxhihiied by the muscles of the 
triradiaiL pharynx of W. el, /»,w.'/Wo/////i/<"'^' did nol 
disappear over a period Qf three homs (Fig- 4} and 
the brightness o\ these muscles, although not as 
inaikedasthal ol the stylets at ihe commencement of 
die i:\pfiimenL remained constant throughout, 

Kcfi'titntlion <>l the stvlch* 

Tardigrades appear to have the capacity to leform 
iheir stylets under favourahle conditions in what 
appears to be a reversal M the dissolulion which 
occurs when they are placed in unfavourable siiua- 
tions, Specimens thai had heen induced to dissolve 
their slyleis m sealed slides and had become motton- 
less were observed to regain their muscular activity 
and reproduce stylets, over a \2 h period, when 
placed in shallow distilled watei in an unsealed Petri 
dish, in ihe company of several small nematodes. 
This reformation did nol always lead lo precise 
realignment of Ihe stylets and the efficiency ol tins 


leiojuiation ami correct realignment may he 



denl on the slalc ot health ol ihc tardigrade at the 
commenn-uK'ni ol the experiment. 

lutt'tvy dixfjt'iMvo S-niv uftttiysts of fh< \M h is 

lAammatton undenhe SEM of the polished carbon 
surfaces holding (he dried- squashed and disrupted 
tardigrade*, clearly show-- whole stylets, or parts 
thcrcol (Fig. 5f. By meansot this technique, it is pOfr 
sihle to obtain suilahle I DX analyses o! the elemeu 
tal composition of the siylei.s. It can be seen from the 
bDX spcUrum (I ig oithat tk-stylets are rich in eal 
t ium. with maximum X-r:n inlcusUy Ctftfilti at i-w^-t- 
gtes matching ihe Ca a at 3.6°-0 keV and the C'a ft al 
4JM2 kcV ehaiueieiist'C \ ray lines 

Hisioc/ttttHstty (<f (fit foU\%lT\ >A"/<A j//f/ft/ji3 \hh'> 


When the tardigrades were genlly squashed tn a 
Iv solution ol ah/arm red S n pH 5-0. the salivar> 
glands summudiug the stylcls eradually hecame a 
<K->:p ied fPfg* "L S) This reaeOon was both marked 
and localised. pro\ ided that the b«>dv ol the tardi- 
grade had been crushed _o allow rapid peneirnlion of 
Ihe stain. These ydaiuK which ate ihoujjhi in Ir- 
responsible for the secretion and dissolution o\ the 
stylets (Kniehin I'HJdf, are apparently rich in eal 
cium as indicated by Iheir reaction to ah/aon ■ r* I S_ 


Infouuaiioii on the namrc ot ihe tardigrade siylcf. 
is sparse and has received little or no altctilion in the 
rwo recent general accounts of Ihesc oigauisms, 
notably a ruosl intorniatKc book on their geiu-ial 
biology b> Kinchin l IW-h and a collection ol papers 
published in the Zoological Journal of the Linncan 
Soeiety in l^'fo ;md edited by Melnnes & Norman, 

Tardigrade slylets were llumtdtl lo have some sun- 
ilariiy with nematode si>kls by Rigem ( l%2 ) uiled 
by Kinchin l ( )94). However, it is clear ibal Jaichgnide 
stylet.-: (Bird l°-%t differ markedly from nematode 
styleis Ipird <V Bird I9VM » both in coinposiiion ami 
Miucture Thus, FA 4,1. a mixture of formaldehyde 
and aceiie aeid and a common fixative tot nema 
todes. causes rapid dissolution of lite tardigrade 
siylets because o\ its low pi I and ihe hrjb leinpeia- 
lure used in the procedure, This Hxative, however 
does not cause dissolulion o( nematode -.lylcts 
Vloyer's medium, a mixture ol gum atabic. chloral 
hydrate and glycerol (Kinchin [*M?4i does not dis- 
solve (ardigrade stylets ami has the added advantage 
of functioning as a eomhined fixative and embedding 
medium, In many illustrations of ihe buccopharyn- 
geal region o^' tardigrades. the stylets are not shown, 
picsumably because they d^olVCj when lUtid. 

Tbe '•mall si/.e ol tardigrade -.tylcfs ( U) pm m 




Fig. 5. Anterior region of dried and fractured Macrobiotus cf. psendohiifelancii (see Materials and Methods) viewed undei 
die NFM showing parts of the two stylets (arrow heads). The site of EDX analysis is eireled. Seale bar = 10 um. 


00 2.00 4.00 G.00 

Energy (keV) 


Fig, 4. Series of photomicrographs viewed under the polarizing mieroseope illustrating the stylets of a living speeimen of 
Miumbiofus ef. pseudohufekmdi exhibiting diminishing birefringence as they gradually dissolve in distilled water on a 
sealed slide as shown in Fig. 1 - Figs 2 & 3. A. At eommeneement. B. After 56 min. C. After % min. D. After 1 13 min. 
E. After 120 min. F. After 136 min. G. After 150 min. H. After 165 min. Arrows indicate parts of stylets. Note that the 
museles in the pharynx do not lose their birefringence as the stylets dissolve. Seale bar = 10 pm. 


length in t\jii< >ohii>tn\ ^[^pwudtiltufcLmdi) and then 
labile nature, have made il difficult hulctcHiunc thch 
composition. We hifyc overcome this by adapting a 
technique used to separate the layers in a nematode's 
cuticle third lV BttUUSCh 1957) which takes Kilo 
Ek&VftiM the adhesive ijMlirtcf Of'ttlflnWl tissues to Hie 
surfaces on which they have heen dried. In this 
inxT.inec. the two surfaces- were polished carbon tods 
mounted on SLM stubs as described in Materials and 
Methods. In some insUmces. tins leehnique lot! to the 
exposure flf cither the Wfiolc 01 large rmnigh [iicGCS 
ol >ivlct to be recognizable f Fig, .M. These were ihen 
easily pltolojjiaphcd aiul sublet led to energy di&per- 
-.iv L - \-ray analysis measurements rtl die Sl:M. These 
aie.isuiemcnts clearly show that die elemental com- 
(..'MiHiii til" the st>lr.fs consists mainly of calcium 
(elements with atomic numbers less than thai ol sodi 
inn ,ire not dcteciahlc)- The rapid dissolution ol the 
stylels in dilute aeids and their marked birch iii'jcikc 
imdci pnlari/cd light. indicate dial they arc com- 
posed ol calcium carbonate, Thus, the eomposu.on 
ul tardigrade siylels dillers horn that }if nem.ilndc 
evicts which an: pmtcin in nature, 

lath uudigiade wjf|p| lies in the lumen ol a stalj- 
\;.i- t L-I.ind (Kinchin IWli. kristeuscn ■ H/7fo i cited 
by Niet-eu IWty-foti -nc-e'-red that sepma'c lol>C6 ol 
[he s.ilivaiy glands ,ia' responsible loi the formation 
nl die stvkM- and tlicir supports in ihe tardigrade 
Itatifhj^'i Whether 01 not ihis applies lo S'1. cl 
fu •tultihnfcUnifH iviuuins to he determined. Wt liaVC 
HOI observed 'he fmmatiuu oi' di--.M-la.oh of i|k 
i i aipporls in tins tardigrade, irnlicattiH' lhal their 
ehetmeal composition is diuVtvm ftom that ol Hie 
\l\ I. «:- Wl! have no inloiniaiion on the oriein and 

Uiciua.d composition ol tto tvte 1 support* lU Wi i?i' 

Qui cspciunems indivaie thai the >i>lets of iff. cl 
ftitetfdtfHtfflatUti ;ne secreted and rcubkfirhcd by the 

suh'.ajs. elands, We have shown lluf (1 iis > Si 
when the turdigrades are gently gnashed ty j dilute 
solution ol the slain alizarin red S al a f>H ol J U Will 
scaled, the cell surrounding each stylet stains as di. 
sivluis breakdown, indicating lhal ealeimn \s Ijty ni 
led arid difTusi thioughoiu two eeJK Ali/anii 
tvd S is recognized as a fusloehemical ie ,'iecnl lot the 
detection of lalcinm (Conn 1477). changing in 
colour rp-mi ydtoft 10 teil 10 puipte <i\ei '*k pit 

lima* *.7-5.Z 

G. McCLl \-\ 

The cradiuil hi-eak down ol 'die stylets in a slij;hily 
acidic and anoxic cm irumuein. US demonstrated in 
this paper, when photographed under polari/cd lighl 
{Fig. 4>. closely resembles dieu siep by step farina- 
lion as depicted in drawing* by Marcus I, l^'M (cited 
alitl redrawn by Kinchin |394) sbuwiui.', stages in 
their lormalion. Although stylets of calcium carbon 
ate are unusual in the animal kingdom and appeal tfl 
|>e both unii|Lie and einmnon to all uudignidcs, some- 
what similar siruciures are tound in other proups ot 
animals r«n instance, many ut' the terresuial pul 
tnonale gastropods shoot darts of calcium carbonale 
into t-*ach other prior io copulaium. This l>ehavioui is 
thought to act as a stiuuilaiil to the sevual aei The 
darl ol Helix osf^rsu is about 8 - 10 mm in lentil 
' l^nipa IVS2), at least ten limes the si/e ol an enia. 
lardiL-rade anil some 200 tunes larger Ihau lift -"' fi l] 
stylet ol Macmhiottis cl. j)setnloi\L<jclttniH. The rtflfl 
which takes about five days to form, is secreted in j 
dart sac thai is part of the rcprodueiivc system. The 
i ompi.silmn nt llu- -..: daits, when subieetcd b.L-neivy 
dispersive X-ray analysis, is identical to the lardi 
^rude stylets with holh Vt\ Ka and t 1 u K|i peak> 
(Hunt l')7°) The evoluitoiuiry sicmlicance ot these 
findines is obscure- 1 lowever. the>' do lend mhiic sup 
run! tit die phylwjjcneta pOSitTOJl '»• ihe lardierad-i 
proposeil. as a result ot sludics on Ihe IN S ribosoinal 
gene sequence, by Moon »fc Kim ( HWfk Thcst 
authors stale "the lardicrade clade appears as an 
independent lineage from the nemalode cl.ulc' and 
lh*_* calcareuiis iiiilua- of the tardigrade r-tyki .. 
opposed to the protciuaccous natuie ot (lie nematode 
siylet- lends support io Ihis hypothesis. Moon & Kim 
(l n( ->(>) limber succ/.esl that die mnhjjradc cSatle is a 
si-aer group of die protoslomc eucoelomate as.^eiM 
bla^.e that emerged bWfft the uiolluses. atuiehds 
arlhrnpods antl sipuuculids cxidved. This would sue 
ecsi independent evoUmou ol the ealcaa'ous slruc- 
liircs in laalivrades anil inolln.-cs. 

Some measure of die caps in din knm\ 'leilge i»l ')>•• 
0| i-iusantl relationships ot the TardiLiiada RS iudiLat 
ed by the lact lhal tardigrades arc not mentioned in a 
book on the orieins ;ind relationships amon^ lovset 
iaverichratcs edited by Morns c/ ql ( I9S5), 
However, some primitive Camhrian fossils spate 
m.iih. Lharaeteristies with holh Onychophora and 
taidiyrad-i. iiichating |>ourl\ articulated limbs end 
\W2 in claws dohopodia), tennjnal iiM'Utlis and the 

liy, 7. VVIioIl* spccmit-ii of \hui>>hi>>nis cl I'^toii'liiijt'litiuii that has been WtrsheJ In permit the miiy ot ili/:ntn red S :ii 
l>ll 5,11 The photoeuiph WIW luken unUCTvishl hi-td Hp|ie> hIumiI *•() mttl ntlL-r llie luhihicik-ChkmiI m! ^taininu. Slunviny 
eye spots tsmall arntwsl ami daws (]i|J£v urn iwsl. The iiicu jmaiul 'liu litrdiurddt cniUaiiiv nuU'.iiai ffifil tUis Iven -.'\ikI 
ed lnnn "he nipluad jmiiiijiI juJ Uie Iwo >ulivary irtiinds lh;il Ik jura .niliiiul in lllc aitl.selikil pharynx .ire heavifv stained. 
Scjlebar^ 100 |mu 

I \fr K Knlnrged poitton i>l Pljg ') -.hovvin); tve «pOW tsniall ;utowm. the sUuued salivary glands iS^i. the muscular phaiyA 
|i>) and material lliat hat; been exiuleil by rupuiriiiv \o permit cmrv ol die stain ilarye jitowsi, Senlc bui - li> [ifii 



■ ^sfi:/v.;. 







s j 






last pair of legs merging with the caudal end of" the 
body (Kinchin 1994). For these reasons it is specu- 
lated by Kinchin (1994) that the Tardigrada and the 
Onvchophora have originated from a lobopodian line 
that diverged from the arthropod line in the 
Cambrian period. However. Nielsen (1996) consid- 
ers that Tardigrada are more closely related to 
Arthropoda than to Onychophora and suggests that 
all three phyla have originated from a group which 
he calls "Hanarthropoda'*. and that the ony- 
chophorans diverged first. Jt is clem' that the origins 
and lineage of the Tardigrada remain obscure and 


that much remains to be discovered about them 
before they can be accurately traced and defined. 


Wc wish to thank J. Bird for constructive criticism 
of the manuscript and P. Kolesik for discussions on 
histochemistry. The senior author is grateful to 
Division of Soils (now Land and Water) CS1RO for 
accommodation and facilities and to the Australian 
Biological Resources Study for providing equipment 
that made this study possible. 


Biro. A. F ( 1996) Studies on the soil-inhabiting tardigrade. 

Macrohiottts cf, psi'tuhhufelututi, from South Australia, 

Thitts. R. Soc. S. Aust. 120. 147-154. 
& BIRD, -I. ( 1991 ) 'The Structure of Nematodes" 

(Academic Press. Sail Diego). 
& DburscH, K. (1957) The structure of the cuticle 

of Ascaris lutuhricoidcs var. vw/.v. Ptmtsiurtogx 47, 319- 

CONN, H. J. (1977) "Biological Stains" (Williams & 
Wilkins. Baltimore). 

Hooi'KR. D.J.(19S6» Handling, fixing, staining and mount- 
ing nematodes pp. 59-80 In Soulhey, J.F (Ed.) 
"Laboratory methods for work with plant and soil nema- 
todes'' (HMSO, London). 

Hunt, S. ( 1979) The structure and composition ol the love 
durt (gypsobelum) in Helix pomatia. Tissue tfc Celt 11. 

KAhSlNCK, A. (1968) "Invertebrate Zoology" Vol. II (John 
Wiley & Sons Inc., New York). 

Kinchin. L M. 0994) "The Biology of Tardigrades" 

(Portland Press, London). 
Kkisjhnsbn. R. M. (1976) On the fine structure of 

BatHfipes nocrrcvungi Krislensen 1976. I. Tegument and 

moulting cycle. '/moI. An:, 197. 129-150. 
MtlNNhS. S. J. & Nokman. D. B. lids (1996) Tardigrade 

Biology" Zooi J. Lion. Soc. 116. 1-243 (Academic 

Press, London). 
Moon, S. Y. & KlM, W. (1996) Phylogenelic position of the 

Tardigrada based on the IS S liposomal RNA gene 

sequences. Ibid. 61-69. 

Moukis, s. c. Geokgh, j. D.. Gibson, R. & Platt. II. M. 

lids (1985) "The Origins and Relationships ol Lower 

Invertebrates" (Clarendon Press, Oxford). 
NiKt.SHN. C. (1996) "Animal Evolution : interrelationships 

of the Irving phyla" (Oxford University Press. Oxford). 
To.Mi'A. A. ( 1982) X-ray radiographic examination of dart 

formation in Heli\ aspersa. Heth J. Zsot 32, 63-71. 



By Alan F. Bird* & Stuart G. McChUREf 


Bird, A. F. & McClure, S. G. (1997) Studies of the eggs of Macrobiotus cf 
pseudohufelandi (Tardigrada) from wheat fields in South Australia. Trans. R. Soc. S. 
Aust. 121(2), 51-57, 30 May, 1997. 

The tardigrade, Macrobiotus cf. pseudohufelandi, and its eggs were isolated from soil 
from a wheat field at Avon in South Australia during the winter of 1996. The surface 
of the eggs was examined under the scanning electron microscope and was shown to 
be highly ornamented and reticulate with numerous "inverted goblet-shaped" 
projections. The dentate margins of the heads of these projections consist of clusters 
of coral-like globules. Statistical analysis of the size of the projections revealed two 
significantly different types of eggs in the samples. Examples of ornamented eggs 
were found in discarded exuvia, a condition which is not in accord with a recently 
proposed hypothesis on tardigrade egg evolution. Observations on feeding behaviour, 
population density and egg laying habits are presented. 

Key Words: Macrobiotus cf. pseudohufelandi, microscopy, tardigrades, eggs, 
morphology, evolution, soil. 

IrmiMHW'tt.s of the Kmul S<>, n'lv •>{ S, AlUtt\ EWH, I21t2). 5 1-5^ 


by Ai an P. Bird* & Stlart G. McCuure 


I Jinn. A. ! : & McCi iu\ . St. i IW) Studies ol ihc eg^s r»f MavmtiioUto ef. pwihinlmfrhmili tlfrrtltgrotfal ftwri 
MMllnll fields tn Suuih Australia. fVtffK A', ,W. V 4tttt 121<-' 5t-37- ?0 May. [W. 

The tardigrade. M>.nrohinlti\ el. pu'iiilt'httfclan-h, and its i*gg> were utoltdccj hoin -.oil Ihhh u wht'it! held fl! 
Avon hi South Australia during the whiter ol' I99fr. The siulacc ol LfaO &tigs WU* examined untlci Ihc scanning 

rhvimn iiiiciosiope and was bllWll L(l be highly ornamented ;i i r<J icliculLite with nilniciOlls "inverted gohlel 
shaped'' pnijcelionv 'I hi* i teiM*llc margins til' Ihc heads tit these pni|cctums consist ul clusters of uuniMikc 
j? lobules. Statistical analysis of Ihc si/e ItT ihc projections revealed (wo significantly different IVpeU rtf BgJW in 
Tin- samples Examples ol onminented eggs were tuimd in discarded exuvia. .1 condition wIik h is noi in accoid 
wiih .1 tceenllv proposed hypothesi;-; on lantiyrade eg% evolution. Observations hh IVcdtnij behaviour, population 
density and e$G hiving habile ate presented. 

Kl \ WukOs: XUurnhiahis el pM-thlnhttjrhiihli microscopy nudit-radc-- fc£gS. inorpiiology. PVWlUllMIl- Sull 


1. injigp ulrs. ol*0 known .is water hears or mcv- 
piglets (Kinchin ISM4I. b&tOug U' a discrete phylum 
11I cosmopolitan distribution Irom diverse h;il>iiat- 
tucludnig murine. fresh wuitfl and semi-terresliinl 
environment;-.. The uudigrades responsible foi the 

; ■;•: ■*. lies*. I (hyd Ifl 1 1 * I " - paper Wele identified a> 

\hta<>liit>(ns ef, p\i'H(l(>iitf{ef(tti(lt lharos l%o In S. 
Claxton (Rifti Im%j ami are semi-lcncstrnif having 
heen isolated from sand\ loam soil ill a wheat field at 
Avon. South Australia. 

The tardigrade egg ^tioH i--, a useful taxonomie 
eaiidc lo species identification, particularly in genera 
such as HifiH tvbwtus where the shell is ornamented. 
\Uh tahUnti.s cf. i>H-Uilninifi-(aiuii, which are only 
ubuuj 500 pin long by 150 pm wide when fully 
'iimvti, l.iv comparatively large circular etigs which 
have highly ornamcnled relieululcd shell surfaces 
with numeroitf "inverted ^nhlel -shaped" prujcctions 
iHnd !'>%,». 

It has been sliown by Bertolani it Kebecchi 1 1^)3) 
thai dill'cieiiecs in ct^ shell morphology in 
Mfi< rohii/fus fmjt'linnh', previously thouuht to be due 
|0 vaiiabihlv wifhin this species, fall inlo seven ilis- 
lincl lypcs thai aiv tvlalcil lo clilfcjvnl animal mor 
photypes. Usin^ egg shell morphohigv. along with 
olhci characlcrs. these worXei-s have described a num 
Ivr of new species from the A7. htifcfatldi croup, fees 
(hat belong lo ihis uroup havw pitied or reiictilaicd 
shelf- with pi'otnidinv proccssi-s shaped like invened 
goblets, chalices, thread spools 01 'Voolin^ UnveiV, 

; 1 1'liivloit) KWhl Mileham S. AtiM. 3U62 
( SIU( ) Land iiikI W:iU-i, PMB 2 t *kti Osnmti.l S. AaM 5(l/vl. 

lierlolam ct ui { l l )s>6> have shiLed lha( ornamented 
ci'i's arc iienerallv laid live in sod 01 water and 
smooth shelled ffgg* are laid in die moulted euliele 
(euiviumi. fhe-e wmkers include the tannly 
Macrobioiidae. lo which M c\\ (twuifohnfikttnii 
bdoipK, in those lainilies dial lay tree, ornamented 
eggs. Berttdam ct ill. ll l H.>6i have pin hwwaid an 
hyp(>ihc--is in which ihe\' cvplain ihe evolution «>l 
tardigrade eggs. 

In Uiis p<iper we examioe Ihe slruelure of Ihe et»e 
shell "I AY, cf, f>\ctt<t<>ltnf,kfinli and measure the 
processes proli tiding Irom the stirlace ol ihe ugg 
shell. We also discuss its ijgg layine hahits in relation 
ISS Ihc hvpoihcsis ol Bertokuu el til. ( L*>%) and kmii- 
menL o\) reedtne behaviour and population density, 

Materials nud Methods 

The lardigratles were colleelcd on 18 July |V96 
from sandy loam soil and Irom Ihe same loculily at 
Avon. South Australia (latitude 34 ' 14' S. lonyilude 
1 35S " 1^ B). as those collected previously, using the 
sampling lechnii|tie described by Bird (W>oj. The 
soil samples were collected in mid-winicr so that the 
sues were wel *md the tai'digrades were feeding, 
reproducing and depositing eggs. 

The lardigrudes and oihei tneiolauna, consisting 
predominantly of nematodes, were isolated from this 
soil over a period of three days using a misting appa- 
ratus (Yeates & Bird l 00 4>. Tardigrndes and nema 
todes were counted and tardigrade eggs were picked 
oul using u denial No. 3 ncn r c broach and examined. 
alive, in distilled water under a covershp. with the 
light microscope Kggs to be examined under die 
scanning electron microscope (SEW) were fixed in 



**• *- 









&% gluteraldehyde in Sorensen's phosphate buffer ai 
pH 7.2 unci at -V C and kepi m this fixative for sev- 
eral tlays. They were I hen washed three times in dis- 
tilled water and sonicated in an F.lma T420 sonieator 
Hi a frequency of 35 kHz for 10 sec or until shown by 
microscopic observation to be free of debus. 

The eggs were frce/e-dried by placing them 
between membrane fillers which were frozen rapidly 
by placing them in a slurry of freon cooled by liquid 
nitmgL-n. These fillers wilh the eggs were quickly 
transferred to a Irce/e drier and free/e-dricd at - 70 : 
C. The dried eggs were picked up with double-sided 
tape winch was ailaehed lo an ShM stub and coaled 
wilh 30 mil of gold. This material was examined and 
photographed in a Cambridge S 250 Mk 3 SPM 
operated at 20 kV using l 7 P4 Plus II ford roll film. 


Nimther\ ami Jtt't/ii)i> 

At die lime of collection the ratio of lardigrades Hi 
nematodes in 50 g of soil was 84:297. Heeding on 
nematodes was also observed during the course of 
this invesligalion and the nematodes held by the 
taidigiades usually did flOl move although once- a 
nematode broke \W<: and moved away. In one 
instance, a tardigrade was observed to be arching its 
back in the manner of a scorpion, during feeding 

/:,ee favin.K ami c,v/,'> 

Uggs that were about 90-100 fim in diameter in the 
living unfixed state (fags |. l\ umi about 00-70 jam in 
the fixed and dehydraied slale (fig. *)\ were laid 
eilhci wiffiiti ryuvia («;as| eiiliclcM ifies L 2) or free 
iPigi 3), figures 2, 3, 4. 5 clearly show thai these 
rges have a morphology of Ihe AY. luticlatnli group 
wilh reticulated shells and characteristically 
uptuincd-clial ice -shaped proLiuding piocesses. 

Although the pattern of die reticulations remains the 
same with the apc-nures on the reticulate surface of 
the egg shell being about 0.25 pm hi diameter (Fig. 
hi, the shape of the protruding processes falls into 
tvvu distinct groups. Type I is shorter than Type H. ls 
narrow at ihe base and has a wide distal head (Pig. 4). 
Conversely, Type II is taller, wider at the base and 
has a narrower distal head than Type I tfeg. 5). For 
10 processes of each type, the differences in ihe 
means are statistically significant with V5% confi- 
dence (Table 1 1 

When the proeesses weie examined mnWr the 
higher magnification of the SPM (Fig. 0). Ihe dentate 
coe-shaped margins of the head were shown 10 con 
sisi of clusters of globules with a structiue resem- 
bling madreporarian corals in appearance and were 
approximately 0.5 urn in diameter. The inudreporarian 
globules and reiieulated shell surfaces are similar in 
both the shell types described above. 


Bertolnni vt ai i \99fr) have proposed An hypothe- 
sis lo explain (he evolution of tardigrade eggs. 
According lo these workers, the eggs of lardigrades 
have evolved as a a\sull of two events, ihe fusl being 
the acquisition of ornamentation and ihe second Ihe 
use of the shed exuviinn as the site for egg laying. 
wilh Ihe subsequent loss ol ornamentation. Thus, the 
ornamented cge's of the Macrobiotidac are thought to 
Iv laid Uvc, Our obsctvalions that Ihe ornamented 
eggs of M. ci\ iK\('H(i(>hii(flitniIi can cither be laid in 
exnvia (Figs 1 . 2) iw free i Pig. 3) do not appear to be 
in accord with this hypothesis. The exu\ia containing 
ornariK-nied eggs were transparent and devoid of 
body eoiiteuh and did Hot appear to |v temalrs ilia' 
had tiled before completing <j^, laying. A possible 
explanalion for the laying of (Ik egys in Ihe exuviiun 
by lite M. cT. i>s<'iuh>httfiLtti<li from the soil ai Avon. 

I Mil I P MiUMiwrnenh t)f tin- /in- t\{>v\ nf ftmiVXW* ITeftittlliiy, fl'rtH itn <y,y sin If vttfmi* W VlacToblDUlf- I /, 
|isctulotiuietandi ( 


No. Height (uni) 

Mean + .SD Khih 

Basal width tUml 
Mf:ni + SD Ranee 

OMal Width lUm) 
Mead ± Sli^i 

fl) 5_M ll.i S.440 4..S 2 3.*J 4X 

lo 0.4 0,4 s7 7-i 5,3 Qi 4,0 5 u 

i |0.'*-|>2.2r> 



s\) - Slumlord Potation, i = Sludenf* nw 


I t 


I 2 -I 

Pifi. I . Pi!U of <\tm>fltbut1us d ffsahlf>fntji.hmt_/i in east l uncle (esuviuiiO. Arrows imlieaic position ul\.ist |eg> w\t\ claws. 

Briulu field uplics. Scafe bar = HJO |un, 
lay, 2 S.mie specimen ,u hiyhu m.mnilkMiii'n Miowinr 4k II punocutiii* I V) covered In cast cimkIc K'l. B rigid field opiie* 

Scale HW= -0 uiTti 

\. F. BIRD & S. G. McCLlIRE 

t*GGS OP MACM)ii?On;s 

is that this may be an adaptation lo the hot, diy SUJU- 
mers experienced there. 

Our siudies on the lardigrades and nematodes in 
soils from wheal fields al Avon in midwinter when 
the soils were wel and the populations o( the meio 
fauna could be expected to be at their peak, show thai 
latdigrsdas make up a substantial component, 
although they are not as numerous as nematodes. The 
ralio of K4 tardigrades to 2 l )7 nematodes per 50 g soil 
found m these experiments varies at other sites where 
the tardigrade numbers per 50 ti soil may be less and 
nematode numbers greater (Bird l ( »o). However, it 
is clear that the tardigrade presence al Avon is wide- 

The lardigrades isolated Iron) soil at Avon feed on 
nematodes and can survive hoi dry summers in an 
auhydrobiotie state (Bird 19% I heeding on nema- 
todes was also observed during the course oj this 
investigation and the nematodes fidd by the tardi- 
grades did not usually appear 10 be moving, although 
ihey were coiled and therefore probably not dead, 
.suggesting to us that they might have been paraly/ed 
by some type of injected narcotic. Since ihete is no 
inloiiiuilion on tardigrade diversity and geographical 
diMribulion in South Australia, further studies are 
warranted- particularly on their leedmg habils, as 
they may have a role in the bioeontrol of the parasitic 
netnaiodts which have been shown to occur in the 
wheal lields ul Uon (Yeaies #: Bird I W>. 

Tin* line ^iructure of lite egg $I1£|] Hi the fl&nuS 
M(U'tttbinm.\ is of great uiMmounc imporlanee. From 
oui observations on the uiirastriietiire of the egg 
shells o\ (he Avon lardigrades. it would seem that 
thete may be two populations of M. el. ftstthfohuft' 

c\- rsnuixvwrrfANDi $ 

huiJt in the Avon soil or there may be two different 
species, neither of which completely resembles those 
described so tar for the hiijehuuh group t Bet'lolani & 
Reheechi ($93: Biserov I996J- The reliculaled sur- 
face of the shell and the structure of the globules on 
the heads of ihe projections are similar in the two 
forms ^y\' eggs described above but the diameter of 
the apertures on the reticulated surface ol the egg 
shell is much less than that shown in die eggs of 
other members of the liufclaiirfi group described by 
Bertolani & Rebeechi <IW>. Kinchin iW4i and 
Biserov ( 19%). To our knou ledge, the glohules on 
the heads of Ihe projections have not been described 
before and might prove, together with the reticulated 
surface, to be useful laxonomic criteria if die hni^- 
Id", h group »s further divided on the basis nf egg 

Clearly further siudies arc required on Ihe taxon- 
omy o! these lardigrades and on then" distribution in 
the semi-arid agricultural areas ot South Australia 
and oilier sinnlai nitons i\\ ihe Australian comment. 


Ihe sept 61 author is grateful for a grant from the 
Australian Biological Resources Study which pio- 
vided facilities that made this research possible MMi 
rtJ CSIRO t.aud and Water Adelaide \~n( aieommodu 
lion and equipment, Dr W. R. Vlillei. Ueparimeni ol 
Biology Southwestern College Winfield Kansas 
USA and Dr W.I,. Nicholas. Department o\' Botany 
and /oology AMI Canberra ACT are thanked loi 
constructive criticism of the manuscript 


Uruiol \\l. K, <V- Kililt'in. I . (IW.M A tvsUinn ol' Ihe 
Mat minoltn hii(ri,ni<h croup ( laixllgrada. 

Mnerobioiidaei. wilh some olwivalions on ih<* ia\oiiom- 
m tlmi;ii.lcr. ft eUtal'itunidev A-o/, < nri(>hi 22. 127 

tV Ci moon. S. k. i \ty&\ Phylogcncltc sic 

taliLiiiKc t»* i_-pv< shell variation in lardigrauYs. /<**!, J 

tin w. lift, i W-l is. 
Kiki), A- F ( 1*996) Studies on lliesoil -jnluihilini: tardigrade. 

Mtirrohi.KtiK cf. p\<-nil>>htifi-ttttuti, from Souih AuMraliu. 

Intits. ft .W. ZAitit 120. U7-I54. 

Him Nov. V Y. i IWfi) laiihgrndes ol ihi- ttiimyr peniitMilu 
with descriptions yj two new ^pteies. An>l 7 Stn 
\\h 2\> 2H7. 

Kincuim, I M 
iPorlluntl Pre: 

i P»w4i Ihe Uiology 
.. London), 

ol TaoligraUVV 

Yi.ot.s. I. \V tV Brno. -V I. ( PW-lt Sonic obNeivalton.s on 
ilie inflaenee of agricultural practices on Ihe nematode 
faunae ol some Smilli Australian \oi!>. funtJtim A/tpt. 

MmaHfl- n. I WUs, 

I'iji V Scanning elcelmn niicro«irjph ol whole L-L'gs ul Miu-rohhHits el. iwtuh>imf<'litntli showing lire inverted jJohM- 

■shapeil pniju'eiioiis of Tvpes t tTI ) and if \'\'2\, Scale bar = 20 pin. 
l-'in a. Seaniiiiii* eleelion inierogiaph of pari ot' the fcgg shell surface of a Type I e^y >Iiowiml' pro|fcrnnis wnh haiTowei 

Ivises anil Uu^er dentate ecm-shaped heads Mom iho'-c olTvpe II. Not-.- similarly ivlieul;it«-Mj silrfticcs ol vu)i shells. Sc;itc 

lxn - 3 mil 




Fig. 5, Scanning electron micrograph of part of the egg shell surface of a Type II egg showing projections with wider bases 
and smaller cog-shaped heads than those of Type I, Note similarly reticulated egg shell surfaces. Scale bar = 5 jam. 

Fig. 6. Scanning electron micrograph of the head of one of the Type II projections at higher magnification showing the 
madrcporarian globules (arrows). Note the size of the apertures on the reticulated surface of the egg shell on the top right 
hand side of the photomicrograph. Scale bar = 1 um. 




By Peter Kolesik* 


Kolesik, P. (1997) Two new species of Asphondylia (Diptera: Cecidomyiidae) from 

Halosarcia spp. (Chenopodiaceae) in South Australia. Trans. R. Soc. S. Aust. 121(2), 

59-66, 30 May, 1997. 

Two new gall midge species are described from South Australia. Asphondylia inflata 

sp. nov. was found at Port Adelaide in swollen branches of Halosarcia pergranulata 

subsp. pergranulata. Asphondylia ericiformis sp. nov. was found at Lyndhurst, at the 

southern edge of the Strzelecki Desert, forming spherical, spiky galls on branches of 

H. indica subsp. leiostachya. Descriptions of the larvae, pupae, males, females and 

galls are given for both species. 

Key Words: Diptera, Cecidomyiidae, Adelaide, Strzelecki Desert, South Australia. 

Transactions of the Hoyal Society of S, Aust. (1997). 121(2), 59-66. 


by Peter Kolrsik* 


Kolbsik, P. (1997) Two new species of Aspliondxlia (Diptera: Cecidomyiidae) from Haiosarcia spp. 
(Chenopodiueeae) in South Australia. Trans, R. Soc. S. Aust 121(2), 59-66. 30 May. 1997. 

Two new gall midge species are described from South Australia. Asphondylia inflata sp. nov. was found at Port 
Adelaide in swollen branches of Haiosarcia pergranuiata subsp. pergrannlata. Asphondylia ericiformis sp. nov. 
was found at Lyndhursl. at the southern edge of the Stivelecki Desert, forming spherical, spiky galls on branches 
of //. indica subsp. leioslachya. Descriptions of the larvae, pupae, males, females and galls are given for both 

KEY Words: Diplera, Cecidomyiidae, Adelaide, Sir/elecki Desert, South Australia. 


Utdosarcia is a plant genus comprising 23 species 
commonly called samphires. The genus is endemic 
to Australia except for H. indica (Willd.) Wilson 
which also occurs in Malaysia and other countries 
bordering the Indian Ocean (Wilson 1986). Two 
species of Haiosarcia were found to be infested by 
two undescribed gall midges (Diptera: 
Cecidomyiidae) collected in South Australia during 
1996. These gall midges are described in the present 
paper. Asphondylia inflata sp. nov. causes swellings 
of branch segments on H. pcrgrannlata (Black) 
Wilson subsp. per^ranalata (Fig. 1) and A. erici- 
formis sp. nov. forms spherical, spiky galls on branch 
segments of H. indica subsp. leiostachya (BenthJ 
Wilson (Fig. 2). 

Haiosarcia pergranulata subsp. pergramdata is a 
shrub about 0.5 m high which grows in southern 
Australia (except Tasmania) associated with coast- 
lines, estuaries, swamps and margins of inland lakes 
(Wilson 1984). The plant forms a substantial part of 
the vegetation cover of saltmarsh flats north-west of 
Adelaide, These saltmarsh flats are areas covered by 
small, hardy bushes that grow on the landward side 
o\' the mangrove swamps. Areas that are regularly 
inundaled by tides are typically dominated by 
Scleroste^ia arhuscula and Sarcocornia tpnnqueflo- 
ra, while areas that are only occasionally Hooded are 
dominated by Mai retina oppositifolia and 
Haiosarcia spp. In May 1996, a large number of 
galls caused by A. inflata sp. nov. was found on //. 
pergranitlala subsp. per^rattalata at Port Adelaide, 
about 400 m south o^i the Torrens Island bridge. 

Haiosarcia indica subsp. leiostachya is a small, 
decumbent to erect shrub widespread along the coasl 
and around inland salt lakes of mainland Australia 
(Wilson 1984). It is a common plant in the Str/eleeki 

Fig. I. Gall of Asphondylia inflata sp. nov. on Haiosarcia 
pergnmnlala (Black) Wilson suhsp. pergtamdata. Scale 

bar = 10 mm. 

: ' Department of Horticulture, Viticulture unci Oenology Faculty or" 
Agricultural and Natural Resource Sciences. University ul 
Adelaide PMB1 Glen Osmond S. Aust. 5064 

Fig. 2. Gall of Asphondylia ericiformis sp. nov. on 
Haiosarcia indica subsp. leiostachya Wilson. Scale bar 
= 10 mm. 




Desert where it grows in a variety of habitats, includ- 
ing sail lake margins, open clay plains and gibbet 
plains, It k one oi' the dominant plants around 
Fyndhursi. where, in l-ebruary 19W\ all examined 
shrubs exhibited a low lo moderate infestation by the 
gall midge A- t>ri<ifi>rwis sp. no*. 

Material and Melhods 

(ialls were sampled horn Hnlusttrcm ituUcu subsp, 
lehistttchya at Fy ndhuiM 1 15.ii. 19%) and Halosiurui 
f)4'r\;naiul(HH Mibsp- (>t j n>ranultiut at Port Adelaide 
(5v.|00n). The gfilfe colleelcd en bolh occasions 
v\ ci l' processed in one of Iwo ways. A small number 
was cut open and the larvae preserved in 7(Kf 
eihauol, A Inrgei number of galls was kept in plastic 
bags, and the larvae Were reared to adults. Pupation 
took place wilhin tlie -ialls. Plastic bags were exam 
mod daily and emerged adulls preserved, together 
Willi their pupal skins, m 70'i tuhanol. Canada bal- 
sam mounts wf type Npecimens for microscopic 
examination were prepared according to ihe tech 
nic|ue outlined b> Kolesjk tF>')5a). All measure- 
rivals refer to type series. The type series and othei 
iniileiial rclained in 70';* ethanol are deposited m the 
South Australian Museum. Adelaide |SAMA| ami 
the Australian National Insect Collection. Canberra 

Genus Axphoiutvlin Loew. 1850 

Loew. IK50. Diptcrologische Beitriige. 1850: 21 and 
37 MW subgenus of fVr hinmvui, 1803) 

Type species: Cl'Lilh/fftyUi .surothumru loew. 1850. 
I.e.: 3N (des. Karsch. 1X77): Revision der 
Gallmuckeii: 15). 

\\f)h(/ud\1iu is a worldwide genus ihal currently 
comprises some 2fi0 described species (Gaguc 
loooj Ii contains species that have a ventro-dislal 
spine on the ffrjtl tarsomerc. the ovipositor with large 
basal lobes, female flagellomeres 12 progress- 
ively shortened, the guiioeoxue with a veitlro- apical 
lobe and a dorsal ly situaled gonoslylus thai is aboul 
as wide as long and bears two nasally merged teeth. 

Axpha/ulytia inflata sp. nov. 
U'IGS I. 3 5. 7-9. 12-14. I6-1K. 21.24-27,31. Hi 

Hi/fotypc: 4 . Port Adelaide. South Australia 13450' 
S. 1 38' 30' E|. emerged o.v.19%. P. Kolesik, reared 
from branch gall on H. pergrLinultiia (Black) Wilson 
subsp. iH-r^icmuiuhi.. gall collected 5.v.|0%, 121283 

font tv{>t j s: I J * . 3 9 V , 2 pupae. I pupal skin 
(SAMAk \r- V? V, I pupa_ I pupal skin |ANK'|. 
all same data but emerged 5.V.-1 3-\i.l l W6: 1 larva 
ISAM A I, collected with holotypc. 

Otht'f mutcnttf; 2V £, 13 pupal skins |SAMA|. col 
levied an ilh Irololvpc 

A/<nV (Figs 3-f\ 7-0. 12. 13 J 

Colour: seleivti/ed parts of body dark brown, nou- 
scleroti/ed parts o\ abdomen orange. 

Head: Anicnnn: scape broadest distally. length 1.5 
\ breadth at distal cm}, I b • l.x \ length pedicel, 
pedicel about as broad as long, fhsl llagcllomere 2.0 

2.2 \ length of scape, flagellomeres evenly cylin- 
drical, ctrcomfila dense, equally distributed along 
segments. Gyc facets close together, hexagonoid. eye 
blidge 6 - 7 facets long. Frojis u ith 12 - IX setae pei 
side. Labcila reduced in M/e. fused, laleraily with 3 
o setae, setulose. Mamillary palpus 3 segmented, 
length of third seumenl. as well as tolal length, sari 

Thorav. Wing length 3.s mm (range 3.4 - 3.5). 
width 14 mm ( 1.4 - L5|. Sc cell pigmented proxi 
mally. Anepisternuni and anepiineron coveted with 
scales- Claws of all legs subec|ual in size, similar in 
shape, as long as enipodia. 

Ahdomen: Genitalia: gonocoxiLe* short, with long 
veniro-apical lobe: gonostylus with 2 unequal apical 
teeth, ventral about 2 x length ol dorsal, aedeagus 
elongate and narrow. 

/r,W<(Figs 14. HV|K T 2F24) 

Frons wilh ^-11 setae per side. Twelfth llagclio- 
mere sometimes fused wilh eleventh. Ciivurnlila 
comprising two longitudinal bands conneciedby two 
.Tiort transverse bands. Wing length 3.4 mm (2,7 
3-7). widih 1.3 mm (Ft - 1.4). Seventh abdominal 
siernite 2.2 x (2.0 - 2A) length ol" sixlh. Genitalia: 
ovipositor 2,2 \ ( 1>J - 2.4 1 length ot' seventh .sLcrnile. 
cerci glabrous. Olher characters as in male. 

Fit-',-. ^ M $ HuihJ ol nuilf Asphondvhtt tttfltthi .sp. nov. .p view. A. Geniliiiiu ut ' »YdK- A,\f/ttottJ\tiu atfhiht sp. nOV. 
hi ilorsiil view. 5. Cnitioslylus of iiiali-' As^h^nJxIiu influx! sp nw\. in posiefior vifw, O Lioliusty lus i\t mt\icAst)h<)tuh'liu 
t-m *j<ntni\ :,p nov. at posiciiiii view. 7. I^fisl ihrcv mr^llamergs ol mule Aspiwiuhlia itsfktia -ip. nov S. Si.Mh flajiel 
kmtLiv ol male A,si>/t<>)ulyliti infhttu sp, nov '). Wttlt Asphuiulyiiit tnfltht $p. nov, I ;>st lai'som^re with cUtw and empiMti- 
uiu. 10 <*t I I . Maxillaiy palpus of iuul<: Asftluinilvlin t'nnfonttt\ sp, nnv 1 \ Hirsi laiHomere ol male A\f>hon\h'Ha iufhf- 
(ftsp, nov J3 Vfn\& \>j[ |TnUo Mphotidxlia mjUua sp, urn State burs= 100 jiiu3.4. 7-12: 50 urn 5. 6. 500 uin 13. 










Prf/w (Figs -Jfi 27.31) 

Colour: abdomen orange, remaining parts dark 
brown. lotal length 3.(1 mm (2 K - 5.3). Aniennal 
horns serrate medial)}. 185 Jim (141 - 191) long 
Upper and lower frontal horns simple. Two pairs of 
papillae on lower lace, one of each pair wilh a sela 
Prolhoracie spiracle broad al base, narrow an distill 
half, curved beyond Irucheal opening al mid-length. 
Abdominal scgmenls 2 - 8 wilh Iwo pairs dmsal 
papillae, two pairs pleural papillae, and one pair ven- 
iral papillae, all papillae seiose. Abdominal dorsal 
spines simple, prominenl pair on lasi segment curved 

bust inshtt iim'H tl ; ie_ 34) 

Colour: orange. Integument eovcted with dritvr 
spieulae. Lenglh ?_() mm Head capsule strongly pig- 
mented, posterolateral evtensioiis shorter than 
length of head capsule Spatula wilh Iwo long, point 
ed anterior teeth, shaft narrowed near middle, 
widened ;ii:;iiii posteriorly, surrounded anteriorly and 
laterally by e\lensi\e pigmented, glabrous a 
Papillai pattern generally as loi Asphctitivftu (Muhn 
1955) eveepi nuly _ lateral papillae on each side of 
spatula and III) terminal papillae visible cm the avail- 
able sperimrn 

t I Minting \ 

J he nam* ' inllatu" is a I aim adjective loi mll.ned 
i\lerrihg to Ihe appearance of die called branch. 

(uill and hit>ii>i:\ 

Hi.ntib segments ol' finh^curiu pn^ftitnilnni 
subsp. p( tinwtnlitiu infested by this gall midge arc 
- 3 times larger than normal in volume, greyish- green 
and hard in contrast to ihe vivid, given colour and 
soli texlurc of uuinfestcd brunch scgmenls (fljk 1) 
fetch yftll has <Hie to three ehanihciv wilh om: forva 
in each chamber, The chamber wall is lined *\iih 
bard pulr-gieem 0.2^ - [J33 mm thick tissue 

Pupation takes place inside Ihe gall. A eituilar 
brown area appear-- on (he lop of the gall hefou ilk- 
pupa cuts an opening with ils antennal httois. On 5 
May |00(>, at Port Adelaide, the ealls appeared veil 
common in the host plant population covering sever- 
al hundreds of in 1 . 

Asplumdstut chcifomm sp, nov, 
(FIGS 2. 6, HI* II, 15. \<). 1ft, 22. 23, 28-3U 



Hirfotxpc; i % Lvndhurst. Soulh Australia \MY 17' S. 
I3cS [I 2I' B|. 2ihiUOU„. p_ Kolesik. reared from 
branch gall on flnh>smrut indica subsp, lao\huh\(\ 
(Benin.) Wilson, gall collected l5.ii.l°-%. I2I2H4 


Parnivpi'.w I :'. IS S?, I pupa. I pupal skin |SAMA|. 
I ■', I £. I pupa. 1 pupal skin [ANiC|, all same dal.t 
but emerged 10%: 3 lurv«ie |SAMA|. 2 lar- 
vae | \\IC|, collected with hololype. 

fJj/tcrtmiti-iifiLl pupae. 4 pupal skins |SAMA| col 
lected with hololype. 

M<//r(Figsb. 10. Mi 

Frons wilh 6 - N selae pet side. Wing length 3.4 
niin(2.l - -1.1). width 1.3 mm ((I y- I or I eye bridge 
- 9 fucei.s long, Ventral tooth on gonosiylus as litrig 
as dnis, ( | h Otherwise as in 1 inflaui 

/<WeiFi«s 15. I y. 20,22. 23) 

Frons wilh 4 R setae per side. \\ ■'nig lencih 3.4 
mm i2.3 33>t, width 1.2 mm (0 ( > III. Seventh 
abdominal slcrmle 2.3 a i2." 2,5) length of si\(h 
Ovipositor I \ 1 1 .X - 2.0) lenglli of seventh sleruile. 
ccrei glabrous, Willi line ol teeth dorsally. Otherwise 

^ in .v Htffttta 

/'H/w(Figs2S -30. 33J 

b»lal length 4.4 mm t4.l - 4,7i, Aniennal lv>m- 
|0S firn ( 154 - 214) lone, I 'pper fronlal horn simple. 
m* othci horn present. Dorsal spmes tafias! abdomi- 
nal segment about same leneih. straight.. 

Uist tnstur Ittnn tFies 33. 35) 

lolal length 3.2 mm <2.4 4.S). SpaUila with lone, 
pointed anterior teeth, shaft short, broad, parnllcl- 
sided. Three lateral papillae on each side ol thoracic 
segments, ftp terminal papillae visible 0)1 iivnilahle 
specimens. Otherwise as in A. infhiln. 


The name erieiloruns "" is ;i eompusrd I .aim adjec- 
live formed from "erieius'* (hedgehoyj and "formic 
referring to the the hedgehog-shaped gall. 

Cmll and bmtrt#y 

I hi s species transforms branch scements *y| 

lius 14-24 14 Female \\/.'ln'iuhlm irtlhttu sp. nov. Oasul I«>(ks on >.»vi|>«>siloi in tlwi-Sjd view 15, Female Asphnihlylia i-ii- 
iiji'iitth sp. llov. Masai lubes ol ovipositor ill dorsal vnw, tt\ Antenna ol fenmle A\pltniulylia tttjlnla sp_ nov. 17 .Si IN. 
VJ.iVillary p.ilpits Of female Ast'honrfylia infltttn a? nov. F) iV JO MaAilltin' palpus ol Ifmale <\\{rhondytt(i trit ih>n>it\ 
sp, nov. 21 , Si.vtli I'layelloinerc ol tcmak* A\plt'>n<fxli,i mfh/tn sp. m>v. 22, Female A,\f>lnnuf\liii <:ikijonui\ sp. nov. hinl 
»>l abdomen in lateral view. ?.V I'cmale AsphorulSlhi W?'i tjorwU fli. ft<A. End ot "ovipositor in lateral view. 24. Female 
As(>lu>tul\lui tttfhtfo ftp. nov. End of oupusilor in luteial vie\v_ Seale bars = 100 pm U-2F 2.^. 24: S0O pin 22. 



kA- ■< 




Hulosurcitt irulkn subsp. h'hiskichxu into spherical. 
Spiky, monothalamous galls, each occupied by one 
larva iLig. 2). Oilier Jiuiiitfier oi g$(] 0-12 mm. 
inner diameter 2.1) - 25 mm. Chnmher wall lined 
Willi haul, brown, 0,15 - (1.33 mm thick tissue. 

Pupalion lakes place inside llie gull. On \S 
behiuaty, 19%. ill Lyndhurst. 10 examined shrubs ol 
the host plum bore a tola! of about 2<H) gulls of the 
new gall midge species. The galls contained larvae oi 


hour species ol \ Asplwittixlia have been previously 
known 10 "ccui hi Australia (Gagne 1\W); Kol<"'k 
W5bk Asplumdxlia d<»doihhitt j . a Soulh Australian 
species common in ihe Adelaide Hills, mal forms term- 
inal brunch Steins and primary leal veins ol \' Dorfomua 
ri\at,\ti (Sapindaccac) (Kolesik 10*J5h> Two species, 
A, /ecu/and A. lubttindu, were described in llie pre- 
vious century from adnlis caught in flight in Sydney, 
New South Wales, and their biology is unknown 
(Skiise ISSS, I MO). The fourth. A. hilii. was 
described from females and pupae bred fiom .m 
unknown plant in Darwin, Norlhern Territory 
tLdwards l l Jlru These las! ihice species, which weie 
described superficially and can nol be compared on 
their descriptions', are not considered in the present 
papa, bul I plan a review of Australian A.spiiendvlttt 
spp. at a later stage. 

Morphological similarities between the two new 
•pci_us and the fact thai their respective hosi 
plains belong 10 the vnrie .uemis. suggest a i low 
relationship. They lorni a distinctive group that 
excludes A. ih'dtnuu j ur \\phan(Jylitt dtnttHUH'm 1 
differs from the two new -.pceics in ihe following 
ivspviK. AdulUhave prominent labclla and scapes 
as long as broad at the distal cuds. The mule lias j 
shori a wntro-apical lobe on the gonocoxile. suudl 
lolv% adjacent !o icclh on ihe gonosiylus ami the 
aedeagus. much shorter than the gonoeoxiies. The 
seventh abdominal Sttil'JKllc ili On? fetmlla ''•■ ilaee 
Iiiiils longer than the si\lh. The pupa ha* no 
frontal bonis, the antennal horns arc triangular ami 
set rated laieoillv mid Ihe prothcuacic spiracle is 
iioi i-oosulcialilv bnurtlcrtn. ihe base. The uicj sur- 

rounding the spatuln in the larva is not pigmented. 
Adults ol the two nevv species differ from each 
other most prominently in the shape of the gono 
stylus and Ihe end al^ the ovipositor The gono- 
stylus bears teeth of unequal length and ihe end of 
the ovipositor lacks external processes in A. inf'ht 
lit. In contrast, the gonosiylus of A. criciformis 
bears teeth of equal length and the end of the 
ovipositor is serrated. More differences arc evi- 
dent in the earlier developmental stages. The pupa 
ol A. iiiflntti has both upper and frontal horns pre- 
sent and a prominent pair of dorsal spines on die 
lasl segment is curved laterally: in A. cncijhnitis 
the pupa has ihe uppe^ horn only and all dorsal 
spines on the lust abdominal sermon arc equally 
strong and straight, The larvae differ m the shape 
of the spatula and the numher of lateral papillae 
two per side in A. itijle/u bin three per side in \. 
crtt 'Ifontti*. The two new species resemble each 
other in the shape of the antenna! horns and pro 
thoracic spiracles m pupae, the antennal segments 
in adults, the veniro-apical lobes on Ihe gonoeox- 
ites in males ami the relative lengths of the sixth 
and seventh abdominal sicruiles m females. I In. 
elongation ol" the ventro-apical lobe on the gono 
eoxile is unique its these two species and disting- 
uishes iheni from the oilier Asphondylid spp. 


I am grateful to R G Wilson, Western Australian 
I lerbarium (Anno for Ihe identification ^\ Htdosumu 
anluii subsp. h , it>swrhxu. R. J. C'hmnock. Soulh 
Australian Herbarium AdeJarde tot the idenufication 
ol Hiilo\ttrcnt pi'tyjutnultini subsp pcixromdati^ 
I, B Rcaulon win* led i South Australian Museum 
collecting trip during which ■Xyphtwdylia crii-ifcrinis 
sp. nov. was discovered, '\. Stark. Malic Germain I'm 
providing copies of Karsch's and Lock's papeis and 
J D Gray, Department o( Horticulture, Vib 
culture and Oenoloey "'iiivcrsity of \delaide and 
R. 1, Systematic Entomology Laboratory 
ILSDA Washington IJC for then comments "n an 
eail\ dm ft of the manuscript 

I !•■ W J ^. 25 ■\\jfliniuh(iuwifahi*\} ii'V . Amkmum part ol pupa view 2ti, \*}thoiu{Yli<i mfh>t« >.p ii"\. Autcrioi 
ran iH \m\\m\ in laieo.1 \ lew 77 A\fht"nl\lta (nffom sp. liov. I'ruihoracic spiiacl- ^1 puiui 2*. \snfwiHf\ha i J ^f< flvwfts 
sp im\, Anicrioi p;iri el pap.i En \k\s, J 1 ', Ait/ffaimtytiil I P^f/f'ffOW s l>- nov. AmcrMH inn! ..I pupa m luU'val vjrvv. 
\\), \y(.'/t<?/ut\h<t i'nnlniwt,- sp. nov. P/rith^ruc'iC 'pirack' ol pupa. M I'upa o|* .\^>it.'H'lylio lufhthi *p nnv_ I ;tsl aluktui 
inal ItugniCUII 111 ilorsal v i*_-\s , J ~ Pupa "I AApAftttitttfR cnvtfonius SP nUVi I H^l ahiloir.niitl ^^^itienl m ilor>.al view, l.i 
I ;iiva "I A>!>!i<>tuhli(t i'rhiftWnk *P< nOV. bast [AMU ubUvinitHtl secmniK h< ilttrsat view. W. \nlefinr pair ortaivuol 
MpfttUttiylOt ttifl'lte sp. rtl'V. m vemral view 3 r \ lit ad ati.l l'n\l Itmraeie se^aienl nl taiva ol \.sph<i/u(\ini rti< IfrOWto sp 
nov. in VL'itlrat vRm-v Vale ftur4 : I(M»|Hm 



Edwards. F. W. ( 1916) Two new Australian Diptera. Ann. 
Mag, Nat. I list. 103 (8th Scries, Vol. 18), 498-502. 

CiACNh. R. .1. (1989) Family Cecidomyiidac pp. 152-163 /// 
Evcnhuis. N. L. (Ed.) "Catalog of the Diptera of the 
Australasian and Oceanian Regions" (Bishop Museum 
Press and R.J. Brill, Honolulu). 

I 1994) "The Gall Midges of the Neotropical Region" 

(Cornell University Press. Ithaca New York). 

KARSCH. F, A. F. (1877) "Revision der Gallmiieken" (E.C. 
Brunn, Minister i. W.). 

Kolhsik. P. (1995a) A new species of Eoeitieticortiia Felt 
(Diptera: Cecidomyiidac) on Eucalyptus fasciculasa in 
South Australia../. Austr. cut. Sac. 34. 147-152. 

(1995b) Asphandylia dodonaeac, a new species of 

Cecidomyiidac (Diptera) damaging leaves and branches 
of hop-hush, Dodonacd viscosa (Sapindaceae) in 
Australia. Thins. X. Sac. S. Anst. 119, 171-176. 

Loew, H. (1850) "Dipterologische Beitrage. Vierter Thcil. 
Die Gallmiieken. Zu der otTenllichen Priifung der 
Schiller" 1 (Das Konigliche Friedrich-Wi I helm- 
Gymnasium zu Poscn, Poscn). 

Mohn, E. (1955) Beitrage zur Systematik der Larven der 
Idonididae (= Cecidomyiidae. Diptera). 1. Teil,: 
Porricondylinae und [tonidinae Mitteleuropas, Zaalagica 
105, 1-247. 

Skuse, F. A. A. (1888) Diptera of Australia. Part I. Vvoc. 
Linn. Sac. N.S.W. (2nd SeriesJ X 17-145. 

( 1 890) Diptera of Australia. Nematoeera - 

Supplement 1. Ibid- 5. 373-412. 

Wilson, P. G. (1984) Family Chenopodiaceae pp. 81-317 
/// George, A.S. (Ed.) "Flora o( Australia." Vol. 4 
(Australian Government Publishing Service. Canberra). 

( 1986) Family Chenopodiaceae (Dysphaniaceae) pp. 

236-3 1 1 In Jessop, J. P. & Toelken. H. R. (Eds) "Flora ol 
South Australia". Part 1 (South Australian Government 
Printing Division, Adelaide). 


By J. B. Jago*\ LwTiAN-RUif, G. Davidson$, 
B. P. J. Stevens^ & C. Bentley* 


Jago, J. B., Lin Tian-Rui, Davidson, G., Stevens, B. P. J. & Bentley, C. (1997) A Late 
Early Cambrian trilobite faunule from the Gnalta Group, Mt Wright, NSW. Trans. R. 
Soc. S. Aust. 121(2), 67-74, 30 May, 1997. 

Trilobites from a new locality within a siltstone of the Cymbric Vale Formation, 
western New South Wales, are described here as Redlichia cf. ziguiensis Lin 1978 
and Hsuaspis cerastes (Opik 1975). The species described as Strenax cerastes Opik 
and Estaingia bilobata Pocock from nearby localities are included in a single 
redefined species, H. cerastes. The genera Pseudichangia Chu & Zhou in Lu et al. 
(1974) and Strenax Opik 1975 are placed in synonymy with Hsuaspis. The fauna 
described here is of late Early Cambrian (Late Botoman) age. 

Key Words: Cambrian, Trilobita, Australia, New South Wales, Hsuaspis, Cymbric 
Vale Formation. 

httitMu iit'if; ofihf jtoyuUhHWty tf/S. \ttst * IVV7). 121(2.), 67-71 



by I IT JAt.ii', Lin Timm-ru 1 , fi. Davidson . M. R J. S'n vkn\« & C. Rinii i v 


lArtO, -I IT. L(N TiAK-Kll, DwiosvtN. £r_, Sn.viw, B. P. J. ,V lilMlfl A, ( '. (IW7) A. L;ilc F»nl\ Cambrian 

inioh,!,- I'uunuiti ItomlbrUwilUt Group, Ml Wri^ NSW, ftw^. A. ft*-. & Artri Q|12')67 74, jtOMuy, W7, 
Iriluhhes from '" U*w locality within a silicone ul ihe Cymbric Vale l-ormaiiun, western New South Wales, 
an- ilcscribeil liCD! us Hi'iHii iiui el'. ziy.itrrn\is I "in l')7X ami ftsua\fv\ trni.Mts (Opik IQ?5)- The species 
described US Snviu/\ anisfi's Opik Ulttl f.*htith,'jii hilnhdut Pocoek from nearby localities ait included in a single 
trdebiicd species. // « m/OrA. The ;_vik-t.t I'mulIx htiH^m ( Itu & /hull lit Lu fl ( // ( 1974 > unil Strung Opd. 
IU75 arc placed in synonymy uilh H\n t /\fn\ (he l;»nn;« itesi ribed here is o| late I ao\ ( 'ambium (I .ale BolOmtlD) 


K) v WitKDs; Cambrian. Trilobiia. Australia. New Small Wales. Hsiuispis. Cymbric Vale* formation. 


Davidson discovered 11 small tuea cii fossilifcious 
sdlslonc of ihc C'ymhrie Vale Formalion in the Ml 
Wright area, western New Sou»h Wales (Tig. I). 
Davidson idenlilied the trilobitcs as Slrctiax cerastes 
Opik and Kxtuittgia hilwbtilu Poeock of Dpih 
t 1975b). More samples were collected recently and 
the results of the examination tiflhp fossils are given 
he low. 

l lie fcurly to early Middle Cambrian Cinallu Group 
comprises three formations, from bottom to top: die 
Muiini Wright Vokanies. the Cymbric Vale 
Foimalion and the Coonigan Formation. The disirib- 
ulion of these formations |tt shown in Fig. I is altei 
Wairis" anil Rose ( l°oN). 

The (.Jnalla Group erops out along a broad valley 
west of Cymhiie Vale homestead and is mostly eon- 
lined by the Ml Wright Fault on the cast and die 
Lawrence Fault on the west. A smaller area <tf 
Cynihne Vale Formation crops out in the core of an 
anticline west of Ihe Lawrence Fault The structure 
of the Gnallu Group is very impeifecily known a.% a 

lif|MilmciH vl Anpllul (it'Ok^'v. Stlun'l H [.nuirnvriiij' 
I in, . in, ,.i SmhiIi AiKlnili i Ilk I . v» 18 S AlKl SUtf?" 
l>r|i;iitiHL-nl nl ! .inli Sen 'luvs. \.in(in;' tltiivvr.ny -- H;mkon Na.iiiuj.' :iiins People'*! Rcpilbiiu nl China. 

ucnii>^ ucpAitiiiiSH. t ni\\ .-I fnsiminitifipo iio\ :s: 

Mniuri lii^. 7I)I)L 
■ (tei^njUOBl Siirws !•! N.-u S;itilli W'ali"-,, 33 Snlphidi. Mitel 

UimVch Mill NSW 288(1, 
1 I (AVttJMW (i * l')KM A t'oiiliilmlnxt lo the jH-nlogv nl ihc VI* 

Wii;jIh aici USc HKmis) iIk-sIs. \usirallan NutHuial t hivcimIn 

(uiipuh >. 

VV U-:i:i',. li..I. < »*J(>7> *rht + sliaiiurapliy ami piilai-onluloMv of 

lIontlWCMCIO New Stuilli Wj.|l*>. PhD lli'.-sjs, S > dtK_v I i)ivl-i-i'.> 

(iilipnh I 

result Of gCOlt^ICftl et'inplcxily (faults and folds), 
poor outcrop in much o( the area and a lack oi bed 
dilljj ni die abundant vokanies of (lie Mt Wuyht 

The siltstiHie to fine saiuixloiie from whieh 
Hie fossils were collected (Fig. h is isolated from 
oilier outcrops by a cover of soil and cobbles repre- 
senting a Caino/oic lay deposit. Hence there is no 
outcrop continuity with any identified Gnalla Group 
formation. The outcrop also cuiitains 110 diagnostic 
rock type. The structure ol the Cmalla (iroup is suITt 
ciendy eomplev to remlerit impossible loeonlidcni 
Jy place the outcrop in a formation. Davidson placed 
this and nearby outcrop*, ill the Cymbric Vale 
I'urmatiiin, while Kruse ( IS)S2) plaeeil the nearby 
outcrops in the CooniL'an horinalion, As shown 
helow, the species described by Opik ( 1975b) fruin 
the C'yinhrie Vale Foiination as i-^ntin^in biU>hma 
and Stn'fitn r< :ru.\tr\ are found at die locality being 
considered here. No Mkh tussils are knuwn lioin 
either the Mt Wriyhl Voleanies 01 the C'oonigan 
I'ormation, Itenee. the outcrop w probably Cymbric 
Vale Formation. 

Previous Work 

the teim Cmalla Group was introduced by Warns 
who assigned an Barly to e;irl\ Middle Cambrian aye 
10 the group, from limestone in the Mi Wfiglti 
Voleanies. Warns identified algae and archacocy 
atlnds. including lluikiuuu \<ithii\ fn/i'liculis Taylor 
From the Cymbric Vale Formation he identified 
/ \Uitnxitt hilnhoHt, Ctflihlist MS sto, and t J a^vii</r\ sp, 
and assigned u middle or late Larly Caiubiian age. 
He idcnlillcil the trilobilcs ftvilltchm i'dOrtcud. 
\xstridunt smut sntttlu. Piiyt'int .st^nijhuus. 
f'c'ionapsis nottfhifu. OrYcHncplutlits sp. and 



pdktt£dic !Ai4iBluri>&. UlulUiJl 6 ajliatV B3iill&tuii£ aluflte, 

Lal= Oevci jn- '"^lim.-i.l jrOUS ftUtinilAlu t-L'rrr.jli' ir, bllfl 

f t — yliiy ■ . | ,- r ,. J|ir r . n»'.i'.|-rr.firj(omnrTitB 

■'.--•..■ - ■■-■.■■ ---■ J, . -- .■- 

i .!,!■,■ - Mi'ii!i.-,'"j t.A'ici itiltvwcna F«niis.troti; 
sgtlttitaUi. '[i.i invlfl sill'jItyW 

! in i : hi th'i rijfU ''V.i, ,■,;,- Wl (■ F&UUtaldrifl 
: inrfsM i I' 'Tii it. 'm • ., ltd -.r-.riit-. lirnSEl 

-- ■ ■ . 

. . .--. •-. j 

IfltfiiltnCjpiitnt.-wiO.npn^an » arniaitan. 

i #s r ari"i Gjirtng/] E ■■> il. il V^Ib F wnrfllbri- 

Li ' \-u ,| ■:, -|.i|.', > .It'.'.' in',. || H'Hl 11 tl ',"->li'H'|i- 

)i , a H i : n */i i italiiaTiTaEi 

i 1 ; i '- ?nd !■ r vote ittle: m'ncft'aftsib-IWwitoHBi 

.... . i j i "■■' ('■ nil . Ij, ; lI'. . niluomJ ri!- in 'tn/t"j ii i. ci ■■!*.■, iniil..l/n.7.,it 
I „ ■■ ,,..■ , It- -f >l> J'" ',. i- ;|.H'|r. i 'rtui IBSflhW 

11 .- . i.\ -v- - ■ , ,-■-.-.. <VV\V*»<f**^ 

. i i I-: r'v in .-!'''■, '* K»iWt*lK 

, . .1 i I. iVi li ii,ii(I hi|i lyi riiV.!.'*! 

■ J ' :: j 

i hi. .in Mr • li,' 

■ n |i 'i i n Wi -..'■•n li i.,v-. i 



'..,„ tilpudH-J Mill*,, s 


■ ML'KHjtjttJuS 


* '.\al -'i- 

Mil! us 

• 1 C-Ml Ll 

Defypyw sp. from the Cooni^an Formation, indicat- 
ing DM early Middk- Cambrian age-. 

In terms of the Cymbric Vale Formation, PlfrtchOt? 
(1964) first recorded archaeocyalha Irorn lenticular 
limestones near ihc base of the unit. Kruxe i 19823 
noted fourarchaeocyath limestone -bearing lenses: he 
considered t\vo 10 he ol Aldabauian 01" Lciian ace and 
I wo tt) be ol' Lenan oge The uppermost Cynibric Vale 
Formation coniains well-bedded sandstones and qur 
bonules with abundant fossils. Opik ( 1 075b) record 
ed 1 1 it* (n)obj(e's Pint \<t\ Ml, vratuiUtsus, P\iti'tt}ut<i 
hilahata. Snvtwx ceww.v. S. fU'lihtn, Scrrodtsm.s 
i///cif ilns. MfniM at hits nn'twtits, Dim t'lncMti a 
frayjitn ami Payytia, die moiioplaeophoian SivncUu 
ivtittila/a. il»e iitarlicLilale bractiiopods liotsfovdia cl. 
taclahi. Lftf$Ht£ll<t iind :V<nht>hi\", <ot nnnid'- anJ 
sponee spicules. 

Vjlt* and Correliilion 

As shown below we consider Ihul the >}>CCi^ 
described hv Opik ( I^Y-Sh) as Sttviiuv tt'/tixh's .uid 
Lsnwtxit! hihtbatu inun jiearby lueulities should 
bivlb be iueJuded wiihin a single redciiocd specie^ 
H\nn\f>j\- (.n-(t\ft\. These lin; ( tlities Op\W considered 
to bcct|uiva1enl to the Sanas1ilyk"c*oi llori/ou ^\ ilie 
AhaySayan region which, its noted by Zhurnvlcv 
and Gravestock (l l .) ( ^». can he roughly unrelated 
with die Boioinan of Siberia. The archaeoeyalha 
liMid lite C \oihr*c Vale 1-oiiiKition which occur 
below Ihc Inlobites desciibed bv Ofiik (t l >75b) were 
correlated with the infonnal archaeocyalhid unit I he 
S\nnvtirnvm<i fttvtis beds by /huravlev and 
Gravesiock (1W4J which w r as considered tt> he 
equivalent to the middle (0 Eate Hotoman by these 
authors. These authors considered thai the 
Syntii>thiH'ina f/ivus beds fall wjlhin the Irilobile 
based Parttmin /V///ae/oue of Jell (in Ben^tson ci at 
N ( )0). Jell suggested that the P. janae /one cmy be 
correlated with the Boioiiiaii and prohablv with du- 
late Bototnan (see alsu hiiel" diseussi.m tiy J40Q 
lft*)6), As noted above, the exact slrati^rapluc posi- 
tion ol the present fauna is a lillle unclear but it prob- 
ably oeeuis hiyh m the C'ymhrie Vale Formation and 
stratigraphically close to the I'aunas described by 
(^pik ( I075h)_ The Coouijtuin Eormaiion which ovei 
lies the (.'ymbiic Vale t-mmation has a verv each 
Middle Cambrian a^e (Opik |973tti Jell |*»75j 
Shergold el ai I9K5). thus providing an upper limit 
lo (he aye of the fauna currently beine described. 
PallUC! and Kowell (19951 note that Hsuaspis is 

Pig. I GcuKtyv i>t the Mi Wri^hi aiva sheiw top duality 
of t'ohSi Is describe J herein Geology Irom W.'im%'- Rtise 
( |V(iKU Davidson'. KfoHq (E>K?i. U. Slevens ami 
A. Eiawlool luiipith. ti^ttJ ttaial- 


However, its si>**j_ outline, slende 


closely related to (Ik* Siberian genera Ber^'tnniid'us 
and Hetwreniaspis which characterise the Botoniaii 
of Siberia. 

In China Rvdlitliia TJ^idensis occurs in the laic 
I. arly Cambrian Palavolcnus Zone of the Yangtze 
GocgC Tiiis is approximately einiiviilenl to pan of 
llic Toy ouiun of the Siberia Platform, 

The above discussion suggests that the faunulc 
described herein is of late Early Cumbrian aye and 
probably ol' Late liotonutn Bgfr, 

Systematic Descriptions 

leiiiiuiology csscuiially follows Harrington et at 
( I 1 )!*))- Specimen nnmlvrs refer lo the palaeontolog- 
ies! colleclion n\ die SoUlll Australian Museum 



KedhcluidaePoulsen IMS 
Redlichimae Poulseu 1927 
Kcdhctna Cossman 1902 
Ki'fliithiit it. zfkufetTXis Lin tV7H 



I97K Ki'flliiltiii ;ii!t<ittni\ 
p, |4?>H 20, r%* 

in in /hou & Lin F'^x, 


One incomplete craiialium, SAMP35344. 


Cninidium -dibreciangular, ahotu 35 mm long 
Cdabella (including occipital Ptlig) about OvB length 
of cranidium. Glabella tapers evenly forward to 
rounded glabellar anterior. Narrow shallow axial fur- 
rows. Three pairs of lulcral glabellar furrows; I p. 
moderately deep, directed slightly to posterioi and 
ineci at centre of glabella to form an evenly posteri- 
orly arched furrow; 2p pair, shallow, directed slight- 
ly i" posterior; $p pair, rcpivseriled by lauil depressions 
on glabellar margins- Occipital furrow deepest later 
ally; shallow medially where il is arched slightly to 
anterior. Anterior border furrow o\ moderate depth: 
anterior horder. very gently convex, and of uniform 
Icugdi (sag.|, Very short (sag I preglabellur Held. 
Small anterior areas of fiyigeoac gently convex, lojlj 
wide palpebral lobes extend from ju st to anterior ol 
3p furrows to level with occipital furrow. Narrow, 
distinct palpebral furrow shallows posteriorly. 
I'alpehiaJ areas o( fixigenae very gently convex. 
Width of palpebral areas of fixigenae at their widest 
is about one-third that ol 'glabella. Preocular sections 
of facial suture markedly divergent. 

l)is< Hssien 

A single incomplete internal mould of cranidium 
makes a definite specific identification difficult. 

glabella, the posi- 
tion of the posterior of the palpebral lobes, the nature 
pi Ihe preglabellar area dm\ Ihe shape c\i Ihe preoeu 
Jar section o\ the facial suture suggest Redtichia 
zi^nietisis Lin \iu Zhou & Lin 1 978, p, I4x PL 20, 
Fig. S|. However, it differs in having a moie tapered 
glabella, a forward arching at the centre of the occip- 
ital ring, and a more shaiply rounded glabellar antei- 

Supei family Lllipsoecphnloidea Matthew IKK7 
family lehangiidae Zhu I9B1 

CeuU-s H t stui\pi\ Chang PJ57 

HsiHisprs Chang l l ^7. p. 45. I.u w uf„ \%% p. S5: 
Zhang ej a!., 19S0. p. 244; Jell /// Bengison et /*/.. 
\Wi p. SHfc Palmer ami Rowell. I 99\ ,,. |„; Ncdui 

P>9_Vp. 3(j 

J\taiin;ia Poeoek 1964. p. 4b2; Opik I975h, p. K). 

A'v tufirfhtiwitt Clui & Zluni in l.U et id., 1974. p. 93: 

Zhu in Zhang et al, I9S0. p, 2frk 

S>n; ia \ Opik 1973h p.13. 

/Jnaielhr Zhang & Zhu in Zhang et uf.. 1980; p 247. 

Tvpe species: £ttffoft>f*J flflWti < bane, l L >>* p. 

\1K PI 2. lags 1-16. 

lell {in Hengtson i'f /</. IWII| placed i-\iuim>iu 
PocoeL 1964 and Zfwxit'tUi Zhang & Zhu {in Zhang 
( j < al. 19801 in synonymy vvHh HsHospis Chang K>57, 
a move supp»>rted hy Palmer & Rowell (l l ) l J-"i). 
although the latter authors expressed some doubt as 
to the suprageneric position oi Hsutispi*. In aoldilion. 
Jeil suggested that Siivmu Opik I°7,S might be 
regarded as a junior synonym of PseittHchanxin Chit 
& Zhou in Lu et a/, 1974 frotn southwestern China. 
The present authors support Jell in placing Snnmy in 
synonymy with Psrutln hanyia bur also consider that 
Pxenriichattyjit is a junior synorry m of Hstui\i}is, thus 
also placing Sftcnu,\ in synonymy with tfoiiOSpfti. 

The lollow'ing species of Psetnlit htin^ui as figured 
in Zhang ft ttL I9S0, f\ t/tunUn -h'um.s (Chang) {PI. 76, 
Pigs 13! 14; PI 77. J-sl R ^t{ivh4$ Zhu (PI. 77, 
Pigs o-S), f\ J ft *.\« f iMs Zhang & Zhu (PI. 134, lag. 
2) and PseudichanKta ( .'1 inwrfithi Zhu. PI. 77. Figs 
9.H), may all belong in a single species, i.e. Hsiunpts 
diwtittneasis (Chang). ZhnxieUo hohiinsis Zhang 2k 
Zhu (see Zhang ant. HW). PL 134. Fig. 3) also 
appears to belong in H. dam fatten sis' as far as can be 
determined from the available tlgure although the 
glabella of damianetisis extends further forwards 
than that Oj'the type species ol \ Hsaaspi\, If. sinensis, 
and the anterior sections of the facial -iiitures of 
sinensis are more divergent than those of datniat'en- 
sis. We would regard these* as specific latlier than 
generic differences. 




-' *>: 


.: -^ 




Fig. 2. A. Uedlithia cf. z'tquictisis Lin I97K, SAMP35344, cranidium. internal mould, xl.5. B-L Hsuaspis cerastes (Opik 
1975). B. SAMP35329, cranidium. external mould. x2. C, SAMP35340. cranidium, internal mould. x5. 
D. SAMP35335. LM-unidium, internal mould. x2. E. SAMP3533L cranidium, external mould. x2. K SAMP35343, crani- 
dium, internal mould, x2. G. SAMP35337. cranidium, internal mould, x2. H. Top. SAMP35327. cranidium, external 
mould. x2. Bottom. SAMP3532S, cranidium. external mould, x2. I. SAMP35342. cranidium. inlernal mould, x2. 
J. SAMP35333, pygidium, external mould, x6. K. SAMP35332. cranidium. external mould. x6, L. SAMP3.S341, par- 
tial thorax, external mould, x2. 


Jell \'w Bengisou W of. I UL M0 queried the taumom- 
it. position of L\fttin»ui hilt>hntd from western New 
Soulh Wales us described hy Opik | WJSW), Jfl Ottt 
VWW die specimens described hy Opik 0975b> Jo 
not belong in Hsuaspis hilnhalti iPocoek 1464 1 
rM cause ihe glabella of Opik's specimens extends 
IlittlK't to iho anterior ih.m does lhal * ■»<* //- hihthaia as 
deseitbed hy Poeoek (1^64). In addition, 111 three of 
Ihe four cramdia figured by Opik. there is a marked 
torwatds expansion of the glabella whereas in the 
type material ttf /Y bilahtiUt figured by Poeoek 
| I 1 'M| i hi* glabella is either tapering forwards or has 
a >he,hl vvaisi, hi our view ihe specimens described 
and figured hy Opik ( P>75b. p. I [ PL I, Pies 1-7) 
belong in die -.ante specif as those described hy 
Opik (1975b, p- 14. PI. 2. Pigs f-fij as SV^.M 
rruisit v- Apart from (lie specimens llumed hctcin. 
iluittetous othct specimens are available and rljeic is 
a complete gradation [htifl specimens such as those 
figured here as Pigs 213. K which are quae similar to 
Ihe cranidium illustrated by Opik ( 1975b. PL 2. P»g 
] ) as Ihe l)n|oty pe of Sftvttux it rash S .geil. el sj>. 0* »> 

10 specimens such as Ihosc Ileum! herein as Tigs %Q t 
M wht&H aie indistinguishable from those figured as 
fMiiifixia bilohitlii by Opik ( P>75h. PI. I }. 

Opik (l ( mK p 16, PL \ Pies 1, 2. text. Fit- S) 
etceted a subgenus of Sn<-nu*. i.e. >/nv«M 
iSnntirixiux) based on (our eranidia with Ihe only 
illustiated cranidium having a length ot only 18 nun. 

11 is clearly an iminaJurc specimen which should not 
lu- |l)c basis of a new (axon 

.lei) (PWl creeled Hwusph <>iripit<>spi?nt winch 
is characterised by a relaiively Itmg preglabellar field 
crossed by a preglabellar median tidge and rbe pHSS- 
Bttf£ id a short slender occipital spine. It is possible 
thai the specimens described hy P.ilmcr A. RowcP 
I \W3\ from Ihe Cenlral Transanlarelic Mountains ys 
Hsttti\pt\ pft II- hth'hum i Poeoek i belong Ml H 
o ( -i'ipitos()iiut. because Ihe Anlarciic specimens have 
;i glabella of similar length to //. in^ipiti^fiiHd as 
well as a similar preglabellar median ridge 
However, the Antarctic specimens have an occipital 
node rather lhan an occipital spine. 

it\(ui,spi\ ivxj.vw (Opik 1975,1 
(Fig. :BL. Pie. 3/ 

I97S Sltvtua <vr«sh>s Op.k IfHfr. P- 14. W. 2, J irs 
1-6, |e\t. 1% 4 

I&75 f-;Muin\;ut bilfhalit Pin.ock. ()pik 1975b- p- It 
PI I I ii:s 17. 

Ahntist twenty eramdia, i»ne pailiat free chrt4 
{SAMP3^3.Hk an jncomj>lete ilioiax (SAMP3>^ U i 
and an incomplete pyeidium (SAMP35333). 



Species ot ttsau,\pi\ wnh dishncll) e\pandcil anler 
ioi p$tt of glabella; length "f ^l.tbell.i (eveludine 
occipital rine) ahoul U.7S0.8 lhal o\ cranidium 
ic.velndine oieipilal riny): octapilal rine beais u 
spme which in some specimens is a small U0tit: m 
iithcrs it is lone and slender. 


Cienlly convex eranivtimn with wiilth sHehlh 
greater than lenttth isat'.l. Deep wide axial furrows 
shallow anteriorly. Cemly ^oiivex elabella ol Icne'h 
teNcludme necjpital i tile' abuut (>,?>() N dial i>f 
cranidium, Glabella increases in width in anterior 
with slight waist near Ip furrows glabella widest at 
eye rul^cs. broadly rounded e.lahellar anterior. Pour 
pairs lateral glabellar Inri'ows; Ip furrows moderate- 
ly deep and each eVend ,tl>uin 1/3 ofjj&liUtti? across 
elabclla and mrcctcd sheiitly It» poslcrior; 2p lurrows 
shallow, extend about 1/3 distance across glabella, 
and duccled \ery slightly to posterior. 3p furrows 
qUlte shallow and diiwK-il almirsl sinuehl acruss 
glabella: 4p lurrows occur .is small mdislinct pits. 
Occipital rine loneest medially; it bears a spine the 
lenelh of which varies considerably Iroin specimen 
i_o specimen. In stvme specimens a small node pre- 
sent; in ulhejs Ihcre is a lone skndrr spiiu- which 
ituiy have a length aboin 0.75 (hat ot elabclla 
texeludine oteipiial ring1 in mature specimens ( 
20) and somewhat longer ui immature specimens 

<2B ( n. 

Occipital ituiow shallows inedi:j|ly. \'v\-\ slmn 
fsau.l *ienliy convex preelabellar field. Anlcrioi bor- 
der gently convex with similar leneth \s»©.] n« 
|ircgj ibellar Held. Poorly-de\ eloped plectrum pre 
sent m si>me spL-cimens (I'ie. 2(i), Shallow anleiior 
border furrow. Cienlly convex palpebral areas ol lix- 
ijjenae have width about same as lhal of glabella. 
Prominent centro-posterioriy placed palpebral lobes 
have a lenelb icxsae.) about 0.55 of elabclla. 
Pnslenor of (lalpebral Ir.hes meet the hroad posk-io- 
lateral border furrows. Well-developed eye 
slighily narrower than palpebral lobes, Shallow 
palpebral furrows. Nanow posterolateral borders. 
Preocular sections of facial suture diverge forwards 
;U about 30 " lo the tnms verse. Short posioenlar set- 
lious of facial suiurc. Nurmw posterolateral border. 

fhe single partial hbrii:enae has (j wnlr l-nnder 
which extends inlo an mcoinplcle genal spine. II has 
a reliculatc ornameuiation. 

I borax with at least nine scemenis- Convex axis 
has width about 0-3 that of segment. Centrally placed 
nodes on 4th and 5ih segments of available thorax. 
Shallow pleural furrows, short broad pleural spines. 
I \\t ihiI\ available pveidiniii is poorly prcsetxe-d. It is 
small and uansveisely elliptical Axis has a wi.lth 
about 0.3 lhal ol pyeidium. Axis extends almost to 



* ft 


-. ''V v - 

Rg 3- tf.\u<txi>t\\ Lvnt\iv.\ (Opik 1975). A. Left SAMP3533& craniUium. inlernaJ mould \3. Right; SAMP35326; > ranid- 
itim, internal mould, x3. B. SAMP3533X. cianidium. internal mould, \3. C. SAMP3533U eranidium, internal mould. 
\2. D SAMP.W3V. eranidium, interna! mould. \2. I*. SAMP35323, eranidium, intnmil mould, \3 I 7 , SAMP35336 
traniditmi. iiiic-rnal mould. x2. CJ. SAMP35334. librigcna. external mould. xJ!. 



[lostei U)t maryin. Axis comprises two axial rings phis 
terminal axial piece Two pleural furrows and one 
interpleural furrow present. Two border spines can be 
seen in available specimen. 


Tbe smallest cianidioni is that figured as 2K, In 
(hts specimen ibe length o\ the L'labcifa tewludui^: 
occipital ring) is 0.7 lliat of crarmlium. In larger 
specimens such as ihosc figured m 3B.D the glabel- 
la has a length ol 0.8-0 85 Iriul of cranidium. The rel- 
ative size Of the spine decreases in small- 
er specimens (ZB«E) Ihe spine luts a relatively thick- 
er base lhan in more mature specimens (3B.D). 

t)iu tnsion. 

Ihe considerable variation in the shape oi the 
glabella, the path ol* Ihe laeial sutUie. the shape and 
length ol" the occipilal ruie. and (he leneth ol the 
occipital spine, of the specimens described and U^ 
tired here as H- tMiixm raises the question as to the 
validity '>l placing all these specimens within a sinyV 
speeies, Howevei. as noted above, iherc appears to be 
a complete morphological varialion present in the 
available specimens and hence we feel the etection ot 
more than one species can not be justified, Ilsnu.s-pis 
eerastcs as described herein diliors from // 
ittuiiiunensis in having a relatively shoi ter glabella, Ihe 
glabella ol //, ti<iniinoe<isis enher reaches or almosi 
reaches Ihe anterior border furrow, whereas dial ol //. 
lemstes stops short of ihe border. However, it is worth 
nohne thai wilh respect lo ihe lenelh ol Ihe occipital 
spine. II Jamutot-nsts. as figured in Zhang et ul 
i, IW). PI. 76 : Hips 13. 14; PI. 77. bigs i-riNiows con- 
siderable vanalton in length of occipital spine as does 
H. ventres. Ihe anterior ol thi: gtabelhi ot //. < t fyfifpi 
is more expanded than is lliat ol H, (hnnunn'nsis. 

ffstwspis oeeipuospim; ol Jell (1090) has a small 
well-developed occipital spine. The spine of H. 
i'emsh'M arises from further back on the occipital ring 
than thai ot' //_ <h npihtspitni: ihe spines on some 
specimens of//, centsies are much longer than the 
biggest spines on //. aceipitospimi- In hL reraste.s the 
ratio oi the length of the glabella (excluding occipi- 
lal ting) to die length o\' the eranidium teveluding 
occipital nngi is 0,75 (&S0". m H, orcipifoywa it 
ranges from O.b.5 0.70. 

The glabella of // sinensis extends further tot- 
wards titan does lliat of//. < trash's. The anterior part 
of the glabella oi' If. tetosn-^ is more expanded than 
//. .sinensis. //. sinensis bears only a small median 
node rather lhan a prominent spine us m many spec 
tmens of//, tentsles. 

The anterior part of the glabella of// eerusles is 
more evpantled than lhal ol \ II. hilobtitn: the glabella 
of//, eerusles is rclalivcly longer lhan thai of//, hilo- 
hatte Ihe glabella of//. c\' II, bilofmni as described 
by Palmer & Rowel I I IWi is shorter lhan that ol //. 


Sample collection was carried out under a licence 
from Ihe New South Wales National lVks and 
Wildlife Service. This paper is published wilh the 
permission o\~ llie Dhvctoj-Geoeial, New South 
Wales Department or Mineral Resources. I. in Tiun- 
Rui was supported by a visiting researcher's grant 
from the University or .South Australia. t)r P.A. Jell 
constructively reviewetl an earlier version of Ihis 
paper. Mr J.S. Bao gave valuable technical ussis- 
lanee. Dr C. Jenkins supervised G Davidson's hon- 
ours protect 


BiNOisoN. N.. r urvwv, Ylouuis. S. ( L'ouir.U* fif, J.. Jilt. P 
A. & Ki nm.oak, U. N. (l'ii)Oi barly Cambrian fossils 
bom Suutli Australia, Metti, Assih'. Ausiiuiltis 

l-n ii III ft. Ik O. llUfvli New liiiiiukml shells IOihi Hie 
l.ovvci Otdovician and middle Palaco/oic rocks ot New 
Siinih Wales. AYr. \it\j Mus. 26. 2K3 : L '4, 

HmvkInoios, It. J t'Utl. ( IWJ) Arl.hropoda I pp I -5"0 //; 
Mtioir. R. C (&J ) 'TrralKC on liiveriehr;uc 
Piilfoniotogv. Pan <> • < reologicul Society of America 
ami KaMsas l ; D»vi.-isU> Piv-ss. I.awrenci. Kansas). 

l\oo_ I. B'. t IWro Soitil i.(Miiiitciii>i tm tlie Cambrian mm.- 
wiilc m ichilmii to receril radiometric dalinc of Australian 
t'amboita rocks pp. l05-b)r» In Linan. E., tiuine/ 
Viiitjiicd, I A. A: 'Jo/oltt. K. 1 1 'ids! "II fit-Id l^onlerence of 
llit* Cumbnan Stage SulnJi\ imimi Working Gioups"", Spain. 
H ?l September. Wfi, field fop Guidf aiul Abslracis. 

Jl 1 1 . I 1 '\. ( IV7S] Ausiraliiai Middle t'ainhriaii codiscoids 
Nvilh a review i,if lite snpetlamdv. fr-iithiwlnf>rophir(-i A 

150. \-<*y, 

tlWtfl Ttilohita pp. :^7 332 In l3eiu;isoa, S„ 

(^onsvav M\<rris, .S.. ("- >oper, B. J.. Jell, P A. & Rnniieear. 
B. N Marly Cniihoan ft^sil.s trom South Australia. 
A/r/». .-Uv"<- .\u\tittiiis fnhn'onlol, -J- 

Khi'SI-.. P b, < 1 1 >7M i New Aivhat'ocvallui I'roni the bairly 
f'ainboan »if the Ml Wright area, New South Wales. 
Meheru^iiZ. 37 47, 

(bJS2i Aretiaeocyailian bitisiiaiiuraph) oi the 
G nail a Gump at Ml Wi tjilif Nt:\v South Wale* 
t'itUwontownphh'o A 177. I 212 



U 1 YANtlOU, ClIANG Wrntang, Cll'tfiN Yivuan, Chu 

Chaoling, Lin, H. L., Zhou. Z. Y., Qian, Y, Zhang, S. 

G. & Yuan. 3. L. (1974) Cambrian trilobites pp. 82-107 

In "Handbook oi' stratigraphy and palaeontology ol' 

southwest China" (Science Press, Beijing). [Chinese, 

English summary]. 
Nhdjn, C. ( 1995) The Emu Bay Shale, a Lower Cambrian 

fossil Lagerstatten, Kangaroo Island, South Australia. 

Mem. Assoc. Ansrralas. Paloeon/oL IS. 31 -40. 
Opfk, A. A. (1975a) Templetonian and Ordian xystridtirid 

trilobhes of Australia. Bur. Min. Res., GeoL Geophys., 

Aust.,BulL 121. L-S4. 
( 1975b) Cymbfic Vale fauna of New South Wales 

and Early Cambrian bioslratigraphy. IhUL 159. I-7S. 
Paumkk. A. R. & Rowkix, A. J. (1995) Early Cambrian 

Trilobites from the Shaekleton Limestone of the Central 

Ttansantarctic Mountains. The PaleontoL Soc, Mem. 45, 

PoGogk, K. .L ( 19o4) Estaingiit, a new trilobite genus from 

the Lower Cambrian of South Australia. Palaeontology 

7. 458-471. 

Rosi:, G, (1968) Broken Hill. 1 :250.000 Geological Series, 
Sheet SH54-I4, Preliminary Edition. (Department of 
Mines, NSW). 

Shfkgoi i). J. H., J ago. J, B., Cooi'KK, R. A. & Laurie, J. R. 
(1985) The Camhrian System in Australia, Antarctica 
and New Zealand, hu. Union GeoL Sci. Puhl. 19, L85. 

Zhang Wkntang, Lu Yan Hog, Zhu Zhaoung. Qian Yi. 
Lin. H., Zhou Zhivi. Zhang Sangui & Yuan Jinuang 
( 1980) Cambrian trilobite faunas of southwestern China. 
Pulaeoniol. sin. set: B 16, 1-497 |Chinese with English 

Zhou. T M. & Lin. T. R. (1978) Early Cambrian trilobiles 
pp. 143-154 In "Sinian to Permian stratigraphy and 
palaeontology of eastern Yangtze Gorges" (Geological 
Publishing House, Beijing). [Chinesef 

Ziiuuavi kv. A. Yu & Gravestogk. D. I. (1994) 
Archaeocyaths from Yorke Peninsula. South Australia 
and archaeocyathan Early Cambrian donation . 
Aicherinxa IS, 1-54. 


Brief Communication 


The Kanmantoo Group (Fig. 1) is a thick (8-10 km) succession of predominantly 
clastic metasedimentary rocks which crop out in an arcuate belt from near Australia 
Plains in the northeastern Mt Lofty Ranges to the western end of Kangaroo Island. On 
Fleurieu Peninsula, the Kanmantoo Group rests unconformably on the Normanville 
Group 1 , the uppermost formation of which, the Heatherdale Shale, contains a poorly 
preserved fauna of possible mid to late Botoman age 2 . The highest well-dated fauna 
from the Normanville Group are archaeocyathids from the Fork Tree Limestone at 
Sellick Hill which indicate an age close to the Atdabanian-Botoman boundary 3 . These 
are at least 400 m below the top of the Heatherdale Shale. The top of the Kanmantoo 
Group is not exposed but lower parts of the group are intruded by the Rathjen Gneiss 
which has an age of 516± 4 Ma 4 , i.e. late Early Cambrian or early Middle Cambrian 
on recent Cambrian time scales 3,6 . The stratigraphy of the Kanmantoo Group is 
described in detail by Daily & Milnes 7,8 . 

'DmiMWtioti.s of tin- Hnvitl W/W\ *</ V -\u.\i. i IW7). I21t ?>.75 T "\ 


The Kanmanloo Group iFJg. Ii is ;! duck (R-IO km I 
succession ul predominant)} clastic mclascdimeniary rocks 
which crop out »n an arcuate bell (mm iieai Auslraha Plains 

iii (he northeastern Mi I .oliv Ranges to the western end 01 

Kangaroo Island. On lleuricu Peninsula, |h« Kanmanloo 
Group reNtsuncouformubtv mi the Norinauville Group . the 
uppetmost formation of \vhich_ the Hcalherdalc Shale, 

contains a poorly preserved fauna of possible mid io late 
Uotomuu Qgc'i The hieiicsi well dated fauna I rot it the 
Nofuianvillc Turnip in atvhacocv;ilhids from (he b'ork TflJC 
lame.sione at Schick Hill which indicate .111 agcdfK^ lo the 
Aldjbaiiinn-Holomau boundary . These are al least 400 111 
below Ihe top ol' Ihe He:iihcrdatc Shale. Hie tOJJ ol ihe 
Kamiiatlloo Group is itol imposed hut lower purls ol the 
•jroup are intruded b> Ihe Raibjcn Gneiss which has art uec 
ol ?ib% 4 Ma 1 , i.e. km- Tally Cumbrian or early Middle 
('ainhnan on recenl Cambrian [fjtKi scales' ', The 
stratigraphy ol the kanmanloo Group is described 111 detail 
by Daily & Y1ilnev\ 

TJlfl Cumbrian sequences us exposed on Fleurieu 
Peninsula. K.ineanMi Island mu\ Yorke Peninsula show 
considerable lateral and vc'ilieal litltological variation'. 
Currently, the limited hiosiiaiiLuaphic inhumation makes in 
dilfieull to cortvlaic accurately from one region lo anolhei 
and hence determine (he tintinp and sequence ol hoih 
stmli^raphic and tectonic events 

One of ihe problems in ihi.s regard has been ihe lack ol 
bioslraltyraphicallv useful bod) fossils in (he Kanmanhi' 
Group. The only such fossils reported to dute aie ihe 
inuriieiilaie bratluopod t ithnttrtln Itom about 10 DO tftxwc 
Ihe base ol die basal unit ol the Kanmantoo Croup, the 
(_arrickalui^a I lead I ormalion. at Ganiekalinya I lead 1 ". uflil 
a possible hyoJilhid from near Ihe lop ol thr Oarrickalmua 
Head l-ormaiion near Blowhole Creek", trace fossils have 
been rcpoiicd Ironi ihe Backstairs Passage Torntation ul 
Accommodation Hill near Hiiro 1 

This note reports ihe presence of further inarticulate 
brachiopods il'iy, JD.I.). from about 100 111 above Ihe base 
itltlie Ciirnekalm^a Head bonnalion at Carriekalinga Head. 
I hey are small and poorly presetted, but appear lo represent 
ul least two species. 

In u quarry near J-'arawu trig. 3). ai 021.551 TI01 reus Vale. 
1:50.000 lopojiiaphic map) on (VtllawoiiL'a Creek Road, the 
authors have found |hr Itrsl known {rilobiles from l|»r 
Kanmanloo Ciroup. The speeimens come from a laminated 
melasilistone probably par) of IheTunkahlla borinabon hui 
possibly upper lapanappa Kormatum. lo date we have 
found aboui 20 specimens, all ol whieli uie uhmtsicomplete, 
ihus mdicaliny low energy deposilional condilions As 

sliown in fig ZAM.C the -»pecimens are potnly preserved. 
lccl<»nieally delormed and of no biosiraltyraphic use 
Although, within ihe limns ol pivservaiton, they seem to 
rej*resenl a ^'nele species. Houcvct. then "presence oiilualfs 
1 Ik pov.siihi hi y ot the eventual discovery ol 
hiosiralijiiaplucallv useful liilobiles from ihe Kanmanloo 









Middleton Sandstone 

Petrel Cove Formation 

Balquhidder Formation 

Tunkalilla Formation 


Tapanappa Formation 

Talisker Calc-siltstone 

Backstairs Passage Formation 









Carrickalinga Head Fm ^) 

Heatherdale Shale 

Fork Tree Limestone 

Sellick Hill Formation 

Wangkonda Formation 

Mount Terrible Formation 

Pig: I- Niraligruphic ouiline of Noimnnville and 
Kanuiantoo Groupp. Tlcuricu Peninsula, showing die 
levels ot the fossils Ikured herein. 


lip. 1 A l'. Irili'liilfK from t|iKirrv near I'urmwi. A. 

Rubbct ca-.i ni oHtenml mOuR BAMP3534?, ?& b_ 

Internal mottl.l. 5AMP3W& vi. C. Rubber easl ftf 
external mouM, SAM1-M5JU7. \.^ OJL Inarticulate 
bruebiopocK Irom C\irricka|in»a lletuJ. IX Internal 
mould*. SAMP35.U8a ana* SAMP3534HH vl.V b 
f'itiii.illy cxtftliatcU specimen. 5?AMP35-349. \Iii, 
.Spec i men numbers refer to the palueonlolostcal 
lollceiion of the South Australian Museum. 





f ; > 


&~$Z. ;> 

■*- " ■ v*-y j 



Peninsula area^ 

Gulf St 


_ 139° E 

Kg, 3, i-oeality map 


'Jago, J.B. (1994) Aust. J. Earth Sci. 41, 445-453. 
2 Jago, J.B. (1996) Some comments on the Cambrian time- 
scale in relation to recent radiometric dating of Australian 
Cambrian rocks pp. 105- 106 In Linan, E., Game/ Vintaned, 
J. A. & Gozolo, R. "II Field Conference of the Cambrian Stage 
Subdivisions Working Groups' 1 Field Trip Guide and Abstracts. 
Zhuravlev, A. Yu & Gravestock, D.I. (1994) Alcheringa 18, 

4 Farrand, M.G. & Preiss, W.V. (1995) Geol. Surv. S. Aust. 
Bull. 54, Vol. 2, 54-57. 

^Young, G.C. & Laurie, J.R. (Eds) (1996) 'An Australian 
Phanerozoic Timescale" (Oxford University Press, Oxford). 
6 l\icker, R.D. & McKerrow, W.S. (1995) Can. J. Earth Sci. 
32, 368-379. 

'Daily, B. & Mines, A.R. (1971) Trans. R. Soc. S. Aust. 95, 

H Daily, B. & Milnes, A.R. ( 1 973) Ibid. 97, 2 1 3-25 1 . 

''Gravestock, D.I. (1995) Geol. Surv. S. Aust. Bull. 54, Vol. 2, 

"Daily, B. (1963) Rec. S. Aust. Mus. 14, 579-601. 

"Gatehouse, C.G., Jago, J.B. & Cooper, B.J. (1990) Geol. 
Soc. Aust. Spec. Pub. 16, 351-368. 

I2 Daily, B., Firman, J.B., Forbes, B.C. & Lindsay, J.M. 

(1976) Geology pp. 5-42 In Twidalc, C.R., Tyler, M.J. & Webb, 
B.P. (Eds) "Natural History of the Adelaide Region" (Royal 
Society of South Australia, Adelaide). 

J.B. JAGO and P.W. HAINES, Department of Applied Geology, School of Engineering University of South Australia The 
Levels S. Aust. 5095. 





Brief Communication 


The pygmy right whale Caperea marginata is a rarely sighted species known primarily 
from standings. The latter occur frequently in South Australia, especially along the 
north coast of Kangaroo Island and near Port Lincoln 1,2 . Migratory patterns are not 
known although it has been suggested that this species moves inshore in spring and 
summer 3,4 . Sekiguchi et aL 5 postulated that such a movement off South Africa may 
coincide with an increase in the abundance of copepods, one of the presumed main 
prey of C. marginata. Spring and summer are also the seasons when juveniles most 
frequently strand 2,4 . There appears to be a broad mating, calving and weaning period 
between June and February 2 . This paper describes two recent sightings of live animals 
off Lincoln National Park, about 10 km east-south-east of Port Lincoln, South 
Australia and summarises past sightings and strandings in that area. Other known 
sightings in the Australasian region are also reviewed. 

tmmuiihmn qftlw Royal Society o/S. Aits!. { PW), 121(2). 79-82. 





The pygmy righi whale Cuperca niar^inata is a rarely 
sighted species known primarily from strandings. The latter 
occur frequently in South Australia, especially along the 
notth coast of Kangaroo Island and near Port Lincoln 1 2 „ 
Migratory palicrns arc nol known although it has been 
suggested that Ihis species moves inshore in spring and 
summer 4 . Sckiguehi <■! </// postulated that such a move- 
meril off South Africa may coincide with an increase in the 
abundance of copepods, one ol' the presumed main prey of 
C. nioi'x/iititii. Spring and summer are also the seasons 
when juveniles most frcqucnily slrnncP '. There appears lo 
be a broad mating, calving and weaning period between 
June and February ■'. This paper describes two recent sight- 
ings of live animals off Lincoln National Park, aboul 10 km 
east-south-east of Purl Lincoln, South Australia and sum- 
marises past sighlings and standings in that area. Other 
known sightings in die Australasian region are also 

While sailing a o-m yacht in Spalding Cove, off I mcoln 
Nalional Park. Iwo of the authors. .1. EL and B, F.. observed 
two pygmy right whales, an adult about 6 m long, accom- 
panied by a calf about 2 m long. Paired blowholes (Fig. I ) 
confirmed that the adult was a baleen whale. Species iden- 
tification was based on the curved jaw-line, medium-grey 
colour, a falcate dorsal fin placed well back on Ihe body and 
the adult's broad back {Figs 1-3). The animals were seen 
belween 1030 and 1 130 1) on 4 January and 13(H) and 1400 
h the following day. On both occasions they were deep in 
(he cove at the far south-western side, about 300 m from 
shore (Fig. 2). Water depth (as determined by a depih- 

soundei ) was about 5 m. water visibility X m, surface wnier 
temperature 20' C and the sea was calm. Spalding Cove is a 
shallow, sloping bay with a sandy hollom and extensive sen- 
grass beds. 

Immediately prior to the sighting on 4 January, a large U- 
shaped swirl about 4-5 m diameter, followed by a curtain of 
bubbles, appeared 2 in from the boat. About 2-3 min lalei 
and 50 m away a large animal, the presumed adult female, 
surfaced and blew. Tins was followed by the blow of the 
calf near the adult. The calf swam slowly around Ihe boai 
coming close lo the bow and turning off. much as dolphins 
often do. It swam with an undulating motion, surfacing fre- 
quently lot air (30-40 sec or as long as 2 mini At this stage 
the boat was under motor at a speed ot 2 3 knots (3.5-5.5 
kph) and the calf showed no sign of being disturbed by this. 
The calf alternated between swimming next lo Ihe boat and 
the nearby adult aboul 20-30 m away and sometimes swam 
above the adult, a common position for dependenl calves of 
the southern right whale F.ithaUu'na tinstrulis (C Kemper 


Fig. I - Adult Copcrca marginuia in Spalding Cove. Soulh 
Australia, 4/5 January I°96. Arrows show the paired 
blowholes and indistinct white bar behind the head 
Nole also the broad hack. Pholo: J. Dulton. 

Fig. 2 - Juvenile Caperea mar^inuia in Spalding Cove, 
Soulh Australia. 4/5 January 19%. The arrow marks the 
prominent, falcate dorsal fin set well back on the body. 
Pholo: J. Dulton, 


pels, ohvi. The calf's prey buck and sometimes iis head 
showed as ii surfaced and blew. Later c.vaiiiinulion of pho- 
tos showed thai Ihe almost while ventral colour exirnded 
WfrN up the side of die body ami dial a distinctive dark lat- 
eral paleli iuleirupted this just behind the head (fig 3*>. A 
.-nnilai -colour pattern has been observed On a South Al'ncan 
juvenile C. lUtttviiiaht' and a recently stranded neonate in 
New Zealand (van Heldeil puis, contuU- 

The adull swam in large circles in the general vicinity o) 
the boat. sometimes criming very elose to ii and at times 
turning on ill side. It showed no sign of being disturbed by 
the presence ol die boat. The adull surfaced to hrcaihe much 
less ollcn than ihe call, about every 3 mm 01 more. Lalei 
examination Ol' photos showed that Ihe adull was a medium- 
grey colour wilh .1 pale hand or chevron just behind the 
head (Fig. I), The belly was lighter Mian Hie back. 
Maisuoka' 1 also described and illustrated light chevrons on 
ihe backs of a luigc giolip of adull C ntannttniu observed 
ai sea (Tabled, 

Trjc animals were m the same pari of the cove during the 
afternoon of the next day when J I) and H K returned 10 
.sail there, A net fisher operating in Spalding Cove during 
late lutiiiary and much ot February 1996 repotted seeing a 
small whale then: on several occasions- His description of 
ihe annual does not allow species identification bul it may 
have heen ( . nutryinnUi. possibly one ol Ihe same animals 
observed in early January by .1. p. and B. F. 

The other a-eeni SouUi Australian sighting was made dur- 
ing ihe afternoon of 7 July \^% by R, M, who observed, 
with the aid ol binoculars, a small whale from the shore 
near Capo Donington lighthouse. Lincoln National Park 
(Hie. -I) It was about 50 m from shore and 60 m from the 
obseiver The weather wa> sunny and the sea calm, with a 
lf» hi southerly to soiiih-weslcrly breeze n( 1-5 knots (2 ( ) 
kph). The animal was moving slowly towards deeper water, 
in a south-south-easterly direction. At limes it swam just 

below the surface with its mouth open. It was not fiossible 
to tell if the animal was (ceding and no obvious signs of 
plankton swarms were visible to the observer Several low, 
thin blows were seen. The animal's colour was dark grey on 
ihe back and light prey underneath, The features which sug- 
gested that this was a C. maixi>i<tii( weie; I ) lighter patches 
between the mouth and the flippers. 2) whitish baleen pla!c> 
which darkened towards the outer edges, ihk\ 3» a small, 
curved doisal fin about -/ of Ihe way along ihe back 
However, without observing the bowed jaw-line. Ihe identi 
liealion as ( itmi^hhtltt could no! be considered confinned 
since minke whales iHciktcnoptvtii at utttro.smua) share 
several i>\ the abovc-tnciilioiicO features. Fslilllaled body 
length oi the animal ft. M. observed was $.5-4-0 in. sug- 
gesting that it was a posMvcaning juvenile , 

Nine slraiidings of (\ nhu^itmla have been rerouted in 
Ihe Port Lincoln area from before I'US lo 1003 liable 1) 
and, iis suggested m 1004 by Hale"', many more have proh 
ably occurred wilhoul being reported. All Ihe reported 
sirandings have been from Pori Lincoln Proper and in. oral 
the enhance to. Spalding Cove (Pig. 4). Roth aie shallow 
seagi ass-covered, sandy/mudllal bays with large tidal 
movements. All stranding.s involved single animal:;, 
although in some cases there may have been a connection 
between certain events occulting within a short lime ol 
each other. Lor example, u .MS m juvenile Was 1ound 
recently dead on 2 February f989 in Spalding Cave and on 
I March 10S0 a decomposed adult (unknown sex) was 
found on flickers Island at the entrance to the cove (strand- 
ing nos 0. 7. Fig, -1). On April 1993 an adult female, with 
no evidence of lactation, washed up near on Ihe south side 
ol Poii I incoln Proper and sin days laler an exiremelv, erna 
eiated .?. Itl m juvenile stranded alive in Spalding Cove 
tslrandiug uo.s X, l ). Fig. 4). Il is possible thai both cases 
involved mother/calf pairs, Of the five adults thai have 
stranded, ihree have heen females anil two of unknown sex. 

ig. 3 Juvenile Capewa IHarginata in Spalding Cove. South Australia. 4/5 January I Wo. The line drawing helps to dis 
lingmsh the bowed jawline. which is slighily distorted hv a wave. The arrow marks a lateral colour pattern of whilr just 

Ivliunl 1I1.' h.-»;nl lnrl «i il.ii'k rvilrh rw»:h»M(ir in llnv Phitln I I ItiUnti 

behind the head and a dark patch posterior to this. Photo: J I Julton 


t MCt tf 1 1 . * r 


_A Nffl I . Hi '..'I fS 

T^iun --^ 

I ig. 4 - Map til" Port Lincoln region showing positions oi 
Mghtings [squares,] ami strandings letrcles) of Cupewa 
fiuiivi'nita. Numbers refer CO ttrandftigs in Tabid I and 

sightings in Table 2. 

These slianding results suggest thai (he region is frequently 
used by fculalcv and calves. although such information can 
he misleading if there is. as in tlii> ease, imieh human acti\ 
ity in ihe region and therefore possibility Ol discovering 

Reported sightings off . margUiatri, summarised in lablc 
2. are? not common in the Australasian region. To our 
knowledge. Ufic Spalding Owe sighting \t{ January |*990 is 
the first time a cow and call* have been sighied and pho- 
tographed anywhere m file southern hemisphere. Nol 
included in Table I are two unconfirmed records found in 
the Australian Nature Conservation Agency sightings data- 
base, one of three animals oil Montague Hand. NSW in 
June l l W (record no. JS7f>> and the other from near Bcrimi 
Island. WA inJune IW3tno. I55N). Since there was no sup- 
porting description ot Ihe animals to allow positive identifi- 
cation and ('. imt/Miimttt can lie confused with the minke 

whale, II aculorosivata. the records have not Keen included 

Many 6f the sightings listed in Table 1 and several Ironi 
South Africa 1 woe made inshore, suggesting that C nuir 
gtntttu inhabits coastal waters, at least for some pari of its 
life or annual Cycle. Lnpuhhshed stranding data show that 
dependent young and recently weaned juveniles (3.0 -3:5 in I 
are more common along the central South Australian and 
wextern Victorian coast ( Kemper unpub. I. hour or the sight- 
ings listed in Table 2 involved dependent young or animals 
that, from their size. Would appear lo have been recently 
Weaned; Three were from the Pari Lincoln area and one was 
from Portland in western Victoria. Larue, protected hays 
with shalhm, sandy boiiom-. and extensive scagrass bette 
may he important calving ami weaning areas lor C '. imo 
vimim. Some cvauiplcs arc Portland Hay. Nepean Has mid 
(he tioston Bay region. Several Mrandmgs of C uua^tnaia 
have been recorded ha the Nepean Say area'- 

The sighting made in Cockbuin Sound in 1 401) hy II, and 
D. Parker tB. Parker perv eomiru merits a special note 
because it records some notable behaviour as well a> |irob 
ahle ftjcUinjl hy the 5-iri animal they observed over a period 
of about one hour, Hie animal was swimming very fast, 
leaving a wake ol water, and nodding ife head noliceaWy. 
The whale came to investigate llie 5-m boat. >t raping itsell 
the lirsi time againsi the bottom of the boat. The seeond 
time it appioaclicd lite boat very quickly, almost in a 
charge. ItTled the boat out ot the watei and almost caused 
the occupants lo be thrown out' 

The authors lo thank II Parker and D Counhran tor 
supplying information on the sighting in roekhutu Sound 
and ihe Australian Nature Conservation Agency \'or seareh 
iug their records for Cnpemt sightings, C Kemper thanks 
al! those who have provided information on pygmy tight 
whale strandings and sightings records, especially South 
Australian National Parks and Wildlife officers ami 
Department o! Primary Industries South Australia 
( fisheries! officers, J. Thurmer suggested and prepared the 
line drawnui lor lm. J. 

T Mil I L Rcuinls af Caperea marginata \ttrfflHfitt$& in ilw rfaft Lincoln area. Stramting Mifnhcr ff?#. fief.) cr< >xs -wan hed 
to AV.v V, ; c\tittutfetl lettiith touted on skeletal measurements'. Museum no, M - specimen in Smith Austnilhm 
\liaenm. S ~ no specimen in South Atotimlion Mustunc 









<l l MX 

SW Port Lincoln 


2 7,t : 


2o.12 P.J.S? 





MM 10 

seen alive before -.(randing 

lh,K| l )nt) 






decomposeil, ma collected 


Spalding ( ove 





ilecom posed 


IVin Lincoln Proper 





washed up dead 

1.2 l<JS<) 

Spalding Cove 




Ml 5024 

very fresh 

1 J/4489 

Bickers Maud Noith 




.M 1 5374 

very decomposed 


1 km SE Horse Kock 




M 1 7 Mil 

washed up dead 


\ 2 km Sft 
Stamford Hill 





alive, very emaciated- returned lo sea 


TABLE 2. Sffthtm&s, including captures hut not sightings immediately followed by straadings, ^/'Caperea manrmala in the 

Australasian region. Latitude and longitude given in degree* and nannies. Hei is reliahlitiv of identification (1 = 
certain. 2 - probable, 3 - doubtful)- A = adult, S = stibadult, J = juvenile. 



Lai. I 



Rel. Size Comments 



4 J $80 

4.i yss 

2<S.ll.l t >8(v 

i iya9 

10 10.1990 

26.1 I J c >92 


north end 46 50 S 

Stewart I.. NZ 168 00 F 

Bruny Island. Tas 43 17 S hay. in 

147 IS E 2-3m water 

Port Lincoln, SA - protected hay 

~ 50 nm SB 3jS 20 S open ocean. 

Cape Howe. NSW 150 20 E oil shell 

Soelu Seainounl, 43 50 S "pen ocean, 

120 nm SE Tas 150 22 L over sea mount 

Portland, Vie. 3S 2i S sajidy. 

141 36 E protected hay 

3447 s shallow, 

135 58 E proteeledha>. 

Cockburn Sound. 32 10 S offshore 

WA 115 44 L 

Spalding Cove. 

420 n m S Cape 
1 ,ceii\vin WA 
Spaldinc Cove. 

Port Lincoln. SA 

4 1 37 S 
115 38 L 
34 47 S 

135 58 E 

34 44S 

1 36 00 E 

open sea 

protected hay. 

edge of 
Spencer Gulf 

mans 2/3 

-80 I 


black fish 



killed by fishers 



no description of 


observed from 




leedini!. no 


description of 



shipping harbour 
possibly feeding 



Pm. 4 icf. no. 2 



photOS suggest 

B. Parkcr 

feeding hehaviour 




three groups near 
each other 



stayed in area at 


least two days, 


hig. ref. no. 3 


Fig. 4 rel". no. 4 



'Kemper. C. M. & Ling, J. K. ( 1991 > Trans. R. Soc. S, 
Aust 115.37-52. 

Pavey. C. R. ( 1992) Aust. Mammal 15. 1-6. 

Davits. J. L. A (iuiler. E. R. (1957) Proc. Zool. Soc 
Lpnd 12<>, 579-5K9. 

Ross, G. J. H.. Best. P. B. & Donnelly, B. <;. (1975) 1 
Fish. Res, Board Can. 32. 1005-1017. 

Sekiguchi, K.. Best, P. B. & Uozena. /.. K. (1992) Mar 
Mam. Sci.8. 2K8-293. 

Matsuoka. K.. Fujise, Y. & Pastene L. (I99(» Ibid- 12. 

Kemper, C. M. V/>/W.iaeeeptedr 

i lector, J. (1X75) Trans Proe. N. /. InsL 7.251-265. 
"(Iuiler, E. R. I I97K) Pap, Proc. R. Soc. Tasm. 112. 1X0- 
Hale, H. M. 1 1904) Rec. S. A. Mus. 14. 314-3 19. 

H Cav\thurne, M. W. (19KI) Rep. lnl. WhaL Comma 31. 
SC/32/Pmyress Report N. /.. 201. 

Blaher, S. ,J. M. ( 19K6) Emu 86. 239-244. 

1 'Arnold, A, ( 1987) Aust. Nal. Hist. 22. 26fv270. 

CATHERINE KEMPER, South Australian Museum. North Terrace Adelaide S. Aust. 5000. JOHN DUTTON, 23 Mitcham 
Ave. Lower Milcrumi S. Aust. 5062. BRIAN FOSTER. 51 Bntainnia Road Nairne S. Aust. 5252 and ROBERT Mt CHJIRE. 
S3 Veran Terrace Port Lincoln S. Aust. 5606. 

VOL. 121, PARTS 3 &4 
28 NOVEMBER, 1997 

Transactions of the 

Royal Society of South 



Bourman, R. P., Martinaitis, P., Prescott, J. R. & Belperio, A. P. The age of 

the Pooraka formation and its implications, with some 
preliminary results from luminescence dating 83 

Vaucher, C. & Beveridge, I. New species of Potorolepis Spasskii (Cestoda : 
Hymenolepididae) parasitic in dasyurid marsupials from 
New Guinea --------- 95 

Littlejohn, M. J. & Wright, J. R. Structure of the acoustic signals of Crinia 
glauerti (Anura : Myobatrachidae) from south-western 
Australia, and comparison with those of C. signifera from 
South Australia --------- 103 

Watson, G. F. & Gerhardt, H. C. The breeding biology and advertisement call 

of Litoria splendida Tyler, Davies & Martin - - - 119 

Barnett, E. J., Harvey, N., Belperio, A. P. & Bourman, R. P. Sea-level 
indicators from a Holocene, tide-dominated coastal 
succession, Port Pirie, South Australia ----- 125 

Gillian, P. J., Cranston, P. S. & Cook, L. G. The response of gall-inducing 
scale insects (Hemiptera : Eriococcidae: Apiomorpha 
Rubsaamen) to the fire history of mallee eucalypts in 
Danggali Conservation Park, South Australia - 137 

Olsen, A. M. An intensive monitoring study of two wetlands of the River 

Murray in South Australia; physico-chemical parameters 
and cyanobacteria concentrations ------ 147 

Kolesik, P., Whittemore, R. & Stace, H. M. Asphondylio anthocercidis, a new 
species of Cecidomyiidae (Diptera) inducing fruit galls on 
Anthocercis littorea (Solanaceae) in Western Australia - - 157 

Brief Communications: 

Wall man. J. F. First record of the Oriental Latrine fly, Chrysomya 
megacephala (Fabricius) (Diptera : Calliphoridae), from 
South Australia --------- 163 

Smales, L. R. The status of Cyclosirongylus medioanmdatus Johnston & 

Mawson, 1940 - - - - - - - 165 






VOL. 121, PART 3 






Bourman, R. P., Martinaitis, P., Prescott, J. R., & Belperio, A. P. (1997) The age of 
the Pooraka Formation and its implications, with some preliminary results from 
luminescence dating. Trans. R. Soc. S. Aust. 121(3), 83-94, 28 November, 1997. 
Stratigraphic relationships, supported by luminescence dating, suggest that the 
Pooraka Formation spans a far greater time interval than previously recognised on the 
basis of radiocarbon dating and stratigraphic analysis of discrete sedimentary sections. 
It extends back as far as the Last Interglacial. Re-evaluation of the radiocarbon ages 
that indicate an interstadial age (i.e. Oxygen Isotope Stage 3; 45 to 30 ka BP) for the 
sediments is required. Alternatively, a considerable time interval for deposition of the 
Pooraka Formation would necessitate that the unit be diachronous across the 
landscape. An age extending back to the Last Interglacial (Oxygen Isotope Substage 
5c; c. 125 ka BP) would provide the appropriate palaeo-climates and palaeo- 
environments for fluvial sedimentation. The revised age has implications for 
landscape evolution, archaeological and palaeomagnetic prospecting as well as the 
antiquity of the Diprotodon in the Adelaide area. 

Key Words: Pooraka Formation, Pleistocene stratigraphy, Last Interglacial, 
luminescence dating. 

ini v<;i : 01 ihi; pooraka FORMATION and its implications, WITH SOME 


by R. P. Bourman , P. M AkiiNAiris , J. R. Pkhscorr & A. P. Bi.i n kio 


Btil KUAft. R. I?.. M vki in Mils, K; Pmsroit.j, K, & BUVEftU), A, P, UW7)Thc ajvofuV I'onraka P-tfraiitDmi 
and Its nnplieatioiis. wilh *W|V prehminaiv result* from kiniiiii.*sceiice daium. 7)y"W> A' ,V//j S T//.W I2I( N V 
•m, 28 Nwumta l'W7. 

Strahtaapliie icI.'iimii.sliipN. supported hy luminescence dating. Wggetit Hut ilu PotVAka l-oonaiion vpvu* a fai 
greater link! imervul than previously recognised on th6 basis ortmlfoearbon dating an! stiahgrapluL auaiwv of 

disuvie htfdlltietttdr} iWClitfiV*- Il eMends bark a,*- lai a* Iht last hueiylacial. Kc evaluation of tin; radioeaibun 
ayes that intltealr an inicrsladial age ({&, ( Kypcn Isotope Sta^e J: 45 Id M) ka RPi lor the sediments W requited. 
■Mirrnamely. a considerable time mterv;il for deposition or l Ik Poniaka I'ojinalion would neees.siiatt* lhat the 
miil be di.iehronous aetoss die landscape. An age tfttendlnu back io thti I j-i Interidacial ({Kygtan Koiopc 
Subsla^c 5e; e. 135 ka HP) would provide ihe appropriate p; laeo climates and palaco-eiivironineMs lor fluvial 
.eiltineiilalion, The revised age has implication, toi kindscapc evolution, archaeological and paLteonwimeiic 
piospeilni^ 3* well a-, (lie anlM|iiily o| [htf />//m>Wt>/J ill (he Adelaide area 

Ki *i VfrfftfW: Pooraka loiiualiuii, Pleasiucene Mrati^rapliy. Last luicr^lacial, linnincNtenec dating 


Lui'gC ardus ni the Adelaide' Plums are underlain by 
Iht- Pooraka I'ormntion (Pirman |96Ti Vtt&\ Callcn 
v! tti. 1 995: Shcard & liownmit IWft). a 
brown coloured Pleistocene alluvial deposit with 
weakly developed calcareous pedoticnic horizons 
ilia) undeilics river terraces and alluvial fans, litis 
unit is also widespread beyond die .Adelaide Plains. 
exlcndm^ DTI U) Ihe Hciuieu Peninsula and inio the 
nnd north iff ihe slate, where it Hanks the Flinders 
and (Jaw lex Ranges. 

The ra\ Coloured sediments dial comprise die 
widespread Pooraka I ornialion have been ascribed 
diUcrcnl names bv workers over time and in dilleicm 
areas, Pot csaniple, Ihcy were originally referred to 
a> the •inaimu-jlile-roiK drill b\ late t \$7Q) becau>e 
skeletal re mams of |hc exlincl. gianl marsupial, 
DipmUnfon itptihtnu were recovered from ihem in 
areas to the vvesl til Ihe city ol Adelaide. Ward ( l^oro 
referred Ki the sedirueius as Ihe Christies Befneh 
boimatnut in die Noarlun^a and Wilkinga suh- 
basms. Twidale f 1 96S ) named them the KICITUIJ 
Sand durtn»j his investii:atiitii of the terraces of Iht* 

Si fund (| i Rth iniiiMrjiicii ;hui KcciL'.'iib.i. vi^h..". nn m i n imv >>i 

i Itj 'tiwLi'inv aftvl Lnvti uhimlmI, tlinvvtsitv nl Scnitll Ausiiali;' 
Wi mil U.I tl lL I i>V| U s \iki S(V5 

t> I' llllll'.Ml :<{ I1l\-ii', KlICl MllllVIMIIIIOl I'll': Il . I il,' I Ml\.'l>il|> 

til va/l.nai' \IW IjtJOfl 

lr,iiM>'f|v Muw\ ;iitd foil _lv Ke-;onio,_-''. Sitinli A.t.vii:il(,< I'd Ui Vy 
lit |>rw*vd s\ \,r.i, JOttf, -.'uiTenlb. MiiiH;iin Ci\\\4 111 
dL.a'sloMi' Nl I iiIMMmi; S ,\i,.,i JlJfiJ 

BCil i ., H b i l'"''lt l.aH.t|.HiMSMii(N.'i nt'iir VieK.t ■ Matbour M'\ 

■Hmi-- "i-.-.r. iiir t ,tn. i ijtj ,-i '\, lucnntHii 

River Torrens. and Homnuin (1%S. |0AJ>*) ivleind 
io ihem as lite Adare Clay wheic Ihey Hank ihe 
Rivers Hindmaish and Inmau in Ihe Viclor Harbor 
area (Loealilies shown on | T ig, 1 1 The red-coloured 
alluviimi heats consistent siraligraphic relationships 
Io a younyer, erey bin k allii\ium IWaldeila 
li.rmalitai ol VVaid (1-9(56). Walkeivtllc Sand Of 
Iwidale (l^o.S) and Ihe Breckan Sand itt BoultTlttll 
( |yfiK)(i which rbmis lower leiiaces and lloodplam^ 
set within valleys carved out ol' ihe red alluvium of 
the Pooraka I ornialion Psuianne shells collected 
fl'OlH within die Waldctla ! ; oruiaLion in the lower 
reaches of ihe Onkapahrnja River (Botiiman I *>72 ) 
relumed a radiocarbon aue of 4oS0 ± 160 ye ( irs \i,l\ 
tHourman l\>7\>>. puring UllA middle Holoccnc lime 
the lower, near coastal reaches of many streams were 
shallow; sheltered estuaries as revealed hy die 
ptehCDCC nf lussdilerous marine deposit.-- at de[>lh up 
\alley This inieiptelation is supptirted hy evidence 
iVoin a locahtv sescral kdomelrcs Irom Ihe coast on 
Hie lower ( >Mkapannea Riv.t where an aboriginal 
l-.iieben midden CiMilailVlll£; cstuarinc shells dated al 
'VSJD i l H) years H.P. (N.H. I'indale pars, comrn. in 
Twidale ft ol Wftl) i 1 - s.ied i)n a well drained sand 
dune site at 20 m asl and adjacent to die farmer, mon. 
eMeiisive estuai^. 

Ages ascribed In the Pooraka Formation 
ami its equivalents 

The Pootaka Pormaiioti has t'cnerally been 
;i-cnbed lo Ihe I. ale Pleistocene, wijh most 
iiumerieal aees. based on radiocarbon dtflilig lallin^ 
within Ihe tanye til S0 0O(,)-2O.I>00 vears '.see res 'iew 



t Study Sites l2^ 
i^- Dry Creek, Pooraka 

l^r - ^ River Torrcns, Walkerville 
liZf^ Hlndmarsh Rivor , Victor Marb 

Fig. i , Location map of sites. 



in C'allcn et at 1995). There has not been universal 
agreement on it) whieh part of the Late Pleistoeene 
the sediments should be alloealed. Twidale (1968) 
assigned the Klemzig Sand to the Late Pleistocene 
and demonsl rated that it must be older than 6,350 
years BP. The distal end of the femur of a giant 
extinct marsupial, recovered from 3.6 m below the 
surface of a lillsLralh terrace cut into the Adare Clay, 
in the Hindmarsh River, was dated at approximately 
12.6(H) years BP (Oak - 2356) (Gill & Bourman 
[972). Firman (1969) and Daily el al. (1976) also 
considered the Pooraka Formation lo be of Late 
Pleistocene age, but younger than the Anadara- 
bearing Glanville Formation, which is now widely 
accepted to be of last inlerglaeial age (c. 125,000 yr 
BP; Murray-Wallace el al. 1988; Murray- Wallace & 
Belperio 1991 ; Murray-Wallace 1995; Belperio c7 <//. 
1995). Confident separation between Late Pleistocene 
Pooraka Formation and earlier Pleistocene alluvial 
sediments is easily achieved in the coastal /one 
where they are separated by coastal facies. Inland, on 
the Adelaide Plains, the Pooraka Formation is readily 
distinguished from underlying Tertiary sands, the 
Keswick Clay and the Hindmarsh Clay. The Pooraka 
Formation is only weakly consolidated, carbonate 
impregnated and mottled in comparison with the 
underlying units (Sheard & Bowman 1996). Ward 
( 1 966) assigned the Christies Beach Formation to the 
Last Inlerglaeial as he considered that the surface on 
it was graded lo the last inlerglaeial shoreline (his 
Lpimonasterian high sea level) at approximately + 3 
m above present sea level. However, at thai time the 
Lasl Inlerglaeial was thought to be considerably 
younger than the present 125,000 years BP. 

From the Dry Creek alluvial Ian, Williams (1969) 
described reddish-brown clay overlying older grey- 
green and red mottled clay, now known to be the 
Keswick Clay (Sheard & Bowman !9S7a,b; M. 
Sheard pers. comm. 1997). Williams (1969) also 
noted a calcareous red-brown earth developed within 
the sediments containing nodules and cylindroids o( 
pedogenie calcium carbonate, A radiocarbon age of 
34,600 ± 2700 years BP on carbonised wood from 
sand 3 m below the land surface was obtained by 
Williams ( 1969). The carbonised wood was regarded 
as delrilal in origin and thus was regarded as a 
reliable representation o\' the time of deposition. 
However, it the carbon were detiital, ils age should 
predate the lime ot sedimentation, which would 
make the ' 'C date somewhat older than the time of 
deposition. The date was taken lo indicate a last 
glacial f Wurm) age lor the sediments. Further 
radiocarbon dates supporting a last glacial (Wurm) 

CJni'in. DJ. (1943) Geological reconnaissance ol' part o! ihe 
I Lundreds ol lineoimler Day ami Onohva. BSc (Hons) thesis, The 
UmveiMly ol' Adelaide (iinpuh.). 

age were derived from a study ot alluvial fans on the 
western side of the Flinders Ranges (Williams 1973). 
Carbonised detrital wood recovered from depths of 
8-9 m and 15 m within the Pooraka Formation 
provided radiocarbon ages of 33,270 (+2130 - 1680 
years) BP and > 37,000 years BP respectively. 

Strati 'graphic observations 

Critical evidence concerning the age of the 
Pooraka Formation occurs al Victor Harbor Here the 
relationships between the last interglacial shoreline 
and the Pooraka equivalent unit, the Adare Clay, 
suggest that the unit is much older than 50,000 years. 
Bourman (1968, 1969 1 ) established that red-coloured 
alluvium forms fill-top terraces along the Inman and 
Hindmarsh Rivers and grades to a shoreline at c. + 6 
m above present sea level. The age of the shoreline is 
considered to be crucial with respect lo the age of the 
terraces and ihe sediments which underlie them. 
Twelve species of shells have been identified from 
this shoreline deposit (Guppy 1943-) and it is 
significant that they contain the sub-fossil Anadara 
trapezia, initially. Bourman (1968, I969') followed 
Sprigs: ( 1952) and assigned these shells to the 
Holocene, Subsequently, the shells were radiocarbon 
dated returning ages of 33,170 + 3.180 -2,270 years 
BP (GaK-5561 ) and >30.320 years BP (GaK-6099). 

Although Ihe above dates are compatible with 
those of Williams ( 1969. 1973) they arc questionable 
because the period around 30.000 years BP was a 
time of low sea level. Furthermore, it is now 
generally accepted that materials whose true ages are 
beyond the range of radiocarbon dating (> 40 ka for 
most laboratories) may yield younger apparent ages, 
due to the diagenic incorporation of low levels ol 
radiocarbon from modern activity. Thus, materials 
with an infinite age by radiocarbon dating techniques 
may yield an apparent age o( 37 ka due to the 
incorporation of 1% "C with a modern activity 
(Gupta & Polach 1985). Gill (1974) checked 
radiocarbon dales oi this age against other dating 
techniques and concluded that radiocarbon dating 
may be reliable for young materials but older 
materials may return ages that are far loo young. 
Similar conclusions were reported by Bowman & 
Harvey ( 1983) and Belperio el al. ( 1984). 

Not only do Anadara shells occur al Ihe + 6 m 
shoreline at the coast at Victor Harbor, but extremely 
large Anadara shells were recovered from a sewer 
trench c. 1 .6 km upstream al a depth of 4 m below the 
surface, within the Pooraka Formation equivalent 
unit and at the same absolute elevation oi' 6 m as al 
the shoreline (Fig. 2), A drilling programme (CSIRO 
Soils Division) further revealed the intimale 
association o\' Anadara shells with the Pooraka 
Formation equivalent unit, demonstrating that here 
the Anadara, lasl interglacial deposits (Glanville 



a last inlei^lacial agC: 


Formation ) and the Pooraku Formation ate 
intercalated coastal and terrestrial equivalents. The 
Aumhhn she IK have heen dated hoih by Uranium- 
Thorium techniques (IDO.iMHMStt.onn y^rfi UPi 

and hv ammo-acid tneemisnlion studies ( bomber & The slratigraphie iolali«msbipx between Uim 

Milnes 19X4 1. which provided results consistent, with inlerglaeial molluscs and (he Pooraka h'ormalion 

o - 

ii n 

^ = 

^ ^natfara Shells (- Glsnvtlle Formation} 


— — . ""-^ 

- 9 - — 

Pooraka Fflnrmlion 

- ? 

Soa Level 

; i iM>Om 

i nj n Sketch section showing, the imci hireling relationships wj [tie Cilam ilk- Koi maliOIl iuu) the Poor.ik.i lortiuiion in ihi; 
lower Hindmursh River at Victor Harbor, 

I -m. I l'(miui:t;i|.h t-l 6 m high river Mull i Ul in Pooraka Formation Mtltfnenis downstream of ihc Bridgewny lluicl tin hiv 
t i.vL- Poomkii MiiMinuin section exposed a 7 in. Red brown wnh wilti ussoaiulsd calcium ewrbunulc www bi "pp** 1 
pari o| A'Clkuj- At 'he huMj ol the sccimhi Mw INn.uku PormuMon rests iinconlormabl\ on an older PJtftUoceiW unit that 
clo'.clv iv.emMes the Titrtllfft Formation nT Want i l%m hui which She-aid (ft Bowman ( IW*uM consider Lfi more lib - 
io be itic Keswick ( lay, 



equivalent, near the coast, at Victor Harbor provided 
strong indications that the Pooraka Formation might 
extend hack to the time of the Last Interglacial. In 
order to test this hypothesis, and to determine 
whether the inland, terrestrial Pooraka Formation 
sediments were of an equivalent age, an attempt was 
made to obtain a numerical age for this formation 
using the technique of luminescence dating. A key 
site chosen tor sample collection for luminescence 
dating was the Dry Creek (Pooraka) locality where 
Williams (1-969] collected samples of detnta! 
carbonised wood and carbonate for radiocarbon 
dating (Fig. 3>. Unfortunately, the steep river bluff at 
this site has now been contoured and landscaped so 
that it was not possible to sample from exactly the 
same site as Williams ( 1969). 

A second site on the River Torrens at Walkerville, 
where a thick Pooraka section had previously been 
exposed, was selected as a subsidiary luminescence 
dating sampling site. As both sites have suffered 
from human modification and landscaping of the 
former eroding river bluff's it was decided to collect 
samples for luminescence dating by drilling using an 
auger drill with an internal push cylinder. This 
method allowed sampling depths to be determined 
and the sample to be collected without exposure to 
light. Two holes were drilled at Pooraka and one at 
Walkerville. Samples were recovered at depths of 3.5 
m, 4.5 m, 4.8 m and 7.5 m (for Pooraka) and 4.5 m, 
5.5 m. 6.0 m, 7.5 m, 9.0 m and 9.3 m (for 
Walkerville). The drilling site at Pooraka was located 
7.74 m above the base of Dry Creek and the second 
site was I 1.68 m above the River Torrens level at 
Walkerville. The same holes were used for both 
sample collection and scintillometry for dose rate 
determination. A summary of data collected For the 

Pooraka and Walkerville samples is shown in Table 
1 . Sample PK I S from a depth of 3.5 m is close to the 
level from which Williams (1969) collected delrital 
carbon for M C dating. 

Luminescence dating ntetheds 

Three methods for luminescence dating (LD) o( 
the sediments were used: selective bleach 
thermoluminesecnee (TL) o\' coarse-grain quartz 
(Prescott & Mojarrabi 1993) and green light 
stimulated-lumineseence (GLSL) of both coarse 
grain quart/ and of fine grain separates (Ailken 
1994; Duller 1996). In the dating of sediments it is 
assumed that exposure to sunlight is the agency that 
resets the luminescence clock and that the sample 
has been exposed to sunlight for a sufficiently long 
lime thaL the stored energy giving rise to the 
luminescence has been reduced to a low, near zero, 
level. This is a reasonable assumption in open sites 
exposed to strong sunlight, but this may not be true 
where there is the possibility that the material was 
deposited by, or under, water, as in the present sites, 
or in a generally eolluvial environment. Both the TL 
selective bleach and GLSL methods seek to 
overcome this uncertainty by making use of an 
easily-bleached component which can be reset to 
zero by short exposure to sunlight. It is assumed that 
this component has, in fact, been reset. Details of the 
methodology are presented in the appendix. 

Dating results 

Pilot TL runs were carried out on coarse grain 
quartz from all samples except PK2S/4.8, WV IS/7.5 
and WV1S/9.3. Such runs are designed to assess 
whether the sample is likely to be datable and. if so. 

TABU: L Summary of collected data for Pooraka and Walkerville samples. 





PPM '111 

PPM U Set. 

PPM Th Set. 


%K Set 

?; h»o 


1 .5 ± 

7.11 ±2 

148 ±0.11 

7.23 ±0.5 


7.12 ±029 

1.08 ±0.03 

0,86 ±0.02 

6.8 ±0.07 


1.1 ±0.3 

7.8 ± 1 

1.26 ±0.1 1 

7.31 ±0.4 


7.68 ±0.29 

0.89 + 0.03 

0.82 ±0.02 

3.7 + 0.06 


1.1 ±0.5 

6.3 ± 1 


6.24 ±0.4 

1.34 + 0.16 

6.76 ±0.28 

0.89 ±0.03 

0.91 ±0.03 

4.4 + 0.07 


1.2 + 0.6 

7.1 ±2 

1.22 + 0.10 

6.91 +0.4 


9.69 + 0.32 

1.20 + 0,04 


12 + 0.07 


1.8 ±0.3 

I2± 1 

1.81 +0.12 

11.7 ±0.5 

1.89 + 0.24 

12.5 + 0.44 

1.85 + 0.06 

1.60 ±0.04 

8.8 ±0.07 

WV IS/5.5 

1 .6 ± 0.6 


1.93 + 0.13 

12.6 ±0.6 

2.33 + 0.24 

10.5 ±0 42 

1.81 +0.05 

1.32 + 0.04 

7.5 ±0.07 

WV IS/0.0 

1 .9 ± 0.5 

10± 1 

1.86 ±0.12 

10.3 ±0.6 

2.47 + 0.25 

12.1 +0.48 

1.59 ±0.05 

1.54 + 0.04 

8.3 + 0.07 

WV IS/7.5 

1.7 ±0.5 

If ±2 

1.66 ±0.12 

11.8 ±0.6 

2.66 ±0.26 

ll.l ±0.47 

1.71 ±0.05 

1 49 ± 0.04 

15.7 ±0.07 


I.N ±0.5 

13+ 1 


134 ±0.6 

2.95 + 0.29 

13.4 ±0 50 

2. 19 ±0.07 

1.71 ±0.04 

18.0 ±0.07 

WV IS/9.3 

1 .2 ± 0.5 

15 ±2 

1.88 ±0.13 

13.0 + 0.6 

2.95 + 0.29 

13.4 ±0.50 

1.98 + 0.06 

1.71 ±0.04 

164 + 0.07 

The first two columns are the results derived from Thick Source Alpha Counting. A DNA was done for uranium only; ppm 
Th (DNA) were obtained by combining the count-rate from thick source alpha counting and the uranium concentration from 
DNA. ppm U Set. ppm Th Set & %K Set were derived from the on-site gumma ray scinlillomelry data. 



lo give a liniiied-aceuraey estimate of the acquired 
luminescence and the sensitivity in radiation, and 
hence plan (he schedule for a complete dating 
procedure. A pilot run consists of eight discs; half of 
the*£ are bleached, alter which two discs each of 
bleached and unbleached are given a radiation dose 
o! 60 Gy. For all samples except PK IS/3.5, the 
shallowest of the Pooruku samples, the TL was 
clearly saturated and no further work on them was 
justified. Although PK IS/3.5 was approaching 
saturation, a full dating procedure was carried out for 
both TL and GLSL. 

Such procedures give the Equivalent Dose. Do, a 
measure in iiravs of the energy absorbed by the 

sample from radiation in the environment since it 
was last reset to zero. The age of the sample is found 
by dividing (he equivalent dose by die dose rale in 
gray per kiloyear (Gy ka '). 

Equivalent doses, dose rates and ages 

TL glow curves are shown in Figure 4a. shine- 
down curves in Figure 4b and corresponding dose 
curves are shown in Figures 5a. b respectively. None 
of the curves is scaled. It is evident that the dose 
curve of quart/ (Fig. 5a I is close to saturation but that 
the growth curve for GLSL on fine grains (Fig. 5b) 
has a different shape and the curve eonlinues to rise 
quasi-lincarly for high doses. Tins is because the 

'. :ni 

, IV, 

Temperature (°C) 

Fig. 4a. G 
In each 

low curves for sample PK I S/3.5. The figures next 
curve indicate the dose in Gv. 


1 / 

o o.O ?on 400 60Q son 

Dose Gy 

Fig, 5a. TL growth curve for sample PK IS/3.5 for the Id 
interval at M)5' C. For Figures 5a and h. the curves arc 
filled hy the "Australian slide" method; the (natural + 
dose) points are shown by crosses: the (bleached + dose) 
points are shown by circles. There is an apparent 
sensitivity change for The TL bleached curve but the 
scaling factor does not differ significantly from unity. 

20 30 40 50 60 70 

Shine Time (in seconds) 


lig 4b GLSL -shine-dmvn curves for sample PK IS/3-5. 
The figures next to each curve indicate the dose in Gy. 

Fig. 5b. GLSL growth curve 
integrated over the first 100 s_ 

lor sample PK IS/3.5 

AGl: Ol Pc >< Jl< \K A l-OKMATION 
Iami.i 2 . htfiitvuli in Haw, tlbse fWtfi ttnti tffcei A" WftJiftfr- nt i S/SS 




I) Lv 

selective hk-in.ii TL 
249 ± 5H 

(il SL 

20s l -U 


> ISO 

dose rait 
t.v ka 


I 90 ±0,06 

LQft ttUM 


2.54 ±0.12 


2.54±0 H 

Weighted mean 



ElUtf kil 


105 i 2 

iti + 6 

> ioo 

undifferentiated line grafts consist o\~ a mixture of 
minerals in which quail/ and feldspar arc dominant. 
Thus although ihc quail/ Component saturates, the 
U-klspai component docs not, Equivalenl dose values 
from all ihrec uielhods appear in the first line o!' 
Table 2. Il will he noieil Ihul the two methods ol 
linding I), give different values; this is characteristic 
ol Ihe methods, 

Two distinct methods for finding the dose rales 
were used (described in the appendix), the aim bring 
to gel two independent values lor each sample, to 
improve the statistical precision and as a check im 
Ihe presence ol radioactive disequilibrium (Prtfscftli 
& I lutton 1 005 1. 'Ihe values obtained are included in 
Tabic 2 As wilh I),. dose rates differ lor coarse and 
line grains The values are in excellent mumal 
agreement ant! this shows thai radioactive 
disequilibrium is ahscnl, 

The weighted mean dose rales are shown on line 3: 
line 4 shows ihe aces derived from the P. using die 
age equation from the appendixThc weighted mean, 
1 16 ± d ka is dominated by ihc GLSI. determination 
lor Hue grains 


Allowing h)\ statistical lilting uncertainties, all 
three ajgeS are in good agreement. Bearing in mind 
that they are based 0|1 different physical processes, 
all of which assume that the luminescence signal was 
set lo /em in Ihe pasl. it shows thai this was very 
likely the case AnA that ihe age being determined is 
indeed die tunc eff deposit Ton If ii is not so. the 
apparent ages will be loo large. 

As already mentioned, the dose growlh curve ul 
Figure 5,i, which is lor qnarl/ TL. shows that the 
luminescence is close lo saturation. Tor ihis reason 
Ihe cslimalc ol relative uncerlainiy given by the 
analysis is large. This is also Irue for CI. SI. ol quarl/. 

Tor GLSI of tine grains, on die other hand, Ihe 
presence ol the fcldspai component allows |> to be 
found with significantly belter precision. 

The best estimate ol ihc time Of deposition ol 
sample PK1S/3.5 is (lie weighted mean age. I K>i 
6 ka In fact, this is dominated by Ihe GLSL 
deiei munition on fine grains. It is probably best lo 
regard ihe quart/ determinations by selective bleach 
and GLSL as being supportive of the GLSL fine 
grain age. Al the two standard deviation level the ace 
exceeds 104 ka. Consistent with these numbers, the 
saturation of the. luminescence of all ihe other 
samples shows thai they me as old or older Ihnn this, 
Therefore, although only one sample has yielded a 
numerical age. There is enough evidence lo establish 
a lower limit lo the age ol the formation and il is in 
support of l lie geomorphological and sedi 
mcnlologieal evidence. 

In addition lo the three methods already described, 
infra-red stimulated luminescence (1RSL) of fine 
grains tWintlc 1004) was tried 1RSL uses only 
feldspars and is less susceptible to saturation, 
although il may be subject to long-term lading. 
Allhough the growlh curve was similar in shape in 
the GLSL curve o\ Figure 5a, Ihe statistical filling 
procedures did not satisfy our criteria for an 
acceptable D,. beyond showing that it was greater 
than 100 ka. Whether this is due to the sample or the 
methodology remains lo be determined. However. 
the result is suffkiemly encouraging io suggest that 
it may be possible lo find ages (rom the lower levels 
at I'ooraka and lor Walkervillc where quart/ methods 
were unsuccessful. 


The location of the dale derived from the uppei 
section of the Pooraka Formation al a deplh of 3.3 ni 
below the surface is m complete accordance with ihe 



numerical uuc obtained ol I J6 i fi kit, \v i I h the Last 
iitlcrglucial ranging Irwin approximately 132-1 18 ka 
to Chen ( 7w/ IWI : LanAwh & Nakuda RWl« Zhu 
,/ of- f*W3; RfjTiiitfz tf <//. IW5* fosenfuner *v ///. 
|*J%). J he ( ICSl technique oilers the possibility ol 
obtaining a luminescence dale lor lowei scciioi^ ol 
rhe Pooraka Ponitiftrori thereby fiutnliitrtitijj the 
calculation *»f Ihc rates of sedimentation during the 
I ,jki Intere.lacial: ii may he that sedimentation during 
waonci. wetter, iasi nilcrvJaciiti limes was quite rapid 
and Ihis technique pflfelN Ihe opportunity to lesi ihis 


I h$Ke result* sut^eM a far gieaiei age rutj£<! tor ilk' 

PoDrdkii tH'M.iaii.»n than hfts prcvtntislj biicn 

< ; nWed In liic past, the auc or the Pooraka 
liuiii.iiion das been poorly constrained m about ?()- 
ill ka HP. possibly heeaiise ol the now-known 
Jiimnaiom- (>t ladiocaibon dating. furthermore, i| 
>|i«ni|d »»e iMt(ei) thai oilier' melliods such us TL, 
' il SI ami IKS!, were not available when niJJiy of 
die oulioc;itho|i dales were tamed out. 

Many observation* and cores indicate thai, m 
pl.h is. die PooiaU I urination appears to in- youmici 
limn Ibe Past iiverylacial < Hum die formation. 
especially in subsidiiu' areas. Por example. 
Ludbruok ( 19701 in die Por! Adelaide area described 
Pooiaka Formation overlying calcieU J J Glanville 
boinnnioii. ulneh in mm overlay ibndinarsh Clay, 
lielperio and Rice IPJN*)) showed thai o| f>2 cores 
troin ihe Gdlnian area. I I record lite sequence of 
llol(tcene overlying calcreled Glanville formation 
withoiii Pooraka Pormaiion. aiul 5 record Holocene 
oveilyin.u Pooiaka winch o\erhes ealcrvlcd Glanville 
formation, The calcreled surfaces are interpreted as 
pedouenic features developed during subnet la I 
exposure and would have lormed prior lo ihe 
deposition of the overly ine sediments ol' the Pooiaka 
formation. In contrast, tour ol the cores record a 
sequence of I toloccnc/Pooraka/Glanvillc ftynijufipm 
withoui a well-developed calereie on ihe Glanville 
pormaiion, and otw records imet mixmv of Pooraka 
and Glanville. These last cores muy indicalc coeval 
(ilanville and Pooraka scdunenlalion or litlle lime 
break between the two, 

l-Alensive work by Sheard & Bow man ( IV$ft)< 
which involved drilling coivs lo depths of 10 m ;n 
170 siies over die Adelaide Plains, intersected 
Pooraka Formation in SO cores. This work 
eoitoboralcs the tiiuline- of tfelpeno & Riee I 1089) 
tor the coastal /one. More landward sue- indicated 
die sequences to he: I loloct-ne/Pooraka/K.esvvick 
CWMnnlmarvh Clay and Hiiluet.-ue/Pooraka/ 

Wi •! »i »m it:. I ci mi lii .V Mro I't'K Hly I i«J t I'I'vSi Apprinlis -. 
Ci ■>!. lii^ 'Mi>.ilntiv, in l|'»|,|t;(^i Shi»iv^ In ti;uiklciMoii t ' 

Hi miti ii 'i'-'» ',- l nurol Wptkj and tur. wiiuii IU(Uia tn 

I IriiiHj I' , Vvihotiiv nj SA. iinipnlr \ 

Kc- v\ fck Clay/Terliitry saivls or Adelaidean rocks, At 
the MO Mies unersectin^ Pooraka Pormaiion 
sediments. Sheard k Bowman il^h) r;i.ei\ 
tSpi.aiem.ed JifFjC uldcs [\] identify ine Pooraka 
Pormation from the flVerfvfng and underlyini' 
uuitctiats iM Shcuid pets, conini. l L 'V7r 

Receii! drilliiu' work by Woodhcad n tit. I l*-)°5 » 
for hiauldershine llornibrook in the Holdfast Shores 
itilcne-l.i! Pala\\aloiifja Uedevelopmeul area) hus 
demonstrated thai deposits ol IVioraka formation 
2-4 in Ihiek occur well offshore bom the 
present KOUSt line. These deposits ate delachal 
fiom moi\ landward deposits either by /ones 
ol nol)-deposttion or by early Holocene erosioa, 
Vanable stratigraphic relalionships are appareni in 
this locality. Pot example. lUc lollowiuy relatiiaisiups 
ot lloloecne/Pooraka/Glauvillc/Hindinaish Clav 
lloloLL-iic/'Gl.aivdtiv' tbinhnat'sli ( la> ctf I Wcfi CRO/ 
tultuun Sand/Pooraka/CilaiiMlle/Hindtnarsh C 'la\ > 
occur within an HOU m ea^t-west seclitm. 

It may be that the Pooraka Pormaiion was 
deposited over a considerable period of lime. Tins 
\\\'\\ is v dso supported by the occurrence t»| 
palaeosols within Pooraka Pormaiion equivalent 
sediments as exposed in Schick^ Creek and Cobblers 
Creek. The luminescence data indicalc an art- 
approximately coincident vvilh the \.,wi hueryl.k «,il 
( I2.*i ka HP), but the results at any one place may be 
influenced by the tcricstrial/innriue interactions, lor 
example, if there is not an active supply ol alluvium 
to Lhe coastline ihcn marine deposition, exposure d\u\ 
wilcrete loimaboii will dominate, m |.»luees 
.subsct|uenlly manlled by Pooraka or Holocene 
sedinienis. It may be dial only where there is u 
sufficient supply ol terrestrial sedinienis to the 
coastline, .such as where streams debouch al tfn 
toastluie in lelativcK constrained valleys (e.e. 
Hindmarsh River al Victor Harbor), Unit Ihe 
i ntei fingering eharaeterislics of lhe two sediment i 
can be deuiousiraied. It is also possible that Pooraka 
Pormaiion sediments do not reach the position ol the 
Glanville formation luuil after it lia.s been calcreled, 
It should bC noted that milch ot the Pooiaka 
formation has been deposited inland, well beyond 
lhe elevation ,)\k\ planimelric posilion of the last 
inUrrcfacml shoreline, so thai in ihese situations iheiv 
is noUcar sirao t eraphic (elalioiist)ip beiween rhe tWtJ 

hnplimti<tt)\ t») tlw Pixtrnka J't>n»ah"h tAhnJiur 
hu<i tf> thr fast hiiri^hjt ml 

II (lie interpretation that Ihe Pooraka fotmahon 
exlends liack lo Ihe Past Inlerelacial is coricci, a re 
evaluation ol the rudioearhon ayes rhtii inihuU>' 
glacial or iiiterstadial aees for the sediments is 
required Ahcrnuuvcly. the conflictm;. 1 ivsnli> iiiiiy 
suggest that there could he sedimeots. who fl 



although appealing similiii, occupy a ri*ngt: ol ugws. 

At .1 number of locations, the Pooraka Formation 
has been nuioJ tn overlie ihc (ilanvillc Formation 
vvilh a variably developed ealcrcli", Wggirtlillj! 
MiHiieL- exposure prior lo burial by younger Pooraka 
hot nun ion sediments Flsewhcre. such as in Ihc 
lowet Hindmarsh Rtvci. where iherc is an mieipluy 
ol coeval coastal and teiicsiiial events- ihr 
slraligraphtc relationships [jCIWeeti [lie PopMKfl tltl<l 
(ilanville Formations may >imply represenl a lucics 
change ami not a geological succession A 
considerable inne interval may be requited tot the 
deposition o( the Pooiaka Formation su tlial in act" 
could be diiktiionte. with deposition i»cuii'uiiii 
dining i he* rise in sea level up ni and beyond thai o\ 
Ibe lust mici>'Jaeial lew), 

A lust inleiglacial age provides appropriate palaeo 
climatic and pabeo-sca level conditions for 
explinnini' the distribution ol the Pooraka Formation. 
CJust? lo the eoast ihc unit relates to a shoreline 
higher than the modern one. while inland ihc welter. 
ilHUJ&Jflcta! i uiiditionv Would have favoured 
aegradatiou of sediments as opposed »o dtiei. gkiC iaj 
conditions lhal would have faedilalcd dis>cciion, 
Thb timing would also ensure sufficient time both 
lur tlw- htnld up of exlciiMVC deposits or ibe Pooraka 
Formation ovo knee .iieas including much of the 
city of Adelaide, and lime during glacials and 
imerMadiuls lor erosion .it ihc Ponoikii I ormniion lo 
develop die extensive terrace -\-.n-m ol die Kiwi 
THnvns (Iwidalc IWN). 

A last inlerglaeial age lor Ibe I'ooraka Formation i, 
ol* signilicancc loi archaeological prospecting. Al l|ie 
pieseni lime theiv is dehaie concerning ihe jntU|u»Ly 
of hmnau.s on the Australian coniincni. Substantia! 
iluii •. in vei'et.ition ''elated to Aboriginal burning 
practices bave been interpreted as resulting ff4Mfl 
human impact and used to infer Ihe arrival ok people 
in Australia Ufi '"a," ago as 1 40.000 veai's fKctshaw 
ft w/. Iv>'>v Kershaw hjijj, PW4.* W5i Ihe-.e 
chums haw Ken ejUe^Lioiicd by various workers (c g- 
Andi-i-MMi r*W; Mope P>V4; Wlntc |4QJi Webb 
ts»*/5l. II the Pooraka Formation i.sol lasl inlerglaeial 
lljEW 11 may present a prospecting opportunity to lesl 
il Abiuigmal colonisation had occurred in southern 
Ausf-ialia pilot to 125JKX1 y«JN UP during Ihe l mc 
• .kibe penullunaLe glacial low sea level. 

A minor p daeomagnetic event, the M3-£Ujlcd 
' Plake l-Aeuf . has been idenlitied m the noiihcrn 
heini.spbeiv in loess sequences (llougho el <// PJ l .'M 
at approximately 120 ka. Given lhal the general ilgt 
ol lite Pooraka Formation is almost certainly ol laM 
inlerglaeial age. and lhal the upper pari t»1 the 
lormalion has been ascribed a GFSt . age of I lb t fi 
ka. Ibere may be opporiunilies to identity ihe Blake 
evenl ill the southern heimspl^ic 

A last mleiglaeial aye lor Ihe Pooraka Poi illation 

has implications bu ihc antiquity Dl Ihe tbtpnXtidttfl 
In Ihe Adelaide area, Oipiolodontid remains have 
been recorded well back into Tertiary sirala in the 
North Kliuders-Callahonna Plains areas (Callcn & 
Tedlord l u 7fr, pledge & Tcdlord P»0: M. J. Sheatd 
pers. eomm. I l Ml.) but [101. so far, in the Ailelaide 
area. Discoveries o{ DiptvMoti remains ha\e been 
made tn the Pooiaku Pomiiilton tlvvidale !%N. N 
Pledge pers. eomm. fljJJ6) of the Adelaide ;m_*a and 
while ibe I)iffmt(.<<l'Ht n*iay have predated and 
.survived well beyond the age of toiS scthmcnlaiv 
una. it was almost certainly roaming the swamp). 
abrading Adelaide Plains some I25JOT Veaiv U^O 

liic age has further implications IY>i landscape 
evolution as there appears lo have been a ittaioi 
erosiomil hiatus of sonte 120.000 years between Ihe 
deposition of the Pooraka Poiniaiion ami ihe gre> 
black Wiikleila Formation, which is of mid-Holoeene 
aye. There were no sea levels higher than Ihe preseni 
in Ihe intervening period so lhal erosion would have 
dominated ibis inteival of time. Alternalively, any 
sediments deposited during ibis lime could have 
Iven lemoveil in late stages of erosion. 

( nnelusinns 

Ihe mam «_onelusion from this work j$ that ihe 
Puoraka rormaliou mu>i span a far yrcalei lime 
\x l iotl iluui picviously reeoymsed. pn»bablv 
eMcndin-i .i-. tar back as Past Inlerglaeial lime i.e. 
125 k:r Il appears lhal frfrc leclomc ami 
eiivironmeiital selling, which influence^ the snppl\ 
of leriesirial sediments lo the coastline, is ol extreme 
impoilance in imerprclmij Ibe -toitigtapliK 
lelaiionships between the Pooraka Pornuhon ami Ihe 
Gkmville Formal ion. Much more luimnescenec 
dahnjj vuiri. «, iciuiieil lo n.ivdrain oritoeitmeni Ihc 
aye ranges in different >ctlmt-"> i.c, in Ihe Vicnn 
Harbor selling where the two are imetrmveil. in 
eontrasi !»• ihe Port Ado/aide se-itme whcie a inajtn 
ealcrete palaeosol separates them Wc may cypect 
luminescence dating leelirnttues to doe-nrnem ages 
from last inlerglaeial tune-., possibly LhtOUgh !•> M)- 
.S0 ka. if the qiiuii/ grains ,ue suffieieiiily tesei. This 
oilers Ihe posMbtlily * »l pownhny a more reliatile 
dating tool than radiocathon leehnit|ues. which still 
snuggle Itl prt»v nle tneamiigdil dales past 4M ka. 

Ihe o'il-notvvii alluvial ^ediiuenis. reiericil Ul as 
Pooraka Formation and et|iii\aleuls, are lousuIcumI 
lo extend back lo at ieasl the Fast Inleiglacial ol e 
125.000 veats \>A: Thts is demonsiraied by the 
Miutiguphica! .eljiionshitv- »,i the red-brown 
sediments lo both younger \uk\ older &CU*iAlC|)tft, the 
inienligiiaiing of demonsiratctl lasl inlerglaeial 
maiine ilept>siis with ihcm ilieii grading to a i- b m 
higher shoreline containing lasl inlerglaeial marine 

L >: 


tl kftsilH at Victor 1 Rubor, anil ifit! fact lhal a higher sea 
level, togelher wilh associated warmer and wellci 
conditions would favour aggradation, whereas 
COldcr, drier glacial Conditions associated wilh lower 
sea levels would httve i'uVMUred erosion. All of iheso 
laelors support a lasl inierglncial ape. 

Hie IttniineseenecdaUi have dciuoilMralcd Viability 
as an independent means of testing ihe hypothesis 
lhal ihe Pooraka l<u maiion is ol last inlerglacial age. 
I In Mai sediments present special piohlenis lor 
luminescence daling because Ihe scdimenis hcing 
waler-deposited. may nol have been exposed Lo 
sutllieht lor sufficiently long to zero the cjtiurt/, 
grains resulting in inherited wtUiralion levels within 
Ihe grains* A I'tii-ihei eomplieahon niay be lhal in ihe 
Adelaide area, (he sediments may have nol been 
transported sufficient distances ftfr the quartz grains 
to have been /crocd. Given these constraints, il is 
exU'emcIv 'jiahGopj lhal i! u;is possible l<> aclue\e a 
reliable luminescence age lor die I'owtaka Formation 
ul l)r\ ('reek, offering (he possibility of furihcr 
advanee in this area 

An aim ol this project was lo establish the 
cKcclivciiess ol luminescence us a daimi! Uvhniipic 

lor Quaternary lluvial sediments m prdci 10 ivsolsv.- 
dilfertng and i<> facilitate 
correlation of river lerraees in different valley 
syslems. Clearly, Ihere is a need for many further 
dales to be Obtained from river terrace and alluvial 
fan deposiis over wider-running areas 10 writs die 
Conclusion* of lire present work. However. Ihe 
implications id an age for ibe I'ooraka Formation 
extending back lo Ihe Lasl Inlerglaeial are so 
significant lhal our preliminary results are presented 
here and provide the basis for turther study. 


Funding tor Ihe drilling of the Pnni'aka Km maiion 
and milial luminescence dating of the sediments WfUj 
provided b) a I inversus of South Australia Research 
Grunt, The Tl. analvscs weie earned out b_\ I* 
Marlinailis under ihe supervision o\ Professor 
heseott. We are gratclu! lo BrC Murtay- Wallace 
and l)i N, Alley for commenting on the text and 10 
M. Mishop for assistance with field w..rk. Wv arc 

appreciative o\ ihe signihcanl conlnhiilions of \\v 
referees Mr M Sheaid and l)i V. Gosiin. 


,\ori\. M. .1. (IVM-t Optuul dating, ii nou :,pceiuhht 
ii-vicu. QiiuL <if<HtuotL iJJunL Si i li,\J 13 S|H-S(i;s 
ii H.SUN, A. ' IWt Continent on .1. Peler While'.-- p'ipet "Sue 

820 hikI she evidence IVff i-.ui> octiupamm in Au^nalia" 

(Jthit .\nsir,tin\«i 12, 30- 1 1. 
hi i |»| RIH; A. I'. Smiiii. H, W.. Pi)] U il. II. A.. Nil IWJI I R. 
C \ t I>iM\mm:. l> I, Phi v on. J. R ll.-uis, .1. R Hi 
tiwsns. V. A- tJVtW) ChnnntlttMiLiil Sm.lie^ of |||..- 
Quaternary murine sedimems ol the northern Spencei 
Gull. SoUtll AiiMmha. Mar. tifttt, 61, iJ65-2Vfl 

&Kirl\Jf. I fP)R9) Slmtii-Riphir 
ihe Gilluiau dusclopiucnl sue. Ron Adelaide rstuuiy 
South .\n\( fh/'i \hitr\ <i>hl I iw'x>\ R, jwri h\ >,>!,. No/o\ 

\h, mrav-WMI \\ i. (.. V. \ C\>ih I II tiWo'lhe 

I uhI sliou-liue in scuiIiltu Aii4irtili;>- 

iiu>;u-:iph)c ^inalion^. hi ,( l.-Mitp.'i'alr i;urKmi*tC 

•ii'iui:. Quai, (ftteriuit, 2t>. 7->9; 

liOt'HMAN. K. P. llOOH) Urrmr:, ol llir Itilll.UI Jiul 

Uuahnarsh Rivfti fntuiut'.il, \7-2\ 

_ < \ { )72) Somr aHpeeis of ihr t"Ieoinnr|^inlu^ ttl thd 
i 'nkapuringij River, fhitl. ". 1 1 M i ; . 

j I 7V) I ( irOliHMplnilttL'ICjl LOIllrtliUlMMIS It) ( 'O.'l' 

niauajji'iiii'iii pp. B0-8M //' "IVoetv.lin^ l"nelw nil illll 
Souihem I Icni^e" (CUiKsUlvuliOn Count U Ml S,\ and 
Department ol Gomi tilling RductUittJl UntVfiMkfy ul 

lui\\M'a.J. G ,V H. ( |:vi v. N. HOXV' RudJ^'lflrboil daiiu; 1 
iiiariu.'^.helU. in S.>i,i|Ii Aio.iriiha. 3'/v7 AllihicHlt 17. \\^ 

G\i 1 1 \. R. A t"v Ti tjioho. \L II i l-» *oi isiu i .niio/ou' 
rOck anils and depositions! eiivironinents. I alt Iomiu 
ulca. South AuMO.Ilu. tulns K. fat V \tnt IIH). \2> 

. Siil'MU). VI. I. iO'.How.M G A: lii i rn-io_ \ p 

lb.i05) AltiivtMl fins jiikI piedmoni «lo|M deposits pp. 
241 :-G hi Drewfe ,1,1. & t'reiss. W.v [Rds] i be 

Gri'liMiy f»t ScUlli \limi [ltd Vt.t 2 The Rhuut-ni/ou 
Sixifft Atl\l tivnt Sun: Ihili 54 

Tin -, .1 H.. Cri!i>\\, H A Womi.B & \\ \\\\ ion tu., I i 
.1. ilWh Prcctst* Gti-ouoloL') ol thf biKl oiK-tL-lucuil 
pciitnl "'I • "Ttt ilutu Iroui IoxmI conit i»vl\ mi |li. 
Huhiunu^f.r-j/ &n \mrr Mi Htf. R2 l *7. 

U\ll\. li l : u y,\\\_ I. IT. Iwum ■,_ II (i \ ! I \HtftM I \! 
H07o,Ge..|oH > pp s V. MTwtclale C K.Tsl-.i M, I 
& Wehh. H.V, (IaI'-i "Natural Ih-aory of ihe Ailelauk' 
ktfgiOlT" ( Soueis olSoulli Auvlrulin. Ackfuik-i 

Pi i i ( r G A. T t looo, faced aewlopmems in 
lumineseeiiie diOfng »•! Gn.itooi.uA SoJoucuts f'nn- 

Oa.Mi sin it A.. Xih . /. K.. { oi t ins. L B,. W \ M(» f . K. II 

& I I- list \l HW, R. ( |W6^ UlL- I.UVl !UlL'IL'I;, C i;i| <., L ;, |0Vp| 

uIiuiil-c: lluw cvak'iKC Iroiu Ihe Abrolhov i*.l:ui<ls \\\>:\ 
Aiisir.iliii Gent, fbwtbil' S5. ft06 ol4. 
I LKMAMj B ( 10071 Slialii!ra|>lty o( lute Gjiuo/.mc tU-posiW 
in Souih \usliulia r^HW A' StJV S Al/Jf, W, IAS I8f) 

i |0fiOt SU'ali*'raphu .jiuI\ms ol mmIs Ueui AdeluuU- 
thhi W : " ~"k 



Ciiii- \i D i 197-Ti Carbon 14 and I raniuni/1*hoiiuril check 
Oil viyj:e>lcd lmers|adi;4 hi«h sea level around -U1O0P 
UP StchrtiS. 211 

& Kooums. K. P. < 1972,1 Research on Quaternary 

Shorelines in Australia and New Zealand - a summary 
report »*l the AN/.AAS Quaternary Shoicltnes 
Citminitia 1 , Report Km South Australia. Ihul. 3 IU3-1U4. 

til l-l A, S. K, & Pm w IL H A- ( I9*S) "Radiocarbon dalin^ 
Practices at \Nt " Handbook (Radiocrabon Laboratory 
Kiscaich School of Pacific SLudic-s, Australian \aimnal 
University. Canberra*. 

H<»v;iui, /ium.. K'Hi'O Tr Sha*. J. & Zhimiim,;, AN 

<t 9\i a detailed paUenaUignfllc refcoftl lor the 'tui 
rntcnjtacial period. frVfrt/l P/dWM Si I /■'" 133. 330- J5|. 

Hoi*. & ( 1 0<>4 1 Comment on ODP Site K20 and the 
interviicc pi L-aily human occupation in Australia. Glint, 
Aitstrnla\t\t 12. $2 33 

ll> Mi i v.nJ..HcrM»N t J. T.&Pkkm -mi. j.R tl<)tij| rht 
stranded beaeh-tlunc sequence ol south-east S« itlll 
Australia: a loM ol iliermolumincsccncc dyiinji- O-N(K) ka 
(Mmt.Sri. AVe. 12, I 20, 

UOOIRIY SVIMII. I), I & ll-VShlll, I 1 .. II. flV&M 
I chl-induced emission spectra from some quarts add 
feldspars, ffitf. fruiks ami Riuttat, Mens. 18. 127 131. 

. Ninskv M K-. Iiiovtsov. J. & Y E 

I I9X0» An impiovcd alpha scintillation counting meliiod 
lot oVtcrrrunainni ol |'h. U, Ra 22(>, Th-2>0 excess aiu' 
Pa - 1 tl excess in marine sediment*. (tin. J fainh Sri. 
23. 959-9%. 

II' iio^n. J \.Sc Phi "si oil J. K i 1992) Field and bbomiorN 
measurement ol" low level Ihonum, utamum ami 
poLissium Nmf, Tim_k\ atUt Umliat. \tt\t\, 20 367 _WU. 

Jk\u-N. M_ Is, & Pttl:S( m i. I K. ( l*W> The Thick Sounv 
Alpha particle Technique: Compaiisons with othei 
Techniques and Solutions to the Problem of 
Oven oundny f>\l'f ' Jma mil ' *), J5 -'*o. 

Kiksowv A P 0993) Palynolorjy. bioslraligr tphy and 
human impact fin Artefact 16. L2-l$. 

O^'Ml Site K20 and the evidence lor tarl> 
occupation in Australia a response (htut AuMralaMo 
12. 24 2SI 

( r0^5J The palacoeeologiral iccord Irom Silc K20 
it lunher response and research developments. fe£f. 13. 
M 2b 

. MiKl.N/.u. (i. M. &. McMlSN. A. (1993) A 

Quaternary vej*etaiion hi%toi> ot nonhea^tetn Queensland 
Horn pollen analysis o1 ODP 4itj2 H20 pp. 107 114 hi 
McKen/ie Ji .v/Oasies, P. J. & l\iln»er JnKin. A, U'-dsi 
$ % 9tH ■'. (h win Unit Ptyjjjuin St i. AYv»//\ 133. 

KiMLtnv, K. W. L & Mil SIS. A, R. i P»M) The e\|ei|l ol 
racomisalion ol amino acids in I luloccnc and Pleistocene 
marine molluscs in Southern South Australia 
Pietmnnary daUl on -■ utnr iiatncworK tor culCl^ 
lornmtn»n./\/is/ ./ hnth Sa 31- 27M-2Ku. 

f.AMItfi't; K. & NaKaDa. M. ( l*»2) Constraints im the Zpt 
.mil duration of the last interflaciul period and on sea 
level variations N«Utn'S51, \2S-\2K 

I.I 0ltr<«w»K- N. M- (l°76) The Cllamille t-ormabon at Port 
Adelaide ,V- Ami- thai Sun Q. OVoA \'ott'\ 57. 4-7 

MtiRRW ,' ' . ('. X liSi.i* Aminostrmijjraphy til 

Qualcn.u; coastal set|ucnces in southern Australia - till 
<ncrvie\v. (Junt. Inurnai. 26. W-XU. 

& BumtO. A, P (19W1 The Last Inlertrlacial 

shoivline in Auslrulia. Qnot St i. Rt\. 10 441-461. 

.KiMBtR.KW I...Bn,i'i.RKLA. P.&Gosiin. V A. 

1 198X1 AnmiONtrali^raph) ol the Last hiici^laciai in 
SouIIkmi Australia- Smirch W, W M\ 

Namhi, K- S V- <t AtTKhW. M. J. (|0M()i AnniiJd tlosi- 
conversion factors lor Tl . anil HSR daiuo- 1 An hut-Witt tt\ 
28. 202 -205, 

Plt-IHT. N. S, & 'If Ot.mi). i<, M. | 19001 Vertebrate Foss.Js 
pp.lMO-l'CW //, Tyler. M. .1.. Twidale. C. K.. Ddi tes. M .< 
Wells. C B- IPds) "Natural HisLory ol the Nnnh Ujst 
Deserts" {Royal Society ol^oulh Australia. Adelaide). 

PMMivn .1. U tSi Ml fO)N' T j, T flWBj Cosmic ray 
contributions to dose raL's lor luminescence and liSR 
daring; large depihs and lone term vatiations. Rtul. 
,V/,,7a: 23. 4*>7 S(M), 

& (l t ) , ).S> P,n\troninei'la1 dose rales and 

radioactive disequilibrium Irom sonic AtlUfaiUfl 
iumuiescence daliii" sites- (Jtn\i Si A A!i"i- (QliQtt 

GvOcHrQn 1 14,43**448. 

& MOMKIUHI. H (iW*) Selective li leach An 

improved "partial bleach" method r»i tmdine ei|uivale»l 
doses lor ihermoluminescencc datine ol quart/. Afhfi'ttt 
Ti,_ 11. 27-30_ 

. Mi viti:i. 0. I <V Ml nUNiLT « l ()t) 3i hsomation 
Of cquUalL'in dose in ibuimoluiniMescciice dating - the 
•'Australian ^lide" nuihoo thut |-S. 

Ki: \ont\o. M. L O^NHi rheimoluomiescence Jaiiny *il 
ijUarl/ m aeohan sedimenls Irom southeastern Australia 
Qucu. V/. f<*\. 7. :>7-2f>l. 

Mil Mti). M. J. & Ko\\\m\, G M_ 1 1 4 >87;0 IX^mitiitn ol the 
Keswick Clay. Adelaide/Golden Orove Lmbaymeni. 
Para and lulen llloiks, Souih Anstnilia. Sit/tib Australia. 
Civttl- Sun- Quart- liml X,>(r\ 103. 4-9. 

cV (!9H7b) Kedellnition i>t the upper 

boundary of Ihc llindinarsh Clay Adelaide Plains Sub- 
Basin and Adelaide/*, iolden (irove F.mbaynietn Ihiri, *>- 

& I I99(i) Soils, siraiipiaphy and 

engineering ^eolt>u> ol near surface maienals ol the 
Adelaitle Plains- Afirtrs untl fMtr^x Smith Attxlmliti 
Report Hook. 94/9 

Shouo. R C. (1952) Tlu geology ol the Soudi Kas1 
Province. Souih Australia, wiih special relerence M' 
Quaternary coastline migrations arid modern beach 
development (U'til. Snr\- Sauth >\n*t Hull- 29, I P0. 

SiiKi.iNf., C. H., Y.sw. 1- M.. MiCm.KO. M, I. & 
I.Avmitk. K. |l4B$) Itigh pavisioii I series Bating ot 
corals from Western Ausirolia and implications ru thc 
limim' and duration ol the Last Interylacial. fzurlh 
Wank ScL Utt. 135. 115-130. 

T\I! . R. I I879l The Anniversary Address ol the President 
irans. A'. SYv S Aum. 2. \v\i\ -Ixxv. 

TwiOAit. C R \ I96K) tfci.iiioipihilojfv ( Ihomav NclstHi 
tAustt Ltd. Melbourne K 

. Dmi.s, B. & Fihman. .1. B. II9W1 Kustatic and 

clinuttic Itistory of the Adelaide ateu, South Anstiallu. A 
discussion. Junt. ileal- 75. 217-242. 

Waki-. W. I. I l9poi CleolovV: ec«*morplioloy> and soils ol 
I he southwestern part ol Couuiv- Adel.ude. South 
Auwwltfi, C'SIKO Soil Publication No 23. 

WtHB. R P fIW5) ODP Site K20 mid Hie 
eoloni.satkitt ot Sahul. Qtnv. Austtakts'm i&)^ IS 



Wnun, J. P. (1994) Site 820 and the evidence for early 
occupation in Australia. Ibid. 12, 21-23. 

Winti.h, A. G. (1994) Infra-red stimulated luminescence 
dating of sediments. Rod, Mens. 23, 607-612. 

Williams, G. E. (1969) GlaciaJ age of the piedmont 
alluvial deposits in the Adelaide area. South Australia. 
AM J. Sci. 32. 257. 

(1973) Late Quaternary piedmont sedimentation, 

soil formation and palaeoclimates in arid South Australia. 
Z Geomorph. 17, 102-125. 

Zhu, Z. R., Wyroll, K.-H.. Collins, L. B. t Chen, J. H., 
Wasserbuug, G.J, & Eislnhauer, A. (1993) High- 
precision U-series dating of Last lnterglacial events by 
mass spectrometry: Houtman Abrolhos Islands, Western 
Australia. Earth Planet. Sci, Lett, 118, 281-293. 

Appendix: Methodology for luminescence dating 

Quartz grains (90-120 urn) were extracted as described in 
Huntley et al. (1993). Briefly, pure quartz of the right size 
was obtained by pretreating with HC1, followed by NaOH 
to break up clay aggregates, sieving, etching with 4098 HF 
for 40 minutes at 20° C, magnetic separation and floating 
on heavy liquid at specific gravity 2.67. For measurement, 
5.0 ±0.1 mg was deposited on stainless steel discs. 
Individual discs were post glow dose normalised with 6 Gy. 

Fine grains (undifferentiated as to mineralogy) were 
separated after the HC1 and NaOH extractions by settling 
from aqueous 0.0IN NaOH, the 4-11 um fraction being 
retained. This was then deposited on aluminium discs from 
acetone suspension, about I mg per disc. Individual discs 
were 0.5 s short-shine normalised. 

As stated in the main text, both dating protocols make use 
of the easily bleached component of luminescence. In the 
case of TL this component is selected for by both 
temperature and wavelength of emission (Prescott Sc 
Mojarrabi 1993). For GLSL it is assumed that the 
stimulated emission comes from the easily-bleached 
component; optical filters also select for this component 
(Huntley et al. 1991). The output is expressed as intensity 
as a function of temperature for TL and as a function of 
shine-down time for GLSL. 

For both protocols the emitted intensity is measured for 
"natural" samples and for samples which have received 
additional doses from a calibrated laboratory beta-source 
(N+B). About half of these samples are exposed to 
laboratory bleaching by sunlight filtered by a 475 nm long- 
wavelength-pass filter (Chris James 101); this bleach 
removes the rapidly bleaching component completely. 
Some bleached discs are also irradiated (YB+B) to provide 
the shape of the "missing" part of the dose curve at doses 
less than the natural dose, The data analysis follows the so- 
called "Australian slide" procedure (Readhead 1988; 
Prescott et al. 1993) and the data output is the equivalent 
dose D L - expressed in grays. Such curves are known as "dose 

Two methods of dose rate determination were used; In 
situ scintillometry (see e.g.. Hutton & Prescott 1992) uses a 
sodium iodide scintillation crystal, 75 mm x 75 mrn 
diameter in the auger hole from which the sample for dating 

is taken. The instrument is calibrated for K, U and Th and, 
independently, for total gamma ray dose. Scintillometry 
gives a completely self-contained measure of dose rate. 

Thick source alpha particle counting (TSAC) (Jensen & 
Prescott 1983; Huntley et al. 1986) gives a value for the 
contribution to the dose rate from U and Th together, and an 
estimate of the U and Th concentrations separately. In fact, 
the dose rate to the sample is effectively determined by the 
total alpha count and is almost independent of the relative 
amounts of U and Th. However, the measured ratio allows 
a (small) adjustment to be made to the dose rate. Combined 
with measurement of K, TSAC gives an independent 
measure of dose rate, X-ray fluorescence spectrometry is 
used to find K. 

In addition, U was found using delayed neutron 
activation (DNA). If this differs significantly from the other 
methods of assay for uranium, it gives an indication of 
radioactive disequilibrium, which was not the case here. It 
is most conveniently combined with the data from alpha 
counting to give the U concentration and hence a more 
accurate value forTh. These are the values shown in Table 

Table i includes the elemental analyses for all samples. 
The dose rates calculated for PK1S/3.5 using the 
conversion factors of Nambi & Aitken (1986) are shown in 
Table 2. The water content measured at the time of 
sampling was used in the dose rate calculations. Cosmic ray 
dose rates have been added in (Prescott & Hutton 1994). All 
data are included in Table 1. even though a numerical age 
was found for only one sample. 

The age calculation is conveniently expressed in terms of 
the so-called "Age Equation"; 

Age (ka) = TL of sample 

TL per unit dose (TL/Gy) x dose rate (Gy ka- 1 ) 

In this equation, "TL of sample" (which measures the 
accumulated energy) and "TL per unit dose" (which defines 
the sensitivity of the material to radiation) are measured in 
the laboratory on quartz or tine grains extracted from the 
sample; and "dose rate" is determined from measurements 
in the field and/or laboratory. 

Transactions of the Royal Society ofS. Aust, (1997), 121(3), 95-102. 


by C, Vaucher" : & I. Bevhridgh 1 


Vaucher. C. & Bhvfridgf, L (1997) New species of Potorolepis Spasskii (Cesioda: Hyrnenolepididae) parasitic 
in dasyurid marsupials from New Guinea. Trans. R. Sac. S. Aust. 121(3). 95-102, 28 November, 1997. 

Potorolepis aruensi.s sp. nov. is described from the small intestine of Myoictis melas from the Aru Islands of 
Irian Jaya. It is most closely related to P. bradleyi from which it differs in mean hook number (17 in P. aruensis, 
13 in P, bradleyi), size of cirrus sac (0,27-0.42 mm in P. aruensis, 0.14-0.25 mm in P. bradleyi) and arrangement 
of testes. Cestodes tentatively allocated to P. aruensis were also found in Antcchinus naso. Potorolepis woolleyae 
sp. nov., from the small intestine of Murexia longicaudatu from Morubc Province, Papua-New Guinea, differs 
from all congeners in having longer rostellar hooks (163-182 um), The generic diagnosis is re-assessed as well 
us the relationships between morphological sub-groups within the genus and the marsupial families they 
parasitise. A key to the species of hymenolepidid cestodes occurring in Australasian marsupials is given. 

Kr.Y Words: Potorolepis, cestodes, Hyrnenolepididae, marsupials, Dasyuridae, New Guinea. 


Cestodes of the family Hyrnenolepididae Ariola, 
1899 are common parasites of birds, rodents and 
inscctivores in most regions of the world (Czaplinski 
& Vaucher 1 994). Vaucher et al. ( 1 984) reviewed the 
species known from Australian marsupials, 
redescribing the three known species and erecting 
five new ones. All were allocated to the genus 
Hymenolepis Weinland, 1858 though it was noted 
that they formed a morphologically distinctive 
subgroup within this large genus. Subsequently, 
Jones & Anderson (1990) described a new species 
from a pcramelid marsupial in New Guinea and 
transferred the other species occurring in marsupials 
to the closely-related genus Vampirolepis Spasskii, 
1954. Spasskii (1994) erected a new genus, 
Potorolepis, to contain most of the species found in 
marsupials, although one, H. cercarteti, was 
transferred to the genus Radentolepis Spasskii, J 954, 
with the implication that it was originally a parasite 
of rodents. Spasskii (1994) was apparently unaware 
of the species erected by Jones & Anderson (1990) 
and did not include it in his new genus. The 
hymenolepidid fauna of Australasian marsupials is 
relatively poorly known (Spratt et al. 1991) and its 
taxonomic and phylogenetic affinities are uncertain. 

In this paper, we describe new species of 
Potorolepis parasitic in dasyurid marsupials in New 
Guinea and re-evaluate the definition of the genus 
proposed by Spasskii (1994) as well as the host- 

Museum d'Histoirc Nature! le, Geneva Switzerland. 
Department of Veterinary Sciences, University of Melbourne 
PuikvilleVic. 3052 

parasite relationships between sub-groups within 
Potorolepis and families of marsupial hosts, a 
relationship first suggested by Vaucher et al. (1984). 

Materials and Methods 

Cestodes from Myoictis melas were collected 
when the host animals were autopsied after a short 
period in captivity at La Trobe University, 
Melbourne. The cestodes were relaxed in water and 
fixed in AFA (Prjtchard & Kruse 1982). Cestodes 
from other hosts were collected in New Guinea. 
Following the death of the host, the entire 
gastrointestinal tract was fixed in 10% formalin and 
the cestodes were subsequently removed and stored 
in 70% ethanol, 

Cestodes were stained in Celestine blue, 
dehydrated in ethanol, cleared in methyl salicylate 
and mounted in Canada balsam. In contracted 
specimens, the tegument and dorsal and ventral 
musculature were removed with a fine scalpel after 
clearing (Jones 1990) to improve the visibility of the 
internal organs. Some scoleces of each species were 
mounted in Berlese's fluid. Serial sections were cut 
at a thickness of 9 um in both longitudinal and 
transverse planes and stained with haematoxylin and 

Measurements are given in millimetres as the 
range, followed by the mean and the number of 
measurements made in parentheses. Drawings were 
made using a drawing tube. 

All specimens studied have been deposited in the 
South Australian Museum, Adelaide (SAMA) or the 
Museum d'Histoire Naturelle, Geneva (MHNG). 

Host nomenclature follows Flannery (1995) and 





Sprall til ul. t{99fk Tabulated morphological data 
wetc derived farm Bevendge & Barker (41*754- 
Vaueher <>i nl (1084) and Jones & Anderson (1990). 
tlit-a distribution dala were derived from Spratt el at. 

Potttmlvpis aruensis sp. nt>\. 

(NGS I 7 1 

/\/>i'\; llolotype from small intestine of M\<ticiis 
mchis wallih'fi Gray, 1858, Koboor Is., Aru goupifv 
IJ'S 134 32'L). Irian Jaya. coll. P A, 
Woollcy, SAM A AHC ?7877: paratypes. 23 who].: 
mounts. H seolcces mounted in Berlcse\ fluid, serial 
sections. SAMA AHC 2787K-27 l )05; 2 wimL- 
moimis MIING Z3407 INVH; additional specimens: 
numerous specimens 2. xii. 1992- SAMA AHC 
305K6-305R7', 3 specimens, 3- \ii- IW. SAMA 
AHC $Q$Kfc numerous specimens l6vi.l L »3, 
SAMA AHC 30589, 

Mttfcritil examined: from Mxou tis mclus 
{ Mii) let. 1 ft40) types. Prom Anrciltinus tfOXtl 
lleritmk. 191 h: 5 specimens, Mt Kaituli P 2CS. 
146 41 'I-). Papua-New Guinea, SAMA AHC 
27864-27876. 30584- W585. 

Oc\< nplictt 

Based on types. Small ccstodes lip to 60 in length 
Scole\ gltiboste, 0.52 0.65 (0.58. n * ID) in diameter 
Suckers sub circular in superficial views, unarmed: 
CUJi shaped in section, wilh openings directed 
anteriorly. 0. 1 34). |6 10 \% n = 1(1) K tt 15(1. 1 K (0. 1 7. 
n = 10). Rostellum muscular, 048 0.26 (0.21) \ 
0.17-0.23 (0.21. n = 10): rostcllar sac 0.26 OSS 
(0.30, ii = 10) k 0.26-0.31 (0.28. n == l()>. Hooks 16- 
IX ( 17. n = Id), aiiauged in single ring wilh broad, 
curved blades; Mender handles prominent: blunt 
guards often with irregular surface: core of hook 
blade hollow. Hooks 0.128-0.147 (0438. n - 10) 
long. Neck variable, 0.74-2.05 ( I 26. n = 10) lorn;. 
Segments craspedole; mature scgilieuts mueli wtdei 
than long, 0.06-0.17 (0.10, n - JO) long x 0.93 1.63 
( I . * I n - 10) wide: near gravid segments longer, but 
slijjhilv narrower, 0.13-6.29 (0.22, n = 10) long. 
0.81-1.22 (1.04. n = 10) wide. Genital pores 
unilateral. Three testes arranged linearly, one pond. 
iwoaporal; Very little \arialion in testis distribution: 
single segment with 4 testes; single segment wilh 2 
testes, single segment with 2 poral. I aporal testes, 
Teste* oval, ol similar size. 0.10 17 (0.15. n = 10) 

long x 0.08 0.1 1 tO.09. n = 10) wide. Vasa efferenMa 
of anliporal testes run along dorsal margin of 
medulla lo elongate, pvrilorm external Wfrtriiiu) 
vesicle 0.06 0.16 (0.10, n = 10> Ion- * 0.04-0.0') 
(0.07. n = 10) wide, anterior and doisal to poral 
testes; distal region of external seminal vesicle 
slender, sinuous, enters elongate cirrus sac 0.27 -D 4 ' 
(0.35. n - 10) long x 0.04 0.05 (0.05. n = MO wide 
Cirrus sac contains elongate internal seminal scsielc 
occupying two-tJiuds of volume of cirrus sac: no 
armature seen on cirrus. Genital duels cmss 
osmoregulatory canals doi's.dly. 

Ovary median, with 3-4 indistinct lobules. 0.06- 
10 (0.08. n == 10) long. 0.10-0.19 (0 14, n = KM 
wide: vitellarium reniform, posterior to ovary 0.03 
0.06 (0.04. u = 10) long x 0,05-0,07 (03)6. n i I0» 
witle. Vagina posterior and venlral lo cirrus sac. 
dilating lo hum sacciform seminal receptacle dorsal 
to ovary; seminal receptacle 0. 1 4 0.24 i0.18.n- H)> 
long x 0.08-04 6 (041. p = 10) wide. I'lcrus 
originates as transverse sue 6fl ventral aspect oi 
medulla- extends lo osmoregulatory canals. 
developing small number of diverticula, never 
becoming reticulate. No segments found wilh fully 
developed eggs in uteri Ventral osmoregulatory 
canal 0.03-03)5 (0.04, a = 10) m diameter, dorsal 
canal narrower. 0.010.03 (0 02. n - 10) in diameter, 
longitudinal sirobilur musculature arranged in two 
concentric rings: outer ting composed of uumeious 
small bundles with only I 3 fibres pet bundle, 
hundles of inner ring larger wilh 5 40 fibres per 

Potorolepis waolleyae sp uov 
{ HICKS 8- 14) 

Tyjws. Iloloiype from small intesiine of Mtmxin 
hftiXittiudtita (Shlegcl. 1866). Mount Missnn (7 
13'S. 146 49 'E). Morobe Province. I'apua New 
Guinea, coll. G Gossek. 24.x. I W4, SAMA AHC 
27sH)6: paratypes. 9 fragmented specimens. I seolex 
mounted in Bcrlcsc's fluid, serial sections. SAM \ 
Al'C 27s>07-27s)l'J. 30590: 2 specimens MMNG 
23408 INVI;. 


Small ccslttdes, largest fragment ^ lony. Sc<»Ic\ 
globose. 0.44 -0.55 (0 50. n - *J) in diameiet. SucKers 
sub circular, unarmed. 0.13 0.21 (0.17. n = 10) long 
\ 040-0 19 (045 n= 10* wide. Rostellum museulai 
o 4 341.20(04 7- n = V) long \ 0.i^ft.38(flilLn=»^| 

li'js 17. Ptilntttfffu* nturii\i> sp. uu\. types. I Sc.»le\ wilh roslellain evcrictl. 2 Stokw wilh n>s(elliiin wutiiinnMi. V 
RuMclkir hiuik in prohle. 4. RoNk' lutuk, view Innn rH>sicnoi nftfott! showing t-ntiirirc-d liuok g^MM, 5. Mjiuit 
vj,'nicnis. b Nearer; iv id . seymenl. 7. Traiisvcisc hishtlu^tuit steittm Of niiiiurt- M^nleoC dorsul DUp?C\ towards mp vl 
p.ifv. State- burv = i).lmm I 2. 5 7; O.UInnn 4 4. I cgt-nd D, Jursal osmttivsukiiory canal: (.S. ^vtfrnal svidiikiI VOick; 
SK scrniiial retepiaete l.ieMis;! uieius V ventral osmoregulatory ewftf- 


' 10 



wide; rostellar sac 0.26-0.29 (0.28, n = 9) long x 
0.20-0.29 (0.26, n = 9) wide. Hooks number 15-19 
(17, n = 6), arranged in single ring; hooks with 
elongate, falcate blades, slender handles and blunt 
guards; core of hook blade hollow. Hooks 0.163- 
0.182 (0.172, n = 10) long. Neck variable, 0.90^1.41 
(L20, n = 8) long. Segments craspedote; mature 
segments much wider than long, 0.0.5-0,06 (0.06, n = 
5) long x 0.76-0.90 (0.85, n = 5) wide; gravid 
segments longer, of approximately the same width, 
0.13-0.35 (0.20, n = 5) long, 0.57-1.22 (0.87, n = 5) 
wide. Genital pores unilateral. Three oval testes 
arranged in triangular array, with 1 poral and 2 
aporal. of similar size. 0.08-0.13 (0.10, n = 5) long x 
0.04-0.05 (0.04, n = 5) wide. Vasa efferentia from 
aporal testes run along dorsal margin of medulla to 
elongate, pyriform. external seminal vesicle 0.13- 
0.18 (0.16, n= 5) long x 0.023-0.049 (0.043. n=5) 
wide, which extends along seminal receptacle. Cirrus 
sac pyriform, 0.14-0.17 (0.16, n=5) long x 0.036-0.042 
(0,040, n=5) wide; cirrus sac contains prominent 
internal seminal vesicle; cirrus unarmed. Genital 
duels cross osmoregulatory canals dorsally. Ovary 
median, with indistinct lobules, c. 0.06 long x 0.14 
wide, on ventral surface of medulla; vitellarium 
reniform. posterior to ovary, 0.04-0.05 (0.05. n a 5) 
long x 0.02-0.03 (0.03, n = 5) wide. Vagina posterior 
and ventral to cirrus sac, dilating to form sacciform 
seminal receptacle dorsal to ovary; seminal 
receptacle 0.10-0.13 (0.12, n = 5) long x 0.05-0.07 
(0.06, n = 5) wide. Uterus originates as transverse sac 
on ventral aspect of medulla, extends to 
osmoregulatory canals, developing few diverticula; 
never becoming reticulate. Eggs spherical. 0.032- 
0.045 (0.035, 1 " n = 5) in ^diameter. Ventral 
osmoregulatory canals 0.03-0.10 (0.05, n = 5) in 
diameter; dorsal osmoregulatory canals 0.01 in 
diameter. Longitudinal muscle arranged in two 
concentric rings; outer ring composed of numerous 
small bundles with few fibres; inner ring composed 
of larger bundles with 10-20 fibres per bundle. 


Spasskii (1994) created the genus Potowlepis for 
several species of cestode from Australian 
marsupials which had previously been allocated to 
Hymenolepis (see Vaucher et al. 1984) or to 
Vampirolepis by Spasskii (1954) and Jones & 
Anderson (1990). Species included in the new genus 
by Spasskii (1994) were P. antechini (Vaucher, 
Beveridge & Spratt, 1984), R aklei (Beveridge & 

Barker, 1975). R bettongiae (Vaucher, Beveridge & 
Spratt, 1984), P. bradieyi (Beveridge & Barker, 
1975), R isoodonlis (Vaucher, Beveridge & Spratt, 
1984) and R peramelidarum (Nybelin, 1917). To 
these should be added P. peroryctis (Jones & 
Anderson 1990) comb. nov. (syn. Vampirolepis 
peroryctis (Jones & Anderson, 1990), the first 
species of the genus known from a New Guinean 
peramelid marsupial, Peroryctes raffrayanus, 
Potowlepis peroryctis is morphologically similar to 
R peramelidarum and P. isoodonlis, also from 
bandicoots, and clearly belongs within the genus 
Potowlepis. The species was presumably overlooked 
by Spasskii (1994) in erecting the new genus. 

Spasskii (1994) characterised his new genus as 
having a rostellum armed with more than 10 hooks 
each longer than 50 um, all with a well-developed, 
elongate blade, longer than the guard and with a 
tubular uterus which may develop diverticula. 

The new species described above possess these key 
morphological characters and are therefore allocated 
to the genus Potowlepis. They are readily 
distinguishable from the known species based on 
hook number and size as well as from the disposition 
of the testes (Table 1). Based on hook number, the 
species of Potowlepis fall into two distinct groups, 
those with 10-23 hooks, all of which are parasitic in 
dasyurid marsupials, and those with 24-40 hooks 
which occur in peramelid and potoroid marsupials. 
The two new species, both from dasyurid marsupials, 
have hook numbers in the range 15-19 and therefore 
most closely resemble P. aklei, P. antechini and P. 
bradieyi. Both P. aruensis and P. woolleyae are 
distinguished from these species in having longer 
hooks. The lengths of hooks of P. woolleyae lie well 
outside the ranges of other species within this sub- 
group, though hook lengths of P. aruensis may 
overlap with those of P. bradieyi. Potowlepis 
aruensis can be distinguished by mean hook number 
(Table I) but also by the cirrus sac which is shorter 
in R bradieyi (0.190 x 0.026 mm) than in P. aruensis 
(0.350 x 0.050 mm). In P. bradieyi, the central testis 
lies dorsal to the ovary (Beveridge & Barker 1975) 
rather than aporal to it as in P. aruensis. For t^esc 
reasons, the specimens described above from 
Myoictis melas are considered close to but distinct 
from P. bradieyi and warrant the erection of a new 
species. None of the specimens was fully gravid, the 
terminal segments instead having developing uteri 
which were only partly filled with jaggs. This is 
surprising since the animals were trai^ported to 
Melbourne following capture and mainlined in the 

Figs 8-14. Potowlepis woolleyae sp. nov. Types. 8. Scolex with rostellum everted. 9. Scolex with rostellum withdrawn. 10. 
Mature segments. 1 1. Gravid segment. 12-13. Rostellar hooks in profile. 14. Transverse histological section of mature 
segment, dorsal aspect towards top of page, Scale bars = 0,1 mm 8-9, 11; 0.2mm 10; 0.01mm 12-13; 0.1 5mm 14. Legend; 
as for Figs 1 -7, 



laboratory until iuili>psy, providing adequate lime for 
eestodes to mature. 

The spec i mens From Antcthinus tut.u> are 
tentatively allocated to this species. They are poorly 
preserved and internal features are difficult lo 
discern, Hook lengths are Identical lo specimens 
from Myoictis. The number ol hooks. IN-22 (20) (n 
= 5), is larger than in specimens from M. ruehis hut 
the range overlaps. They may represent a distinct hut 
very similar species, although the current evidence is 

The eestodes from Muivxiu lon^icaiulata were 
quite severely contracted, limiting (he morphological 
details which were visible in whole mounts. 
Nevertheless. I hey represent a new species based on 
the features ol (he roslellar hooks atone. While they 
resemble P. arurnsis in terms of mean hook number, 
the si/c of hooks immediately distinguishes the 
material from all congeners. In having the three 
testes arranged in a triangular fashion. P. uaoKcyae 
most closely resembles P. aklei and P. ttnfechini. 

Apart from adding to the hymenolepidid fauna 
known from marsupials in New Guinea, the new 

species described support the erection of the genus 
Potorolt'pis by Spasskii ( 1994) in providing 
additional species which conform with the proposed 
diagnosis. Spasskii (1994) provisionally included in 
his diagnosis the character genital ducts crossing 
osmoregulatory canals dorsal ly". This is 
unequivocally the case in /* akiei, /! bradtvyi, P. 
anu'ihsis, P. woolleyae and P p?ror\ilis and probably 
is similar in the remaining species of the genus. His 
generic definition (Spasskii 1994) also needs lo be 
amended to allow for testes in either a linear or 
triangular array and for cirri which arc either armed 
or unarmed. Apart from these minor modi Ileal ions, 
(he generic definition provided by Spasskii (1994) 
appears to be reliable- 

The description of the new species also provides 
evidence in support of the suggestion first made by 
Vaucher el td. (1984) that each family of marsupials 
was parasitised by a distinctive morphological group 
of hymenolepidid eestodes. Sprall el til. (1991) 
reported P. peramclidarum from Antechinus 
wtiimonii which would represent a potential 
exception. However, the identification was tentative 

T\m i I . Mctt,\utrnwtiis and Am WMpkotogh a! features af XptH its of Potorolepis frfim marsupials. 





No, of 


Species trom dasvurid hosts 

AiUwhittus sp- l undeserved)* 
A. flttvipt's. Psfiuliwurhmus 
hilanti^ Sitiiuttut(>si\ Itno/nts 




/' l>nt<Ilr\t 
I'. t murium 
P aruensis 
P wanlU'vtw 

Anift (units sp. (undeiscribcd) 




Anwclunus swuhisonit 

56 S9(5X) 



My wet is twias 

I2X-I47 (I3X) 


tl iieai 

Muirxut htmu'i uiidala 

163-1X2 d7:> 



Species from pcramelid hosts 

P pcnmwlutanwi Peranwfr\ mmilt* 

P. xuntui 
IsawJon aht\ulns 

yuioi <9K> 



/• tit}&damf.\ 

P {U'tiH ViV'S 

/sootion t*b<rsukts 
Ptanyt tt\ raffrtntiiM: 





Species from poioroid hosts 

(', fumuin htlonuts ithlatixttis 

98-103 d02) 

>4J 53 

linear or 

P hiuou^iitc 

Ht'Utmi<iii (jaJtMUft/i 

79-91 (X6> 

'4 27 


forineilv identified as Anifchuiu.s stuarta (see Strahan 1995) 



and based an incomplete specimens. This dubious 
record has therefore been eliminated from 
consideration until more material is collected and the 
host record confirmed. The species found in 
dnsytnids. P uktei, P. utitfthini, P. amett\is. P 
hraJIcyi and P nooHeyae. belong to a group of 
species Willi •> small number ol roslellar hooks ( 10- 
23) compared with 33-40 hooks in P. t\so<nl<mli\, P. 
ju'iiimeiuhium and /' peronctis from peramehd 
hosts and 24-33 hooks in P, fiotemi and P. bvttoniiiue 
from potoroid hosts. The relative si/e of the ovary 
also separates the first tv\o groups, the ratio of width 
o( ovary to segment width being 9-15''/ in species 
from dasyurid hosts compared with 2b- W;> in those 
Irom pcramelids In (he first yroup the uterus 
contains relatively tew eggs, but it is clearly hilohed. 
Mine data are needed from hymenolepidids 
parasitising pcramelids ami potoroids to confirm the 
utility ol this character. The observations of Jones & 
Anderson ( l°90j on a species from a New Guinean 
peramelid marsupial and the current description ol 
new species from dasyurid marsupials from New 
Guinea provide additional support for the hypothesis 
advanced by Vaucher el of. tl u X4). The data also 
suggest that the hymenolepidids of dasyurids from 
Australia are similar to those of New Guinea, as are 
the comparabte cestodes of peramelid marsupials, 
finally, we agree with Spasskn iWH) in allocating 
tfywentolipbi cetrarwii Vaucher, Beveridge & 
Spratt. 19.84 t" lite genus Kotleiitoltpis. 

2. Hooks number 20- MX 1 J-tH uin for* 

Nodtnfnlrpix nunu 

Hunks 17 12. 1702 urn long.. RtHlrnfdU'fns cenwrtcti 

}. fewer Hi. hi 22 hooks m. il 22 luniks present, 

hooks >|f)U un» long ........ 4 

More ihan 22 hooks oi. if 22 hooks present, hooks 
<l(Mlpm lotifir - - - — - - 7 

4. Hooks shoner ihan or equal 10 1(H) um 

Hooks longer than 100 uin 

P</luttflf/n\ uktrt 

5. Hooks 163-182 um iqng pnu>r<>Uph wuvlkytU 

Hooks less than 130 um in length .{? 

6. Hooks 10.VI2S pin long. 10 15 in nuinhci 

FaTbrnh'p'tt bnuiUvt 

Hooks I2K-147 |nn long- 16-22 in number 

Fammlepis untemth 

7. fewer than ?S honks — .. - ,.*8 

More ihan 2H hooks 9 

8. Hooks 5ft -59 um long. 32-23 in number 
hnorultpts anti*chm 

Hooks 79-41 um long. 24-27 in number 

.- _ -- PiihmtlitpU hrtlimiiiih 

9. Hooks 124 192 um loni», 40 in number 

- „ , - - .f'tttttr/tlcfv\ perm snis 

Hooks less ihnn 120 um long, fewer than 40 in number 

10. Hooks shorter ihan K? uni Po\on>lvpis tuunJ t niti\ 

Hooks longer ihan K5 pm - _ I I 

I I HtMtks number 29-33 _ _ Pnlnnilcpis pofinft 

Hooks number 35 -3S P/mttolt'pts pcrameliihtrwn 

Key to the species of hymenolepidid cestodes 

occurring in Australasian marsupials based on 

rostt liar hooks 

Hooks sffiall. shotiei than 50 um Innu, f rale moid in 
shape 2 

Hooks linger, length greater ihan 50 pan long, not 
fratcmniil m shape- — • -— — * 


We are grateful it* Dt P. A. Woollcy who collected 
material used in this study and who also made 
specimens collected by others from New Guinea 
available to lis. 


Hi viwiua. f & HAKkhK. I- K- (1975f Acuailid. capillarud 
antl hvuienokpitlid parasites of the dasyuiid marsupial, 
Antrihintts Mtumii Mucleav. ISJl. lr<Pin southeastern 
Australia / Itrttirinthot 49, 2 M t21. 

CVam ISSKI R & Vai t nut, C. ( I9 l >4) family 
Hymcuolepididac Armla. 1H99 pp. 595 663 tit Khalil, L. 
I -.. Jones, A, & Bray. U. A. tLds) "Keys to the Os&Hlc 
Par. isiris of Wiielnaies" ifAIJ Iniei naooiuil. 

l-i \\\i in, 1 K I IW5) "Mammals of New Guinea" (Reed 
Hooks. Chaiswoodi 

Josus. .A ( 19901 leehniques fitr hantJ-sectiornng thick 
IhhIu-U Platvheiminllis. Sx\t PtiniMh'l. 15, 21 1 SfiL 

& ANOhksoN. T< I C- ilWH) Helminths rtl 

rodents uni} marsupials I'rOffl f'apua New Gmnca. wilh 
the deseripiion Ut iwt) new species. EihtfUKsitmui 
iihyrtiiptwaf- n. sp. (Digenew : Kehinosioniandaei 
and Vutnpittt frjfl \ pinttMfis n sp iCcshtda 
llymcnolepidklaei Sy\f. I\mi\h»t(- 15, 223-237- 

Priuiiako. M H &. Kktsi_. Ci. O W_ (I9K:i Thr 
Collection and Piescrvalion Of Ammat Paiasifex" 
t University ol Nebraska Vss. Lincoln i. 

Si'asskii. A. A (1954) ICIassilkalioii oi' \Uv 
hymenolepidids of mammals.) tnuh txib. tiet mint, 7, 
120 Io7(InRussuio. 



(1994) [On the systematic position of 

hymenolepidids from Australian marsupials.] 
Parasitologiya 28, 66-69 (In Russian). 

Spratt. D. ML, Beveridge, I. & Walter, E. L. (1991) A 
catalogue of Australasian monotremes and marsupials 
and their recorded helminth parasites. Rec. S. Aust. Mas., 
Monogr. Set: No. 1, 1-105. 

Strahan, R. (1995) "The Mammals of Australia" (Reed 
Books, Chatswood). 

Vaucher, C, Beveridge, I. & Spratt, D. M, (1984) 
Cestodes du genre Hymenolepis Weinland, 1858 (sensu 
lata) parasites de marsupiaux australiens et description 
de cinq especes nouvelles. Rev. Suisse Zool. 91, 443-458. 

Transactions of the Royal Society of S. Aust, (1997), 121(3), 103-117. 




by Murray J. Littlejohn* & John R. Wright" 


Lrm.nJOHN, M. J. &, J. R. (1997) Structure of the acoustic signals of Crinia glauerti (Anura; 
Myobatrachidae) from south-western Australia, and comparison with those of C. signifera from South Australia. 
Trans R. Sot: S. Am 121(3), 103-1 17, 28 November, 1997. 

Advertisement calls of 51 males of the Australian myobatrachid frog, Crinia glauerti, from five sites, and 45 
males of the presumed cognate species C. signifera from four sites, were analysed and compared, Patterns of 
geographical variation in the advertisement calls of C, glauerti were explored and a dine was found in pulse 
rate. The structure and geographical variation in frequency of occurrence of another signal, the squelching call, 
of unknown function, in the acoustic repertoire of C. glauerti, were investigated. The findings are consistent with 
the earlier reports of more frequent occurrence of the squelching call in the south. Because of possible similarity 
in function, the squelching call of C. gtauerti was compared with the encounter call of C. signifera. 

Key Words: Crinia glauerti, Crinia signifera, advertisement cal 
structure, spectral composition, geographical variation. 

encounter call, acoustic analyses, pulse 


The commonly produced call (= advertisement 
call sensii Littlejohn 1977; Wells 1977) of the 
endemic south-western Australian species Crinia 
glauerti (Loveridge) was subjectively described by 
Main (1957) as: "A prolonged rattling call." Main 
(1957) also noted: ^Adjacent to Perth the call of this 
species is constant and is predominantly a rattle, but 
occasionally a short squelching sound is made. On 
the south coast ... [away from related species], the 
squelching call is far more prominent ... " This 
observation of inter-populational variation in the 
calls of C. gktuerti was considered by Brown & 
Wilson (1956) as a possible example of character 
displacement, with the squelching calls being more 
frequent in the populations that were allopatric to C. 
insignifera (Moore), a species which also has a 
squelch-like call (see Littlejohn 1959). 

Although some values were cited by Brown & 
Wilson (1956) from A.R. Main (in litt., based on 
analyses by M.J, Littlejohn), the first published 
objective description, which included an oscillogram 
(= waveform) and an audiospectrogram, of the 
rattling call of C. glauerti was provided by Littlejohn 
(1959). This account was derived from the physical 
analysis of magnetic tape recordings of the calls of 
12 males obtained at only one locality, South Perth 
(1I5°52' E, 31°59' S), near the northern limit of 

' Department of Zootomy, University of Melbourne Parkville Vic. 

Hawe. S. M. (1970) Calling behaviour and Territoriality in Males 

of Two species of Crinia (Anura: Leptodactylidac), BSe (Hons) 

ihesis. Department of Zoology. University of Melbourne 


geographic distribution (Littlejohn unpub.), so that 
there was no consideration of geographic variation in 
call structure within this species. Littlejohn (1959) 
also noted "occasional call variation in the form of a 
compressed series of pulses" which may be equated 
to the squelching call of Main (1957) and an 
oscillogram of one of these pulse trains was 

The disjunct allopatric south-eastern Australian 
taxon C. signifera (Girard) is presumed to be the 
cognate (sister) species of C. glauerti, with which it 
constitutes the signifera superspecies (Main 1957; 
Main et al. 1958). The advertisement call of this 
taxon is a short, rapidly repeated pulse train and the 
first published objective description, and an 
oscillogram, were provided by Littlejohn (1958). 
Littlejohn (1959, 1961) supported the proposed 
affinity of C. glauerti and C. signifera, on the basis 
of the much lower pulse rates of the advertisement 
calls, when compared with those in calls of the then 
recognized members of the related insignifera 
superspecies (Main 1957; Main et al. 1958). 
Quantitative descriptions of the calls of C. signifera 
were also provided by Littlejohn (1964. 1970), 
Littlejohn & Martin (1965), Hawe 1 , Littlejohn et al. 
(1985) and Odendaal et al. (1986). Interpopulational 
variation in advertisement calls of C, signifera was 
considered by Littlejohn (1959, 1964), and by 
Odendaal et al. (1986), Straughan and Main (1966), 
through choice playback experiments in which tape- 
recorded advertisement calls of C. signifera and C. 
parinsignifera (Main) were offered as alternative 
stimuli, demonstrated that breeding females of C. 
signifera exhibited positive phonotaxis only to the 
conspecific calls. Encounter calls (sensit Wells 1977) 


m. j. urn. i john & j. k. WKinm 

|= calk litllejohn ft ai 1985) o\' C. 
\i\>nijera were identified through Held playback 
experiments by Httwc' <m\ Liltlejohn ri tii (19X5). 
The squelching calls o\' C. ghuu-iti may also have a 
territorial function but this has yet U> be determined. 
In the absence erf experimental documentation, the 
term "squelching call"' (Main 1957) will be retained 
lor the compressed series of pulses. 

Tape-recorded samples of advertisement calls of 
51 individuals were obtained at live localities across 
(lie geographic range of C. gttmerti (Fig. I , Tabic I ) 
ID provide a measure of geographical variation 
Mundaring Weir (Site 1) is the type locality for C 
glattenj (Cogger el al, 1983), Squelching calls were 

produced in the recorded sequences of 15 of foesc 
individuals. Sufficient squelching calls wcic 
obtained to provide a preliminary description of this 
type of signal for comparison with the eonspecific 
advertisement calk and with the encounter calls of C- 
sigftif&nt, and to allow un estimation of geographical 
variation in the frequency of production relative to 
the advertisement call. Tape recordings ol 
advertisement calls of 45 individuals from the closest 
populations Of C. siwufera were also obtained, 
namely from four localities in southern South 
Australia (Fig. 2. Table 1) at or near the western 
limits of the extensive geographic distribution of this 
species (see maps given by Brook 19X3, 19X4; Tyler 






1 1:- I. Geographic diminution t based on Tyler ti ttf l ou 4t pig 2. (toogvaphic distribunon (based on Brook I9S4) of 
of Crima xfauerti, and" locutions- uf recording sites (sec Cnnia sii*nifimt in South Australia and locutions n| 

Table 1 1. recording sites (see Table 1 1. 

Ta.ui I |. Dehiih nf n'nmtin\> Un ufiiitw. 

All sites are in Western Australia Uvrt'rinut flltMtirfi and m South Australia for C ^^uiffnt. 



Dales nt recording 






O xlnueni 



1 lelena River, at base 

rtf Mumlaring Weir 


1 16 10' 


06.vii.X6..1Lvii-X , > 

2\ km souih-suulh-wesi 

ul Busseltnn 


I 1 s to* 



Diamond Ttec mtlway sidmy 

M 22' 

1 If. IK." 


24. 25.vii.86 

3 km east Of Nornulup 

34' 5 ( >' 

!lo Ml* 


m. lU.vui.X" 

1 J km east-north east ol Albany 


MS 110* 

l' sivn'/tro 


18 viii.KC: l5.ViU.HM 

9 3 km soulhofWaitilla 

»4 36' 

135 40* 


03. 04.vm\X9 

Velttmmait Creek. 3 6 km 

Konlli-rasl gf Mclrnse 

V2 si' 



2H. 2y.vii.90 

9 km north-north-west of 

Vicioi Harbour 





X 3 km north norlh-casi 

nf Milliccni 

37 11' 




19X5: Ctfggpr 1092). The description of ihe 
encounter call (territorial call) of C siyiifera is 
based Ofl accounts Oi lluwe 1 and l.illlcjohn el al. 
(IV85, unpiib.). 

Materials mid Methods 

Rentnljni> o) r calls 

Tape ret ordings were obtained al ihe breeding sites 
(Pies I. 2. Tattle 1 1 wuh an open-reel recorder (Nagra 
IV-S) and a cardioid dynamic microphone (Beyer M 
88). Males of both species call from a variety Of 
sites: on (tie hanks adjacent to water, while silting in 
shallow water, or while floating and supported by 
emergent vegetation. The effective temperatures al 
Ihe calling sites (sUrlaec water, wet- bulb air. 
depending on Ihe calling position of Ihe frog) were 
i hen measured with an electronic ( thermistor) 
ihermomcler (Takara Digimulli Model D61I, Willi 
sensor type S/.L-M). The mean was used <^s ihe 
effective temperature when an individual was al the 
interface between air and water Where possible, the 
recorded males of C ^laaerti were collected, 
euihanased. preserved in Tylers fixative (Tyler 
|0n2). stored in 70% elhanol and lodged with the 
Western Australian Museum. It Ihe specimen was 
preserved, the measurement of snouFurostyle lenglh 
was later made with dial calipers (to 0.1 nun. 
rounded to 05 mini, Otherwise, it shon plastic ruler 
( 15 cm) was placed alone Ihe dorsal surface of Ihe 
living speeimen held in a thin, clear plastic bag. and 
the snout uroslyle lenglh determined to the nearest 
5 mm 

Ac/wstic athilysis oj r ret nnlintfs 

lor acoustic analysis, tape recordings w r erc 
replayed on an open reel tape recorder tRcvox B 77 
Mkll or Sony TC 510-2). Advertisement calls were 
analysed by using a digital audio spectrograph (Kay 
Elemetrics Model DSP 5500 Sona Graph), Where 
ihcrc were high levels of low-lreijuency noise, a 
passive filter (Allison 2B. high pass, cut-off set at 
$ 12 Hz) w;i.\ inserted between the output o\ the tape 
recorder and the input of the audio spectrograph. 
Stati-vtical procedures were carried out wnh 
SYSTAT. Version 5.03 (Systal Inc.. P.vanstont. 

Clutracfcrislics of 'equipment used in rcconlwg anil 

Tape tran>port speeds ( I*; cm s ' ) o\' the recorders 
involved in recording aiul playback were periodically 
checked against & locally piodtked slandaid 1000 
kll/ calibration tape and a frequency counter (Neath 
SchlumbergerSM I I8A or Good Will GFC -80I0G) 
(Revox, Sony), or against the 50 H/ AC mains 
fiequeiicy by a buill-in stroboscope (Nagra); overall 
variations in tape speed through recording and 

playback are estimated n be less than ±0 5%. The 
frequency responses of all electronic eonrponeiils 
used in recording and analysis are presumed to be 
close lo linear within the narrow range \)\ frequencies 
of interest (c. 2.0 5.0 kHz), based on manufacturers' 
specifications. The relatively high sampling rates 
used in the digital analyses <>44 kHz) preclude the 
production ol artifacts from aliasing. 

Siruenitv and aeausite attnluaes aj calls 

The calls of the iwo species are of simple slruclure 
and each consists of a group ot discrete damped 
oscillations (Tigs $-5). For convenience, and 
following previous usage, each of these oscillations 
is referred to as a "pulse." and the group (i.e. the 
pulse train) as a call (Figs 3. 4). The number of 
pulses in a call was deiermincd by direct inspection 
of ihe displayed waveform The depth ol amplitude 
modulation of the last two pulses in a call was 
sometimes less than UtYA ; in such cases, separate 
pulses were recognized if the depth of modulation 
exceeded aboul 7YA The duration (lo ncaresl ms) of 
a call was taken as ihe mteival from the peak of the 
first pulse to the peak ot the last pulse {= "peak-peak 
duration"). The puKes m the advertisement calls arc 
produced in a quasi-periodic fashion, and the pulse 
rale (as p s ') was calculated over a complete call as 
(n - I pulses) x 1000/pea^-peak duration in ms. 
Dominant frequencies were measured as the peaks in 
a power spectrum t^\ the whole call. To describe the 
temporal and spectral properties of pulses in 
advertisement calls, a tape reeoiding of one call ot 
each ol three individuals of each species (C. glanerti. 


400 600 




Fig. X Waveforms of advertise me ill calls Upper ( rutin 
K'litiHTfiz Rcteivikv R4J9 7, Sue I: effective lempcraiurv. 
wcl-bulb uir — I \.H t. Lower C. Mintifcm, KetciOiKv 
R'408-2. Site R dftviivt* temperatures. vvl( bulb ;ur - 
I MT,, water = J2 5 C 



Site 1 ; C. signifera, Site 6) was replayed into a digital 
sound card (Sound Blaster SB 16, Creative Labs 
Milpitas), installed in a desktop computer (IBM-PC 
compatible). The sampling rate was set at 44,100 Hz 
and the sample size at 16 bits. Files were prepared in 
the WAVE format with the Creative WaveStudio 
Version 2.0 software (Creative Technology 
Singapore), and further processed with this package 
and with Spectra Plus Professional, Version 3.0 
(Pioneer Hill Software Poulsbo). The figures of the 
waveforms and spectra (Figs 3-5) were prepared by 
the same procedure. One pulse (the middle pulse or 
next after the midpoint of a call if an even number) 
of each call was used for analysis. 

Conventional rise and decay times of pulses (from 
10 - 90%, and 90-10%, respectively, of maximum 
amplitude) were not calculated because it would 
have been necessary to extrapolate between peaks of 
the carrier frequency (see below). Accordingly, the 
number of either positive or negative half cycles to 
reach maximum amplitude was counted and the time 
interval from background noise level to the peak 

estimated. The pulse duration was measured as the 
interval from approximately 10% of the maximum 
amplitude (either positive or negative) at the start to 
the same level at the end of a pulse. Two dominant 
spectral peaks are present in most of the calls of both 
species (Fig. 4). There is a well-defined sinusoidal 
fundamental frequency within the wave train (Fig. 4) 
and this is referred to as the carrier frequency (OF), 
by analogy with amplitude modulation in 

Rates of production of advertisement call were 
determined by playback of original recordings on a 
Sony TC-510-2 recorder. For C. glauerti, the 
durations of five consecutive cycles of calls and 
intervals were measured; for C. signifera, 10 cycles 
were measured. The following protocols were 
employed to arrive at the sequences of calls selected 
for measurement: C. glauerti ~ the first six clear calls 
in the recorded sequence; C. signifera - the last 16 
calls in the sequence were digitised, then the last five 
discarded. The interval from the end of the first call 
to the end of the sixth (C. glauerti) or tenth (C. 

-i 1 1 r 

3 4 5 6 

Time (rms) 

3 4 5 

Frequency (kHz) 




4 5 

Time (ms) 

3 4 5 

Frequency (kHz) 

Fig. 4. Structure of pulses in advertisement calls. Upper panels, (a). Expanded waveform, (b). Frequency spectrum for 
Crinia glauerti (Reference: R439-9, Site 1 ; effective temperature = 12.2° C). Lower panels, (c). Expanded waveform, (d). 
Frequency spectrum for C. signifera (Reference: R408-5, Site 6; effective temperatures, wet bulb air = 10.9° C, water = 

12.7° C). 






400 600 



Fig. 5. Waveforms of squelching calls of Crinia glauerti. 
(a). R441-5, Site 5, effective temperature =11.8° C. (b). 
R44M, Site 5, effective temperature c 13.TC. (c). 
R4G4-2, Site 2. effective temperature = 9.6° C. (d). R44 1 - 
4, Site 5 V effective temperature = 13.1° C. (e). R407-6, 
Site 3, effective temperature = 1 1.1° C. (f). R404-3, Site 
3. effective temperature = 9.8° C. (g). R407-7. Site 3, 
effective temperature = 10.7° C. 

signifera) call was then measured to the nearest 

Numbers of advertisement calls of individuals 

The number of advertisement calls of each 
individual to be analysed was determined as follows. 
Both species produce advertisement calls in long and 
regular sequences. In C. glauerti, production of the 
longer advertisement calls is slow - about 20% of the 
rate of C. signifera (see below). For C. glauerti, as 
the first step, four clear calls of each individual (n = 
10) from Site 1 were chosen at random from the 
recorded sequence. If four clear calls could not be 
obtained, that individual was discarded from the 
analysis. For C. signifera, all individuals from Site 8 
(n = 10) were used and data were obtained from three 
successive calls: the 12th, 13th and 14th (or the 
nearest clear call if there was an overlap) from the 
start of a natural sequence, or after recording had 
commenced. For each attribute of the call and for 
each species, an analysis of variance was carried out 
with comparison between individuals. The variance 
was partitioned and the proportion of the variation 
due to within-individual effects and that due to 
between-individual effects was calculated. For both 
species and all variables, variation between 
individuals was far greater than that within 
individuals. For C. glauerti, the within-individual 
variation accounted for 13-20% of the variation in 
the data. For C, signifera, the within-individual 
variation accounted for 24-25% of the variation. 
Based on these results of the analyses of variance, it 
was decided that for C. glauerti, the average of two 
calls per individual would provide a representative 
sample for that individual. As the calls of C. signifera 
were slightly more variable, it was decided to use the 
average of three calls for each individual. Mean 
values for individuals are used in the subsequent 
treatment of these data. 

Effects of temperature 

Linear regression analyses of the full data set for 
advertisement calls (Table 2) indicated that for C. 
glauerti there was a significant (p<0,05) relationship 
between effective temperature and duration and 
between effective temperature and pulse rate. The 
linear regression analyses also indicated that there was 
a significant relationship between dominant frequency 
and effective temperature for advertisement calls of 
each species. Accordingly, values of the dependent 
variables were corrected to 1 1 .0° C, the nearest integer 
to the pooled mean for effective temperature (C. 
glauerti, mean = 11.1° C; C. signifera, mean = 10.8° 
C); these values were used in subsequent calculations. 
Where the slope for the combined samples for each 
species was non-significant, the raw data were used in 



Tahi l 2 InfftH'H r >>f effect /tv temperature unjoin uttnhute\ nf the iidvertisement mils qf Cnma "lauerli fn - 5/ J .uulC. 

signitem in = 4$) 

Results of analyses Uy linear regression. Sec Tabic 4 tor paogca ot temperatures. 

Attribute »-1 call 



Significance ut 
slope {p i 

CoclTicicni ol 
iMenitirution tr- '» 


C. vUinerri 




(\ M}tnifiru 




N umbci *»t pulses 

( i-httHTll 




C \f\>ittfrni 




Pulse rale 

C- \{ftitUTtt 




c . \i$mf»ru 




Upper tlorninaiii frequenc v 

C. uluittrti 

KK) 107 



i sii\mftTti 


-.4) (MM 


the subsequent analyses. 

Occurrence uj ' si/m'h hing calls ofC glauetti 

The presence uj squelching calls was determined 
subjectively, by replaying the tape-recorded 
Sequenced pf each individual. In this analysis, three 
types ol call were recognised: advertisement calls, 
squelching calls, and transitional calls - as it was not 
possible to assign some to either category. The 
squelching calls of t.\ i>lutterti also consisL of pulse 
trains (Fig. 5). Because these calls are highly 
variable in temporal structure, and were produced 
only during the recorded sequences of annul 50% pf 
the individuals, they are only briefly described in a 
subjective way. 


Strut tare oj f pulses in tu/vertisement culls 

The pulses in calls of both species are similar, each 
with a sharp attack and a gradual (negative 
exponential) decay (hig. 4), Estimated durations are 
4 bin for C gtaucfti and 4-7 ms for C. si^ntjera. The 
maximum ampliludc is reached within three positive 
or negative peaks of the carrier frequency, that is, in 
about I ms for both Una. rixamiuatmit ol the 
expanded wave form of each pulse indicated the 
presence of a clear sinusoid ( = I'undamental/cariicu 
with a frequency that is close to the Upper peak of the 
spectrum (Table 3 ). Accordingly, only ihc upper peak 
(= currier frequency) was used in subsequent 
calculations The frequency bandwidth at Hi dB 

below the peak is about 1200 11/ lor both specie-. 
(Fig. 4). The envelope ol the pulse is amplitude 
modulated to a depth ot about M) hiV/t (hased on the 
first cycle) with the envelope modulating frequency 
U?MR within a range of <V-M to K20 Hi for C 
S'iatterti, and of 505 lo 885 rfe fOf C si^nifera (Hg. 
4). These values arc close lo the difference between 
the upper and lower peak oi' the spectrum lor cacti 
individual (Fig. 4. Table 3). It is suggested that the 
lower peak that is present in the spectrum of the 
advertisement calls o\ some individuals ol each 
species (Fig- 4) is a sideband of the carrier frequency 
(i.e. the upper dominant frequency). The level of the 
R>wcr [Teak is about 4-0 dB below that ol the upper 
peak (Table 3 1. 

Slrttcturc {'(the advertisement calls 

Both species produce advertisement calls in long 
and regular sequences. for C\ \>lnucrti from Site I, 
calls were produced at a mean rale ol 26.5 calls mm ' 
(range = 22.9 - 2M.8; n = 9) at a mean cllcelive 
temperature of 12.1 C (range s 11.4 - 12.8). Plot ( 
signifera from Site H, the mean rate was 124.7 calls 
tftirr 1 (range a Ml. I - 187,1; a- ID *fl •' mean 
effective temperature of I1.4 3 C (range = 112 
1 F7)_ Values for three primary attributes (duration, 
numbei uJ pulses and carrier frequency) and the one 
derived attribute (pulse rate) for five samples of ( 
statu rti and Tour samples of C. stgnifcra. corrceied 
for the effect ol temperature where appropriate, are 

I mu i 3. Spectral thorn* wrist ics t»t pulses in advertise me in cults o/'Criniu glaucrli and C. signiteia 
All values arc in Mi. See ICX1 Jbf explanation. 

Species- Ind 











spectral peak 

spectral peak 


he l ween CF 

J-ielween LMI- 






spectral peaks 



and t'sp 

and DSP 

( . vluticrti 





at us 








a 1 34 












< \i\nufem 










67 1 
















T\r.i i 4- Physical liuimftrrhHrx t\f mh'vrthetiiem calk ofCvln'm glauerli ami C. slgmfcja, annta! ta an pffiu-tiVf icm 

fHTtitnft of I i.ft (\ \\ : heir ■■\ppropriuie iw lahlc 2) 

Foi each celt, the mean jihI standard deviaiion arc given on lite upper line, and range an paretilhesesi on the lower line. 

Species Sue 



Call duration 

N Limber of 

Pulse rale 



temperature ( "C» 



(p si 

frequency (ll/i 

C~ yhfiicrlt 1 


124 7. 47 

738. 87 

9.70. 1.21 

1 1.80. 1.94 


(11.4- I2.8i 


.7,5- 11.5. 

(9.4- 16.3. 

(35«4 4430i 



J 04 4. 0.64 

724. 153 

9.83, 1.50 

4279. 285 

(9.2 - 11.01 


18.0- 12.0) 

i 12.0- 13.61 




10.27, 1.03 

705. 115 

11.50. 1.88 

14.94, 1 SO 

4068. 215 

(7.7 11,5) 

(547 -935> 

(9.0 |5.5) 

ill.S- 18.3) 




10.56. 0.57 

526. 82 

10,06. 1,84 

16.91, L48 


(9.6- ||.4) 

(438- 655J 

(8.5- 13.0) 

(14.2 ISO) 

(3250- 4370) 



598, 85 

18.20, 23)0 


(iio 13,1) 


(8.0- 16.0) 

(134 -23,3) 




1140. I.IK 

664. 130 

10.57. 13)2 


4098. M)2 


(7.7- 134) 


(7.5- 16.0) 

(9.4 -2^^) 

(3250 4570) 

< ' \i<aii/trti 6 


272. 77 

6.28. 1.54 

19.08. 3 41 

2790. 325 

(10.4- 12.2) 


(4.0- [0.3} 





10.64. 0.7s 


5 31. 141 

293)5, 7.56 


(V...2- 11.2) 

( ( )| 272) 



(2092 3871) 



11.43.04 8 


4.87, 0.69 

22.50. 4.5 i 

2569, 87 

(11.2- 11.7) 

(101 -2371 


(17.5- 293') 

(2471 -2710) 



9.25, 0.27 

158, 20 


21.32. 1.9 1 

2825. 169 

(S3)- 9.ft 

( [25 - 192i 

(43) -5.0) 


(2549 3103) 



10.75. 1.02 


5.26. 1.34 


2645, 279 


(S3)- 12.2) 

(91 -437) 

(43) 10.3) 



1000 r 











. -as . ; ■ 






Fig. 6. Box plots lor durations of advertisement calls at recording sites, ol' Crinitt xlatterii (hatched boxes) and C- sigtiifi?ra 
(open boxes). Values are corrected to 1 I" C where slopes are significant. The box indicates the interquartile range and the 
included horizontal line is the median. The vertical lines outside the boxes (whiskers) counecl to the last data points within 
± 1.5 s the mtciquartilc range (the fences). The asterisks indicate outliers (values lying between ft 1.5 and 3.0 * the 
interquarlile range) and Ihe open circles indicate extreme outliers (values beyond 3 x the interquartile range). 










1 II 

f 64 

J I I I I I I L 

2 3 4 5 6 7 




Fig. 7. Box plots for numbers of pulses in advertisement calls at recording sites of Crinia glauerti and C. signifcra. See Fig. 
6 for explanation. 




50 r 



i2 20 - 

10 - 

T ^ 


4 5 6 



Fig. &. Box plots for pulse rates of advertisement calls at recording sites of Crinia glauerti and C, sfgfiifefa. Values are 
corrected to 1 1 ° C where slopes are significant. See Fig. 6 for explanation. 



presented in Tabic 4. Values lor combined samples of 
each species are also presented in Table 4. Box plols 
ol' these values at each sile are presenled in Figs 6-9-, 

Correlations oj f attributes of advertisement calls with 
body length 

Calling males ol C. glwterti are smaller than those of ( I 
si^nifcni (Table 5), with mean snout-urostyle lenglhs e>i' 
1631 (ranges 14.0- 19.0; n = 2 l ». and 21.31 ( range = MK.O 
-25.0: n = In) mm respectively (l-tesl; /> < 0.001 ). There 
is no significant correlation of earner frequency (corrected 

io 1 1.0" C; Table 2) with snout-urostyle length when all 
collected specimens of C. ^ianerti are ineluded (r = - 
0.297; p = 0.1 IS; n a 29); but there is a signilicanl 
negative correlation when only those specimens thai were 
measured following preservation are considered (r = 
-0.440; p - 0.032; n = 24). For C. signifera. however, there 
is a highly significant negative correlation between earner 
frequency and snout-urostyle length (r = -0.796; p > 
0.001; u = 16, all speeimens were alive when measured). 
A scallergram of the conelation of snout-urostyle length 
mid carrier frequency is presented in Fig. 10. 

5000 r 







*= 3000 





1 6 


r 1 











Fig r 9. Box plots for currier frequencies of advertisement CilIlS al recording sites of Crinifl ^Imicrn and ('. si^nifera. Value*, 
arc corrected to I i C ivhere slopes are significant. See Pig. () (or explanation. 

Iaiiii 5. \%thtt1 (in mm) for mout-uiVXtyte [Cftftthii "{ 'watts nj Cnnia glaut-rti and C. smnifcia tolltrltd a/hi tin a 
athriliunwttt tails had been m onkti 



Sample si/e 




Condi Hon ol' 


( '_ xhiih ni 




10.5 - IM.U 






14.0- 1 5.5 






15.5 - 17.0 






15.0- 16.0 






17.0- Itf.O 






14.0- IV.U 


( . wen'/erw 




22.0- 25.0 



l l >.7 







ISO- 25.0 

: 'i 



Geographical variation in advent semen! valh of C. 

Analysis o( variance, with Site as the grouping 
factor, indicaied that there are no significant 
differences lor number o\ pulses (p = 0,074 ) and 
earner frequency (p = 0.41 I). There are. however, 
significant differences for the means (adjusted to 1 1° 
C) o\ duration (/> < 0.00 1 ) and pulse rale (/; < 0.001 ). 
Across the distance of about 450 km covered by the 
fi\e sites (Fig. I), there is no consistent trend in 
duration (Fig, 6, Table 4). although a multiple 
cuniparison flukey test I indicated thai the means for 
Sites 1-3 are significantly lower than those at Site 4. 
and Site I also differed from Site 5. There is a cline 
of increasing values for pulse rates (Fig, 8, Table 4). 
\ lukey test showed that the following means for 
pulse rate differ significantly: Site I from Sites 3. 4, 
and 5; Site 2 from Sites 4 and 5; and Site 3 from Site 

(.hnuriipiiua! variation in advertisement rails ofC. 

Sites 7. X and 9 are within the continuous 
dislribulion yf £ si^iutera (see maps given by Brook 
1983; Tyler 19S5) and are spaced at about 300 km 
intervals. Site 6 is in the isolate on Eyre Peninsula 
and about 300 km from Sites 7 and S (Fie. 2). 

Analyses of variance, with Site as the grouping 
factor, indicated that there were significant 
differences for all lour attributes of the calls (p < 
0.003), Multiple comparisons (Tukey Lest I revealed 
the following significaul differences in means 
between sites; call duration - Site 6 from Sites 7-9; 
pulse number - Site 6 from Sites X and 9; dominant 
frequency (adjusted to 1 \ C) - Site 7 from Sites o 
and 9: pulse rale - Site 7 from Sites 6. 8 and 9. 

Comparison of advertisement calls ofC. glauerti and 
C, signifera 

Results of analyses ol variance, with Species at the 
grouping factor, indicated that the advertisement 
calls of the two species differed significantly (p < 
0.001 ) in all tour attributes. From a consideration of 
the combined samples for each species {Table 4). 
calls of C. glauerti are much longer, with means of 
durations differing by 3.4 times, and with no overlap 
in ranges ol variation (Fig. 6). The mean value for 
number of pulses in advertisement calls of C 
ylanerti is twice that o\' C, signifera but there is an 
overlap of ranges between 7.5 and 10.3 (Fig, 7. Table 
4). For pulse rales, although the mean for C. glaiterti 
is only 64'/r of (hat for C. st\>nifern. the ranges 
overlap extensively (Fig. 8. Table 4). particularly lor 
die closes! samples (Sites 5. 6). The mean for carrier 












• • 



• • 






• • 

• • 






e c 











1 1 


14 16 18 20 22 

Snout-urostyle length (mm) 



Fig. I0_ Seatleryrarn showing the correlation between carrier frequency ami snoiiMiroMyle length for mules of Crinifi 
f*l(liii rti and C. Jflwrfjfi'rti lhal were collected after then Lalls had been recorded. 

( AU.SOrrAV.YM f.7,W7.A'//.\NI)( SlGNIl I HA II? 

'i'Mil t <>- f l h\suulth«tat-!t'>'s}u \ ,.f mwh iqttelMflX c«//.\ of Cl'iitia elatieni; //W( the .\twtc set oft nils pn si-ittviUti tifjttW $ 




Number al 

Low pulse 

HiL'h pulse 

Overall pulse 


(unci Lapc 

( C C> 



rate ( p s ' 1 


raw tjj# v i 

I requeue) ( Hy i 






43 2 




h. (R44I-I) 



■j i 





c (K404-2) 












$6 1 

H7 2 



L-_fK407 (,) 







1. (R4H4 M 








g. (R407-7) 












57 Ml 


54 55 



'M> 13 i 

43 K 832 

2| 511 

3 7 46.7 

61 7 -205.9 103.2 

£7fl3 +392 

: I.iuit ih.m lim puKc uif bCCUUtt ol the loin lone brCdb in the linii 

frequencies is higher in C. .uliittctii. by 1452 Hz. bill 

wiih m\ overlap ol reuses between 3229 and 35$8 H/ 
(Fig & Tabic 4). 

Structure of the $ifuehhtit# vail if d tzluuerii 

Those calls are highly variable, as is indicated by 
I he selection of waveforms presented in Fig. 5 The 
pulses arc sometimes in groups within a call, and ihe 
pulse rale can vary greatly through u call (Fig. 5). 
Values for four attributes of the seven calls presented 
in Pig. 5 aiv giycil in Tabic 6. The pulses are ol 
sinulai structure to lho&0 oi the ailvcrtiscnicnt call 
and (here arc also two peaks in the frequency ppcctrfc 
as in the advertisement call. For the seven calls 
presented in Fig. 5, ihe lollowme- dalu apply {set 
section m) puKcs of advcrtiscinenl calls for 
methodology): duration - mean = 3.77 ins (range = 
5. 15.1 ): carrier frequency - mean = 3 L )55 1 1/ t r&rlgC 
= 3481 - 4427); upper frequency peak- mean = 3976 
(range = J703 - 4414); envelope modulating 
frequency - mean = K77 Hz (n = 5; range = 735 - 
I062& lower dominant Frequency peak- mean = 3260 
Hz (n = 5; range = 3100 • 3464), The dil'lerence 
bctWCCD Ihe means of peaks of upper and lower 
frequencies Ol 716 Hz is consistent with the 
explanation advanced lor the pulses m the 
advertisement calls - of the lower sideband of an 
envelope modulating frequency. The pulses may be 
grouped within a call (Fig. 5) m^ the pulse rales can 
differ considerably beiwecn groups ill one call (Table 

lU-it^mphh of variation in~ frequency o) ocvurrciHi 
of ihe siftirh hint* mil o/C. glauerli 

Recorded sequences of calling by 25 of the 51 
individuals included squckhmg calls. Because Of 
confusion from Ihe calling by two or more close 
individuals, only those of 21 could be reliably 
assessed, classified and counted (Table 7). A 
geographical trend is evident, with the frequency of 
production o\' squelching calls by individuals being 
higher in the southern and south -eastern samples 

(Sites 4. 5, Table St. No attempt was made LO 
determine the extent ol variation in the squelching 
calls ol individuals, nor over time for a population. 

Slnulitrc of the amnmtcr <olh of'C. signilcra 

Liulejolin el uL < 1985). by playback <>( 
advertisement calls of C. puiin\i\>nii< 'iv and C. 
ftignjfisra at peak sound pressure levels (0 dfi re 20 
fal'a) above c. 98 and 104 dl> respectively, evoked 
encounter calls (= territorial calls SO/W Hawe 1 
Littlejohn |s>77) from eight males of C, &i#ftiftiM ill 
Willowmavm m south central Victoria (57 16' S. 
144 '54' I-). The encounter call thig. 1 It is also || 
pulse train, vvith a regular pulse rate (Hawe 1 . 

Taihi 7. \iiinhirs f\f tklwtftxemtMt t'Xilh. innnurJiutr 
((nin\ili>t>tul) inlh and M/nckhin^ titll\, tout proportion^ <>/ 
liw hater f(ij f off itiifs af Unit huli\icliuifi, pwiitu tul hx 21 
maks i'l I rinia glaittiti. 

N.'ie thai because nfoveilap wall neigh bui)rV calls ot 4> could not be analvscd. 

Site Individual 












su tit' Whin;: 



. . II 














II S3 
























(1 iy 






























fl Ift 


1 1 























































)i i M.;.uni.rjoiiN& j. rw rich it 

Tabu 8. Munhrrs iuuf pHtpttnttws of umics of < riniu gluucrii fimditfittlft ytflttkhitjg Kiih m pacjt sin 


Number ol nmlos 

Niuiihci of males 
pit klucinti wucsichiirg talis 

Propuilion of male--, 
producing squelching Cilll* 



Liltlejohn l"77> A waveform of an encounter call 
from Site 5 is piesenled in Fig. I F Values for means 
and ranges from the Original data lor advertisement 
calls and evoked cneoiinlct calls of (he same fjyjS 
individuals discussed hy I iltlejohri cf u(. U885) are 
presented in (able 0. 

Otiti(Hui\on of squelching etdlofC. glauerti and 
terraoriul call o/C. signifera 

The variable squelching call olC'. glauerti (Fit;- S. 
Table o) is longer, contains more pulses, is of highci 
average pulse rale, und ol higher dominant frequency 
(Iran the lerrilorial call of C signifera (Fig. I I, Table 
Oi Rvcn so, they are bold pulse trains which arc ol 
similar earner frequency bul ol longer duration, 
conlain more pulses, and are ol Irighel pulse rate than 
(heir respective advertisement calls (Fig. 3. Table 4). 


200 300 



Fig- II. Wawloim o) an encounter call ot C v^ntl'tra 
< Reference: K40X .9; Sile 0: elTechvc lt j fni>eraUiics_ wct- 
bulb jii I I.VC. waler L2.5 1 C). 

1 \ui j </ t vmpiifixtW <>f nttnhttnw at ailvv ruse mini mlh 

ami mronntvi mils a) Crinia sigmlera jnun Wil/aunuiYin 

Yiri/'ttn tfat/n data *>f lJttU'j<>hn c( al. l$&3). 

Means und range* tin parenthesis j are eaven oi =5). Data 

flffi not corrected lur pov-ahle effects ol temperature. 

P.Ucclive lemperatuic; raimocl from 10.7-1 \.2^ € (mean = 



Advertisement call 

Em oiilitcr call 

Duration- Ons I 




<I7 C >-2|0) 


a 3 


(4 5) 

iKi-l l >) 

Pulse rate 



fp s ') 


(74.2 100. It 

Upper dominanl 



iVequoiicN (H/l 



Production oj encounter calls hv innU s of C 
signi fern 

In a subjective evaluation during playback ol the 
tape recordings ol* C. si^nifem, involving S3 
individuals and sonic 3527 advertisemeni culls, three 
interactions, presumed to involve production ol 
cncounier calls, were noted. Otherwise, the recorded 
sequences Of 47 individuals consisted only of 
advertisement calls. 

Sfmefnn' oj the advent wuiettt aids of both speeics 

The advertisement calls of both species aie of 
.similar structure, each consisting ot a quasi- periodic 
pulse train. The pulses are short damped oscillations, 
each with a sharp attack d\\i\ an exponential decay. 
There are more pulses in the calls of C glauerti. and 
these aie produced al a lower repetition rale. The 
frequency spectra arc of similar shape, with most 
individuals having two peaks, the upper being 
allribnled to the fundamental i- canter frequency), 
while the lower is presumed lo be produced as the 
side band of the envelope modulating Irequencv. The 
carrier frequency is higher in C, i>tauem< a\u\ Ibis 
may be correlated with ihe smallet si/.c o\ males ot 
this species - a usual characteristic of the calls o\' 
anurans ie.g. Robertson 1986). There is a trend oi 
increasing pulse rale from north to south-east in the 
samples ol C glaueni and the durations arc lower in 
the samples from Sites 4 and 5. Values for samples o\ 
the other two attributes display no obvious pattern. 

Advertisement calls from tStte 6 in Ihe 
geographical isolate ol <". signifera on lower Fyre 
Peninsula arc longer and conlain more pulses than 
those lo the cast in the main distribution o| r 
signifera. Thus the westernmost sample ol ( 
\i,unif''rti (from Sile U\ is more similar lo those of C 
ii/iinerrl than are the others. The populalions of f 
sixni/'tra on F.yre Peninsula have probably been 
separated from the main distribution since ihe sea 
rose to its preseni level at the close ol Ihe last glacial 
period of the Pleistocene Fpoch (from c 1 2.000 to c 
6.000 years ago; sec Liulejohn *'/ ///. IW for a 
summary and references) and this isolation may have 
contributed to the divergence. 

I'M I. Mil CA'/VMi,/ \l 

Odcndaal tt «/; <l u Nnj recorded a sample Of 
advrfUscnvm calls of C tfffrttft'/a Runl the same 
local ion on >ci|nwm<tn Creek (their Recording Sue 
No. 5: our Site No. 7) jmd over a comparable range 
trl effective iwalerl temperatures * 1 0.S 12,2 < v. 
*■' .2 112 Cr this study). Fbl till lour uiinbules. 
IHAVCVCfi lllGJl VffiUittis a e higher than those obtained 
in the present stud}. The use ol oiber protocols ami techniques may account for mm tic ol ttlC 
dilleicnees- htfl at this stage, no explanation can be 

FlfVg/tfVlriJ t'fcttrfunlrr fiil/s fri C. ghuierti 

Moie-iceem observations by Lilllejohn Umpub.) 
unhealed that C. gkwarti occurs in syntopy with ( 
suhitist^mjrra tL ittlejohn) near (he eastern Until td 
its distribution U*,g. Site 5; Fig, 1 1- O'ww 
snhtitu$nil'er<t has an advertisement call that sounds 
like a "long low-pikhed squelch" < Lilllejohn $57. 
l&jWp. lor two temporal attributes ot (lie 
adveHfeCWteni tfill« &f -^ individuals of C 
\uhitfsri:niJ<r<L collected to an ctloctive temperature 
of HP C [he mean duration is 54(1 ins - 420 
6n0r and the mean pulse rate is 174 (range = 12°-- 
2l(»Kl ittlejohn l%l). Asihese ranges overlap those 
Of the squelching Will tf <' vhaivrti (Table 6), 
explanations olliei llian reproductive character 
displacement must now be .sought Tor Hie htgJicr 
IVei|iK'iu'ie'; ol occurrence of squelching calls in the 
south unci south-east but none can be provided al 
present. Clearly. there is a need tot fun her 
investigations, including playback experiments with 
advertisement calls and squelching calls as .stimuli 
under controlled conditions. Such studies should be 
preceded by Ihe docunentuhou uj frequency ol 
octunenee of squelching calls in natural 
assemblages, and ihe context in winch they arc 
produced, The measurement ol lite sound prcssure 
IcvcK o) ealb- ol coiispccifie neighbours is ulso 
required so thai the appropriate stimuli can be 
applied (Little John rrai 19X5). By varying the levels 
of stimulation, thresholds could then be determined 
.iii.i -vographica! patterns may be revealed. 

ttcU(ti<>n\hii^ >>l C, idatierti 

I torn l he presented data, it appears that the nearest 
populations of ( . yhntvrii and C. .-lixniftm t Sites 5, 
6) have the most similarly structured advertisement 
calls. The main difference betvveen the calls o\ Ihc 
two species is in earlier frequency which may be 
accounted lor by Ihe difference in sizes of the two 
Ui\a (Table 5. Fig, 1.0). The simitariiies in eall 
sirueU.m- are consistent wild the postulated clost 
relationship ol the two lava suggested by: Man 
HVS7V. and subsequently supported by |Q£ 
muhtvariate numerical analyses pf morphology rift 
leatutes ol lite history carried out by Blake < l u 72i« 

7A'/MMU', S/<VV///7>'i 


and Thompson th»sl>. Although conststeni in 
.showing a close ussm huhm between diem. 
molecular studies <U) not help m resolving the 
relationships iff ( qtuitvrti ami C . si^nijcrn. The 
.ilhumin immunological analysts of Daugheiiy and 
Mavson |I9§31 fJ^OC* £ sn^tiftTu closest lo ('. 
ripariai)^ ID units), followed hv C vhtumi And ( ' 
futnnsi^Dift'ru (both 24 If) units) mh\ C, Rtiijrgimtl 
CM ID uinis). The cLtdistic auulvsis ot allo/ymes 
carricct otii by Barendse (l°K4) oflers seveial 
inlerprelJtions and appeals iiMuvlnsive about the 
icliiiionships: in one scenano C. ^laiirrii and C 
li&iijfcud ittB grouped wiili C yrni^nitm, Baa'titlse 
<1 1 >X4) tliil n»>t mcIuJe (. tiiuuut in hts sniiJ\ 
Kotx-Hs Si Watson t IW3) have reviewed the iceenl 
hieiatuie on relationships within some croups <il" 
Australian Irocvs. 

Three ol the species ol CviUtH described since the 
\\i'il <»t Main ( !°o7) - C InJhi^ita Martin. Tyler £ 
Davies (Murtm vi »{ \ABBt* C nuwui lylci & 
Paiiet (Tyler & Parker l l >74) and (". rif'O'iO 
Ulllcjelhn Si Mariin (I itilciohn & Manin l%5) - 
have clearly pulsaiile advertisement calls. I\ kr *S. 
ParKci ( 1°?4> noted the similatity oi the 
adverliserncnt calls of C rt'itittla and ( yUiiHftt. but 
Ihey did not provide infornialion -about the recording 
temperatures As Ihe recording trt die call o! C 
rcmola Wih obtained at Vlorchead. Papuu New 
Crimea in January, the anihicnt temperature*^ 
prcsutnably were much higher than those ;ippl)iue U) 
recording.s analysed in the present study. Hence, u 
direct t.nmp;irtM>u cannot be made with the calls ol 
C gfat/prti obtained H\ tetnpr'amrcs ol' LVI C ami 
lower. Blake t I97S) noted thai irtnui fifiutiu lacks n 
tympanum and columella and placed the ia\on into a 
diiYerent species group along with C ftjflWJtffWtf 
t(nmllier). Because o) the lack ol a distinct 
tympanum in (". tvmotu t'lvler ^V Parker I l >74i, it i% 
suggested that ihere may uoi be a close ielati"i)sliip 
hclvscen this species and C, xluHrrti 

,\s ihe name indicates, the advertisement tall \>\ t 
hiliugtat is strongly hipflask (M.uiin rt ,il l l )X0). In 
thi.s species, males commence a ealbtig Muniener 
with short calls (<Mfi ms) ol big'h pulse ralet>7(> p fi ' |, 
and then gradualh change over lo long calls <>5X0 
ms) ol low pulse ratjf « 54 p K ' l< Aeain, (he high 
recording temperatures twethulb air = 2.T4 2rvfV 
O uiean that it is not possible to make a ptoper 
comparison with the advertisement call of C 
qliitti'/lt. l:\en st>, tlu- iiiitliospectrogriim otthe shot! 
call appears (" K* sinnlar to some M' the variations in 
the squelching calls ot C, yjmurti- Martin ft at 
i I^S(J) considered the functional significance ol the 
two distinct calls of C bifittftna and raised tile 
possibility Hut the slioit calls uf higher pulse rale art 
mating- calls, and that the long calls of lower pulse 
rati- are iLititorial calls, ihe converse ol ihe MUiulinn 

I 16 

m .1 i nil hjmiK&j. r.wrjuiii 

for pillft rale ill < stymjVM Tin- Ioiijj call d!C. 
hilin^itu contains akuil DWlCfc as many pulses u-?7) 
as mi the adu'lliscmeill Call §i ( ^Inurrf, (<$&)'. I| 
allowance is in,i<tc fo? the dillciciicc til lenipcialure 
(b> Hsmy u O M ol 2).. the pulse rate raft' l'i/invu<i ,n 
1 1" C won 111 foe tiboui 2D p * ' and eauld hvffrluji thi! 
pulse uies in the advertisement calls off, $lciucrlL 
Crini,( htlitt\>n<i pO&sCSSCs a tympanum hm possible 
i i" i' villiiiitio with c. iitiutcni were not considered 

l>\ Miirtin p/fi/ i IMSO). 


Thu field studies in Western Aastraliu were curried 
inn under licence Numbers 1274 1 lash) ,md SF l*' 1 
(19MV), issued by Ihe Department oi trtnaeryaituji 

and Land Manage me nl. The lie Id si i lilies were 
t.irned mil during Held re scare h leave from I he 
I'liivursilv til Melhtamie. hquipmeut was provided 
by firUMW from ihe Australian Keseart h ( QUTlcU 
I \RC| Hnd thC I 'nocisttv of Melbourne. Recordings 

oi (\ sjviuf.'ia lo>m South Austiaha were ohlaine.I 
iluini'j ihe lenuie ol a research mini from ilie AKC. 
Number A 1883131*1 ( 1989). Some irf Ihe acoustk 
and sialislieal analyses, and preparation Of pan-. Of 
Ihe iiiauuseripl, were carried oul during Ihe tenure ol 
Other gnCiHtS bom the ARC. Suiiill grants NJieme 
(firanl Numbers SCi (W355I4, S 0KsM757X) m«i lire 
procedures dial were developed have provided 
protocols lor acoustic and statistical analytical 
tcelniit|uesut' wider application. Mrs \\ (J, Lilllejohn 
assisted with recoidme al all ol ihe siies in Western 
Australia mid ai two of the South Australian 
localities. I )| T. Ci. lilllejohn assisted with I'CCOftll fly 
al the other iwo Soulh AuMiulian localities, Mi I.. A 
Smith, western Australian Museum iirrunped lor ihe 
euration ol Ihe preserved specimen-, ami kuallv made 
Ihem available tor subsequent csarmnation and 
measurement during a subsequent visit, |)i M I 
Kecutgh provided :idvive 01] statistical analyses and 
Or M C, BcnUCt-QIwlk provided an interpretation ol 
Hte spccltnl ami temporal vlruchuv ol pulses in the 
advertisement calls 


H ■! i 'i W [|VH4) Spcci.ilion in the gOlltth inuia 
• Anuia \l',Mt- mi H in MMtilic-rn Australia \ 
ph> liiL-uiieiK ;i 1 1 li lysis ni allo/\tnc dan supporting 
1'ihl. 'imc stieeintn-tli m southwestern Australia tfiw/yftVwi 
38. IHs lift) 

Hi kKl *- ' P- ll*>73) T«xt)nu»if mid reliniwnshlp* iir 

imohairacltinc lro>-. 1 1 e|>r-iLaO ,\ (Uimoneul 

►tpproudr, Aiur, I a><4. 21. Ikm i<> 

UlbUttNi A J llWl 'Atlas If!' Aieaialian Amii'.i 

(l)epailineni tfl KooUfgy. t Iniver-.iis i»l Melhuunie 

KV»4J Ml^ "' R*up uf Soulh Auarali.r 

iDepailnicnl "I /uo|op\. ( iiim-isiiv Ift MelhiMirn, 

Bwjwk*. W- i £ WiL+uN. i:. (). tl^jfi) riiLi.j,!!.-. 

tli-pkici'mriii \v\t 7jmI 5, 49-riJ 

I ■ ii. 'a iv tl (i, ilvvji kepiilcs ami Amphihiui>« ol 
Ausltalia' >l\\ ciio (Kced Hooks, (."li.a'-.wuoJ). 

, Camiuov I-, L & C'h,oik-_ II, M, il^S.V) 

'Xooloiiiiail Ciiialoiitic o| \ti-iirulia. Vol. I. Amphibia and 
Reptilia' (Ausiralian Oovcinmont PllhlLshino Sci^lei.'. 

h.Mia.M.-u.C, H. cV Mwst-M. I., K, t l l >Hi) A luoLlvmir;,! 

mwtssnwiir oi tl u? cvukainn nj inwihntratliiiic IttiMi 

Lil Hi iMii:s_\t. I (l')>7)Ancw s|Vck--.a it..: 1 o! 
t ritttfl. Hew Autf, Nui to. Ifl !3 

(I*>SK) A MUM -.|KCif. c>1 d«>u ol the i;ni^ VrJMO 

NcUutli li-Min -,oHlh LMfleoi .'XLraulKL /■>,»■ linn, Soi\ 
\.\ W S3, 22?. rib. 

H*)s«>) (;i|i (linerentiation in a compTes ttl wwpn 

,pVi uM'I'f '/>r/ c ; i \nuia. I ei'lod.K'Ohik.LM kwttttitm W 

as? 4'-K, 

ii l »(»ii Aye mill orpjiii of n-jiTiy ^Hiirhwc usm 

AusiKiliiin -.peciL's ol Cuiii.i ( \naia: l.e|Uotiactv luhu 1 ) 

pp 514-53^/n BtoSjtW.l tTJL} VpHoftniw SpocMOO 

(1 ni\et-al> ol fiWHiS PtGx.'j, Au>tlll). 

I WWJ rJi'M^raptlU CoKiholl .intl iicHmih »;il| 

JtiKiLiai.a ia f'tf/ifii \i'itifr>,i iCwhttim IH. 202 

I h'?t)l i iMIHi htWUttth HSIS i Ullli;! 

Ltf^odfietylute) f!i?utfmfi|iU disirihuiitw. niaian! a\\\ 
slnielnie. ttnd telalionstiips. fare, Utitt. V< . N S W 'M, 
KW 127. 

ity?/) l.oiiuijii^L- JKinisiie coitmutnicalioii m 

.iiiuuais: Ait ink* "mm! .ui<l cvoluiionarv approaJi pp 
[6? 2'M in Taylor. D II. vV (iuiuiun S. I (HilSj I la- 
Keproihk-itu' Bijilog) ol" Amphibian ' (I'Iciiuim I'i. ■--,, 
Nctt Vorki 

. llAKfttf*ON, I 1 . A. & M\rwnv R T (' tpJN'o 
litiuisprtilK ;k«mi-.Ik' inlfi actions in svinpaiiu 
popnkihons ol' Rounlviln S'tgivfciti iMlil A' pt)tfrt\tglf(lnw 
<Ainir;t; I , l\ h-lat t pp'. 2%1 2VA /// ChtgK CI 

Shine, K, (<: Ehmaiin, II. dais) 'Hhifogy nF AusliahiHian 

1 |oh s kl | 1( J Kopiiles' (Royal /-ooIo;jil;iI Soliciv ol New 
South Wales. Sydney: & Suoe\ licaily ami Son-.. 
C'liippini; Norlon), 

A M\l/iis. A. A i |»Wi5) \ .k-w -pcics ol rrf^iw 
tAnuta; LepioUaciylul.iO) iVom Soulh Auslniliu Utmtfu 
i%5. M'M.U. 

Kom ins, J 13.. \V m->(.-.;. (I |, & \yy : s I,-,, M, | |wn;;, 

1. K Mvob.ariu'hi.lae pp. -II-..S7 /// (il.i^hs. ( . I Rum 

CI J II. & Keesley. I! I . (Eds) I au..,. ofAusiialia Vol 
IA Aniphihia ami Kephli.i' i Aitsti jli.m Cios^TMtneiii 
t'uhhshin,^ Service, CinhL-oak 

M.\rs. A. K. (1957) Sllldtei in Ausfralian Aimia 1 llu 
j/CtlUM CW/t^i Tselnuli in south -wcaern A»S(Tfl.lki antl 
KunTC species from souih-csi-slern Ansu.ilij. \/m/. ,/. ZurV 
5,30 55, 

. L i . A. K. t^ lai n i joiis, M, J. ( |V58J Evu-tliliun in 

three eeneui nf \usiralkin Iro^s I yol\tlh>n 12, -^ 1-233. 
Mo-ii\. A A., TVI Wt M. J, cV'Dv/iis. M. ((M8(l) A m-u 
species ul' RanttU'flu lAnnra: I.epiotLietvlnlae) hem 
noilhwesteoi Australia, Ov'Ww l'>80. 1 ' J ,-^o 



OdendaaL, F. J., BULL. C, M. & Telford. S. R. (1986) 
Influence of the acoustic environment on the distribution 
Of the frog Ranide/la riparia. Anim. Behav. 34, 1836- 

Roberts, J. D. & WATSON, G. F. ( 1993) Biogeography and 
phylogeny of the Anura pp. 35-40 //; Glasby, C. J., Ross, 
G. J. B. & Beesley, P. L. (Eds) 'Fauna of Australia. Vol. 
2 A Amphibia and Reptilia' ( Australian Government 
Publishing Service, Canberra). 

Robertson, J. G. M. (1986) Female choice, male strategies 
and the role of vocalizations in the Australian frog 
Uperoleia rugosa. Anim. Behav. 34, 773-784. 

Siraughan. 1. R. & Main. A. R. (1966) Speciation and 
polymorphism in the genus Crinia Tschudi (Anura. 
Leptodactylidae) in Queensland. Proe. R. Soe. QUI 78, 

Thompson. M. B. ( 1981) The systematic status of the genus 

Australocrinia Heyer & Liem (Anura: Leptodactylidae). 

Aust.J.Zool. 29,93-102. 
Tyler, M. J. ( 1962) On the preservation of anuran tadpoles. 

Ami I Set. 25,222. 
- (1985) Biogeography pp. 225-229 In Twidale. C. R.. 
Tyler, M. J. & Davies, M. (Eds) 'Natural History of Eyre 

Peninsula' (Royal Society of South Australia. Adelaide). 
& Parker, E (1974) New species of hylid and 

leptodactylid frocs from southern New Guinea. Trans, R, 

Soc. S.Aust. 98,71-77. 
, Smith, L. A. & Johnstone, R. E. (1994) 'Frojjs of 

Western Australia' (Western Australian Museum, Perth), 
Wells. K. D. (1977) The social behaviour of anuran 
amphibians. Anim. Behav. 25, 666-693. 




By Graeme F. Watson* & H. Carl GERHARDTf 


Watson, G. F. & Gerhardt, H. C. (1997) The breeding biology and advertisement call 

of Litoria splendida Tyler, Davies & Martin. Trans. R. Soc. S. Aust. 121(3), 119-124, 

28 November, 1997. 

Breeding biology and description of the advertisement call of Litoria splendida are 

presented. Analysis of a call of L. caerulea is also provided and shows that these 

similar, syntopic species have structurally similar calls and thus presumably show 

significant acoustic interactions in mixed choruses. 

Key Words: Litoria splendida, Litoria caerulea, frogs, calls, breeding biology. 



by CrRAt-AU KW\!'SiiN & H. CaRI Ch RHARrrt 


W,\isoy(i.l*. & (_ii:UitMiOT. H.Ci |9tf?) "titer breeding hinlouv and advertisement call wl iJ!nii<i-\}>trniiithr\'\ter, 
I3WVIW& Marrm. T/W/fv. fc \<v. ,V. Am 121(3), H*> 124, 28 November. 1W. 
Bidding biology anJ description of the advertisement thill ol Uttnui spt&tdkfa are presented. Analysis of j 

L.tll til /„ tuLiiilvti is jlsit piuvuJcd and shows that these similar, synlopic species have •aiLtUurally >iniil;ir calls 
and Uuin presumably show siiiniruJiiil acoustic interactions in mixed choruses 

Ki v Woims: Uittiiu tyh'lHlfrilt, l.fii.ffu itiviuiru. hops, calls, breeding hiolugy. 


f.itoriii splentlkki lylefc, Davies & Martin, K'77, is 
one of the largest [.ultilt body length X2 - 10ft ntni 
('lylci l'W2)| and most beautiful Australian frogs. 
The species is thought tn he sparsely distributed 
throughout ihc widespread escarpment country ol the 
Kiniberley Division of northern Western Ausiialia 
riyler \^)2). I.ttnriti splendiihi is notable |nr: 
ttypctlroplned paroloid iind rostral elands (Tyler & 
Davies W*). a characteristic it shares with 
senescent members of the closely related Utftifid 
ntt'jvh'u* an ability to iitili/e glandular secretions is 
a waictpiot'l covering (Tyler & Davies W3), and 
the prodnclton of pharmacologically at live eaerins in 
the skin secretions i Tyler fk. Davies \W$). 

What litlle is known ol the reproductive biology of 
I si>lr*ttl(<lti is based on reproduction by captive 
individuals (Tyler 1994). In an aquarium, a female 
laid 2000 eggx, depositing them in discrete clumps ol 
up lo 200 eggs, winch Tyler ilW2i has suggested 
may reflect ait adaptation of females in the wild to 
lay batches of eggs in several temporary ponds lo 
maximize llie likelihood ol al least some ol the 
offspring surviving the larval stage. Maximum 
length of the tadpoles was 54 mm (Tyler \W2). 

I.ituriit Kpicmliitu is known to call, and presumably 
breed, after heavy rains in the early wet season (C. 
Done, Department ot Conservation and Land 
Management Kununurra unpub. obs.t. During Ihc 
unmet ous field trips lo Ihc Kiinberley Division 
undertaken by one of us (< iPW) and colleagues from 
the University of Adelaide over the past 20 years. 
however, calls tit the species have never been heard 
nor has breeding been observed. Jn fact, (he species 

l\-p; t iliil.-ML ot /(hMul') University Ul Melbourne* Purkvill-r Vie. 

OivKinn itl HiiiliH'.iL'.'l Siii'in r rmvip.nv n| Missouri < olnmbM 


is rarely encountered except when associated with 
artilicial hahilais. lor example loilct blocks, where 
cool, moist sites aiv readily available i Tyler 19521, 

Dining January, |997. we visited the Kuiuiniura 
area and entounteicd a breeding chorus oi" L. 
s/flctulttla. Here we describe the call frjf the species 
and provide brief notes on associated behaviour, and 
also a desi. nphoii o| (lie call nf synlopic L cuenili'it, 
a phylogenetically closely related and ecologically 
similar species. 


Recordings ot calls of /.. sphiulida were made 
using a Sony TCD-5PRO cassette tecoulei (tape 
speed 4.76 cm s ' ) and Beyer M-KK cardioid dynamic 
microphone. For comparative purposes, one call ol a 
synlopic L, i-ncrnica was obtained Irom a video 
sequence of die breeding chorus (Canon Digital 
camcorder; Hi H mm tape) Air wet-bulb 
lemperaiures (the effective temperature of frogs 
calling on land) were measured at the calling site of 
each individual usinjj an electronic thermistor 
llterntomeler (Takara Digimulli Model Dbl I). 
Recordings were analysed on a DSP 5500 digital 
Sona-Graph (Kay h'lemclrics Corp.) ustnii the in- 
built set-up #10 | sampling rale (samples sec 1 ): 
10240: frequency range' 4 kH/| with playback on a 
Nakamiclii Dragon cassette recorder, Overall 
variations in lapc speed (i.e. from recording lo 
playback) are estimated at less than flJ&% and 
frequency responses of all audio-electronic 
components are close lo linear within the relevant 
frequency range (based on manufacturers' 
sped Heat ions) 

For each call, three primary attributes were 
determined: (i) duration, as the interval from the 
beginning of the first pulse lo Ihc end of the last pulse 
(ms): (ii) number ol pulses per note (direct count): 
and Oil) dominant frequency (H/). as the maximum 



value oi the spcciruu of powci Qi ro& the whole cull, 

In addition, a derived characteristic, pulse repetition 
iatc (pulses s 1 ). was calculated a* MHJO (irl 
pulses )/duraLion in tns. Levels of resolution were less 
than I His for tOWpOftll characicristies and |$ig ili.m 
10 11/ lor frequency 

because (here is no possibility of misidendfyinj: 
diese lactic and distinctive rrOgs and in the interests 
ol their conservation, voucher specimens ot recorded 
males were ntn* taken. Video and 35-ltUU- 
pholographic records eff I he calling males were 

Results and Discussion 

Btvedihg stw uwd bwetlfttg Uekmtnu 

(Ovedinj: and calling were observed, following 
liea-.> afternoon lain, on the nighl of IX.U9ST7 in an 
area ol sandstone escarpment adjacent to a large, 
temporary pond near Ihe main car park within the 
Hidden Valley National Park. Kununurra VVA. The 
pOIld WW foimcd within tin i" defined watercourse 
thai ran along the base of the dills and was led by 
njn-crtT from a number of temporary waterfalls thai 
flow down the cliff face after heavy rainfall. 
Occasional calls Iff males at other nearby sites were 
hcaid bui choi'K behaviour was confined to this one 
silo, lour species o\' Iroys. l.ii<>hu spfciuhilu, /., 
. titntlm. I- mhtlla and ijnwodxnuxtcs (UTItfiWfi 
were calliny atound the pond. Ampleelanl pans ol /.. 
siilitulithi and /. ttunilt-a were observed in arid 
AfOUnd ihv pond. No paiix of the other two species 
were seen, although no exhaustive search was earned 
out. Ihe toamv egg masses of /. nrmtrn.\ were 
scattered over ihe pond as were large, floating, 
single-layered sheets of hylid egi:s, presumably 
those of/., Spiefiilithx and /.. manlcu. although we 
did not observe any pairs ol 'cither species depositing 
eggs. Ncverlhflcss. contrary 10 the suggestion ut 
Tyli i ' I9i)2). ^>n this occasion all L splvntlidti would. 
<\i necessity, have deposited their entire egg 

I oinplemciit in tlie 0I1C p"i)d because no othCJ 
auaialic habitats were available in the vicmily ol the 

Males o\' /-. \pU'ttJiJu called Irom e\poscil 
posiiiotts eiiher on the near vertical clilf \\\c^ or on 
natural ledpes upon the rocky surface. Ihe two 
recorded males were calling approximately 1.5 t mi\ 2 m. 
respectively, above the pond, Several othei 
individuals and ampleelaiU pairs of / tpktuh'tfri 
WCPC observed in similar positions Males of / 
aierttkui called from similar sites on Ihe lock face as 
well as from elevated positions in surrounding trees 
and on the ground near ihe pond. litorin nihclhi 
called from grouinUevel sites near ihe pond .aid 
l.wtnoth/utsltw urttitlits called whilst floalme in I lie 

Although we did uoi observe putt formal n»n 01 
egg-laying, pairs nf /„. ypleuditln a\\) remain hi ihe 
maimg embrace lor prolonged periods, at least tip to 
24 h. We observed several ampleelanl pairs silling in 
the open, or in cliff-face crevices, iluoughout 
daylight hours before the nigtol o{ chorus activity 
described above. Picsuniabb these bogs had m!uvd 
amplexus during the previous ni;jhl 

Pcsffipliatt of till! 

A wave form display and spectrogram ol the call 
o\' L ,\f>lcn(luia are shown til Fig. I. Tlikk I IteW the 
values of measured call attributes. The call is ;i Ion-. 
pulsed and apparently well -tuned call that is 
regularly repeated (maximum call rate observed was 
s6 calls min ' ). The call is broad band bin has a tonal 
quality because its relatively high pulse rate (which 
exceeds the temporal resolution of Ihe human 
auditory system and hence our ability to delect 
pulses ill this rale) is perceived as a complex tone. 
The call is characterized by a very slow rise in 
amplitude with a rapid cut-off after maximum 
intensity is reached The calls of holh individuals 
displayed a number ol frequency peaks in ihe pbWBl 
spectrum (Pig. 2. Table I ) wnh an mier-peak interval 

I'vui i I. Stttnuhirv of ( till t>ttrihnli\ t>J I Jlm'nt splendid.! n-roaltd ffl Iftdiicit Vuift's \nfiomil t'iitL f\nimtmn<i WrsUin 

VilUttS are bated en analysis of live calls ot two individual-.. I n^is were ealling on a sandsmne vlill lace between 1.5 and 
: in ,,how tl point, temperatures ul (he calling site were A, v = 25.5" C and A, A = 25.1 ' C Values for the firs! three attributes 
show ihe mean iind raiiL-e on paieiilhe.scM. 

( all 


No. of 



Rjic i p s i 



Other NoiuMe 


i\U i 

t en frl 

! hi & 



(64-7 -S3 ]> 

( "'-MOi 

■-. . ! 

1 218 

i 14.5 
I io" -s i tu.m 

i S8(J 


400.520,^(1 7X0 
020. 1040. 1 160, 
1440. 1500, u-so 

>:n.M0.7N). sso. 
1000. I2'»0. 1.520 
IP40. I760_ |XSo 



4 ^ 

3 - 


x 2- 


— ^ #>* 






Fig, I, Wave-form (upper) and audiospectrogrum (lower) of the call of l.itoria splcndida recorded in Hidden Valley, 
Kuniiniirra Western Australia. Wet-bulb air temperature al the calling site, 25. V C. Note that the ordinate of the wave- 
form display is not labelled because it depicts a relative linear scale in volts. The apparent vertical discontinuity at around 
600 ins on the audiospeelrogram is an artifact of the printing process. 

I. Mil I 2. Attributes af a repic.saituiiir call oj Litoria caerulea recorded in Hidden Vafh'y National Park, Kuninnaxi 
Western Australia, 
Temperatures at the calling site were A,.- 25.5' C and A w - 25.1' C. 



No. of 

Rate (p s 1 ) 




Other Notable 





\ 47 <. 


440, 580. 720. 
1 140. 1300. I5S0 




Fig. 2. Power spectra of the call Of Litt/ria \f>Utnlhiu across two ranges of (requeue) : - A kl I/, in slum ileiuils ul CO^fg) 
pi-iiks ami X KM/, to sJimv thai there is relatively litlle energy in the call above 4 kHz. 

ol approximately 120 Ha Many of these frequency 
components result from amplitude modulation of a 
harmonic series generated by ihe vocal cords 
because the interval between components is nearly 
identical to the pulse repetition tale. Because of 
resonating and filtering characteristics of Ihe sound- 
producing structures of the emitter, some of these 
Ircqucucy bands are emphasized, particularly those 
around 520 (7 Ihe fundamental frequency o\ the call), 
640,900, I2S0. 1420 and 1540 H/ (slight variations 
around modal values occur because frequency values 
are measured in 20 Hz steps on the digital 
sonagraph). These spectral modifications make it 
difficult lo determine confidently which components 
arc pail ol the harmonic scries and which are side 
bands arising from amplitude modulation. The 
fieqtlcncy band with most energy (dominant 
Ircquency) differed between the two recorded male-. 
(I2K0 for male #1 and 1400 for male H2). vvilh no 
w itliin-individuul variation (within the resolution Of 
this analysis) lound in die five calls of CttCll male thai 
were analysed. Although the call includes .spcchal 
energy peaks across a large frequency ranee (from 
around 5110 to 4ouo Hz), little energy is pttesem 
above 4 kll/ (Fig, 2». 

( 'lutms \irm IHtr 

Although «>nly llitec oilier species were calling in 
chorus with /., splt'iul'ultt, Ihe chorus structure is of 
particular interesl because the morphologically. 
behavit'urally and ecologically similar species. / 

< iH>mifti, was u conspicuous component fFig. 3), 
Attributes ol a representative call o\' syntopic L. 
tttt'ruli'ti are listed in Table 2 and a wave-form 



\ * J / \*£M ^ is** 3 


Fjgj l Audiivspectrogram of part of the chorus of fun 
species recorded in Hidden VatUy. Kuiuauirra Western 
Australia, hi this lecordiiiL.' Ufnritt SplpfldtdU tllA lite 
lOUite&l call, With hackgonnJ talk i»l / cuwulvft / 
tnhi iht ami (he Very slion c.ill ol! LimftfldyfKtXWx tinuttffb. 
KflcUivc temperatures fur culling males were: wei-hulb 
nit Lentpctalute (/., \f>ktufi(hi, I.. <,ua>U><, I . ntinHnK 

25.1 ' C and water icjnporulurc (/ . onintu\\ 26.3 C\ 



mlmfa fiJtfik 

00 200 




3 - 


x 2 


J 00 




Pig. 4. Waveform, audiospeclrogram and power spectrum of the call tjf Utoiia cacrulcn recorded in Hidden Valley 
Kunuiiurra Western Auslralia. Wet-bulb air temperature at the calling site. 25. I 1 C. Note: (i) die different temporal Scales 
on the wave-form and spectrograph^ displays; (iij that Ihe ordinate of the wave-form display is not labelled because it 
depicts a relative linear scale in volts. 

display, spectrogram and power spectrum arc shown 
in Fig. 4. The call has a similar dominant frequency 
and broad spread of peaks of energy as thai 61 L 
spiemlida, but ii is considerably shorter, has fewer 
pulses, a mtieh faster rise lime and less abrupt cut- 
off, as well as a higher call repetition rate ( 130 calls 
mm '). To the human ear. the call of /.. ctwndca has 
a harsher, less well-tuned quality. Nevertheless, 
because of the broad spectral overlap between these 
two large species and their use of similar calling 
positions there is the potential iov significant 
acoustic interference between ihem. 
Our observation.-* of calling and breeding in L 

splethlidu do not support the previous speculation 
that this species breeds only in the early wet season. 
Although our observations were made in mid- 
January, the wet season of 1996^97 was well 
established, two cyclones/rain depressions having 
already passed over the Kumiiuirru area in ihe 
preceding four weeks (pers. obsT During our visit, 
heavy afternoon rains fell on most days and this 
stimulus appeared to trigger calling and breeding in 
L, splendida. Successful reproduction also requires a 
continuous aquatic habitat for larvae to complete 
their development and it is likely that /.. splendida 
will successfully recruit new individuals to the 



population only during wet seasons that have 
sufficient regular rainfalls to maintain temporary 
ponds. Although we have no information on the 
ultimate fate of larvae from the breeding episode 
reported here, it is likely that the pond in which 
breeding took place remained in existence for much 
of this season, which was marked by substantial and 
regular rainfall. This outcome contrasts with the 
calling and possible breeding reported by C. Done 
from a nearby site in Hidden Valley. When this site 
was visited by one of us (GFW) a short time 
afterwards, no free water was present and larval 
developmenl would have been impossible. From this 

experience of the unpredictable rainfall patterns o\' 
this area, even in the "wet" season, it is possible that 
successful reproduction in L, splendida is a relatively 
uncommon event. 


We wish to thank D. Glanz and B. Watson for 
assistance in the field and J. Wright for analysing the 
calls. The study was funded by an Australian 
Research Council grant (S1971 1493). The work was 
undertaken under Department of Conservation and 
Land Management Licence No. SF002003. 


Tyler, M. J. [1992) "Encyclopedia of Australian Animals. 

Frogs" (Collins Angus & Robertson Publishers Pty Lid. 

(1994) "Australian Frogs, A Natural History" 

i, Reed Books Australia. Sydney). 
& Daviks, M. (1993) Family Hylidae pp. 58-63 

hi Glasby. C. J.. Ross, G. J. B. & Beesley. P. L. (Eds) 
"'Fauna of Australia. Vol. 2A Amphibia & Reptilia" 
(Australian Government Publishing Service. Canberra). 
. & Martin. A. A. (1977) A new 

species of large, green tree frog from northern Western 
Australia. Trans. R. Soc. S. Aust. 101. 133-138. 




VOL. 121, PART 4 



ByE. J. Barnett*, N. Harvey*, A. P. BELPERiot & 7?. P. Bouraian$ 


Barnett, E. J., Harvey, N., Belperio, A. P. & Bouiman, R. P. (1997) Sea-Level 
indicators from a Holocene, tide-dominated coastal succession, Port Pirie, South 
Australia. Trans. R. Soc. S. Aust. 121(4), 125-135, 28 November, 1997. 
Peritidal Holocene sediments at Port Pirie in the northern Spencer Gulf of South 
Australia contain several indicators of sea-level change over the last 7,000 years BP. 
The elevations of present subtidal, intertidal and supratidal environments and 
corresponding sediment facies were surveyed in order to establish critical boundaries 
relative to the tidal spectrum. The subtidal Posidonia facies occurs at or below mean 
low water spring (MLWS) tide; intertidal sandflat, mangrove and samphire facies 
occur over specific intervals between MLWS tide and mean high water spring 
(MHWS) tide. Each facies is clearly identifiable in the subsurface, with intertidal 
sandflat facies particularly characterised by in situ articulated bivalves Anapella 
cycladae and Katelysia scalarina or K. peromi. A combination of several palaeosea- 
level indicators from different tidal facies best defines local sea-level change over the 
millennial timescale. 

Key Words: Holocene sea-level indicators, tidal zonation, prograding coastal 
sequence, facies boundaries. 

ftffw«fvmw "/ ifo t<>>Y«! $<H.'ig(\ >>i v \um (1W7), 121U), 125-1 ij 

si; all vkl indicators from a holocene, tide-dominated coastal 


bv I:. I. B\r\ktt\ N. IIarvly A. P. Rii n.Rio & R. P. BoUftMAN* 


Hauni 1 1, K, ,1, , II \t!\iv, N., Uu i'i.kio, A. P. & Bookman. R, I' I I 1 ' 1 )?) Sou Level LndieaLws Irooi a MnhtLvnc, 
tide dominated I'oastul succession. Purl Pine. Smith Aiisii.ih.i ftg/M. A'. Sor, S, Aust. 121(4). 125-1*5. 2K 
November. 1947. 

I-Vriihlj|l llolo L vnr sediments :ii Putt Pioe in iltc* northern Spriu of (rull ol South Australia contain several 
indicator; Of sea level ehuniic OVW Ihc last 7,1)01) yen < HP. Tin; eK-v^iions ol present HtiblldaJ, mterlidjil ,nld 
Miprjinliil .nvitunineiUs .mil corresponding sediment facies wwe surveyed in order a> establish crrilivfll 
boundaries relative to the tiUul RpOflmnu The sublidal ft>\ith>tw< laeies occurs ( il or below mean [pvi valL'r Spring 
iMI.WSi tide: laiurlulal saildllut. muilCTOVe and samphire lacies occur owi -pceilic tnleivuls hcl vveco Ml WS 
lidc :md iiifiin hieli \\iilci ^ | m i u : _- iMHYVSi liuV Riled lacies ta clearly identifiable in the subsurface, with 
inkrhdal sundHal licics paihculatly characterised by yj t/fri ;ir(iu nlnlL-tl bivalve:-. Afiflfteffa < ytludm Mfd 
Krttv(ystti u ulunnu or A pemtifo A combination of several palaeosca level indicators from dillcrciil lidal lacies 
best deline-, local sea-level change over the millennial limcscak- 

Ki V. WoMos: llolocuic sitfM lilVtfl indicators. Lidal /omiiiou plo^raJm^ co:is(a| sequence, lacies bouiaLiocv 


Tide-dominated coastlines commonly gftnPV»le 
prograding loasial sequences With excellent 

preservation of intcrtfdM and shallow suhtidul 

scdmicnlaiy lacies (Belperio rl al. |988i Jo Boor et 
w/. I4SS; PJctchcr *•/ ui. I W), Such sequences can 
reveal [ligh-rCKoIutkm records tM' past sedimentation 
often containing a variely ol' palaoosca-lcvcl 
indicators l Icrwiudl 1989). A thorough 
nn Jorstuiuli rtij ol Ihc rclaliouships ol present 
indicators and sea level, hi inundation level, ifi 
required if correct inioiprclalionc of past relative \y-\i 
levels arc di he .ichieved. Wilb critical appraisal ol 
Ihc present day distribulion Of intcilidal lacies. flora 
and launa. paloeosea level IunIoia from subsurface 
sonography can be more confident I) interpreted, 

The northern Spencer (lull, Souih Australia (Pig: 
I), provides an excellent example ol a wide, 
progradimi sequence in a niesoltdal 
environment with an identifiable /nnutuni nf 
I loloceue const a I deposdimiid ciiviiiHiineiUs A 
number <\l* coaxial studies lias been conducted 
previously in this area (Firman 1°<Vv. lUime 1982; 
Hume A Cukvell I9>,2; Belperio el ill. lOSda.b. 
|<>KK;<;oMiu </,</, r%4. P)NK; Norrisho/o/, 1986), 

Mnw Uwdtuiti ( '> wri lai i ifvimattttmul .smaiL'-. t riivifi iiij 

(.1 AUi'IhTiL Vit i ■■(>!)■', 

| oiintilv Mir--- im.j i i. .■■. |<e«TyjievV' -Soull* AiwwJhi PltBm 

im I ..-.pauihI N Ana '-<■><:'■ I IUVMtlj MilH'|:ilir ClUJll ''.' 

oimt a. tin* Si hill.iiloit S. A«*t. WViT, 

SfllMiOol taiMiciiinonial an-l KetB lUlMI Vhi*Wprtm-Hl T I wn\\$ ' v ' 

i n in. m .iini r^MviTonrottiti i i(v»i«rti rf St^oili taritLi i 

'A'.MH/i.a. Kd Ilk I evfilH s \i ,1 tW3 

In particular, Burne ( 1982) idemilied sc\cral 
unpoitant palac">ca- level Indicators t'loni beach 
tidies, ihc Lop of the sublidal Pt/xMowio -•caiirass 
lacies and base of the mteriidal ^andllai i'acics. }UtU 
Belperio e/ o/. I IQH4b) demonstrated the presence ol 
a Well-defined boundary between Posj(hflttf l&lgtU'tt 
and mlertidal sandllal Tactcs. Related sliatiei'aphic 
studies in nearby (mil Si Vincent include those by 
Cann $ Goslin MQ85). Belperio el al. ( l*>H6. 1 ( >SS(. 
mk\ Belperio ( 199*. 1995) At Port Adelaide in Gull 
Si Vincent. Belperio (1993) confirmed lhai the 
boundary between the inlerlidal sandllal and 
nningfUVC lacies WHS a icbable pat.ieosea-level 
indiealor. Prom all Ihese siudies. ii is apparent that 
dlGTc UVC lOCal and reiiional dillerences in die 
reliability ami distribution ol various sea-level 
indicators. This paper provides a critical appraisal ol 
thedillercnl palaeosea le\el indicatots in a ntesotidal 
tMp nonmciil 

The wide proeradm^ sedimentary sequence ol the 
ooithi-rn Spencer Gulf region, which lorms the apex 
ol' a laree relatively shallow tmeisc (t»t uc^ativci 
c-au.iry is .i direeL response to the modern coastal 
envirouiucni Warm temperatures ;in<l low rainlall in 
the region promote high rales o! evaporation and 

5s ilivs which are ol'ien higher ihan Jiveraee lor 

seawalcr. in excess ol Mfjfa and as much as dX',, 
(Bye IMSI; Nuncs & l.ennon I l >S()|. Seawakt 
tcni|-ieraiiiivs for the nortliern yull' vary typically 
between I 2 and "!4 " Ci Amirs *V Lcnnon l L 'SOi I lit 
titles are mostly semi dun tial. with Spring and m-ap 
tidal ranges at Port Pirie ol 1& in and 0.4 m 
ivspeeli\ely_ \)\\c to the leneLh of the gulf and 
telativclv --low mean seadevel OSCllkliOJlS. Wind 
.-■tress can lurlhci luciease die ttStroUOhltCul lldv 



Fig. 1. Location diagram of the study area in the upper Spencer Gulf. South Australia, showing the Port Pirie coastal zone, 
sampling sites marked by infilled circles, the tide gauge and Broken Hill Associated Smelters (BHAS). 



significantly. Iti Spencer Gull and much of ihc 
southern coast ol Australia (Nuncs & l.cnnon I^Kfil, 
ihCTC is LI iiU'gc -.pring-neap tidal modulation due to 
the nearly equal lunar and solar serriidim nai 
».uii'.niiii.-iiis i .unship a dodge luic oino every 
loitnighl when lihlc lidal variation occurs- Schluicr 

'7 tit, il*H*5i hrtvc postulated thw this FiftltiP 

amplitude ol I he major semi diurnal lidal 
consliUienls giVBS rise to parlicular shallow water 
lidal interactions m the tappet glllK, which promote 
samphire and mangrove colonisation. 

CjivOti Ihc programing, tidal sequence and (lie close 
proviuuly of a litle gauge with reliable long-term 
Miouls at Port ftrio ilus study was undertaken to 
determine the elevations of coastal sedimentary 
foncs mtativc 10 modern sea 1* vol and tQ Identify 
appropriate modem sediments as analogues ol 
subsurface Holocene sedimentary laeies. This 
appioach provides an opportunity lo identify the 
mosl reliable palacosca- level indicators in a 
mesotidul enviioiimciu. ami I" develop a 
methodology lor subsequent studies ol relative 
laud/sea movements at sites where historic tidal data 
exist. The study is uiucjiie in thai il highlights a 
number ol' sedimentary facie.s surveyed relative to 
modem Sfrfl level, mid identifies critical cqiliv. i 
indicators in the geological record. It does noi 
depend on one uidiealoi m isolation, hill uses a sinte 
ol' subtidal lo supralidal indicators 10 idenbiv sea 
level i. hangc. 


Ii was necessary lo survey ihc modern coastal 
eii\ iioitiueiits in detail to establish elevation 
differences ol' the tidal /tines relative lo local lidal 
datum. Tins was done using both lasei and automatic 
spirit levelling utstiunKiiis The first had an efltjl * i 
less than I em OVtS 'ill) m. and mcasiiremrnis wcct 
kepi lo within 400 m 'i.e. ± 0,02 nil. L vmy stpjnl 
levelling. Ihe distance between each reading was lev, 
than 1 0() m, which generally kept vertical 
measuremeiii errors |0 less (halt ± O.Ol m I I 
in, ,l sites within the sandllai, maiiiUnc and 
s.iiupimr ,*.im-, W£t£ suiveycd lo thud -oidci 
AuMiahan Height Datum (AIlO) benchmarks. The 
1 tin, eying was conducted mostly in Ihc Porl Pirie 
uwstal vicinity a* well as Uuthcr to the northeast 
wuhii! the Tl-Iowic Beach eoastal legion, in order lo 
ac^es.-. all "I the modem day tidal settings ibig. 1 ,). 
I lie pa'svnl-day levels ol die scagras.s and sandllar 
zones .a Port Pirie were measured from water levels 
i> ! lDVC U> the hail Pine tide gauge. At Poll Pine, 
/cio udal datum (TO) is correlated with the lowest 
asironomieal tide and rehaed lo MIL) using a 
eoireuion ol I W in (surveyed 1 7.0^. I *>X ^ South 
Air.iiaha Potts Vuhoniy I. Sea- level indicator. 

i lie hid my seagrass. shell and mangrove remains 
wilhm and ai the UTp tff each lidal /one were 
identified and recorded lor later comparison with 
subsurface equivalents. 

In order lo establish and sample lite subsurface 
stratigraphy, u total ol" thirty- live sites was selected 
within the broad coastal flats of I'on Pirie dig- I ). A 
vibrocorer was used to obtain cores 7? imn m 
diameter and up lo4 rn in depth, All ol (he vibrocorcs 
were corrected lot sediment compaction by 
recording penetration depih versus core reeoverv 
length and applvme <\ I OiTCCfiOfl ladoi to the 
IhiekncNs of the scthmenls. Coring peripheral lo and 
within mangrove woodlands was uuned "tit using a 
peal auger. A baek-hoc was used to cxcavalc 
sediments in the suptatidal region lUmj dii- 
method, no correction lor •.ainplmg compaction wa> 
neeessurN. Surface and subsurface elevations in laud 
based cores and c\ea\ aiions were >airveycd to AH1). 
Vtjirine-hased eeuvs were surveyed lo TO and Wkefl 
wilhm a lew kiloinclics ol the lidc gauge lo reduce 
the elfeclsol tidal lag and meteorological conditions. 
Samples were taken back lo the laboratory, where the 
sediments weie lltggpcl with partieular attcnlitm 
being given to the elevations of laeics boundaries and 
tlic (ireseuec of -.ea-lcvcl iudicut<trs. 

\hiilei u depusiiunul lidal eiiv inuiiiienls in 
the Vt\ri Pirie nrc?» 

The toaslal cnvimnmcm a<ljaeent lo P<ai Pirie 
i Pip, 2 1 is a tidally-dominaied lowland. Subtidal. 
interlitlal and Mipraiidal /ones wciv distinguished by 
the extent of marine influence en- exposing md bv 
their vegetation assemblages and sedimentary laeics. 
Broad, shallow subtidal seagrass meadows pass 
lateral I > shoreward iulo intcnidal saudltuiN, 
mangrove woodlands, sjinphiie-algal matshes anil 
siipratidal evaporite llai i iwiroi.menis. this 
association \4 penlidal environnienls and Iheit 
\egetntion /ones has. to ii hirge eMeni. gtmlmiloj the 
successive developmeni <\i the coastal plain around 
Toil I'iiiL* A •icliein.iiic sumiiiaiN of (he lidal /oiu",. 
associated vegetaiioii ami (heir ivLuionsltips (0 
elevation oi | daluHi levels is 'joen m t luuie .v 

The Mthlit/ul zi»w 

In the broad. sluiHow-uuti lite eiiviioiuueni 
northwest i»f Port Puic township, e.r 1:1 is meadows 
composed largely ol ttyitkuttu iwsnuli.s guAv from 
around mean loft water spring (MlAVSt title level 
i./cro I'lJ) lo III m below I U < I ig.s _V 4» t?>wl0}iti 
ilittitf'tllh eau only survoe limited penods t-l 
emeivenee so thai, ul its upper growth lirmh H In 
generally pair by and reslrieted to below 0.23 in TD. 
Al dcpihs below 4 in 11V PtfwtfrtiltN tlTRlHSa 
dv. inmates the iffa^J • '--seniblage, I'osiiU'nut leal 



sheaths and rhizomes arc resistant to decomposition. 
and seagrass fibres become incorporated iota and 
hind sediments. A highly distinctive sediment facies 
results, producing calcareous mud and sand bound 
by masses of pale cellulose fibre. High sediment 
production and the binding and baffling action of 
seagrass contribute to the rapid accumulation ol 
sediments in this environment. 

The intertidu} zone 

The region extending from MIAVS tide to mean 
high water spring (MHWS) tide is defined as the 
inlerlidal /one (Fig. 3). This /one is characterised by 
periodic emergence and inundation during neap to 
spring high tides. At their most seaward boundary, 
broad sandflats have developed upon which some 

seagTasses can grow above MLWS. Hosithmia 
uttsimlis struggles to survive and is replaced by 
Zostent mUeiteri Further shoreward, bare sandflats 
are dominant. These sandflats are host to numerous 
epibenlhie organisms including the inlet tidal 
molluscs Batillana sp_, Veneridae sp . Tellina sp,. 
Chtftcnlus sp.. Anapella cyrlackw and Katelysiti 
scaUirina or K. peronii and foraminifera that live on 
and beueath the sandllal surface. These organisms 
occasionally accumulate in shallow tidal channels. 

Intertidal sandflats are replaced by mangrove 
woodlands above I 32 m TD. Only one species i)( 
mangrove, Avkeimia marina var. resiiiijeru. has been 
recorded growing in South Australia (Butler et al. 
1977; Gostin et aL 1984; Cwm & Gostin 1985) (Fig 
5}. Around Port Pirie. mangroves have fanned dense 

Fig. 2. Aerial photograph of the Port Pine coastal /one. The subtidal and intertidal sandflats have been colonised b\ 
seagrusses (seagrass meadows), Further inshore, mangroves form dense woodlands along the coastal margin and grow 
along dendritic lidal channels. Samphire communities occur more landward In the intertidal to supratidal zone. In the 
supralidal /one, the vegetation cover is sparse in between broad expanses of saltpans. The photograph covers an area 
approximately 10 km x 10 km. The aerial photograph as been reproduced with the permission of the Department &( 
Natural Resources. South Australia. Map land, telephone (OS) 8226 4M46. 



communities within clearly defined tidal limits. 
While the lower limit of mangroves is close to mean 
sea level ( 1 .15 m TD at Port Pirie), their actual lower 
limit, 1.32 m TD at Port Pirie, can be significantly 
different. Their distribution is controlled 
fundamentally by their root system since the 
vertically protruding pneumatophores require both 
exposure to air and Hushing of precipitated salts 
(Chapman 1 975 ). A mangrove-algal association 
occurs at seaward levels of mangrove growth and 
along exposed tidal channels. Cyanobaeterial mats 
also extend on to wide sandflals and into samphire 
areas in intertidal and supratidal zones. Numerous 
other organisms are associated with mangrove 

woodlands, including the small mud crab, Helice 
haswellkuitis, which burrows into the substrate and 
promotes oxidation of the upper sediments. 
Gastropods, bivalves, polyehaetes, decapods and 
other crustaceans, foraminifera and diatoms also 
occupy this zone. 

Landward of the mangroves at elevations above 
2.6 m TD, are broad, flat, gently undulating plains 
upon which samphire-algal communities grow (Fig. 
6). Stircocornia quinquejlora, Sclewstegia 
arbusctila, Halosarcia haloawmo'ules and Suaeda 
cutstralis are the main samphire communities present 
in the Port Pirie environment, followed by minor 
occurrences of Maireana opposliifotia and 

,- 2 

- -2 



Fig. 3. Present tidal and vegetation zonation along core transect relative to the Port Pirie tidal datum (TD) and Australian 
Height Datum ( AHD). MLWS - mean low water spring tide; MSL - mean sea level; MH WS - mean high wuler spring tide 

Fig-4- Shallow!}' submerged seagrass meadow of Posidonia 
aaslraiis in the subtidal /one photographed during low 
lide. Width of field approximately 3 m. 

h'ig. 5. Landward intertidal mangrove margin with intertidal 
samphire communities. Only one species of mangrove. 
Avice/tnia marina var. resinifera, grows in this southern 
temperate latitude. Dieback of mature trees along the 
landward margin ean be observed, which generally 
indicates marine regression. The dead mangrove in left 
centre of the photograph is approximately 1.2 m tall. 

I \d L..I, UAKM.I I N HAKVI ■> A 

HctlGXftrctH nulim toward thq supralidal margin. 

lilr fijpWfjdtrf "IK 

Above MHWN licit clcvalion (3,2 m TD). *he 
supralidal /one iF\£ 3) is flooded on only the lew 
occasions when either high or king titles combine 
witlt Htnrm Surge activity, predominantly limn the 
foulhwcsi, or Online and shortly after extended 
periods of rainfall, Consequently* this /one is 
doiilinaled hy evaporative pioeesses and their 
UtfSOcitiUid sedimenls In some ponded areas, algal 
itKii-, ;iu: vm-II i--,i,ihli-.hed and form cvatiobacleria 
Hals. Although Ihis region eonsisl.s mainly ol hare, 
poorly draining saline mm.1 gypsiferou^ Hats, some 

samphires and sallbttshcs survive (Fig. 7). Ol Ihese, 
thsiosan in tm It u ■tivmt>idt'\ i Afriph \ pdlmfosn. 
HtifasdltUi tn<h\-« WKl AtriftleA iiMcufiu are moM 
abundant, VVillnn this /one, variations in elevation 
Lire ueaied by aeoliaii deflation and formation ol 
^ypsil'erous dunes heivveen remnant channel',. 

I |j| ft litU'iiul;il Minipltire /one includm; i nffl/fl 

t/MMfueftonti $t'tcrp\tef*t& utlntMnln. Huhxattin 
hulttt m ttwnit ■■-, mill Sitat'iki ttn&tafttv, The ruaFiiume in 
I he ni'lil Ir-K-'iinind is ;ip|nmuii:iU'lv > in Ineh and the 
samphire hushes ;ire up to 3U cm in height. 

I il'. 7 Sii|'i,iihlal s;itiipliiK' /one it il tailing ocujsmiiiil 
ff'ih>\-in'iii lf(ilintirtnt>iJi \, A (ri/fh \\ l>tihtift>><», 

Htlltiwnriu ititihti and A triplex wxiattui unJ hiae 
rxpansOJol fclltfltti, Ihi -.imphiivs]". i |'l't 'Hum 

hi iwiybt 

P Mil ,1'EiOO* R, 1*. BOURMAN 

Lunettes have also formed on ihe leeward margins of 
sahkha flats Of salt lakes, 

The distribution ol' coastal environments around 
Port Pirie is shown in Figure 8. Si\ distinctive udal 
/ones transuding Ihe coast have been identified, 
From seaward to landward, these are; i) suhlidul 
seuyruss meadows (no* shown in figure ). ii> low 
inlertidul bare or /m/r/v/ covered sandllats. in) 
uileriidal mangrove woodlands, iv) lueh mterltdal 
samphire aleal marshes, vj siipratidal evaporative 
Hals, and m) supralidal and cxltalidal clay and 
e\p.-.eous dunes and lunettes Aerial photographic 
interpretation of Ihe mangrove WOOdlantl reveals thai 
only ininpr ehunec ill its disirihut ion is apparent lor 
the lasl 40 years or so ( I057-IW3). Manproves have 
prograded seaward ittiu inleflidal sca^tass/sandllai 
areas on the northwest peninsula o|" the PArl Pine 
River, heivveen Fil'Sl and Second Creek and aloinj llie 
margins d\' ihe l J ori Pirie River itself, this is in 
I onlr.iM to rapid seaward uiaueowc colonisation lhat 
has oeeurreil al Port Gawler ((.'aim & Ciostin |*W5) 
and landward colonisation in the Pori Adelaide 
region (Burton 19X2; lielpeno I9B), 

l'\idenee nf deposirinnal tidal sediments 
in the subsurface 

Much ol ihe sedimeniaiv stratigraphy al Port Pint 
represents aggradation and pmerudaiion ol 
sedimenls in periiidal environments since ihe near 
siabilisaiion ami slight (all in SCO level In mi 7.00(1 
yeais HP lo preseni ilielperio 1*995 )< Holocene 
sedimenls .\n\\ Pleistocene alluvial sediments of the 
Pooraka lormalion underlie moa i)\' ihe area, 
forming an undulating boundary with ihe oveilviii- 
hdal sequence, In some places, the upper sections ol 
Ihe Pooraka Formation show eviilence of being 
altered oi gleyed bj marine pttrewuwrs. The tmsiai 
sediments record an upward change m srdiniem 
facies that corresponds with rhe hiiL-ml t hmtgc m ili« 
tidal /ones 

The siihlidal POSklotiiiH faclCS >s the mosl e.MensiM' 
Holocene tidal facies m the legion, Il consists o\ 
mosl I y erey, poorly sorted terrigenous and 
calcareous sandy mud. w-Vth numerous [IKre.s of 

Pvsidauiu ntt.Mrn/i\ and Iraimienlary molluscs (t*.g 
SfNMila sp,. I'ltwiitiiu Hit sp . ( tiiiiiifii "idn\ <p . 

l.h'unia sp. and ttnttlh/ritt sp.) .nnl fora ifcrd Its 

ihick ncss varies Iron) ^iiealer llian 4 m in the piesenl 
>ubtidal /one but Ihi ns inland underlying interiidal 
and supralidal sedimenls lo between and 2 m 
depeniijng on undulations in ihe surface ol the 
ttndci'lying Pooraka bomialioii. The landward exieul 
ol (his facies indicates that much of the prcscm 
coastal environmeni was a shallow maiine 
environment durinc, the e.n l\ to mid Hoto< O^C 
I he intertidal sandllal facies is a ercy to h:'hi jjrey, 



pimrlv sorted, terrigenous arid ealeareous shelly 
muddy sand. It pd3Ul> extensively inland beneadi 
niuehol'the *uidv area aavme developed in response 
lo upwind sIi^mImuj of | he siihfidal xedimenlarv 
environment. In mosl «l the Foil Pine region, ihe 
uiiertidal sandtlal I'neies i* overlain by samphire 
laeies. This is in eonlrasl In ihe presenl-duy Lidtil 
/.oiialioii Wflcrc ." transition Ironi sandflai to 
mangrove woodland generally oeeiu v 

The inieriidal mant/rove laeies eonsisls ol brown ov 
bluish ^'lev. mostly nonealeaieous sediments wiih 
f\ rments ol' roois. sheaths and iihies. It is lairelv 
reslneled lo Ihe present day distribution Ol IWfflgtW C 
woodlands, i.e proeradaiiomd development and 
pie-M'i viitmn ol stratti have been limited and 
\\<i<iii)Hii nuniUtt v;ir. tcswijct<i woodlands appeal 
I" have developed 111 relatively letem limes Wlvie ii 
to undeveloped, modem mangrove roots penetrate 
into the uiideiivni£ laeies, 

The StCdintCin laeies ol the samphnv Hal hums u 
ihm u-ueer over i \irn--.ive areas ol' sand flat faeies of 
Ihe eoasial plain. It eon-isis ttf pale brown to Light 

^re>. often mottled ealeaieous and terrigenous Ja> 
rich muds with oeea>ional small gaslropods. bisahe?- 
and loraiumilern. Small platfl fibres ami thin uibulat 
mots are apparent in some regions but absent in 
other-.. depending 011 wheiher plant mailer vv^ 
originally present and/or preserved. ( lypsum eouteiH 
is variable, iliie largely to elevation and evaporalion 
hi-aoiy, with eypsaiemle dune sediments preset ved 
it) i ho highest elevations of ihe supraiidal /one. Thcie 
is lillle distmetion between inteitidal and SlipnUiiill 
samphire sediment laeies. mk\ the two are eonsidered 
10 lorm a single unit. While partieulur samphire 
speeies ean be identified dWWtng m eiiher the 
inlertidal or supialidal /oik-. Ifl lb* iUbSHl tate 
samphire loollels and remains cannot he identified to 
speeies level. 

In addition to the sediment laeies above, several 
uueioeusuoi)iih:ii|s Ol aihtaeies oo m i>. ilu- rejiiun 
that have conlempniaiy analogues. In parlieular. 
pockets of vyaiiotxieieii.'i ladies are evident 
ihiouL'houi the mtorlidal 111 supiatidal /ones. 

vvheivvci cyanabftcicftal intro* »w present in the 

ST KIL DA FORMATION (UndiHf>r en iiated 
Hoftusene marine and coastal marine sediments; 

Suprstidal ana extratldal ctav and 
gypsum dunos and lunettes' 

\>. Mangrove woodland. 
_ .i.'jXS Organic. tero'rjRnous mud 

-■X-; '-:-'■] Supratidat flats. Gypseous day: 

Stranded bearn ridges aflfl eoa&lal dunes 
Shello/lt, shelly &and end lino sand 

Samphire-algal marsh 

/ n.nn.Hif anc terrigenous muds 

Bare Dr 2ftste£3-pa[anfeed low tntertidal sand 
or mud Hat Mix£d shell and quartz muddy &sric 

Organic/sh<eliy sand/mud of mangrove 
woodland and samphlre-algal marsh 

ABC Range Ouartnte 

Fiji H ( (fend -■■ itli ■, i> nil .ii tfu l*'*" Ptrii 

Kvvnr.i-- N'hiiIi r\u4uliu. 

HI 1 1 ; I ion 1 . Ilk- South Ausliallnii t.enli -yy UtlUlfoaHC MllHf iml I n > 


intertidal or supralidal /one there arc active sites "I 
sediment aggradation* In flic inlcrlidal /one. storm 
ridge facies, or cheniers when developed over muddy 
sediments, have been formed during periods of 
combined high or king tides and storm events. 
Radges ate generally aligned parallel lo the shoreline. 
Only one storm ridge is preserved in the western Pori 
Pjrje area, although several others oeeur lo the easi. 
The nonherh orientation of the coastline gencr.illv 
protects ihe area from dominant southeasterly sioim- 
ndgc forming events. 

I'uiat'osta-levtl indicators 

The special signiheancc ol the northern Spencer 
dull is that the peritidal coastal succession cOfttalUS 
a uell-piescivcd record ol palaeosea-levcl change 
The sediments include various palaeosea- level 
indicators that have been used, vvilh appropriaic 
elevation tftuti, to iccon^troci palacosca levels. 
VIllntLrjIi present da> tidal environments and 
equivalent sediment fiicies may range OVCJ 
sigmncant vcttjcal elevations, ihe contaei between 
each sediment laeie.s is generally lltOfC asp'ii I d 
Nuhsuilace fades contacts can provide relatiulv 
precise estimates ol" palaeose.i levels given accurate 

surveying ot Ihe verlieal exient ol present 
sedimentary facies and their contacts. Once the 
elevation range of a particular sedimentary coniacl is 
known, a height correction for lhal contact can be 
made relative lo pieseni sea level. This establishes 
Ihe elevation at ihe lime of deposition and indieates 
whether sea level has subsequently risen or fallen. 

We have established that, in the Port Pirie urea, the 
boundary between Ptisidptiid facies and overlying 
shelly intertidal sandflat facies provides a palaensea- 
level datum corresponding to an upper limit of 0.25 
± it. 25 in TD (Fig. 9). Consequently, the subsurface 
occurrence of distinctive, massed. Ilbious PvxtdontU 
facies in land-hased sedimenls al elevations hiehci 
lhan 25 ± 0.25 m TD implies lhal t chili vv sea level 
was previously higher than at prcsenl. 

In a similar fashion, the intertidal sandflat facies 
generully occurs between 0.25 and 2 1 m IT) relative 
lo prcsenl day s.m |cVel lln\ve\er. a more precise 
palacosea-level estimate is provided by the sharp 
contact between sandflat facies and 
mvrlyiiie mangrove facies lhal equates to 1.32 ±0.2 m 
it) (Fig. s>). Al Con Pirie, ihe maneiove faeies 
mostly OCCUrs ducclly toitalh the present mangro\e 
woodland, and eotilldence in using us contact with 
the top of die sandflat lacies is greiilesi where 






flat facies 








D-E contact 

C-D contact 
B-D contact 

B-C contact 


(Tidal Datum m) 

3 010.4 

2.6 ±0.4 

2.2 + 0.5 

1.32 ±02 

General Description 
of sediment contacts 

Change from calcareous clay to gypseouc, 
clay-pellet, structureless sediment 

Change Irom organic, rooted, peaty clav 
to cream calcareous clay, laminated to 
weakly rooted 

Change from coarse shell sand or 
coqulna to cream, calcareous clay 

Sharp change from coarse shell sand m 
coqulna to organic, rooted, peaty clay 

Sharp change 1rom poorly sorted, fibrous. 
A-B contact 0.25 ± 0.25 shelly sand and mud to cleaner, better 

sorted shelly sand 

1 1$. m Palacosea-tevcl mtL-iia i\x ii»e iwi hm- 1 oat*iuJ t'i ""■'" 

massed articulated valves gf Anaitelfu rultit/tir tirul 
hftt* Ivsui \V(jli}tititi bl t\. fHromi ate [W^SUIIL, 

indicating W tffyi poM-norlt'OI piescrvMioii WhClV 

the mangrove luties is ahsenl. I he coniacl between 
the saudHui and ^innhire fades is also slurp, 
although the ptcscnt-day boundary heivvecn l|sc 
uilci tidal s.Hulll; ( i aid samphire /ones is DO* *'tfN 
delincd in (he immediate vicinity of IVil PifTC The 
uppei limit if [he syncHW lacies with samphire 
lacies occurs around 2 2±.0.5 in TD (Fig. u >. 

WhitC tflC present Port Pjritf tidal /oue> evhiho a 
transition from mangrove woodlands 10 sainphtu- 
mardies al I'.n t 0,4 ni Tu, this is not comne'iib, 
observed in lilt subsurface sediments due to the lick 
ul piuL'M'lMu.tuil ji \ e'opmcni fcrf 'his slruiiarapfnc 
hot i /on. Consequently. I he level a! which the 
sMtnplritV IWPJWi occurs in Ihe suhsuifaee provides 
only an approximate estimate ol piUatto&ua-ltfVCl 
relative to LM present elevation ranee tif 2,2 l£> eUl in 
TO, Although different •.auiphitc spates ate vlosvly 
lelaicd to small elrvuoon chanec-. these are uol 
ohscived at the macro-level in Hie snbsurface. 

1 liscf issi^m 

Several filwtoTfi must hfi addic.sscd y hfttl 
inierpreiinj.* the evidence lor palaeosca level change 
horn ptOgftldsng perilii.lal sfftJUfctictJ-N In particular 
Ihe relationship of each indicator t* t sea level at the 
lime ol its formalion must b^ established. Al Port 
Pine, the subsurface presence ol /// kilU fibrous 
lenuiins of ihe scagiass Posiihntiit ititsfrah* indicates 
thai sea level w;.w above this rtfie al ihe time ol 
dcposiiion. Ihe transition beKvccii t'csuloinu Uac- 
and o\crlying intcrlulul sandfluL lacies is a more 
powciful indicator of palaeosca level, corresponding 
to 0.25 ± 0.25 m present day II.). Similarly, 
mangrove-, ^mw wilhm a fairly 'mmd mlertidal 
range, bill Iheii contact wilh Ihe inlcrlidal sandllal 
lacies provides adaimn of 1.32 -Hi. 2 m. Mangroves 
have previously been ciled as one oi' the more 
reliable lived, m \ttf\ palaeosea-Ievcl indicators 
[fft'rpfej & Thou. IVRit Thorn & Roy |9&J>. Koi 
noilhem Spencer Gull, Burne (l'JN2) reported a 
range in the elevation ol seaward mangrove 
eolonisaliou from 15 <o 2.U ni I'D (-0.4 lo 1,0 m 
AUD) and as previoicdv mentioned we record a 
lovvei level ol maiu'rove eolonisaliou al \.JL1.±.\S..2 hi 
TD. nearly; Iho level ol seaward colonisation or 
mangroves Uej»end^ i»rimari|v on local C0tt5tAl 
dynamics or caslal orienlalion, -and will ijcenr at .i 
variety ol' elevali<-)ns relative to the lidal sped mm 
(Allen ! U( )5i 1 hereioiv, it follows thai ihe height off 
ihe eontacl between sandllal and mangrove lacies 
will also vary. It is apparent from the difference-, in 
elevation ih;ii llie Use of nuiugioves as palaensea 
level liidicaloi-s can only be ipplicd lucally. wliciv 

MA li'Vn. IMJll "MiikS AT P(HO IMkll I" 

pre.seni-day clevaticais of maniinne -eaward lt* *v\ th 
are welt defined Hven in this ease iIil 
palacoenvironmenl may have differed from ihe 
Un-deiti environment, producing different lidal 
ranges and mangrove distnhulion>. 

Illfi boundary beiwecn •-aiidfl-U ami cither 
mangiove oi samphire lacies has the potential to 
define palaeo.scii -level, paiiicularlv since its eoniact 
in I he suhsuiTucc issharo. HOvVuv^r, the c<nuiiiiliiin> 
ai Prfrl I'irie is that while ihe pn^enl-dav /onalion 
FntiD sandllal to manerovc wondland is cMcnsive. 
\\S$ nnnsition r- i>,»i easdv observed in Ihe 
suhsurfacc t ; ur'|ienii'>ie. v^ lute the po-seiHil;i> 
i/anstiion between fttncMlkl aiul viniphire /.ones is nol 
well icprcNcnied at Port Pirie. this contact in 
tlltoUlfflOC SC^fJlV W : widespreatl Near Port Pirie. 
the pre-.cnt elevation ol Ihe saudl'lat/sampliue 
hounilatv is 1.2 ± 0.5 m TP, To the northeast al 
Tclowie t?eaeb. this boundary occius at 2.5 1 0,3 m 
TD. a slightly higher elevalion (hnn Um Port Pine 
l»nssibly due to local gcomoiphu bichirs , t >i»i 
sedimentary processes in Ihe lee of Weeroona t.vlaml 
tl-'iy. I), the elevalion o\' the top of the sandllal 
varies depending on whcthei it is succeeded by 
mangroves oi samphire, line-. aMh*>nch ilu- omiau 
can be used as a dclcrrumanl of sea-level ehanee. 
iliac \\ n wide \\\u$ic in its elevation. This problem 
nmy be minimised by careful held surveying ol the 

local region, 

Beach ridges and Ihe lop ol /V.v/Wo/i/c sea.eMv-, 
deposits are relatively good indicators of palncosea 
levels. However as wiih mangroves, beach ridge 
elevalion daia cunnol he used on a regional basis since 
the elevations to which such ridges an- const! ueicd are 
highly dependent on local wave regimes. In regard to 
seagiass as u sea-level indicator. Piwdtttfib >m.\imln 
presently grows lo 0.25 m TD (l.oKm AllDtal Port 
Pirie-, hitl elsewhere in ni>rlhern Spencer Gull, an 

elevation i^' 0,1 in ID (<■! \\) AHD) has been 
ohserved (Runic PJS2) These dilferences may be best 
explained by varying coastal orientation, wave regime 
and coastal circulation pallems. Ahhough a s(nne- 
liiiiallrl /onaiion ^i scdimenis and vcgciation is 
Lomitioit thouigluai! the northern gulf, each scduiK-nt 
^iid lloral oi fauna! coniinuruly, whether f'ositltnnu 
dominated seagi asses, \napv(i<\ 0| f\utci\siit sp. 
bivalves, mangroves or halophylcs and sahbushes. has 
a broad a-giona! range in elevation, Hence, u is 
imperative that local elevalion controls and local 
conditions he used in assessing sea-level data rathei 
than applying regional Vulues, 

Cii>'cn that Ihe elevation range ot tidal Ukics and 
sea level indicators can vary, grealer accuracy m 
reconstruction i)\ palaeo^ea-level is achieved il 
seveial different indieaUtrs are used, bach indicator, 
either relational or fixed, will provide evidence that 
oilbct supports ov challenges indicaiors from oihei 



horizons. By using such an approach, some of the 
problems associated with tidal indicators, the 
elevations of which are influenced by local 
geomorphic and climatic variations, may be reduced. 
This study indicates that a combination of palaeosea- 
level indicators from the top of the Posidonia facies 
and the contact between sandflat and either 
mangrove or samphire facies is the most reliable 
method for establishing sea-level change in the Port 
Pirie area. 

A further factor to consider in the reconstruction of 
palaeosea-levels is whether tectonic activity or 
subsidence, due to sediment compaction, has 
occurred subsequent to deposition. There is little 
evidence of local tectonism in the northern Spencer 
Gulf during the Holocenc. but rather, the region has 
been uplifted in response to isostatic adjustment of 
the Earth's crust due to eustatic sea-level rise 
(Belperio 1995)- The effects of sediment compaction 
in the region are less clear. While little compaction 
has most probably occurred in either the thin veneer 
of samphire facies or within the sandflat facies, it is 
feasible that the extensive, muddy, Posidonia facies 
has undergone some compaction. If this has 
occurred, it would affect elevation corrections 
relative to present sea level, acting to decrease the 
apparent height of former palaeosea-levels. 


The tide-dominated coastal plain around Port Pirie 
has resulted from sediment aggradation, coastal 
progradation and relative sea-level regression 
associated with slight sea-level fall following 
stabilisation around 7,000 years BP. It consists 
predominantly of sublidal Posidonia and interlidal 
sandflat facies. These facies occur throughout the 
coastal stratigraphy and underlie present-day 
intertidal mangrove and supratidal samphire zones. 

A tidal-vegetation-sediment relationship exists for 
each of the Holoeene facies deposited within the 
coastal zone. In the upper subtidal zone. Posidonia 
australis dominates the seagrass community and 
binds the sediment. The intertidal zone is composed 
of bare or Zostera -covered sandflats that are replaced 
by Avicennia marina var. resinifera toward the shore. 
Further landward in the intertidal to supratidal 
samphire zone, Halosarcia, Sarcocornia and 
Atriplex communities have become established in 
between sabkha-like., bare supratidal flats. 
Associated cyanobacterial mats grow within 

mangrove, samphire and supratidal environments. 

For each sediment facies, biological palaeosea- 
level indicators are defined by their growth positions 
in relation to the tide. At Port Pirie, Posidonia 
australis represents the subtidal environment from 
just above mean low water spring (MLWS) tide (0.25 
± 0.25 m TD) to depths greater than 4 m TD. In situ 
articulated shells such as Anapella cycladae and 
Katelysia scalarina or AT. peronii are representative 
of the intertidal sandflat environment from 0.25 ± 
0.25 m to 1 .32 ± 0.5 m TD, and mangrove facies 
represent deposition between 1.32 ± 0.2 and 2.6 ± 
0.4 m TD. 

Good precision in palaeosea-level interpretation 
can be obtained from peritidal sediments that reveal 
clear and consistent transitions and contacts from 
one facies to another. This study has established that 
the transition from Posidonia to sandflat facies and 
the sharp contact between sandflat and mangrove 
facies are the best palaeosea-level indicators in this 
environment. The contact between the sandflat and 
samphire facies can also be used to establish sea- 
level change, although only in areas where its present 
elevation can be established. Dangers are apparent in 
the broader, regional use of facies boundaries due to 
the often patchy and variable development of 
different facies along the coast. 

The use of tidally-dominated sediment contacts as 
palaeosea-level indicators depends primarily on an 
accurate determination of their present-day elevation 
ranges relative to tidal datum. Our research has 
demonstrated that in order best to define palaeosea- 
level, fieldwork must be carried out at the local scale 
and take into account coastal processes that have 
been operating over the long or short-term in ihe 


This research has been supported by a National 
Greenhouse Advisory Committee Grant. Field and 
technical support were provided by R. Rice, S. 
Rowe. B. Logan and J. Cann. D. Folheringham 
assisted with surveying and vegetation identification. 
Access into Pasminco Melals/BHAS was granted by 
A. Gilbert. The aerial photograph was supplied by 
the Department oi Lands. South Australia. The 
authors acknowledge K. Gowlett Holmes from the 
South Australian Museum for her assistance in the 
identification of shell material, and S. Proferes for 
drafting the figures. 

m \ i bveL indicators a1 poki pikii 


Aiu.%. .1. K. I.. (P> 1 >5> Sah-rtuusli growth and IUkUkHmi^ 
sea k-vol. implications Of il simulation model for 
I landnan coastal stratigraphy and peal-based sea Itfvcl 
cunvs. ,W (ni>L 100. >M5 

Bui ntkio. A. P. { I W) Land subsidence and sea-level rw 
in the Pod Adelaide estuary: implications tor monitoring 
the grcenhftUKc cfl'ccl Aosl J. Earth Set 40. 350- J6X. 
__ 1 1005) I he Quaternary, pp. 21V-2W to Drexel. .!. F. 

& Prctss. VV, V, (Ld-d "The Geology of Soulli Ausli.ilia' 
Vol. 2. The Phanero/oic. Geological Survey of South 
tUMruliu Bulletin 54. (Mines and Lnergy. Adelaide!. 

. Smm.i. B, W,. I'mi.v ii. II. A.. Nimkoi IK, c. V 
DiVIamik. I). J. Puisrnn. I, R., Ilvus. J. R. & 
tiiiMi^j. V A tlOX4ai Ghionolouieiil sluthes o! the 
Quaternary marine sediments ol" northern Spencer < Kilf 
South Aushaha. Mm Un>t. 61, 2f>5-29C>. 

II ,n s. J R. QqSXJN. V. A. & Pol vol It A 

i l l »x thi The stratigraphy ttf coastal earhonale hanks and 
Holoeene sea levels ol northern Spencer fiUlU, South 
Australia. INd. 297-313. 
= Cann. I II, & tiosTiN, V. A. H9861 Quaternary 

Mratierupliie -.cdimcntiTV environments, northeastern 
Gull Si Vincent, South Australia pp. 83-98 In Parker. A 
J. Ilul.) "One day excursions «jf llie Adelaide region' 1 

Eighth Australian Gco.o^ieal Convention. Geological 

Sonets ol Australia (South Australian I >i\ Tsjon ). 

, Cios'IlV V. A . CAHH, -I- II- >V Vlt rrav-WAi i mi. 

C. V. (1988) Sedimcnt-oipanisni mortal ion and the 

evolution ol Holoeene tidal scu,ueiKes ill soUlliem 
Australia pp 475-47 hi de Boer. II I ... van Gelder. A. & 
Nio. S. I). T its! Tide-Influenced Sedimentary 
Inviroiiiiietils and l : ;.eies" ( D. Reidel Publishing 
Company, I >ordtcchn, 
B1 UNI . K V. ( I0S2) Relative tall of Holoeene sea level and 
coastal proyradalion. northeastern Spencer Gull, South 

Australia. BMRJ. Aunt (nul. Geephvs. 7. 3545. 

& Cotwrti. J B (I0K2> Temperate carbonate 

sediiucms * *l Nonhcrn Spencer Gull, Souih Australia: a 

high salinity Torumol' province. Stxtimcflt&l, 2*>. 223-23B, 
Bi KTON. T Iv l |V82) ManL-rove chances recorded north ol 

Adelaide 1935-1982. I'nms. H. W. S. Ausl. 106. 183 

I HO, 
Bi it (K. A. J . !>(■) PER.S, A VI., Mi Kll 1 1 f\ S C. .VTlhAiAS. 

I) I* (|977| Distribution and sediments of mangrove 

fprwb '" Soulli Australia. //«'</. Mil. )f> It 
H-YK J A I ( I *->K I ) r\chaui!C Processes for uppei Spencer 

Gull, SouUl Australia. IhU 105. 5<*-66, 

CVNN. I. H..V: GnsilN.V. A ( I0X5.iCou.suiI sedimentary facies 
and loraminiferal biofacies oi the Si Kilda P'ormulion at 
Pon Gawlci, Soulli Australia, (hut, 109. 121 -142. 

Chapman. V. .1. U9751 "Mangrove Vegetation," 
(J. CTanmter. Cieiuiunvi. 

in Bt)t;u. P. L. van Gi i.dlk. A cV Nm, S, |) it-\K) (l'>KK) 
'fide lull uenced Sedimeniary Fan ironments and 
Facrcs" (D. Reidel Publishing Company. Dordrecht). 

PiiuiAN. J. B, (1965) late Caino/oic sedimentation in 
northern Spencer Gulf. South Australia Trans. /V ,SV', V 
Aiist.m. 125-131. 

pi rinu.K.C. H.. Van Pi i r. ,1. L., Bui sir G. S. cV Sin k\i\s. 
J. {\ { W) Tidal Wetland record ol Holoeene sea level 
movement* and cllntaic history. ftjjMftt$eofir. 
ftiltwvvlimtrtvl ft*Artt*wW- Mtt 177-21 \. 

Gosh:-,. V. A.. Mmi s. I, R. & Biii'mim. A P. il , )N4)'lhe 
sedinientarv tiaiuework of northern Speneer Gull. South 
Australia. A/m; Owl 61. I I I HN. 

_. Br.t.prRio, A. P.. & Cann. .). H. (1088) The 
lliiloeene non-tnipicul coastal and shell cuiboualc 
province of southern AusiraUa. Sni Gffi>t, 60. 51 70. 

Hunt V. D. ^ TltuM. B. (J. tiyS.lf AustruliiiM sea levels in the 
laS! I5.(WK) years: a review pp. .V26 in Hopley. D. (Ed.) 
'V\usnuiiail Sea Levels in the Lust ]5.0(Hneais: A Review" 
James Cook Cniversiiy Depurlmeiti ol Gettgraphy 
Monograph Scries a iJames Cook University. TovvnsvilJe). 

Nouimii, K.. Rossi u, M & Wauuin. L ,1 il'iHdi A 
yeoehemical study of ntetals piUSOnl ill H'ditncnts Irom 
Spc ncer Gulf. South ;\u.stralia..\,"/'/. iio-iium. I, I 17-1 50. 

Nl'NPX R. A. St Ll -\M>\. G W t I *4S(S i Physical pioperly 
distribultous and seasonal trend* in Spencer GulT. South 
Auslialia; an tnveise esiualv. A//.W. ,t Mttl. It* \An. A'r-.. 
M 3*>-53 

Stun ii k. C dvft. J, A. T. tS; Hakhimin. P. (199SJ 1 "he 
tno-a vigonnis Soulli Austiitlian tide Tnnts. H. Voe .V, 
AM, 1P>. 123 132 

Iikaindi, J, II. .1. (IMSS) reconstructions ol 
tnshoiv tidal dcposiliona! cnviroiiinciits pp, 233-2b> /// 
de Boer. P. 1. . van Gelde.. A, Ik Nta S, D- 0-dsi " 'fitle- 
Influenced Sedimentary Environ menis and Facies" 
(D, Reidel Publishinti Cornpaiiv. Dordrecht), 

Iiiom. B. O. & Rov. RS. i l L JK.i) Sea level change in New 
South Wales o\ei the past 15 (MM) years pp. (>4-84 /// 
Hopley. D. 1 Mel. > 'Australian Sea Levels in the Last 
15.000 Years: A Review". Department of Geography. 
Jajnes Cook University ol North Queensland Moiiojuaph 
Series 3 (James Cook. University, Towiisvillel 





By P. J. GuLLAN*f. P. S. CRANSTONf* & L. G. Cook* 


Gullan, P. J., Cranston, P. S. & Cook, L. G. (1997) The response of gall-inducing 
scale insects (Hemiptera; Eriococcidae: Apiomorpha Rtibsaamen) to the fire history 
of mallee eucalypts in Danggali Conservation Park, South Australia. Trans. R. Soc. S. 
Aust 121(4), 137-146, 28 November, 1997. 

Mallee communities, especially the plant components, are often considered to be fire- 
adapted but there is no information on how effectively any phytophagous insects re- 
establish their populations after a wildfire. We addressed this issue by studying the 
scale insect genus Apiomorpha Rtibsaamen, in which species induce conspicuous, 
sexually dimorphic galls of species-specific morphology on Eucalyptus species. In 
early 1996 we surveyed the species richness and abundance of Apiomorpha galls in 
relation to fire history and species of host eucalypt in mallee vegetation at Danggali 
Conservation Park, South Australia. Half of the fourteen sites surveyed had been 
burnt by wildfire in late 1985, whereas the other seven sites had not been burnt for at 
least 45 years. Only the two commonest of nine Apiomorpha species showed little or 
no host-plant specificity. Long-unburnt sites did not differ in species richness nor in 
total abundance of Apiomorpha galls from sites burnt in 1985, although the two 
commonest Apiomorpha species differed in their responses to fire history. 
Key Words: Fire history, mallee, galls, Eucalyptus, Coccoidea, Apiomorpha. 




by P. J. Ol I LAN :| '. P. S. Ck/WMoV ' & L. ( ■ ( 'nuk' 


( ,i i i ,v-. P. i.i (.'H\r-.iM\ |* S. Al C'ihUv. L G ( 10071 The le-sponsc of i-all-iniJueuijj w.*oJc insect* I Heunpu-i.i 
I NCJUHy.'feltli!! Ai>iohh>rt>htt Kiihsaanieu) to the tire histoiy ol mallec eticalvpls hi Danyeali Cnnsvivnlton Pad 
South /V.tstrjl.a !nm\. K. .\7« , V. .\/iW. 121(4), 137 I4h. 3ft Moveinlvr f<W 

Malice euiiMttuiiiiics especially lhti plnni eoniponeiis. .in.- "lun ioiisnlt*risl Rl [re fire adapted but (here tj no 
information on how ellcciivclv any phvtophasMiis i isllis iv-L'slahlish then populations tiller a wiklbiv We 
uldri'ssoJ this imiu! by studying the seali hiked gtfnift -V""""/"'''' Rubsaamen. in which kptfeic- imlticc 
hhU|hVuuuh. M-uially diHHii|>iiic grills ol spa IW spe< die Morphology rtri f.tu\ilyptu\ species, hi early l!?9ti WC 
fi vfu-il llir spcucs riL-hm^'s •Mnl abnndanti; u1" 'ipiftmurftliU culls hi rulalioil hi hrt! history tffld sprues «„'l host 
; ik.ilypl ill malice voieialmnai [Mnytiah Conscrvnlion Park, South Australia, Half otitic foiuleen vilOit -urveyed 
had been burnt by wildfire in tale 10X5, wliueas the uthei scVCfll MTes had nw llUftn burnt lor al jeUfil 4^ \caiv 
Onh the two commonest of nine \piouuuphn species showed hide or no host -plain specificity Liuuj-unhuriit 
ale* did nol diffu ai -avcics nchnos nor m Wild abundance ol Apfanhlt'filW fcHflfo bom sites hnmi in |*W£ 
.ilthouuli IHtf iwo Lutiumtnost .\pu>t)H'tph<i f$e£\&& dillered in their responses to lire history. ii.UK of t. 
'inilU V\i>'i'li( 'Julian were ct|tmHv abnnilanl al lon^-nnburnl aikl buritl sUcv. whciL-as yalK ol A ''"' t'ttmivs 
I k-ppL-n wcTfonaV'-i'i^t' loiuuiiii^ mow ubumJanl :tl lon^-uiibuiiil than ul btuiil sile.s; llnsclillcrcnec ma\ rctah: 
lo ilifleivnlinl tlifipe ^al strJl^t?^ ol the UlsHlisUn Itymphv, \\V CTmcludo thai nialk-'f wihllires at illler\als \^\' 
uiotv lhaii Hi yaiTt wouhl l\' uiiliKcly lo impac! (Irlriinenlallv on ihfi lon^-lerni survival ofa ilivcrsv.* A(itt MMlpki 
avsL'Oibla;-v piovulo'l ihal tfpfiC lony -urm tinbiifiH aiv^is (relu^ia) remain lu Nerve as sourees lot llie eolonisiny 
it\ niplo. 

Ki \ Wokds: 1'iiv hiNU>ry. mallec, gull .. ulvpru-x, c oce^ulea. iptwwrptlUi 


lite is ii siriiilicatil facUn in Anslraliaii eeoUn;y 
and in ntLit'b nl'thecinuinem the flont is J'ttv-udaptcd 
iKatiow l l )KI ). Many plants, incMuJin.i! mosl 
l\n<ulvi>fif\ l.'Her s)xx ios (Myilaeeae). pivsses^ fnv 
pn^eeleJ slrutUnvs tcpiLtumic buds or li»uiuUibers) 
Iroin which new grmvih sprouls, or liave seeds Lhal 
gurmtmiie alter ilrcs (dill IMSIa.b; Hod^kinsou <Kc 
Co-illin l l )H2; Noble |9S?>- Sneh ivueneraliuti 
abililies arc parlienlarh chaniclerislie ol plaiils in 
mallec vegetation that is. woodland comnunnlies 
dominated by iniilit-sicmmctl ciicalvpls. wliicb aic 
diemselvcs also called mallec (Noble ! ( >82> l-oi 
many cucalypls, especially inallces. fire can faciblulc 
seed eerminalion (Weltin.iilon l u S'*> oi ebmniaU 
plivlo|iha^(nis insects (N^bie I0S2) and paraMlie 
irvLsl lelocs (Cjill I^KIa . UtiCiil^pls dominalc tittfsl ol 
Anslraliit's Toresied ecosysLems and ba\e coniplev 
iclaiioii-.lnps wilb uatisc ainnials. nicludin,^ many 
niNcels (Givensladc & New \W\ \, Habuat 
maitagehicnl nsinc fire wbelbct consislcnl Willi 
hiNtiiiy (.0,^ "" uvo/aU-' poifnaicd pjist Ahorii»inal 
land manyjjcmctU praclicc) "i l**r hazard rcduclion. 
has eliecls are relatively well understood loi 

i.iiM-iuti ill h.m:hiv and ?Jw||igjj i in* luwruliiin NMriwrcil 

Li.isitV.O ('.i.ilvn.iAC'TnroO, 

1 \.llll!.ll Kl"iM|I|L;'X *IMt| lflMt|i\rlMlV I 'l ■ -:-i III! |>,VIM",(«<| 

I ftuuHAti£) f MKmil'DH.A I /IKK unN \("l' 'iifll 

vegotiKilVii bul die implicalions loi many oihei 
impiHt.unl or-eanisms, espi.-ci.illv invertehratcv are 
largely unKitown il^radslock el w/. 1^5; Krtcnd & 
Williams lsi'Jb)- The ffw Auslrahan studies of the 
eliecls ol fire on inscricbiaics have conccntratetl on 
soil and litter -dwelling oreaniMns (oe. Campbell tS- 
•Janlon MB1; Ncumami ^ T |.»lbui-.l L%j;York W4| 
or have sampled a wider in\ertcbrale assemblage 
umii^i only pilfall Maps tc.-:. Friend & William. 
Pttftj, '\rborcal insccis pntbably survive fires lew 
well than epigetc auvl liypn^ejc •-.prcics (Wbektn 
FW5) and, provided sampling biases can he- 
addressed, should be good candidates tor Miidviny 
I he effects ol hie on imcri cbrate populations. In the 
picseut study w.- examine the cIIclk oi fift bistury 

Cui oink-mic Aoslnihao stale oisods ( Hvilnplura 
C oeeoidea) lhal live OTftlj in die canopy Ol mealy pi fi 
Scale insects (0 Ihc ^cnusApitwitifpfnt Rubsaatucu 
1 1 nococeidael live wilhin ^alls lhal they induce on 
(heir eucalypt hosts tGullan l°^ In 
■\pinmtnplui. each adult lemalc resides in a lar^e. 
often symmetrical and woody pi\\. usually on the 
stem but .sometimes on leaves, buds or fruits oi the 
hus| etiealypl (GlllllUl I^X4,i) I hesi- an- i:,is.b, 
recognised as cocc<nd yalls because (here is a v .iuall 
il(ncul orifice Ihtou^h which Ihc lemalc can ehminalc 
her waste honeydew and also mate with the male 
Hei offspring or tirsi-iustai nymphs, called crawler 
make ihc it exit front the maternal call through this 
same opening and then disperse io initiate new gulls 


I' ' • I IIAVI'.N,' KA.\MMN\ I ,| K 

• Mi .i.llUihk- Ibli^gC, TllO grfk "' 'Hairs ;.ur ninth 

smaller th»tl l&OSfc t>F Ihe Icmales. rarely more than 
uric centimetre Inng, nnd are luhular with an finical 
otilicc and sometimes an outwardly-directed Mange 
at fhC apev fhr shape of thr -all o] the fCflWle LS 
pcculiat CO Ihc ApiVfitf/rfHiu species lhal induces it. 
rceardlcss of I hi identity of llie host cucalypl Most 

\t>:- nwtfilhi species show* sonic debtee Ml spccificiiy 
|0 a restricted range ol A7/< ,//v/>/nv species (Cuban 

|^S4ak \pii»HU>tj*ln{ has 3° described species 
(Clullai I^X-ia; (iullan cV loncs |0Wf, each nl which 
I trims ehaiai'tcTisiir, sexually -dimorphic galK 
i ■• n.ially. Apioiunrplui can be identified ttl specie - 

level In ihc iTcid even b) rKHH>£# totfw wlih 

minimal trainer 

SljIu msecis aie often claimed to have low v.igilm 
I lor review, see Hanks & Deiiuo IWti because Ihc 
kVjftgitttft adult females usually spend thoiv entire 
lues <>,, a single hoM plant and OViptVSll there, adull 
mall's are shod lived and weak-flying, and nil 
dispeisal is due to the movement of the craw In, 
which usually icmain on their ualul hosl il conditions 
HA ivoiurahle iCrealhcad 1990: NcSlvM i'f d/ l*#| 
hi Af>tt>fnoiph<i. our field observations tn sclcroph\ II 
woodlands h»VC suggested I hat I here is lower 
di\vi:-,jiv and abundance ol galls m ansis ,iih|rrted to 
frequent or seven: burning. This reduction scents 

unrelated to the Rultabillty of poKt-flro encrypts foi 

gall dc\clopnicui. since ihc cpiconnic Mush ol 
rhllttgu 'hai follows H (ire resembles, ihc preferred 
plant matenal utilised by ,\pi<>ttior(>ltit in l>>ng 
unhurnt areas, tt is more likely ilui after a IJrfl kills 
Ihc calls and I hen occupants, it lakes tunc loi 
ret olonisntion by ctawiers io occur and if lues aie 
licquent. 01 poienual sources ol colonics ute distant. 
local populations may not re csLahlish. 'This 
hypothesis rentauis speculative in the absence ol 
cjuanlihealion ol any differences In species diversity 
and abundance between bumi and long -uuhunii 
areas, We investigated a* colonisation alter lire hy 
surveying ihc species richness and abundance ol 
[pitmtUFftlM m relation to both die fire history and 
the species ol host cucalypt it 1 malice vcgetaMon jji 

& l \nslialia. 

This study was undertaken in malice lor several 
tVitfOlto- Firstly, mallei- is a Ly pica IK Australian 
u-yruninn thai has been m sciious tfcCJfoc fbl iftC 
past 15/) vears tfuouyh land eJearancc and other 
tonus ol' dcgradalion thand Conservalion Council 
I0.S7: ChCHl \W9\ Harris 1^0). Secondly, il ha>, 
been hypothesised that luallee plain and animal 
communities aiv nuunttunetl by episodic lire iNohle IOSVk but nn re.seareli has been iUum: M 
mscMieate liow criecosely any phviophue*'us insects 
it establish their populations in malice allci Cue, 
Iknlb. U is easier lo louuI the c.'Ms Oil Ikv 

cucalypts than t»u the taller b>rcsl and w^oUland 

Sealc insects itl ApiWtiQffmA are eood CUll^ldiUC* 
lor slutlyintr the elTeels \\[ wildfire because, unlike 
tuoic mobile insects iWhclan fit ai I 1 >S0l they aie 
not able lo mow Irom ihc iheir host plant-. 10 a\oui 
the Hamcs and. ahhou^h the ihick wall <»t frji | Ulll 1 
ha^ been su^ycsled lo be an avlaptalion ft>| ffnc 
proteetiiin ( Koteja I986H the liijth itttcnsiis oi malice 
wildliies usually kills ihc ovcrsioic\ UlUa^i 

i,Hr;.dslock lOVtli, In .s.nirast n» Iht- Wvll-k \i 

biases o| pillalhne Irappine and t»lhci methods ol 
sampling soil or liller lauiu (see Whelan IW51. "iir 
suivcy inelhod hn* Aphmunphu potentially assess 
Ihc total number ol ealls on each dec burih, i 

.u\\ nu:h'i-s i.f ihc use \\\ \ffhvuayphft galls h that 
Iheir abundance varies linle \Mih season ol' siu\c\ 
miico the Icmales of "most species ptobably live ml 
much longer lhan a year (L/l, Cook unpuh. i ,iu-.l 
gnjls a\y remain on the trees I'or several Viae, ,itn*' 
the death Ol the occupant, lhus providing a rec(»o.J Ol 
the presence ol Ihe species at a Mle. Lastly, in ordci 
lo elucidate causal relationship* between 
invertebrate abundance palteins aiu! inc .selccied 
invertebrate groups nei'd tn he examined at i linct 
level of laxonomic reMilultou than Ihe order oi 
lamilv level lhal is used in most studies i bnciid t\ 
Williams [9UI}), I iotn this pffCjx\T&iWG„Ap{fflntttphtt 
is an ideal suidy eenu> because ^alls can hi] 
fdClttiflcd 'CiKiils t(i species level. tVfi !hc lidd, 


The wt»a vw.s can led out |fl Daue^ab 
Conseivalion I'atk which was cslablished in H>7(\ 
hi'iame Auslhllia's llisl linispheie Keseive in IV77 
and is now pan ol Ihe Bookmark Biosphere Reserve 
This quarter of a million hcciare ivscne is ahoiU 90 
km north of Kcnrnaik in South Ausliali.t and lies m 
the northein hall ol die Miuiay basin, adjacent lo the 
Ne\\ South Wales border Urn. 1). Il includes boih 
arid and malice lurid systems and allows access lo 
-some speclaculai old en-wili malice. Of panieitlar 

I'liikirancc to our sludy is ihe documenlcd lite 
hisntiv of 1 >;ui*iijal i Conservation Park (finin 
National Parks and Wildlife Service. South 
AUStralliA). hue has \\0\ |»ceu used as a iiiaiia^cmciit 
tool in the maintenance of its malice for 
conservation: recorded wildliies either have been 
naturally -occurring, following lightning strikes. HI 
lh'* ivsult uf human accidents. Mans 1 areas have un 
evidence of burnmii lor a considerable (Wind 
perhaps for over a ceniury or jt least not since either 

I*' I I oi |?5| whei'Ms trust vc w dd lues bnon Oh Ol 

the region. However, ol special relevance lo l|v 
piesent stud\ is li major lightning induced lire that 
bumi ;i l.ii'ec i.eiUial area of die iesei\e m I >cecmhei 



Humes! cad 

niirMMimMMtllM11IM>i i tsf ,,,,, Mi iS ^u ^ ^1^1 ■ . ■ - 1 ■ ■ .. .r m* , - ;Hn;-: ; : sr.«j n i 

"""i Park boundary 

—* Road or irauk 


Study siu- 
;xli:nl of W4 fire 
IB fcxtenl of I'm fire 
1~1 Kxlenl of I "85 fire 

5 10 

Fig, I . Miip of Dan^gali Conservation Park. South Australia, showing the extent of recent wildfires and the locations of the 
14 study sites where galls of Apiomerpha were surveyed. Inset shows locution of Danggali Conservation Park. (Figure 
based on map and aerial photographs from the National Parks and Wildlife Service. South Australia^ 


l> .MM I.I AN. I* S ( KANSTONcV l.,(i.t dOK 

1985 dig. I). Since then. Ihe burnt eiualypts have 
regenerated From (heir malice liguoiuhcrs, although 
the dmiJ stag* i»f punt substantial liinbs siiii protrude 
Irnm ihcs uow-lloiirisliing malice I'^mwih. 

We nude ct>iii[>arisons bclween the Aftmmor/tfiH 
galls associated wilh cuealypis in the old-grovvlh 
malice | M iotlp-imhiH'»t" sites) and in ihe iTWllCC thftl 
had regenerated alter tiv |985 lnv ("hunt" sites). 
AM was cOlldliOlctl in April |W& FmirtftCtf 
iltC* were chosen (Tig, 1 1, seven in each treatment 
lypc Ouuni versus long-unkiiuU, wilh the site Co- 
ordinate's as given in Tabic I. All cucdvpls at the 
burnt Mies had been burnt completely [fl ihc I9SS fire 
and sites 6-9 were completely surrounded by puxl- 
litc regcncialed malice. We only used bttrnl silcs in 
die central area ol the reserve we Were 
certain thai vegetation in Ibis area had been 
completely dcMioycd in I VMS, wheteas the 
vcgciati'ui atound Tippcrary Dam and "Morgan 
Vale" Ruins and in Ihc CW0 smaller areas also bnrnl 
in I'W.s on the periphery ol' the reserve tfig, I) 
appeared tO have- lveu mote paleliily burnt. Five 
burni .aid flVC k>h£ unburn I sites had an undcistoiev 
wilh porcupine grass, Triadw irritate R. Hi. 
(Citamincac). sometimes as ihc dommanl ground 
lavei planl. Hie olhci loin sites either had reasonably 
ban: gfrmtttl-OUl mixture of shrub species. 

I he inatlec enealypl species at the study sites were 
I. ut a/v(?/u\ t>h mwiifti Sieber c\ LX'. (syn, ti <oxtnrn 
f Mu.-ll, & hchi c\ K Much K A, <iuti)n\,t Glintl, c\ 

Oh ley, /. tfrovilti V Mncli . /■ oleum !-. .YUicil. e\ 
Miq, .\\k\ L. toi in/is h Much c\ MU|. IdjMittfTcaiiim 

was made using rostermans il9°4). cvecpt dua the 
naiiH' / ilwraxXUUi (syn. /•'. Ctiftfxtftk) is relained (M.I. 
Brookcr pcrs. conum), The pioporltons ol each 
species varied according to locality bul usually ihice 
"I Hie five malice eucalypls were present at all sites 
( f'ablr I U We chose sites in long iinbiinii or posl lire 
regenerating patches to provitle 10 multi-stemmed 

malice eucalypls. of between 3 and 6 m in height, per 
site- These 10 trees were chosen k|H representative ill 
the proportions ofe-.ich spi\ ics m the local area I hi 
height was imposed by ihc need lo inspect the total 
foliage both front the ground .\\k\ from a ? m ladder 
placed in the ventre <»l the tree 

[•or each tree we counted the numhci of galls 
conlaining live and dead AphmmrphU females. The 
surveyed •jails varied in >i/e lnnu a lew mm (young 
or aborted) lo more than 4 cm long, and in shape 
from cylindrical and buddike lo urn shaped or ovoid, 
Galls were recorded as containing dead Icmales jl 
(hey were okj and brown tir showed signs ol attack 
by predators or parasiloids l dually, the presence of 
a living coccoid was confirmed by Ihc presence of 
white powdery was at the gall orifice. Only VKichci 
material and galls ol uncertuin idcnliiy were 
Collected, so the survey vvms iclativcly ntin 
deslriniive. Voucher specimens o| -jails and slide 
mounted insects n| Apiottiotphu buve bLL-n d^nsiied 
in the Australian National Insect Collection (ANK'i 
(7SIRQ, Cacitoara. 

All data analyses pertain lo gaJIs OonUUfljllfe! live 
plus dead female insects, unless otherwise slated 
The golly "I dead insecls were incliidrd in counts, 
because any successful initiation of a ['all was 
evidence that ilflScul had icached the site ami ihat the 
tice was a sutlable host Host specdictly within 
'\f}fttthoip/m was examined by calculable the 
pereenla;:c ol the surveyed trees ol each cucalvpl 
species dial supported galls ol' Icmales of e.u I, 
Affioiutnpfm species. 'I he response of AfiWnwrphu 
species to fire was evaluated using one wa> , -i ■-. 
ol variance (ANOVA* lo compaie burnt and long 
unburnl sites in terms of die total numhers ol ' ; »alls of 
temale- iof all species summedl total numbers of 
galls of die seven least common species (i.e galls ol 
A. nutlh Viwola (Julian and A, uvitltlwctex ('leppcn 
excluded), and total Aphmtofphd species found at 

t via i.i, ,S7/r hn ttlrtrrx <fn»u c/'s midiHM) with /'/<* htnw \ //""f tmimm tf-u) \r>><t\ huna in /Vi\3 i'b)\ <uul the mwiln . 

r./ hrr\ ,>j ,,uh K^/vm umiplvd ut nuh v//e I-. «l - \ iftfltHWU. F-. g " l_. ^.ac.h-, )-. i. E im l ,-mII.l I,..i, \_ 
u\vt)\n, \ s _ [. M>ciahs, NwtfhctS til ihr (onf qf <yrh v/.e< i> > CtlluMU nfrr In ihr tt>t,if Httutlxr of in. -\ wrr,, \, W ftH 
lli'fl \/u> U>&, 

Sni 1 li an ■ 



1 ,1 

F. e. 


1 i 

i n 



1 I-u 


1-10 -}S'\ (" 


2 I-u 

33 171 r 





i b 

.^ r '17'16" 

UO 3.^'UJ" 





4- h 

n u iTW 

140 3s OS 





$ I-u 

3.i IU'jo' 

140 j.vir 





33ld 0V 

MO «'lf 






7 h 

33 uriM" 






M h 


i mavos 11 




" h 

J3 IftW 

140 4257' 




1(1 l-ti 


14(1' 4 V S|" 




II I-u 

sr 15'i)?' 

140 42'4*T 




\2 In 

33 IV \*r 

140 IV IP 





n i-u 

13 I7W 

I lo _iyw 






14 h 

3? c l»9'20" 

140 35' JO" 







1 / 




tiiich Nile 1 14 units totuli Analysis was curried QUI at 
the site leviil, not the live level, because there were 
Iwo w more differenl tree species per sile and many 
trees had ni) or few galls. Two species. A. 
tnallccaado and .4. tnict>loidcs. were common 
CflOUgll lo examine their individual responses to Ure- 
al the sile level using ANOVA bill the dalu were 
irauslormcd [I,. lx+1)] to correct Tor skewness. The 
responses of ,4. mattecaadu and A, ovitf/loidts to 
lire also were evaluated using individual trees as the 
UOlls Of analysis. In trns ease, binary presences mm\ 
absences were analysed using Chi Square tests 
because galls were not abundant or widespread 
enough to satisfy underlying statistical assumptions 
;ii this level. All analyses were carried out using 
.IMP- 10 <S AS [jislilHte Inc.. © Kftft-fl), 


Apioinorpha species recorded 

A total Of nine species ol Apiotnorpha A as 
recorded from the sites surveyed These were: A. 
culwina (Tepper), A. densispinosa Gillian. A. 
kttrsthi Riibsaainen. A. itndlceticohi, A. ntttndu 
nndtccnsfs (iullan. A. ovi< <doidc\. \. n^idoris 
iTeppcr). A, \hvnthylo\<t (Tepper) and A. mnaln 
< leppcrr All of these species have been collected 
ptcviously from mallee vegetation in souiluin 
Aiisiralia (Tepper 1X9-3: Gillian IW4a). 

HiKst-plam specificity /'/Apioinorpha 

One species. ,4. den\i\pinosu. was recorded Irom 
just three sites and solely Of! £ dtimusii but only SIX 
galls were found and none ol these contained a live 
coccoid. Six other of the nine species ol Apiamorpfto 
showed some degree o! host -plant specificity (Fig, 
2). Galls ol A. atlycinti were found on lour of the 
seven surveyed trees of /:, im nissoto as well as oil 
two other euealypl species. Galls ol .4. ho\ehi also 
occurred on ihree eticalypt species, wheieas A. 
muniki nudleensis, A, re^itfari.s, A. xtmmbytostt .'Hid 
.4. iiriuifi.s each were recorded from only two 
CUCalypt species, However, 13 of the 14 trees that 
supported galls of A unudis belonged to /:'. gntclli* 
and nine of the 10 trees with galls of A sitt>tithylo\ii 
were L. socudts. Only one species. A, ovuolnides 
was recorded on all five species of eucalypl; it was 
the commonest species, occurring on 52 of ihe 140 
surveyed trees. The next most common species wUH 
A, nitdieemoUt which was lound on four species ol 
eucalypl and on 37 of the 140 surveyed trees. 

Specie* richness und ohionktnce (>j Apiomorpha in 
relation lo fin- fdsK'ry 

All nine Apnnmnpha species were lound at both 
burnt and long-unburnt sites, liurni and long-unhurut 
sites did not differ significantly In the number of 
Apiomorpha species recorded on survey trees 
ih |t =3.57. ,>=0.0K) (Table 2). ANOVA of the total 

I xui i. 2. Means per siif ± I SIX fM" >">/' nmf& in parendnyfs. and siyiifhame <>l dii'iavm i\ dw tnfitt hfotirry "t tin *T#fo 
(*/ Apunnorphj species, mmbcr ttfgu&s oj ail Apinmnrphu tpecfcx, ntinfoer of $alh <>f A. inalleewcitai mnnhet uj $alk 

fi/ \ nvieoloides and number tf'g&Us of nil vpevics 6XchuUng A. rnalltvacolu and A ovicnloides. {for A mulleeacula 
and A. micoloidcs, ntrans and fun$es are ftmi the miv dam hut F and p wiuas arv from uonsjannvd data. M 


indi< atrd b\ > }. 

Lone -unburnl 




n (sites! 

Mean number of 
spCcicS per site 

Mean namber of 
yalls per sate 

Mean number of 
\ mtdhranikt 
galls pei site 

Mean nuniber ol 
,\, t»ui>ioidc\ 
galls per site 

Mran tiunihvi iff 


galls per sile 

with \. maltiiuu oia 

ami A. avit ahtid<'\ 


5.3 ± I.I 

M 4 ± A 


13,3 t(f-M)) 


4 0+14 

te.5 ± I4.K 

7.6 ift-21 1 


12,7+ 14.1 




It U 









A. calydna 

h ;■; ',: 

r T^ 

6 £ 5, E /. E o, B 
Eucalyptus species 

1 HT i 1 1 

E d E g, £ L £ a r q 

Eucalyptus species 

4. maUaaacoki 

A. munitt! n:-i!k\n$!$ 


; -,( 1 - 

- ■ - 





__ __. 




1 I 

E d E, g. E i. E. a E. 
Eucalyptus species 

i t r 1 

E d E. g. E. I E o. B. s 
Eucalyptus species 

A oviLZrkiitiv;; 





30 ~ 

20 -: 

E d. E g. E. I E. a. E. s. 
Eucalyptus species 

A. regularis 

Q. - 


| 20 ~ 


g IS- 

n i 

% 10- 


3 c - 

5 ; 

I 1 1 


d E 9. E (- £ a E 
Eucalyptus species 

A, strombyiosa 


r - — i 1 1 1 

E. d E. g. E. i. E. o. E. s. 

Eucalyptus species 





30 ^ 


15 4 






1 1 I 1 1 
d E. g. E- l E- o. E. s- 



tfo& species 



numbers 0] fittlDi ol females UlVp atul dead) n&nrd&l 
from t-iich she indicated that the lire history of the 
Mlirs did iiol affccL gall abundances t\V ApHtoWpffft 
specivs IT.,j=3.*fir, /MU>7> liable 2), Btirnl and 
tone unburn) \iic.s Jul nol diller in the percentage til' 
live 10 dead Apttntit'tpiui: 'lV/< ot gulls ;ii kmg- 
uiihiirnt sites and 24 c /< lit galb a" burnt sites were 
estimated iu contain live coeenkk hvcept Fur I 
mtilUt'UvnUi ;inU V or/i o/ou/o. I here were 
insulfieienl doiil lor statistical analyses based on 
individual scenes, although (or the thud mo.a 
ahund-anl species , I n/n.ili\- there were compar.jbli- 
populations :ii hoih burnt .n>d Imig-unbuiut \\fe$ t 

tin V nuttlecttntiiL ANC.iVA ill the iransformed 
abundance |T« i\+l>| at cadi sue indieated thai 
number, of galls did HUI diller between btirni and 
loug-unnumi siiesfh, p =0.40 T /»=0.>i) (Table 2). Chi 
Square analyses ol presence-absence dala loi ,\, 
ni<t!ltctici'lti on individual trees alstl indieated thai 
lite Ibsjorv diil mil affect die likelihood of finding A. 
mtitifivtoid galbi (X", i^OXW, />=0.S4). wilh ga|U 
being present on 2"/'/ ol all bees sampled in long 
HiiHurni sites and 2o'v or all trees sampled al luirnl 
sites. IT lives nf /.. ntrru*\<it(i. on which A, 
iti(i/lrca<ol(t was never found, were exeluded, iho 
icsttlls remained very similar \/J ( ■ Pl -H,2 (4, /r=o,M: pieseul Oil *<>', : > »'l lone unburnl trees and JfWI 
ol bmui trees). Burnt and long-unburnt sites ititVcreit 
only slightly in the peivcniagc ol live to dead A. 
ittatfcauofti. i$.1$ ol vails ul long-unburni sites and 
*9',i ol galls ul burnt sites were estimated in COMQil 
live cotAtudv Among the btnnl sites surveyed, there 
was no evidenee that y alls oil tiuillvcw >i,i wctc 
mote abundant ai site^- elose to i' range 1-U wnh 
mean of M oalls per site), compared wilh iJc-.t.mi 
ffOffl ir.ittvr M I I with mean ol K.3 galls per -uko, 
long uuburrii ate. is, 

In contrast, tor ,\. nnmlnuks, ANOVA tif the 
transformed abundanee |l„ (x + lll at eaeh siu- 
indieated iliat number-- id galls vuav sigtnfieaittly 
Uvet al burnt Mies (F, p =| j *7 ( ^=<M)l) (Table 21, 
Burnt and long unburnl siles differed only slightly in 
Ihe percentage of live lo dcadvl. oyitfj/ou/t'\: iM'f of 
galls al long-unburnt sues ;md 14'^ of i-alls at burni 
sites were csiimated lo contain live coceoids. Chi 
Square analyses ol piesence-abscuee- data for _V 
iiVttoloidi J \i>n individual livo-aKo mdieaied dial tire 
history affected the likelihood of finding \ 
owcoAi/fAw gtilts 'x\iis =l) « f ^- /?=OM>03), with ealls 
beiny pivseni on 4M'./ ot all lives KaMpfej] in loug- 
nnburnt sues, but on tmly 2A^c ofbllrnl trees. Among 
the burni sues surveyed. p|.H of A. ovictiloitleK wen 

sli,-:htlv more searee at siu> IIMI-2(M) m Ironi unburnt 
vegeiaiion (range 0-3 widt mean ol'2.0 ealls per site* ul sites seveial km dist;nH loan lone unburn! 
areas (ranue I -8 wdh niea.u of 4..^ yalls. pet siteK 

If both \- iHultccuittht and -\. uvllvlRitlvs were 
excluded Ironi the analysis l>\ lofal numbers of uulk 
ANOVA ofihe gall abundanee at eaeh site indie.tted 
lhat uumbets ol ealls of (tie other speeies eianbined 
did not diller between burnt and lon^-utibunu site^ 
tb, (1 =ILU). /^i).54) ( Tahle2). 

M'rfttlitv f>trttfr\ 

The original oeeupanls ul inauy ol the Afinttiifipfu) 
galls that we recorded during uttr survey euhet had 
been killed by parasiloids. probably wa.-sps. oi 
removed by predators, probably eork ( ibn.s ^nd 
patfots. S'tme ealls had a sintile, large exit bole m the 
wall, providing evidence of the emergence of udart- 
lailed wasp CborWWtwfUi Oallu Torre (Pleiomalidaet 
(Tillyatd l l >2b; ^Joueek l*>XS; N;i.tmatm (WIk and 
Otbci eaIN had many fmy etnergeuee h(»les. Twelve 
galls had one side removed which is the typi. i\\ 
ap|ie.iranee ot a gall opened by a bud. Mortality ol 
the laiter kind was twice as common in die Jong 
unburnt sites «iv in the horm sties, Inn theie were loo 
few galls damaged m this way lo determine wheihei 
the difference had staiistical signilicanee. Many 
iMhei galls weie brown and obviously dead bui 
generally we could nol determine the cause <»1 dealb 
A fe:\v olhor galls were ikfnimed by im|iuhucs. thai 
js, nihcr insects bad occupied Ihe gall tissue or the 
eavilv but bad nol dtreelly killed the Apiomorpha 
lemale. The idenlity of the itK|uilines was noi 
deiennined because those Mill occupying the gall 
vVL-.e enlier dipteran or byntenopleran larvae and no 
rcanue lo adults was atleinpted- 


In the evenl o\ lite, season \M burn is believed io 
have the grealesl influence on ihe plant composition 
of m:illee communities, wilh frequent autumn fires 
causing aibslaiuial nt.>H:tliiv o\ muilee eucalypls 
(Noble I9H3, IMS9). Conlinuotis cantipy go^wib <a 
malice eucalvpts can t>ceiu aftrt ,i summer wddbre 
and may be due to the absence ol phytophagous 
insects (Noble ls»N2l, I ire frequency (the interval 
beLwevn llresi is helieved to have the most important 
lon»-iertn effect (m malice fauna because most 
animals adapl nol to Tire itself bui lo the fk>risiie and 
struelural leatures of the plain eommunibes thai 
resaill from differem lite legimes t laud riu^ervalion 

Ptg 2. Hosl-platil specilK-nv ot Af'ii'tnorphu plotted as ilv percenter ol I he slirveVLvl trcc^ til eaeh enealvpt sjveies llun 
siipjvortffd j:alls of lemal.r- ol cash r\pt&tnMjJhti 5pecic&; l:nvol\ptm speeics; /.. ./. - /- tfNNftaxn fe e = '* ^nit'tJlt t, >. 


' cil ION?), |,,i' HWcrtchrates, v\ hich ecncraJIv 

1 nil seasonal acli\il>, mappropnale nnm^ - * t "^ ' 
pjiiii nl.trly frequency of fire can have danvitiuiu: 
i "ir eminences populations (land Conservation 

I ■ il I *>S 7 FYlCinl tit Williams l ( K'f»». ftjr ai le\tsl 

"iiil- (n,uliii malice conunnnitic 1 ., h Nil? toCCT 
.1 £gi?HlPi1 thnl I he luiltinil fine frequency is unlikely 
in escecd moiv thou one lire every 15*30 year* 
hecause of ihe lime required lor fuel loads CU 

u# liiic (Noble i^S'j). At Dan^ah Cqrrcctvaiinn 

l\nk. many ol Hie mallee site-- (foul WO HUdJctJ Lid _ L 
hitultti understorcy and il had heen 10. 5 years 
wildluv had destroyed our hurni sites. Tims, il 
Kip .iiH.ic. ut \ni<>})n>rpiui species iM fojTlU Mies 
could be shown lo have iceovcivd (it piv-biir.i V in -u 
ihe prohable naiural lice frequency of IS-2U yew iH 
JTHUii would be mi likely lo have any lime-iurm 
ikinmenlal effects on populations o[ Apffmnphci. 

Hie- main lltulll^*)' "'" s "' Kc '>' ■ V,:IV " ul t,K * Il '"' : " 
ruibuint sites dnl nol tuTlei in species richness m 

total ahuudame oi \fliMhtHJ?ku jnrtls Iffm h »rnl 

10 i >eai% ae.-. but thai the medium-leim elicit nl 

hit- in v ".. M.llcreut \piomtnphti -.pccics. L-Vll 

die I\v" s]-"'oes (hat m>civ common cnomjti it. .illow 

analvsis i tt thc»r abundance in iclation lo lliv liNlon. 

1111c il, itiiilhonoki) war-, equally abundant in burnt 

ahA long-unhuinl sites, whereas the other ( \ 

r ii |1 ,■/,.■'.../'. ■>,! w,is siLMiifieauily less ahuitJuni ul the 

burn i ill left. Indeed, lor A, c\ n nh'nh's. 0J- 

,-Uilitistiment al hunil sites was low even where a 

source ol poiemial colonists was jusi across u HI m 

w ijc load I or fltwi species with slow oi limited post 

file le-cslablishmcul, the eliccl of auothet lire in ihe 

il.- x i Hvtt io 10 years mmhl be virtual extinction. 

especially il HO old-inowth malices, thai may serve 

js lire i'clui!ia loi sueh insects., survive the lire The 

conseivnlion of kmy unhiunl areas [)[ mallee should 

be a manaeemeul pi unity. Sumlaib. I net id & 

Williams ( l')06i have emphasised Ihe importance ol 

die manaecnvnl U» proteel nre-seusiiiw specie* nnd 

habitais Irum lou-tieo,ueni tires in mullce-liealh 

shrublantJs ol south wcsleru ,\tistialiu 

In eeinlraa. .\ tmdUvtuohi antl the ihird must 

common species, V mj/^/i'm, had re -established 

popuLilions etjtiivalenl lo those recorded in loii^ 

iiiiImhiu sites at sites butnl 10..^ years previously 

(although the number or records lor A umalis weie 

iusullieient lor statisikal itualysis), In addilion, ihere 

were' no obvious Inc elTeels on re-eslabhshmeni ot 

lite olhei si^ \t*nut)orplu\ species bill numbets ol" 

call-. »'bser\ed. at both burnt atul lone-unhunU sites 

lor eaeb specie,-, t?eneralt> weie low eumpared wilh 

\ ituflli'vutulu Re-etilomsation had oceuocii ftl 

btirm sites dial were 10 km or more tltsiaut from the 

neaies! st.mrls ol lonji unhurnt malice, b'or vaeile 

>iK,ie'. the posi lire tltish o\' ^rowih mav ho 

beoeOeiiil lo rail establishment. This sueciesiuMi is 

supported by the observation ihat 111 iflltel 
en\ in.nmenls, ^alU nt' Apiom^rpfut appeal io \Mi 
more abundant "ii yumiL* and rCgl'Wth eiiealvpts 
ihtltl on Ihe lohagc ol oklei bees il.el'welon ••: 
Vaarwerk !W; P. 1. (aillati pets. nt>s.i, AllhoiiLh 
ihis impression may be cuviied by sampling bias ills 
il reasiet to search tow foliage ot saplinys and small 
trees ihan the canopy ol more ivialurc lice-.), pm 
jlasshouse iearin;j, has shown that Ihe first tiisl.n 
nymphs will initiate giflfa only un the new fnliaye ol 
at lively gfovvme shocas. Jn anolhct study (Yen 
|W>). (he abtmdaiue of phytoiihaeoiis iiiseei-- 
especially sap-sucktnt* <|?t;tiw>. has been show n tcv he 
higher till cojipiee thun mauiie malice, perhaps 
because yoiine leaves jnd shoolsare iikmv rmvriClou^ 
than old lohiiL-e .mo C0ppj< C tftC5 have more Vuiimj 
iiiuwih IhcLti inalutc malices. 

Sot ne Apumtnrphtt species exhibit host-planl 
prelerenees lor cerlam etiealyp! speen's, Hoth \ 
rr^nlntis and A. \net)ih\ iosn uccuireit onl\ (»n i\'.o 
eueatypl species, /•. .uniati.s an<" £ Of^iU which are 
closely related species - bolh are in Ihe series 
Subulatac o| ' Lm-nl\pttts (C hi[>pendaie IMMS) C'uills 
of \. mttiiitn tthtlUitixn were lound only on / 
ihiftwn -mil /.. V''"' 'i'tA. wlii.Ti are m difl'eient i -eMe-- 
(fVlltppeitdvtle ( *>S^ >. fWO u.-i\ eU.sely eelalC.I 

\i>tt»H(!iphit species A rtilyiiatl and H. ttnuihs 
(Caillari l')X4a. L.C (o ( »k unpub.i. showed wrx 
l ii0._reul host prcfeienecs. Thirieen ol the l-l 
stirvcyd lives with gulls ol A U*Wfa WW* I 
•;/om//\. whereLis ealls ol A. lytyiWU ^Cjv "m-., 
eommoii im 8 Uu -rowani ami ne-vei ocurrcd on / 
vrm ili\ Since Ihe iwo mosi common AfttMUtlFfttm 
-peues. A, tmtil((<t<nla ami A. t>\ /< o/o/ f /ev also had 
Hie broadesl liosl-planl ranees. diHeicnces in the 
ocLurience and abundance ol Ihese t\vo spccie.s in 
relation \t.) ihe lire history ol sites eannoi be 
alliiboied lo any \ariaiion in the composition o. 
eucalypl species anione. sties. Instead, population 

dilfeieneeN Uimmg Api<>tf!<>rf>lhi species in lelalion io 
ihe lire hisloty of sites may be best cvplauu-d bv 
dilTereiKes in theit pio|,'cusilv lo disperse. The 
eiawlers of some Apnmunpfm species may dtsjieise 
more readily to new lives than those of oilier species 
There is ample evidence thai lusl-inslar scale insects 
o| othei groups arc ilispcrsed passively t>y the wind 
and. even though nie»italiiy is \er\ high, oiay be 

carried lor distances o\ a lew in lo several km ■ 

more rarely a lew hundred km. from Ihe natal lives 
(reviewed by l 3 ednley l°X2. (ireathead l ( »U; Hanks 
^ I >cnno 10« : ),^), Some scale insects eiawlers hau- 
been reported to orientate downwind and stand ou 
theit him! Ic^js wilh juLcnuac and lore lr: 
<tulslretchcd ( Washburn t^ Washburn I oS \ 
Washburn Sl hankie I'JSs. f.renthead \WH>) Sueh 
behaviour probably ensures (heir dislode.emrm ftfld 
dispersal by w ami. The crawlers ol » lew sjiecies o' 



\pjO)lhiFphu have been observed displaying similar 
behaviour under elasshousc conditions (I.. G. Cook 
unptih. i. Afti, itmnpini species may JillVr in ifictl 
propensity cither Lft disperse actively tfuiJJ the hosl 
plmU 01 remain off the <iatal lav. In Ciintrust It) I he 
Mui|onl> itl ApiDinorfthii species, ii is extreme I v 
difficult hi establish inlcMatious AC A malltctu hhh 
A mfwaui and A. <muit>\ by telcasine LTWfavOM 10 
polled cucalypts in a gju^tolrtai <L, (i, C0ufc 
unpuKi; ihis gliggue&s thai a! leued Bfftae crawlers of 
Ihe latter species may exhibit obligatory dispersal 
behaviuiir and, m ihe elasshousc. may suicide hy 
uctivclv depailiiu 1 from the only siiilahle bust phml. 
I'ndci natural eoinJiiio|i,s, however, nalal bees 
pmbahU would he surrounded by other suilabte 
ho^ls, especially in malice vegetation. 

Uispeisal ability may relate Lo LtiiM'pbOlOgiCul 
adaptations as well QS behavioural ones. The 
flattened bodies and two or more lomj. fikuneniotis 
caudal setae of scale insect eiuvvler.s arc believed to 
enhance their dispersal potential tWainhousv |#j£0* 
Peddcy IW2J Thus diiWiKcs ui budy si/.c ami 
shape in. ( y partly account lor differential dispersal 
atpouy species, The crawlers i)\ Aftinnu/rpha have 
llallened, oval lo siibeiivulai bodies (ringed with a 
continuous tliw ul marginal setae (Gullan t*>84W; It) 
addition, the surface area of each marginal seta w 
rM ended by a I Inn sheet ol wa*.y secret mui 
(analogous in eencral appearance lo the vatic from 
Ihe shall of a fcathen. The firsl-iustat nymphs of A. 
<>vtiolohlt'\ are about the same length as. bul 
narrower [ [^5-225 pill al widest [ft Hi than, those ol 
V tiuilivcacoiu i?fo-:so pm wide), although the 
mareinal setae are npproMinalely etpial in leimili 
I 14 44 pint nn boih specie* < I -. Ci Cook uupuh.J, it 

dispersal ability 01 dpitUmtytki is v undated with the 
surface area ( »f pic budy of the crawlers, the 
differences in abundances of A. imtllcvucnta and /I. 
luvVf.'/o/Wev in burnt and long-unburni sites may be 
nrtt ibuted at least in part lo diflerenee.s in ihe si/e and 
shape of their crawlers. 

Sdcetion lor both active dispersal behaviour and 
bnd> morphology thai favours passive drift may 
occur in scale oisect species lhat occupy 
unpredictable or temporary habitats, as has been 
Mi-L-esled for armoured scale msccls that teed on 
short-lived versus lon^ lived hosl plants iCireathead 
LWOii If this hypothesis is valid Hit kflhttmifpUit, 
then xome species, such as A. m<tlkeitrt>i<L tan lv 
posiulatedto be better adapted n» the vaeaiicsolTne 
in thi: malice cnviruumcni, 


VVe wish to thank M. T CollolV lot assistance in lite 
field and Ml Osboinc, the Ranyci at Dane-jali 
( onset valion Patk. foi help and inha-mainm 
provided during our ficldvvork, K K. Pullcn kindly 
arranged, and the National Parks and Wildlife 
Service ol South Australia provided, the scientific 
pei mil for (his study. The Calpci uin/Bookmark 
Sutvey of Invertebrates > J mje.< I funded b\ o\ the 
Australian Nature Conservation Agency (now 
bin iK'hiucnl Australia) provided the travel funding 
tor this work, P. L>. ( V.npci vavc adVkfl on and ft G l,_ 
MtKie carried tail ihe sUuistieal analvsiv \, J\ 
Austin, U. K. Blanche, U. M C4arke. M, ) ("olloff 
and R, B. fluyd made useful (.omments on earlier 
versions oi the manuscript 


bvumw |i A MWDTV Aiistnthan Hot;,; ie. ori-'in and 
.•vdImUuii pp_ ^s 7s In M | tola wf AtuiiMha. Volunk- |, 
iiie-t'lueiHai' i.AuMraliaii Gin Vffi rill ieni I'ublishinp 
Ser\k-. ( jnlvo.o 

13 i 1 1 1.. /. riywj) Ausi t -ai ;i M;in Cbikltuiloid a 

(I lyniviiopU'iai. A Bii^y-k-nialk Revision 111 t.eneia 01 
IouOl-l-ii liiitnlies, with a Reelassil'ieiuion nt Species" 
K'vti Ininnauniiak Walliiu'lordi. 
lUMitsioi k. K. A. i P^O) Kelatiimships bctw&OTl fnc 
ivL-iiik-s. pliiMi sjuviev. ;mj fuels ill malice eommiiiuiies 
)ip :iX >23 hi NolMe. It.. i0S<, I' .!. L V Junes, Xj, K 
fjivlM] 'file Mallee P-iiilK A (uns^ivaUoii PeispeLlive" 
i('Sll^) PuliltALilM.ns. l-.;isl MrlhoMtne). 

. Kriui, P ( \, t v \y iU, b D, MW51 1'hl- uutl 
eons, iviHKHi; iitipt rativ es and teiisiiaiiUs in DlUJlLUillg 
)<>r Oiver-.iiy pp. ?£i 3.i3 tn Btadsrocfc- R. Aj,v AuW f 
\t. Kc-ah. H A KitiMsiuri!, R T. Lunncy. f). fy 
Siveosen \X V (H<ls) Cunsei ■vnii: BitultvCruiy: llneat.s 
ilULl Siilniums iSunev Kiatue ( V. Ne-ns Piv Lltl hi 
assOfilflion with NSW NP\\\S. C^nppiii.-' Nutlon, 

(".'■vvii-hf I l. C;. & I ■vHins. M. T (Wfrlj tlVL-fkasia Inc 
on Ihe iiiveilebralc laiin.i ot suit ,nul lau-r ul ;t i-nralvpl 
hav^l pp. _M5-2at h, ( (l |f \ M, firpves. K. H.S 
Nt'ble. ! H iVxh.S "piiv and iIk Pan,.' 
eAusltnlijii Ai_MuVnty til Setenei'. s ;inlu'iiii,'. 

(ill Al.. D. C ( t L >XW) S(r.ueyii\ fnt eonserr me eu-ninumnies 
fltld spueies pp 4(\4 4?(> hi Noble. ,1 C t & tliailsit-ek K. 
V, i [5< Medie,-iTiineai. 1 .aidseiipes in \iisir;ili;t" 
MKO PubhtalMMiv Lnsl Melbourner 

Chm'oi sjum i - Ci M f|VK8) "LtuuiiY/vm. Anfflfihtnv 
iMyrinccac). Flflrj Hi" AiMralia Vulunit IT lAiislraliau 
Guvcninicni Pubhslanu StrVli*% Timberm) 

t^isiikM.vv,. L. (twar-'livi^ ii| Vieiona ami Adjnininy 
Ait_j-r ifuslerniiiiis PllWishulg, Kranksluil). 

I-ioi sjD, (;. R, & Wiri.rvMs, M K l l«W linpael ut tire mi 
inveilebrale cniiumiiiiiies ui nialleedieaih shnildaiKb nl 
.'•"uilnveMern Ausiialiii. l\uiil < 'amen: Witt, 2 2\ I-3M 

(Jar, A. M- (l')Kbo Adaptive responses ol Austialiiin 
Vaseular plan) species mfire- pp.2rto>27-| tuiiMI. A M.. 
(iroves. k, H. cV Ne.btt- t. H. it-\ls) ant! ihr 
\('s|raliail BitMll fAfU^ftan Aeadetnv i)| Scien^. 


l> I (il'Ll AN. P S. CRANSTON & I ("..COOK 

\ |081b) Coping wiih iiil- pp. 65-H7 />/ Pat*. .1. S. A 

MeCnmb, A, J, (EdK) The Biology ot Ausiulian Plants' 
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Ciui vi in. \i». D. J, ( I WO') Crawler behaviour .mil dispersal 
pp. 3115-308 tn Rosen. D. (Ed.J "World Crop Pcsls, Vol, 
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GuLLAN, I* I I. MJS4u) A revision o! tlie gallTorniing 
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09S4h) laxononiy and biology ol Australian 

gall loimini^ Coecoiue.i. Writ, SIEEC A'.. Bnthpffl 
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,V lt)\i s. M. Li. I 1989) A new species .if 

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1 1 asks. L. M. <V Dimv, K. I IIW) rFie role ol demit 
jLl.iptniion in colonization and spread ot seale inseel 
populations pp. 3*43-41 I ht Kim. K. C, iV MePheion, B 
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Variation" Uohn Wilev & Sons, 1 1 ic , New Vorfc); 

HARRIS, C. R OWO) The history ol' ntallee land 
Aboriginal and Luropcan pp 147 1*>I />; Noble. J. C. 
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Hoih.kimson, K. C. & (il<it-MN. (J I- (IWCl Adapt. itioii ot 
shrub species to (ires in the arid /one pp, 145-152 /" 
Barker, VV. R. & Grecnsiude. I*. I, VI. (t-ds) involution of 
Ihe Flora and Fauna of Arid Australia" (Peacock 
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Koiiia .!. [1*>8GJ Lire has given birth to complex plant 

vails Holh Uih Em. agr, Filt^m Sihcxtrt 43. 35-3^ 

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Malice Area Review" (Land Conservation Coimeil. 

|.lHki:l<>\, M. Si VAAftWfeRK. M (1W3> Miscellaneous 

Holes on iMuucrous Xphumtt'plHi spp. t llomoptcra: 

l.riueoceidaci and their hosi plants m N.S.W. Sydney 

finsiii Naluhilis] 2, l-l 
NAUMANN, I- D. iP»',t|, Hymenoptera pp, 5)16-1000 In 

CSIRO "Hi' Insects ol Ausiiah.i 7n*l cdn iMelbouttic 

liuiversiis PrBKh, Carltoti). 

Nisni. I). CtHibN. II., Sahiiih. N.. Km in. M. A: Mim.ii , 
/ !l l '*>5) Spatial disttdtution ot stale iiiseet'.: 
Loni|'aiaiive siud\ Using TUyJUP-h power law. Hwlr, I nr. 

24. sue, si: 

Nil mans. V. G & [V'l.HlWT, K 1 1491) Llleels ol I'uel 
icdiKtion btinun^' "ii cpigeuj ailhropods and cailhwornis 
indiv scleniphvlleuealvpi fores! olWest-ccntud Vu loo a 
Ami J, fool l>»- J15-330. 

Noun . .1. C. i PJK2) The sipnilieanee irf lire in the hioloyy 
and evolutionary ecology of malice im •tlxpttn 
populations pp. 153-159 ///Barker. W. R. <V (ipamJ ii 
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and uialk-e coppice dynamics pp. lo.N-IKI! (ft Nohle. J 
C & Hutdstork, R_ \ il T iNi •'VKdiu-iTanean Landscapes 
in AuMralia ,, (CSIRO Puhlieahonv Last MclhoiiiiKO. 

INomMA. [>_ B- OVS2) "Windborne Pests and Diseases 
Meteorology ot Airborn.- Organisms" (Libs Uurwood 
I Id. Cluehesler) 

Thvi-k. J. Q, O. ( ISO^) Descriptions, of Soudi Auxtratiun 
btaehyseehd galls, fhtits. li, fiif S.AtlSl 17, 265-2SO. 

1 1. i \ \k'i.', R. J. (IV20) 'Insects ol Austral^ and fcitfU 
Zealand'* ( Anens Robertson Ltd. Sydney). 

W Aisiioi si.. D. O l )K(t) Dispersal of In si itisiar lai\ae ol the 
felled beech scale, CrypttitVL't'UX /<"fAue>' - ' "/'/ ;/ - /: '"' 

WeaniikN. .1. tf t V Ikankii. 0. W. i,l«JS5) Biological 
si i idies ot iccplant scales. PuMtidHrihi 
nwsi'mbrYtinttH't'ti anil I'nhithiriti detotSo\ (llomoplera: 
CocLidac), in California. Hilt'ctnlnt 53. 1-27. 

Si WAKiimikN, L. t IMMT) Active aerial dispctsal ol 

minute windless arthropods: eAploitatioii of boundaiv 
lj>er velociiy gradients, ,s ( a7/ce223. lOKM-IOXM. 

Wi 1 1 tMiiiiN, A. B. (!9H9) Seedling regenerati<»n and die 
population dynamics oleucalvpts pp IS5-U.7 //> Noble. 
.1 C. & Biadstock. R. A. tlHds) "Medtlciraiieai. 
L.andscapes in Australia" tCSIRO Publications. Bust 
Melbourne i. 

Win l an, R. J ( 1995) 'Ihe F.eology of Lire" (Cambridge 
Cuivetsitv Press, Cambridge). 

-, LANlfttUYK. W. ft PASHBY, A S. i lOHOlThe fitfccls 
of wildltre mi arthropod populatie-iv. in Jatrab fi<nd\<>i 
woodland, li! 4iisr, Nut. 14.214-220. 

Yl'K, A, L. (IQ89) Overslorev invertcbnties in ihe Uii: 
De-.ert. Victoria pp 2^5-297 /" NoUiJ. J. C. t*v 
Brad>(ock. R. A. (l ; ds) ■"Mcditciuuicaii Landscapes in 
Ausiralia" ICSIRO Publicalioiv.. l : ast Mclbi.uineL 

Ni ir:K. A. ( D94) The long-term effects of lire on lorest hOI 
etunrnunities. management mi]>liealions lor ihe 
conservation ofbioilivei-ity, Mem. L*<l Wwi" 3*», 2-^1 -230. 





By A. M. Olsen* 


Olsen, A. M. (1997) An intensive monitoring study of two wetlands of the River 
Murray in South Australia; physico-chemical parameters and cyanobactena 
concentrations. Trans. R. Soc. S. Aust. 121(4), 147-155, 28 November, 1997. 
Quantitive data were collected on physico-chemical characteristics of surface water 
temperatures, pH, turbidity, conductivity, dissolved ammonia, dissolved reactive 
nitrate and total phosphate of Banrock and Loch Luna wetlands from 46 samplings in 
each wetland over a 20-month intensive monitoring study. Concentrations of the 
various physico-chemical parameters were within the ranges found in similar 
freshwater River Murray wetlands. 

Key Words: Wetlands, River Murray, monitoring physico-chemical parameters, 
cyanobacteria, South Australia. 

TnmMWtums of the Royiil Sacwxy oj'S, Aitxt, |!W), 12K-4 1, 147-155, 




by A. M, Oispn 


Oi si \, A. M. ( l u s>7i An intrusive monitoring study erf two wvilamk nf the River Murray in Noinh Australia: 
phvsico-chcuncul pi urn nttiers und eyanobacierw goneentniiionK, Trotu. h\ S<n\ 5- Aw*f- 121(4). 147-155. 2X 
November. 1997. 

(,)n. motive itjtj wi.u' collected on pjiysicd-dirimica] diaiaeu-ristics i»l Miituce water lejiipcfauirev pi I. 
turbidity, conductivity, dissolved ammonia, dissolved reactive nitrate and total pliosphulo of lianmcK and I .och 
l.itna wetlands from 46 samplings in each wetland o\c.f a 20-month intensive monitoring study. Conccniraiions 
oi' t\\e various physico-chemical parameters were within die ranges found in similai i'rcshwalei Rivei Murray 

Weather factors, such as strung winds, heavy rain runutf and liehlimu!. prodtuvd pertuhaUons ill ttirbtdily. 
conductivity, dissolved reactive nitrate and lolal phosphate levels in the two wetlands. 

Nutrient concentrations in excess of 036 tnel ' total phosphate and 4.0 mgl dissolved reactive nitrate with 
jr-in;j water temperatures were related IQ rapid cell multiplication of the cyiinobflCTCTEJ Anohatmi Rgp, Ihree 
Siutlhti'thi Kpp were predominant in the two wclUuuk ,wm\ reached their greatest numhers (23,700 t'Clh nil ' ) hi 
J .och Luna from lale December [994 to mid - January iw.s, 

Ki v Wmkpv Wetlands. Kivet Mm ray. monitoring physico-chemical parameters, eyunobacleriu. South Australia, 


in 1990 the Murray Darling Basin Commission 

through its Natural Resources Management Strategy 
(NRMS) funded a pielimmury 1 study pf the water 
chemistry and uqualic invertebrates anil land 
veriehrates of 10 wel lands oi" the River Muiray 
floodplains in South Australia (Goonan et at. 1092.}. 
The survey was conducted during May June l u «)0. 
Between May lyyQ anxJ February 1^92, a second 
more detailed physico-chemical and biological 
survey of cighl alkaline freshwater wetlands was 
carried out on Ihe above - mentioned lloodplains. 
Five of Ihe eight vvellands were located between 
Clover Lake (Calperum area) und (he Bern 
I'vaporalion Basin and the remainine three were 
between R&mco Lftgoop (Waikerie) and hike Carlet 
(6 km upstream of Munnum) (Fit;. I }. These results 
were reported by Sulci vi id. [ l uo ~3). 

Banrock Station Hoodplain (Section 662, 6KI anil 
6X2 - Hundred of Moorook) and associated wetland 
lie in aboul ihe middle oi the 100 km (npprov.) 
stretch of Ihe river between Ihe live upper and three 
lOWCr wellands studied by Sutcr ft at. 1 1943). 

How regulation of the River Murray al Lock 3 in 
1025 helped create two permanent freshwater 
wrtlands in the region, one located in the tloodplnin 
o\ Banrock Station (34 " OS' S, 140 20' E) and the 
uihci in the Loch Lum:i VA'ciIaud ( oiuplej I }A I I S 

140 22' E) opposite Banrock Station and aboul one 
km upstream o[' Lock 3 weir (ANCA L'996), Banrock 
Station wetland has a 90 to 130 lia area depending on 
water deplh (20 cm - 1.1 ml and was created in the 
1 050s by damming the upper section of Banrock 
Creek. The wetland behind the dam wall is gravity 

Z\ r 



- 1 




-34 a S 1 



• hi; 

: ■ 

J Murray rtfl 
Waltote ' -' y 

South Australia 

Marmuii % --' 





- J 

l4fl'F Shale (kml 

1 1 <ih t^'i'i tirovo Newton s, \u-.i m\ia 

Iijj. 1. Localion map of wetlands studied Riut Murray, SA, 



kx\ by a channel from Lock 3 Pool. Water levels are 
maintained by controls at the inlet and outlet points 
and discharges flow into Lock 2 Pool. European carp 
control structures were erected at the inlet in 1994. 

The study described below was undertaken to 
determine the cyclical changes in the physico- 
chemical characteristics of the waters of the two 
wetlands and their influence on the rise and decline 
of those cyanobacterial species likely to produce 
toxic outbreaks. In the summer of 199 1-2 there had 
been visible blue green algal blooms in both 

Materials and Methods 


This monitoring study of the two wetlands of the 
River Murray began in November 1994 and ended in 

June 1996. For 19 months water from only the River 
Murray catchment and upstream storages flowed 
through Lock 3 Pool. The Darling River did not How 
because of the four-year drought in its drainage 
basin. Mixed River Murray - Darling River water 
flowed into Lock 3 Pool in June 1996 as a result of 
floods in the Darling River. 

In each wetland, five sites were selected for water 
sampling for reasons of accessibility and 
representativeness and for sampling any increase in 
cyanobacterial concentrations irrespective of wind 
direction (Fig. 2). All water samples and temperature 
readings were taken in the morning and as near as 
possible to the same time at each visit. Wherever 
possible. Loch Luna was sampled first followed by 
Banrock wetland within 2 h. Collections of water 
samples were made weekly from October to December, 
fortnightly from January to March and monthly from 




1 -5 

Lock 3 


(34° 08' S, 140° 20' E) 

Loch Luna 
(34° 12' S, 140° 22' E) 

34° 10' S 

Loch Luna 

Sampling Stations 

1 -5 

^ 4# ^v Luna 

7\S^ « 

Scale (Km) 

Inlet Pipe 

140° 20' E 

i \ 

Kingston on Murray 

Fig, 2, Banrock and Loch Luna wetlands showing sampling sites. Note direction of How. 


Tabi.I' I. Numbers of cells of Anabaena spfi. and Tahu; 2. Numbers of i-ells of Anabaena spp, ami 

t ouceutratious of dissolved reactive nit rote and taiid coucentratiinis of dissolved rem 'tire nitrate and total 
phosphate - Loch Luna }994~9&\ phosphate - Banrock L994-M6, 




Total phosphate 



DRNitrate T 

otal phosphate 

(mgl 1 ) 

(mgj M 

(mgl ') 

fmg! ') 


Ongl ') 

Nov 3. I'-m 



Nov }. 1994 



Nov 8 

0. 1 3 

Nov 8 



Nov 15 

0. 1 7 

Nov 15 




Nov 22. 




Nov 22 


Nov 29 




Nov 29 




Dec 6 




Dec 6 




Dee 1 3 

1 .230 


Dee 13 




Dee 20 




Dec 20 




Dec 27 




Dec 27 



Jan X l$$5 




Jan 3, 1995 




Jan 17 




Jan 17 




Jan 31 




Jan 31 


2 i 


Feb 14 




Feb 14 



Feb 28 




Feb 28 




Mar 14 



Mai" 14 

88 "< 



Mar 28 




Mar 28 



Apr 18 




Apr 18 



May 16 



May 16 




June 13 




J Line 13 

wetland drained 

July IS 



July 18 



Aug Ift 




Aug 16 


Sept 5 



Sept 5 



Sept I 1 ) 




Sepl 19 




Oct 3 



Oct 3 


Oct 10 



Oct 10 


Oct 17 




Cfcl 17 



Oct 24 



Oct 24 


Oct M 




Oct 31 




Nov 7 



Nov 7 




Nov 14 




Nov 14 



Nov 21 



Nov 2 1 



Nov 28 



Nov 28 




Dee 5 




Dee 5 




Dee 12 




Dec 12 



Dec 19 




Dee 19 




Dec 26 


2 2 


Dee 26 




Jan 9, \9% 




Jan 9, 1996 




Jan 23 




Jan 23 







Feb 6 



Feb 20 




Feb 20 



Mar 5 




Mar 5 



Mar 19 



Mar 19 



Apr 14 



Apr 14 



May 14 


May 14 


lun 9 


.lun 9 


I ,0 

A M I 'I SI N 

April Li | September on l he dales gjVGn 1(1 Tables I am! 


Al tMltfh site il : ic'. 2) a one-litre sin lace watci 
sample was LlikctI and bulked in a five litre plastn 
bottle wllll -.amples taken from die OthOI '""' «ttC« 
Aliquots nl' the bulked sample were Itaiistetred to 
aii I ref 500 nil polycarbonate plastic scrcwlop 
hollies for subsequent plu si. :i)H hcnucal una Uses al 
(lie ScJCIM Section, (ilos.sop High School The onc- 
hlre walci samples lor eouuluie cyauobacleiia were 
iiunsfened to I -5 I plastic scrcwlop hollies leaving 
u 25(1 ml headspaec. The samples were kept chilled 
until dcitV^T) 10 I hC Australian Centre loi Wain 
OuahU Kc^eaich. Hohvar SA bw enumeration Of 
I vanohacicria cells MIMSO l n V0». The Kpeciul I ,2? I 
plastic hollies lor die water HltfllptlJS Were supplied 
from the Waici Qiialily I .abotalory. 

a floating L Dor Grunc Pvnkf No 742s. bine 

alcohol column llli/ITI ctet was used to io«>i.l ilu- 

•■mi lace waiet tenipcralnres at each site A mean 
surlacc Witter lemperuuire was tllEll ealt ufned for 
each wetland. A plastic hodied minimum - nia\urium 
IhciUH'mctci. i 81 h50 ( I with pressure adjustment 
lor indicalors <iii die mercury eolumii was used to 
Aeiauil minimum and maximum waler temperalures 
h| .leu limes *il eonsceuliNC sampling seals, ll was |S < ut below the surface- 

flume was no |j June 1995 Collection Mi tile 
Hnnmek wetland because this wclland was drained 
Kit titauiienauce work on the initiation pumps used 
n-r highland vineyard irrigation. The Locli I una 5 h l'-.» l Jd sample was discarded because ol 
'.ir.pcctcd contamination 

r In -inictif untitystw 

Waiei samples were held at 4 C until icqmrcdaud 
III most iir-.UHH.iS were unalysed within 4S h. 
MiasLiierucnls ol turbidity, dissolved ammonia, 
lIishuIvbcI teaetive nitrate ^Kt loLal phosphate wvie 
IffiUfc W^ittH tl HAC'll DftkU/5 instrument and 
prcmi\cd reagents iHaeh I-V84). PiTT dissolveil 
.aniuoma each sample was filtered thiouedi a DuilMv 
Kol^-s 201 tiller paper lo remove suspended solids A 
IS ml aliquot was mcaMiicd into a < lean lI;*:^ samplr 
cull and I ml ol Nesslei lea^enl added (Uaeh |QS>£j 

I he midline was left lor colour lo develop, In I his 
case hie blank was distilled water (2S ml) with I ml 
i-l Nesslei icaecni included, Measiireiueiiis were 
made nl 425 nm ancl recorded as mgl ' after applying 

. • uiivciMi.n factor of mithiplicaiion b\ 1 ,29 (Maeh 

lor dissolved leuelive nitrate tIJRN) each sample 
e..e littered through a I >oublc Rings 201 paper to 
icmove cl.iy particles- A 25 ml aliquot was m<-asurcd 

Nui \w<akitW\ 

into a clean elnss sample cell and the contents ol one 
Toil suctiel Ol prenuxed Nilraver 5 reagent added, 
this mixture was rapidly aeilateil Tor one minute 
(hen leii lor five nuniiles loi colour to develop. 
Another tillered 25 ml sample was used as a blank-. 
Measurements weie made al 500 lWl anil recorded .i, 
rn^l ' alier applying a eonveision laciot ..j 

mtiltipircmion by 4.4 diach l*W2l 

f ; or total pbt)MphatU (TP) a di^estise pmtXBfl 
used W convert all lorms ol phosphate to the s(»luldc 
oilhophosphale form (llach l'">2), hib> ml ol Ihe 
sample wro: me.isureLl into a clean \23 ml 
ralenmeyer Husk aitADjJ with 4 ml 5*5 N UNO, and 
l wo sachets ol K.S.O. i llach (W2) The iiumiio- 
was heated in a boihu.c water baih ha 1^1 millUM 
allowed to cool lo room letnperaluie and then •! ml ? N 
NiiOlI were added I hfi SUmplii was -,plii inu> uvo 15 
ml portions in clean ulass sample cells. Out' (oil 
,.u lui *)) pivmixeil I'hosver 5 reaeent was added H) 
one conlauici anc! the eoloui allowed lo develop, I he 
other was used as a blank lot speelmrikiric analysis. 
Measuretneuls were taken at 700 nm ;h\l\ recorded .is 
ryigl ' abet applying a eonwisinn factor (division by 
,i). Ihese DKN .wm\ II' resuh-; uanplenieitt data 
collected lot eit'hl SA wetlands by Sulci el {it 

pH was incaMuei! usinj.- a Manna HI R424L pH 

ItlCtOt The piulv was rinsctl in distilled wuiu 
between each le^t and lelt m the -ample uniil ih.- 
Iiighesl stable rciklme was reached. 

biir turbidity measurements, each sample was 
agitated vigorously and a 23 nil aliquot was added to 
a clean elass sample cell and placed in the l-JRFI /5 
speclrophoiometec YlcaMiicnlcuts were made al 450 
am iiiul O'curded as Nephelometric TurbnhU (NT1 
turtle, A blank ol distilled svuler was UhUd lf» 'vyo the 

t ..n.Juctivity measureinenis were made Using a 
LIT] IT 20o'eonduelivily meler calibrated h> Nl \ (_\.uducli\ ily (IT') units between each u#v 
I he calibration solulion wa.s prepareil by diltiliut 
74.53 g crt iwen • Jrftfd KCI m I000 nil ilistilled 
Wilier. The probe was rinsed with distilled waic piHQi 
In each use and the maximum reading taken eatb 
lime il uas nsrtl Kesidis Were in H( ' unils 


Mian surface wutcr icmt>c*vnuv\ 

Thiouehoui this study the mean suilaee wntei 
temperatures in the two wetlands showed almost 
identical trends In eeneral. water temperalures wem 
higher m the uioie ^|«»sal shalhAv Haiimek wellan-l 
than in the sliehlly more pn»leeted Li»eh t UDil 
weiland. The higliesi surface water lempcTatures Hi 
HuuiHck- :.S-5 ' , and 2S.4" C were rccrded m 
the summer ol IV*J4 3, corresponding leiufieiatures 



in Luch Luna were 27.4 s , 28.5 and 25.3 q C (Fig. 3). 
The lowest mean surface waler temperatures were 
10.3 ' C in Banrock and 10.2" C in Loch Luna on IS 
July 1995. 

In Ranroek the greatest range in minimum and 
maximum water temperatures between consecutive 
visits was 20° C (15" - 35" C) between 3 and 17 
January 1995 and the greatest range in Loch Luna 
was 14.5 C (14 - 2K.5 D C) between 22 and 29 
November 1994. The monthly ranges in 
summer/autumn were often 15' C or greater, 
whereas in winter, they were 5" C or less. The least 
dilTerenee between minimum and maximum water 
temperatures between consecutive visits in Loch 
Luna was 0.5 Con I X July LW5, 

Lock 3 Pool surface waler temperature 50 m 
upstream of the Banrock Station Intake in 1996 was 
slightly higher than the mean temperatures in 
Ranroek and Loch Luna (Fift. 3). 


• ! '■■! 

p.., , Ml ii. i. 



!t« t-nu Mar 


■II- i ■''■' 'J - N » n ' 

Fig. '" Comparison of nflcatl surface u.tier temperature* at 
Banroek and Loeli Luna. Noveaibcr inuajtme 1996 and 
Rher Munav. October L995June Wo. 


The pH values ranged up to 9-5fi at Banrock (27 
December 1994) and 9.04 at Loch Luna (3 January 
19951 (Fig, 4). Lock 3 Pool water registered a 
maximum pi I K.o2 on 14 November 1995 and a 
minimum pH 7.51 on 2b December 1995. 


The shallower Banrock wetland mostly recorded 
higher turbidity Values than those of Loch Luna (Fig. 
5). Highest turbidity in Banrock was 200 NT units on 
3 and 10 October 1995 and the lowest 30 NT units on 
14 March and 28 November 1995 while in Loch 
Luna the highest mrhidhy was 170 NT units on 5 
September 1995 and the lowest 10 NT units on II 
March 1995. Turbidity in Lock 3 Pool ranged 70 NT 
units on 14 May 1996 to 20 NT units on 26 
December 1995. The mixed River Murray - Darling 
water had a lurbidity value ol "4K NT' units Oil 9 June 
1 996. 


VUc initial high conductivity o\' IS98 EC units in 
Ranroek on 3 November 1994 decreased to S71 EC 
units in 6 weeks and more slowly thereafter to 500 
J3C units by 9 June (Fig. 6). In Loch Luna the initial 
condticliviiy of 1095 EC units on 3 November 1994 
decreased to 6S7 EC units on 17 January 1995, rose 
slightly before falling to a minimum value of 327 EC 
units on 19 September 1995, VVilhin a fortnight there 
were two sharp increases in concentrations (2720 EC 
units on 3 Oelober 1995 and 1759 IX' units on 31 
October) before conduct n it v values decreased to 
511 EC units on 9 June 1996, 

The conductivity values o! the River Murray water 
decreased slowly from 686 EC units on 31 Oclobei 
1995 to 51 1 EC units on 14 Mav 1996. The mixed 


i \ \ 

... : ■■ ■!...■•■- p 

pai a m. ■ " i 

Fig. 4. pit. Banrock and Inch Lnnu. November IUO4. 
I4 l Ki and River Murray. October I99fkiune W%. 


1 -..1 .1 ■ ..I- 

Mil Mill 1, 


., , .■.'•.' I)!! t.t.., 

I 1- 3|. Turbidity, banrock and 
iW-June 19% and ftrvpr Mu 


, 1 i,.. 

Loch Luna. 
rrav. October 


.,, V , II 




River Murray 
June 1996. 

Darling water was 438 EC units On 9 


Weather factors, such us heavy rain and strong 
winds, were found to influence some physical 
parameters in the two wetlands. The average annual 
rainfall at Barmera is 245 mm. The Barmera rainfall 
was considered lo be represenlalive of the area and 
its daily rainfall records for the 20-month period of 
the study showed that rainfall was irregular with 
intermittent occasional heavy falls of 27 mm on 4 
January 1995. 2°- mm on I May. 53 mm on 23 
October 1995. 3S.5 mm on 2 January 1996. 25 mm 
OH 27-2K February and 20.1 mm on 3-4 June 1996. 

Dissolved ammonia 

The patterns of dissolved ammonia (Fig. 7) were 
similar in both wetlands. The range in Banrock was 
from 0.17 mgi 1 on 28 Novcmbe/1995 to 3.10 mgl ' 
on 7 October 1995 and in Loch Luna from 0.30 mgl ' 
on 3 January, 2N February and 31 October 1995 to 
3.48 mgl ' on 21 November 1995. 

Dissolved Reactive Nitrate 0RN) 

In Banrock there were occasional high DRN 
concentrations, exceeding 4 mgl '. in the period 
November to March 1994-5 and again on 3 1 October 
1995 and 9 January 1996. At most other times DRN 
was below detection. In Loch Luna there was high 
DRN on 17 January 1995 and between October and 
February 1095-6 there were four occasions when the 
concentrations exceeded 4.0 mgl ' (Fig. 8). 

Total Phosphate (TP) 

The highest IP concentrations recorded tor Loch 
Luna were 0.95 mgl on 24 October 1995 and 0.67 
mgl I on 19 December 1995 (Fig. 9). At Banrock TP 
peaked at 0.7 mgl ' on 20 December 1994. On 13 
December 1994 and on 5 March 1996 the next 
highest Banrock values were 0.6 mgl '. The mean TP 
for both wetlands was similar at about 0.3 nml ' 

|u Ml M. 

'■••--'• . ./ v -^-.^ 

Fig. 6. Conductivity. Hanroek and Loch Luna. Nove-inbei 

1994-June 1996 and River Murray. October lOOs-Jnnr 

Lm lunn 
Fl .: 

r ' 

1 ': '. . i ■;:,,,. 

-i : ; * ■ 

•it. ajtc j«. i Fit Mil A# rj)u, Jun im -'.m Eam O..I ndu Dfe. Jan Feb "Wi AD HMy ' 
3* I -ii i all 

MiTiti Dkii 

Fig. 7. Dissolved ammonia. Banrock and Loch Luna. 
November L994-Jillte IWo, 


When water sampling started in Banrock on 3 
November 1994 the total cyanobacterial count was 
14 cell ml 1 (Fig. 10). By 6 December 1994 
cyanobacterial cells peaked in Banrock at 24S0 cells 
ml ' and then declined steadily to 5 cells ml ' by 2X 
March. The second but smaller cyanobacterial cell 
multiplication in Banrock occurred a year later 
between 7 November 1995 and 9 January 1996. Cell 
numbers rose from 24 cells ml ' to a maximum erf 
2300 cells ml ' before declining to 71 cells nil ' on 5 
March 1996 (Tabic 2i. Three Anahiu na species were 
predominant in the Banrock wetland. 

In Loch Luna between 1994 and 1995 the same 

■■♦—Vv'-- -A-r . 

■I ., i , i Mm Ami H»» 

Fi*:. H. Dissolved reactive nitrate, Banrock and t oc4i Luna. 
November 1994-June IWff, 




Nm Dec Jan Feb Mar Apr May Jun Jul Aug Sep. Oct Nnv Dec Jan Feb Ma' Apr May J UN 

Other cyanobacterial species identified in the water 
samples, although occurring only in low numbers, 
were Anabaenopsis elenkini (6 December 1 994 - 20 
February 1995), Aphanizomenon sp. (22 November 
1994 - 13 June 1995, 23 January - 19 March 1996). 
Oscillaloria sp. (3 November 1994 - 18 July 1995, 9 
January - 19 March 1996). Cylindrospermopsis 
raciborski, Piankfothrix spp., Arthrospira spp.. 
Microcystis aeruginosa and Pseudoanabacna spp. 
were identified from time to time. 


Fig, 9, Total phosphate. Banroek and Loch Luna. 
November 1994-June 1996. 

I ucN LdhS 

4 IA r J'A 

Nov Dec JaN Feb Mat Apr ' May Jun Jul Aug Sep Ocl Nov Dec JaN Feb Mar Apr May JUN 

Fig. 10. Cyanobacteria {blue green algae). Banroek and 
Loch Luna, November 1994-June 1996. 

three Aiutbacna species were predominant through 
l lie rise and subsequent decline of high cell 
multiplication (incipient blue green algal "bloom""). 
During November 1994 the number of 
cyanobacterial cells reached a low of 41 cells ml ' but 
after 29 November their numbers increased rapidly 
to peak at 23.700 ceils ml ' on 27 December 1994 
before declining to 97 cells nil ' two months later 
(Fig. 10, Table I). Anitbaena coiled species was 
predominant until 3 January 1995 after which 
Anabacna circinalis displaced it until mid - 
February. The coiled species again became 
predominant until June 1995 when all three 
Aitabaena species were present in low numbers until 
late November 1995 when multiplication of 
Anabacna coiled species began again. This species 
peaked at 2530 cells mL l in early February and a 
month later cell numbers fell to 63 cells mi ' (Table 
1 ). Cell numbers rose slightly to 444 cells ml ' on 19 
March 1996 but no cells were detected on 9 June 
1 996. 

Surface water temperatures. pH, turbidity and 
conductivity levels followed similar trends in both 
Banroek and Loch Luna wetlands and were 
comparable with the values recorded between 1990 - 
1993 for the eight floodplain wetlands of the River 
Murray in South Australia by Suter ct ai. (1993). 

River Murray turbidity values were highest (70 NT 
units) on 14 May 1996 and lowest (20 NT units) on 
26 December 1995. Mixed waters of River Murray 
and Darling River (Lock 3 Pool) registered 48 NT 
units on 9 June 1996. In the 10-year period 1978-88 
Lock 3 Pool water averaged 60 NT units (Mackay & 
Eastburn 1990). 

The high conductivity value of 1898 EC units in 
Banroek on 3 November 1994 was caused by a 
blocked inlet pipe into Banroek; with the clearing o\' 
the blockage conductivity values in one week 
dropped to 1507 EC units. Turbidity in Banroek 
increased from 104 to 155 NT units between 5 and 8 
November 1994 due to turbulence from the rush of 
water following clearing of the blockage. Seven 
weeks for conductivity values and five weeks for 
turbidity values were required to reach equivalence 
with Loch Luna values. 

In Loch Luna conductivity levels rose from 327 to 
2720 EC units on 3 October 1995 after spring rains 
and strong runoff. At Barmera, 13 mm of rain fell on 
31 August, 9.6 mm on 5 September, 9.8 mm on 25 
September, 10 mm on 3 October and 53 mm on 23 
October 1995. Seiche effects in Lake Bouncy and 
added runoffs caused the high conductivity water 
from Lake Bonney to flow through Chambers Creek 
into Loch Luna wetland. The outward movement of 
the high conductivity water (2970 EC units) from 
Lake Bonney was traced from data recorded on 6 
October 1995 at position POL5I7 in Noekburra 
Creek, a tributary of Chambers Creek. Six weeks 
elapsed before the high conductivity water from 
Lake Bonney had been diluted to 721 EC units (14 
November 1995). 

The range of DRN concentrations in Banroek was 
- 7.9 mgl ' and in Loch Luna - 7.48 mgl '. The 
high registrations Occurred mainly after heavy rains 
but there are also nitrate contributions from time to 

IS -I 


little bom waici It.tin fhc Rocr Murray ami lrom 
;mriculliir;il drums, town el fluents ami sfcwiige 
iU-.i har ; vs a-, well ,is localised ailiolvtir bicakdown 
of nitrogen - fixing blue ^rrcn ujgne [AtltlhtiVHtf 
spp K I hvre |s an unknown DRN uuiiieui input (twin 
huec Hocks ot" peitcund <> 10(H) birds!, swans, 
cormorants ami ducks and lessor numbers ol othei 
water birds in Baittuck and Lcutl LutTti 
wcilamls and on (he hanks of the River Murray 
Nlitrnlcs art tU*0 produced bv Itghtlltltg (Nmiih 

The IttglftesL TP level (O.V>5 myl ) recorded in I tu h 
Luna ucisutted on 14 Oetohet \ i ) i >> rfw duy URcr 53 
linn ol iatn Tell ai Buiiucra and 5S mm ai rianrock 

SU ft I. Kolnkieli pet-., eo , IW). Such 

hce.y ram ami conscipicnl runoff eanse bottom 
disturbances in shallow wetlands whit h ^'distribute 
i lo.-.ol ved ui ^aiiic phosphorus compounds atttl 
itioiLiauie phosphorus bound lo suspended or 
il'-tihU-.l bottom ui^allic particulates in ihe water 
column, lin;:;-. cl til, H°S5| have also drawn 
aik-nnon to the complexity ofehenuLnl relationships 
withm wetlands ariil ihe effects of wvalher factors, 
such us Winds and temperature, on water chemistry 

NO MoGWh H I $ I'.u.leiui weie iftsorYftj i]i 

Uanrock wetland, Ihe iriulliplkaiiou ol 
i \unohuctriinl m'IIs in Kaiimek during December 
!■'"! was hulled by Ihe increased water inflow removal oflllC bl»KkilgC 111 ihe fdlCI ptpC. 
The ivanohaeicna were Hushed oul prevcniint! any 
further development ol' a cynuobn^irital hfhnifll ill 
Uanrock weilund lhai year 

lii loth UUIU! I ■v.ntohnetonal COlVt (cached a 
maximum ol 23,700 cells ml ' on 27 Deecmtvr 
Ivl ore declining to zero by 14 March |QV5 Alter ihe 
I'ollapse ol Hie cyaiiobaUemil populiiiiuu III Loch 
I. una mi December I WML a miall but visible hint 1 
iiLVh ..b '.tl hlooin developed downsticain in 
lebiwury ami March |0M5 alone the eastern bank ol 
Ihe River .uli.u uii to I oek ? weir li is likely 
thai (hi:, bloom had ils yenesis in ihe November 
December l l,l| 4 inrll iniilopheaiion in I ,oeh Luna 
iicaie.ini m 1. 1 mi ihe same side ill ihe River Murray 
hintlll blue eaern ulgul blooms had occurred m iU-, 
v'lii' location in previous vivc (I (« k 5 -.i.iii pets, 
ioinm. 1^95), 

Bowling (IM441] repnrietl Ihe occunencc mill 
possiUle e.iuse-. ol* ;i sevnv Anuhitnui t jo tt/nfn 
bloom m Lake CV.iecllmo NSW m IVWMJJ 
whiii cell uunthcrsLAoecuod U"MU«M) 1 ells ml ' Ihe 
ptivsien clieimeitl Jevels in ihe lake in 199(1 had 

1 ■>■-. >'lose 10 the ! L > 4 >4 A \;ihies III Loch I uiui loi 

1- -nip. -mures, pll, lurl.ntlil> ;md eonduebvity 

but lower m;jviin.i m coiicemouioiis ot'I L aiul (DUN 

'lui- ttftfC IbiUVj for I "eh I I Ho\« Iiiim ( 1004) 

1 \ picked the \ it^N 'iboul the L;ike f ;ueelh;'() bU»oiu 
thai ";illhitueh ^eviral umlei lying eaiisivs ol rhf> 

bloom are piob.ible. Hie elevaled nmrkoi 
eoneentralions. espeeinllv ol imal phosphoms. weie 
major Taeiors dial eonlribulcd Lo il" He di\ U 
attention 10 die kai ih.n most plush 1 hcttiftll 
sUulie^ ok cvanobacteiia blooms weie stattcil altn 
Ihe blooms hud oeeuned. 

In this study ot th«- pliysico-eheiiucal properiio ot 
the tvvo wetlands a search lor bloom - foimine tosie 
eyiinob.ieteriii species WfiH M.n'led bebue au> cell 
uuiltipliealion had commenced and a seasonal 
piiLleiu is described- ("vanobaeierial cells niav 
irmain donnani in eohl Wiiiers ami erow besi ui 
WULUI ti'inpcialures e\ceedine 15' I . vvilh opiim..l 
growth rales at 25' C "l hejher (K»»bails & /oluuv 
l l »S5l, The ellecl ol waler k-mpeitiiuivs on 
esanohaclerial cell numbers in Loch Lima, is shown 
Willi data bom Lihlc I which show thai blooms only 
oeeuned between November and Lchruary Nnmlv rs 
ot Xtutinwiui spp. tost- -.(uldeuly Inmi ato on \ c > 
Novembe. I'-W (20.1 C) lo 23,71/0 nil I on 11 
Deeeniber |M 1 U i2-J_2 C) mm\ ihen hcuufl SUbhOhilg. 
cweu diouvh the water temperatures were hii:b. 
perhaps hecause ol exhausiioii ol nuirienis_ 

The risv and (all in eel! numbers may abo be 
relaied to ihe concentrations ol' the two nutrients, 
|)RN and 1 1' recorded during the L-iowth and decline 
ot the l ( ><-)4_5 and l l H),S-o blue eiecn alyal ouibreak, 
liable 1 1. However since Ihe concciHiaiioii ol 
dissolved reactive phosphoms is nol known, these 
lelabonships must be ueauxl with cannon, 

There wen: lour occasions (20 December I9 ( )4, \ 
and 31 Liuu.irv b^5 ,\\\A Ls \pril |p05l ^ ben IT 
eoneentralions in Loch Luna were at or above 0. Ui 
inel ' ,u)i\ DRN was present e\eti In low 
eoiiLjeiniainHis (W. 2.2 Ulgl ' ), AllCt each olthe.e 
e\ .. mi ■-. i here was a use m the numbers ot* 
c\anobaelcrial cells- In October |\WSimean lo.o ( | 

iftClXi were Tour r rr eoneentralions between " -1 

0. 1 )5 myl I but DRN was low or absent and no cell 
inulliphcaliim tlcu-lnpeil 

A pane" ii o 'Inted lo uiurieni a\ai lability and 
temperature can be seen III carl> I Wo, Alter an 
increase lo JI<MJ cells on 23 January l«/tfi the cell 

bus bCg8l1 lo dec line pOSfltbl) due lo very low 

ciMicenlraliou^. ol DRN and Ihicluaime values ofTP 

in the waiei coh i, I lu i-yanobaciena population 

reached zero on bl Mav lb l Ki Thr mean wan-, 
lemperature nn lhai ilnie wms I V f " v/Jnub i^ near the 
minimum temperature rattle I'm .erowth ofmuuv \~tel 
lb mi" evanobaeteria species. 

trom the tiff ru obiamed in ihis study it is siieeesied 
ibal outbreaks of eyanobjcierial blooms did i|oi 
oeeur in (he summers <.(' 1^)1 SilUtl P)0S-o iv^nr, 
there Wftrti uiadetjuale eoiieenlralious ol IT aid 
DRN ui Ihe waki cohunn durine the perisid ol 
la\ourablc yrowlh for cyanobaeleri.i f>ata liom this 
s|ud> imlkate that I P coucentialioii aluwe 0,M\ nt , l 



urjd DRN concentration at or above 4.0 mgl ' in the 
wetland may provide lor continuous growth in the 
Anabiiena species. 


This study was funded by the Murray - Darling 
Commission through the Natural Resources 
Management Strategy, Project S3I2X. 

I am extremely grateful for the assistance from the 
following contractors: Mr M. C\ SchullA Glossop 
High School Tor the physico-chemical analyses. Mr 
P. Christy and staff. Australian Centre for Water 
Quality Research, PMB Salisbury SA for 
identification and enumeration of eyanobactcrial 
species and Messrs R. I. Gropler and J. A. Pillar for 

their regular and supportive help with field sampling 
from a powered dinghy. 

My thanks to Mrs B. Page. Tilley Murphy and 
Hughes, for accounting assistance and payment of 
accounts. Mrs J- A. Rourke. Adelaide lor preparing 
the manuscript and Mr G, Wright, SARD1 West 
Beach for the preparation o\ the figures. 

Mr B. T. and Mrs E, L. Engel former owners of 
Banrock Station and BRL Hardy Lid, the present 
owner are thanked for allowing access to the 
wetland. Mr C. 1- Rohrlaeh, Manager Banrock 
Station and his staff gave valuable assistance 
throughout the study. 

The author thanks the editor. Mrs J. Bird and the 
two unnamed referees for their constructive 
comments on an earlier draft. 


\i\ik\m\ Nmikl Conservation Acilno <IWi) "A 
dictionary of importani wetlands in Australia' 1 2nd edn 
(ANC'A. Canberra). 

B0W| ini.. I- ( 1994) Occurrence and possible causes of a 
seven. - cyanohaclerial bloom in Luke Cargellico. New 
South Wales. Ausi. I. Mur. Freshu: Res. 45. 737-745. 

Bkm.s. S- V., Maiiir. M T. & CwnNfFR. S. M. (J9S5) 
l.ininological studios of waterfowl habitat in souih- 
wcsii-m N.SAV. I . Water chemistry. Ihiti. 36. 59-67. 

Goonan. P. M.. Bllk. J, A.. Thompson, T B. & Sun r. K 
J. t I W, Wetlands of the River Murray floodplain. South 
Australia, I - Preliminary survey of the biota and physktl- 
ehemislry o\' ten wetlands from Chowilla to Muiinum. 
Trans, ft Soc, S. Ansr. 116, 8 1 -04. 

IJAUI (l l )H4t "Instrument Manual" 1st edn (Hach. 

(}W2) "Water analyse. handbook" 2nd edn ( I Inch. 


Hmso l 1940) Enumeration of algae, estimation of cell 
volume, and use in bioussuys 1990" In the series: 
Methods for the examination of waters and associated 
materials (HMSO, London). 

Mackay. N. & t-ASTiiURx. D. (Eds) (1990) "The Murray" 
(Murray - Darling Basin Commission. Canberra). 

Robaris. R. D. & //in artv. T. ( \%7) Temperature effects 
on photosynihehe capacity, respiration and growth rales 
of bloom-forming cyanobacieria. N/.J. Mar. Freshw; lies 
21, 301 -399. 

Smith. R. L. (1*W6) "Ecology and Field Biology" 5th edn 
(Harper Collins College Publishers Inc. New York i. 

Si. ihR. P. J.. OODWAN, B M-. Bli-.r. J. A. & Iuomcsov T B. 
t(993) "A biological and physico-chemical monitoring 
study of wetlands from die River Murray floodplain in 
South Australia" Murray - Darling Natural Resources 
Management Strategy Wetlands Management Monitoring 
Program. Project No. S0(V_ Pinul Report. Australian 
Ccnlrc for Water Quality Research Report No. 7/U.L 





By Peter Kolesik*, Rebecca WHiTTEMOREf & Helen M. SiACEf 


Kolesik, P., Whittemore, R., & Stace, H. M. (1997) Asphondylia anthocercidis, a new 

species of Cecidomyiidae (Diptera) inducing fruit galls on Anthocercis littorea 

(Solanaceae) in Western Australia. Trans. R. Soc. S. Aust. 121(4), 157-161, 28 

November, 1997. 

The fruit galls on the Western Australian yellow tailflower, Anthocercis littorea 

(Solanaceae), reduce the reproductive potential of this plant, but their causative agent 

has, until now, been unknown. Our research has shown that a new gall midge species, 

Asphondylia anthocercidis, induces these galls. The larva, pupa, male and female of 

the new species are described and illustrated. 

Key Words: Diptera, Cecidomyiidae, Asphondylia anthocercidis, Anthocercis littorea, 

Western Australia. 

MttmtA ft? Hie Kt'Vttl fatttfl W V, kwti, (l'»7). 1 3 1 1 4 »ft7 Id I, 






Koi lsil. P A Whii II mori. R.- & Srvo*. II. M. \\W) A>i>h<>ml\Iin wnhwwSiti. tt new spivics (if 

( .vniomviidiiL- iDipk-iu) inducing Iruil gulkun rt/wVwfWft fimtrm fSbttwwuHC'i ta Western AuMrulia. low* 

ft ft*', ,V A/ftrf. Bll4)i 157- W»l. 2.N iWo.ibe.. 1 0*1 

The huu tkuTs »m Lhe Western Atwiraftap yetltw uuiiituvc Afltfv«.v/vtf< totiorwi Labill. ( Sol;tn;<cc;ic» reduce 

ill,- reproductive potential nfthto plain, bill theii causa|iu- ;ie- nl has. until mt\v hocri I nknown. Our research lias 
-In-wii Ihai a new call midge •(pooie*. Anphotutyhti unihtn mult*, induces these »alK The larva, |>upa, male und 
Irmule ol'ihe new qweks UltJ described and illuslralcd. 

Kn Wouos; t»ipl<_*i t < Vcidun»yiit.Lic. .\.\fhtinti\hu tt/ttluHriihlis, AwharflVh lifKuvti, WcsLciu Australia 


11k gull midge fauna of Western Australia fs 
poorly Known, with only two species h;ivin t u boon 
described previously (Gagne \HJBiY OjVtS ol them, 
lj>om\t<i honwrnissrui Colless, is a species which 
presumably leeds on lungi growing ifl lhe soil and in 
leal liUer (Colhss I9hu The biology of lhe second 
species, li.'cini ini'tnia <msiral<tsnti' hell is 
unknown (FteJl I9I5K although this species is likely 
lo be a plant leeder considering ihat its congener, K 
nudthskii Kolesik. causes galls on Km uh puts 
{<i\<ttttlo,s(i in Smith Australia (Kolesik 1995a), 

The yellow taiHIower. Anllhhrnts litlnrcu Labill 
iSolanaccac). U shrub which grows lo 3 m. is 
endemic lo the south-west coast of Western 
AuMialiu, primarily on calcaicous sands in disturbs 
habilals such as recently burnt areas, roadsides. fi(t 
breaks and cleared lols iPunfiu el al. t982 
Wfctanujrrfc 1 r The (tuil galls rtfl A* hnatva have beer 
known lor some lime (Purdie ct al. 1982 ) bill Iheii 
causative aecrU has remained unknown. In July N9o. 
one ol us (R. W.» collected IVmi galls from A liHoren 
coniain'mg larvae and pupae, from which adulls were 
icaicd, The gull-mducer proved to be a new species 
ol' gall midge winch is described below. The 
development ol lhe galls and their impact o\\ lhe 
reproduction of A, IhUava are described by 
W hittenu've 1 . 

The genus ,\\('fiont/yliu in lhe context Of this paper 

|) L |iaiiia.;ai ill I IttriiiJlluirc. Vilieiilliuc iUk! Ocin,i|jj£} liuuh\ Qj 

■V*ri< ttllnnil and KtftfUHti Sue-tun.". Tin I ItWHrffl) Ul 

^Udiinle I'Miu ni,.M n-.Mh.n.i s vi^j, siwo 

1 1. ■ ( i.ii-i!ui-ni it] |-l.:kiii;,_ l_ rivvisily i'l' Wl-Mcrn Ausluili.i Ntu'kiiul- 

W Anxi htftlT. 

Wkll'l rj MVi R < WWt) As|u-lK ol lhe itisct (•HulutvU I mil gull-* 

an>l ivpdHlik'liw Fmilltg) \f| irtUufn'tt'U tfaflfftl ' >• d;to i 

RSf (IIuiiv)IIicmv. i (ItVW 113 "' WiIMWT \ti-,iial(a iimpiilv). 

is Uelined by Kolesik 1 1997) The new species is lo 
heuuribuied 10 I' K 

Material and Methods 

Stems ol Aothuufrvh itH<>rc« beanos Iruil ga!K 
were collected at llillarys. about 20 km north easi ol 
Perth, on 23.vii.l996. A small number ol" palls was 
dissi^ctetl and the larvae and pupae preserved in 70' v 
ctlianol. Larvae and pupae retained within galls sure 
ivared lo adults on slenis which were kept in plaslic 
vials. Larvae pupaled within lhe L'ulls. nnierged -jail 
midges were preserved logethei with pupal skins in 
70'.' elhanol, Canada balsam mounts ol the lypc 
series ror microsc<>pic examinatii'U were prepared 
H< cord j ny lo lhe lechnic|ue ouiliried by Kolesik 
ll9^5al. All measurements teler to lhe type sciics. 
The type specimens and other material regained in 
70S elhanol are deposited in the South Australian 
Museum. Adelaide tSAMA) and Australian National 
Insect Collection, Canberra (ANICl. 

Asphottdvlia unthoevrcidis sp. \\o\ 
(1IC.S lis) 

liniotxpc J* llillarys. Western Australia | M 4K' S. 
115 45' C|, emergetl 2K.vif.l3M, R. Whitlemore. 
ivurcd froill larva I'rom Iruil gall on Anthinrra- 
li/fotvu Labill,. gall collected 23 vii 1996. I2IW 
I SAM A 1. 

r<tnn\t>i<\: 2 ■: rj, 3 r Ll 3 pupal skins |SAM\|. > 
.' ,;. 3 ? v . 4 pupal skins |ANIC|. all same data but 
emerged 2H-30.vii. IW>; 2 larvae |SAMA|. 2 larvae 
|ANIC|. all collected with holotype, 

Other ttuih'tntl l*\\ SAMA|: 2 ijrf, 5 

1 pupal 

skins, a pupat-, all same dala as paratypes: 2 larvae. 
c(»llccled with holotypO 



Figs 1-8. Asphoiulxtiu aiithocenidLs sp. nov.: I - 4 mule; 5 8 female. I. Head in frontal view. 2. Last three flagellomeres. 
3, Genitalia in dorsal view. 4. Gonostylus in posterior view. 5. Basal lobes on ovipositor in dorsal view. ft. Hud of 
ovipositor in lateral view. 7. Lasi five flagellomeres. 8. End of abdomen in lateral view. Scale bars = 100 ^im 1-3, 5-7: 
50 pm 4; 500 mil S. 





Figs 9 !5. Asphotuixlia <iiuh<Hvrculis sp. nnv.: 9 - II, 13 pupa; 12, 14 larva; 15 infestation. 9. Anterior part in venlral view. 
10. Anterior part in liiteral view. I I. Prolhoraeie spiracle. 12, Head and first thoracic segment in ventral view. 13. LaSl 
abdominal segment in ventral view. 14. Last two abdominal segments in dorsal view. 1 5. Fruit gall on Anlhocercis littoivii 
Labill. Iredrawn from Rippcy & Rowland (1095)1. Scale bars = 500 uni 9.10; 50 mn II; 100 um 12-14: 10mm 15. 

I M.I 



Xhttr 1 1 iys l-4> 

Colour; sclcrou/cd p.trls of body reddish brown 
■ion *cleroii/ed parts of abdomen giey. 

Mi_ad. Anicnna: scape hmadcsi dislalh, [,7-2,0 * 
breadth at distal cud, 2.<v2.7 \ length nf pedieel: 
pedicel width 1,2 1.4 \ lenglh; first llagellomcrc I.X- 
2.1 \ length ol scape, flagcllnineres evenly 
cylindrical: cucumfda dense, equally distributed 
along I'lagcllomcres. I ye facets hoxagonoid. c\c 
bridge .S-°- laeets wide. Irons with 10-17 setae per 
*uie. I.abella piomincul. lalerally wilh 7-10 setae 
sclulose. Maxillary pulpus 3-segmcnled. scumeui 
successively and progressively longer. 

lliono, WjW length XA mm (range 3 1-^.7) 
widlh 1,3 dint 1 1.1-1.4), Sc cell pigmented 
proMinally. C laws (if all legs subequal in si/e and 
similat in shape, as long as cinpodia. 

Abdomen. Genitalia: gtinmiylus wilh two large. 
apieai iceth ol same length; aedeagLis elongate and 
narrow, reachuiL middle o\' gonostyhts. 

tti»uit (Pigs 5 Xf 

FfOIU With 9-Gfl selae per side, labella with 7 -0 
M.-iae laterally, Circumfila comprising two 
longitudinal and two short transverse bands- Wing 
length *jft mm (3.3-33», width 1,1 mm (1.2 I. M. 
Seventh abdominal siermie LS I 1.6-2,2) \ length ol 
sixth. (Jcnitalia' ovipositor l. ( > K (I.K -2.0) length of 
seventh Mcrnitc; basal Johes on ovipositor broad M1 
doMaJ view, divided in posterior thud medially; 
luscd ccrei glabrous. 

Pufni iFign' 9 It. 130 

Colour brown. Total length 4.0 mm (3-b4._S). 
Amennal horns not serrated, 242 uni (237-247) long. 
One upper and three lower Irorital horns. Prolhoraeie 
horn slitrhtly emved, basal part ahoui 2 v widlh ol 
lermmal third, lerminal third setose. Abdominal 
dorsal spines simple, straight, with 2-3 pails Ull last 
segment curved laterally. 

Mature lurvu (FigS 12. 1 4) 

Colour: yellowish-while. 'lotal length 3.4 mm <2.b- 
4,1). Head capsule strongly pigmented, postero- 
lateral extensions not developed. Spatula with four 
anterior teeth, inner pair smaller Lhan outer, shall 
narrow, broadened both at middength and base. 
MiiuHindcd anteriorly and lalerally by B&tciwlvu 
pigmented aica, hach side of spalula with triplet and 
pair ol lateral papillae, all setose. Siv lerminal 
papillae present, one pair coriulorm, 2 pairs with 
4iort -eiai- other papillae .i\ loi A.\f>h<>niivliti (Mohn 

Coll tnuf hn>h>i>\ 
This gall undue induces deformation v\ fruits of 

Antimenk tiUowa The ttiticarpeltow ovaries m 

translori ned into glabrous, spherical W OVflti 
nipplcdgalls, 7-IN mm loiig and 7 13 mm wide (I ig. 
15) -wm\ bright green to purple in colour Inside the 
yall a chamber, about 3 mm long and 2 mm wide. IS 
occupied by one larva. The chamber is always lined 
with lungal mvcelia. Although Ihe fttllgUS was 
abundant in the many galls examined, no sexual 
stages were observed and the fungus remains 
unidentified, Viable seeds are rarely produced in 
jjalN although pollinahon is essential to Main (he 
gall on the plant. The numbers of galls in A. littorea 
|)o|>ulaiions are often vciy high, with (he galls 
outnumbering ihe urn malls developed fruits by up 01 
^X times (Wlnllemoic I. 

Pupation takes plaee within Ihe gall. At the end i>f 
its develo[>meni ihe pupa cuts an opening in the gall 
and lifts most of its body outside the gall. I he pupal 
skin then splits open aiul ihe adult emerges. A\ 
llillarys in 19W*. the adults emerged throughout die 
entire host plant flowering period, i.e, from April in 


A^ftwuthli't (inthtntiviili.\ sp. nov. fa sympulrn 
wilh .3. iiitorai across the emire geographic 
disinbuiion of the host plant. v\lnch ranges from 
Kalbarri |27"5<>' S. I WW H| m the nOflll tO 
Israelite Bay | U 27' S. U^2.V \\ in the soulli 
(Whitlemoii- 1 ), 

Tlic u;ime is derived from Llic generic name Dflhv 

hiisl [ilanf 


A\f>!t<>(hlxlia is a worldwide genus wilh six .species 
previously described from Ausiralia. The life histoiy 
Of ihree of ttiem is known: A lk/ituttanw Kolesik 
induces jialls on leaves of fhuh'tutta v/v.o.Vi/ Jacq. 
subsp srtitlmhtht (Smtdn Wcsi. nm\ A tnfliiut 
Kolestk and A. criiijonnis Kolcstk induce galb oh 
branch segments ol Hulosurc'ut {H'tynmitUtUt (Black ) 
Wilstui subs[i. fH-rxmiittiuni and //. itul'na >ul»sp, 
Mtotttvtiya (Benih.l Wllsotl, respeelivelv I Kolcsik 
]£>95b, 1 097K l-i'e 1 hisloties ol two oilier species ,\ 
h>n\i Skuse and ,3. >nhttwul<i Slutse. are unkuowii 
(Skuse I SSM, hVMM, The temainine specie-.. A. hilli 
pLtwards. has been reported to mdnee i-alls on die 
sieni ot' an unidenldied plant <l:dwaoK IMKo 
Aspoiulxlia tiflti. A. iorwi and -\. ruhkmukt are m»| 
eitnsidered in the present paper The deseriplions ol 
these three species were superficial ami therefore ii i,\ 



no! possible Lfl compare them vvjih each oilier wr with 
y\, (If'tlotuwtiv, A. inflate; A. ericiformis or A 
umhfU'i'tridLi. A review of the Australian species ot 
this genus is planned by P. K. 

The new species differs from AfphOUifytht 
Jfn/otnu'tw in the longer adult scape, the wilier teeth 
on the gonostylus. the nnserraled nnlemial horns and 
the presence of holh upper and lower frontal horns in 
die pupa, the shafted spatula and the presence of a 
pigmented area around the larval spatula. The new 
species can be distinguished from both A. injlctia and 
A, crii-ifttrnii.s by several characters. In A, 
(tntluKt'tr'nlis. the aedeagus reaches the middle of 
gonoslylus. pupae have three lower frontal horns, the 
prothoraeic horn is setose at the distal third and is 
about twice as wide at ihe base as is the distal thud. 
Al least two ol the dorsal spines on the last pupal 
segment are curved laterally and the spatula has Wmv 
anterior tcclh. In holh A. injhifn and ,1 crictfonnis, 
(he iiedeugus evtends beyond the middle o\ Ihe 
liotiusiylus. Ihe pupa ol A. inflate has one lower 

frontal horn antl that of A. chcifitmiis has none. In 
both species, the prothoraeic horns are asetose and 
about four times wider al the base than al the 
terminal third. In the pupa of A. inflahu only the 
prominent pair of abdominal dorsal spines on the last 
segment is curved laterally: in A. ericijorniis all 
spines are straight. In both species Ihe spatula has 
two anterior teeth. 


The field work was supported by the University of 
Western Australia while R.W. was completing hei 
Honours degree. We are grateful to J. D. Gray. 
Department o\' Horticulture, Viticulture and 
Oenology University of Adelaide and R. J. Gagne. 
Systematic Entomology Laboratory US DA 
Washington DC for their comments on ;in early draft 
o\ the manuscript. 


O'Miss. I). II tW65) An extraordinary Aufflratiao 
( 'c'Lidoinyiid. possibly related io ilie \ymphomyiidae 
iDipler.i. NenutoccinV /'»vr_ A' nil. S,u .' /"//</. IB* 34* 
145-1 4<3 

I-uwakos. F. W! I I u Kt1 Iv.u new Ausinihiin Dipiera. Antl 
Mug. Nul Hisl, 1(U 49K-501 

Ti ri, \i. P, ( 1 9 I*5j New ge leru and species of uall iUhIlvs. 
ftw. OS rutin. A/^.48. 145-211- 

CiAtiNlf, R. T ( IW>) Family Cceidomyiidae pp. I52-1W /</ 
Fvenhuis, N, L. (Be!.} "Catalog oi Ihe Diplera til the 
Australasian ami Oceanian Regions" fBishop Museum 
Press am! F J Brill. Honolulu). 

BCOI I SIK. P. ( 1995a) A new species of Eiuim-ticttrtiHi Felt 
flJipu-ia: CYvtdomviidacl on Ematspftts fasvuulosit in 
Solitj! Australia. ./, Ausl, ait. S<n\ 34, 147-152. 

( t995bj Atiphwidylitl i/ftlomivnc, u new species of 

Cecidomviidae (Diplera) damaging leaves and 
hnutehes of hop-hus|i. Doi/otitii'ii \ l L\COSU (Sapmdateae) 

ill Australia. Tmm. ii. .w. s. aum, nv. i7M7n. 

— < 1 1-J47) Two new species o\' .Xsphinulyiiu 
(Oiptera: Ceeidomyiidael from Hnlt>\aicia spp. 
iChenopoduieeaei in Smith Australia, thiti 121. 59-66, 

Momn. B. (1055) Beilrage /ur Syslemalik der Iarven der 
Ilonididae (=Ce<jidomyiidue, Oiptera) I. Teil: 
Porrieomlyhnae and Itonidiiiac .Vliiteletiropas, "foitrfvgit'fl 
105. 1-247. 

PCRDlb. R. W.. Symon, D, B, & H\i:r;i. I.. H°S2l family 
Solanaeeae pp. I-20K In George, A.S. d-d.) "Flora of 
Australia" Vol. 2 { ) (Australian Government Publishing 
Service; Canberra). 

RiiM'hY. E. A: Rowland. bV 0995) "Plants i)\ Ihe IVnli 

Coasl and Islunds" (I ^niversily of Western Australia 

Press. Nedlandsi. 
Skusr. F. A. A. (1NS8) Diptaa ol Ausiralia Purl 1, frvr. 

Lmn. S<«. N.S.W. (2nd Series) J. 17-145. 
— ( !N°0) Diptera of Australia, Nematocerj - 

Supplement I. Ibid. 5, 373 4 12 




Brief Communication 


Blowflies are well known for their ecological, veterinary and forensic importance 1 but 
they are also significant medically as mechanical vectors of dangerous pathogens 2 . 
The Oriental Latrine Fly, Chrysomya megacephala Fabricius, 1794 3 , is notorious in 
this regard. Adults of the synanthropic form of this species (see below) are attracted to 
foodstuffs, human and animal faeces and carrion 4 and have been implicated in the 
transmission of viruses 5 , protozoans 6 , enteric bacteria 7 and helminths 8 . The larvae are 
also known as facultative parasites in traumatic lesions in humans and other animals 4 . 

iHMs.UiHmsnt tin ftnval Wrf/v r </ K A/fiW. [WPlV I21< Il W3 Ihl, 

BRjbf Communication 

mrs i record of the orient vi , i atrine fly, chrysomw megachphali 


KlmvTlicN are well known fbr Hum efcplngieaL veienuar. 
and lorcnsic importance 1 hut ihr\ arc also sii.-nilV.nii 
medically .in mechanical vectors ol dangerous pathogens 
J In- Oriental Latrine II). Chiwnmv,! im ^uvphulu 
I abricms. I ,Nl\ in noi.HtoUs in this, retiard. AvitiJis of rhe 

N\Mai>llllOplC lorill 111' !hK SpCClCS <SCC IvIoW'l LUC aitracied 

to loodstulfs, human ,iii<l animal faeces .mil eairjfm* and 
haw been nuphculed in die transmission ol viiuscsl 
I >n n it/nans 1 . cnitilitc luck-tu Qnd helminth -A The lai ;v«u arc 
BbtU known as tacullame paraviics in liaumaltc IfttffMK in 
humans .nnl uihei animals' 

I irpnii here iho lirsi rccoid ulC uhx-.t, tphitttt in South 
AuMrali.i, It w«t cauelu mi |upj<c number, in a liver-baited 
hap .Hi ilir Nmih Iciruec cJimpUN of tin- Unhcr-aiv rll 
vli kude in April, I W. ) NcnJ H(iE ftr^viou^lj encountered il 
Ul litis sue. dcspiic periodic trapping • >! blowllics ove. the 
ptccedine seven years Nctlhet had I encountered il 
cKcwIu'iv in South Anorak". There are lie specimen* < C 
t»n</u'i'pti/iln trout this slate in the entomological ca.lleL-iiou 
H ilir South Australian Museum anil the closest iceorcl ill' h 
10 South A iisti, .ha is lhat <»l three |c males at 
Murriinihuieniau. New South Wales Ml 5H' S, I 4W 03' r ). 
approximately 950 km to bic casi ol Adelaide <K R. ISorris 
Divis.tmol r ; ii!oino|oi>v C'SIKO pels. eomm. 1<' ( J7c 

lUewhere in Australia, C. nnvu< c/'W// occurs across the 
lull north Hi the continent. down ihe cast coasts of 
Uneensland kind New South Wales and m south-west 
We lirii Australia 11 has not ken recorded ftVUII Vicieaia or 
fnsiitum;i '. 

( 'hn\"in\<t mvy,tu r/tfmln r, piobahlv a leevni 
introduction io South Australia, fcfr it fe aeli\ely expanding 
its rinv-c in iftlner pints ol' lite world. Siikv the !')7tK ii ha- 
invaded New /valand 1 ". several parts o! Aii'ica unJflTCTn; 
ol S^tilb 1 - * Vnlral 1 ' and Notlh Aineru M 1 . It is also found 
ill Japan and is widespread Ihnut^hoiil tllf OrK-nlul tc^ion 
and the AustroM,iiay,tn and Polynesian suhiv^mus ol the 
Austoil.isian ivi'ion 1 ", 

('lny\i)tit\'ti Hh'tyh rfUutft' 4>lviiis 111 two lonm. \\hnh aiv 
morphologically and eeol(^;jicallv ildhreni ]'hes have hah n 
JrlMhtl bs Kiiiiha'-hi' as the* "notni.d" and "derived' 
(onus I'll*: tioimal Ibrm U VGWjitltol 'o Iroptcal loivOs on 
South Pacific iSlotld> honi ihe Bismarck Atehipclayo to 
Samoa. Il i% tulnvcd it. In* Lhe plcsiomoiphi.. loim ol the 
spi'ues. flic derived form i'^ synandiropL aiitt dispcrsivt 
and \h ihoindii to have originated in [\ipua New Ctiiinea on 

the western boundary ,>| H,r Npectes 1 
disinburion 1 " The individuals collet ted from Adcla.dc an 
• »r ihis form. 

Ihe derivet! hunt can k distiireui-hed from the normal 
form b> the :jrcally^-ul:avcd ommal.dia in die appci Iwi- 
thirds ol' the eye in ingles I'be notnial lorm has rtrtlj 
■.lichdv cnlai;jcd omnialidia ill this reeion 1 Swkv i\w two 
1'orms di lie i* so miakedl> in iheir isolosieal pid'canLCs >\\hl 
In Wllie aspects nl their morphology "hev ina> be 
sHlTicicutly divei>;cni jdeuet;cally U> warnml siibhpeeilie or 

CSCM specific sljlu:,. Thr mc-'li- br K'v,\kd b\ bio: heillKal 

ai)al\sis. as earned Hill recently lor othui cloudy-it'laled 
bli»wClie L - ol' lite yeniis Ctillip/u'rn 1 '. alibtaiph a detailed 
morphological onupansoti would also be required betoie a 

h " aboui ihr-ir statu- could Ik- ieached_ 

l.h.lh lorui:-, ol( mi qt&Cpkllti *-Att be dislmyUislied trout 

all other known Australian species ol Ckrywinytt hy (Iw 

tolli-\\ii\e moipholoy-Kat characters; hairs on preulut knob 
black. Ioiujci lhan hm^rhi of knob: anienoi tho.aciL 
•piiaclee blackish hnHVli; legs black, sLibvibnssal sciiikie 
black: eyes m males with ommatidia in Ihe upp't iy\"-thirds 
enlareed and sharply deniatcaled trom the small ones in the 
lower third; Irons in temnles widei in ihe middle" 1 ' |B . 
However the species , s ^,miku phys|o|.,^ie,,||\ to othei 

species of t'lin,u>fn\ii m that it is the plultc. and 

llicrefoie. m South Auslratia. il will i>iCMii>t,d>l> be AeliW 
only ikirini 1 ihe warniei months ol the >e t n. Hut is beiween 
October and April. 

BecjLise ol its habits. ( mcktHVptuJlit ilcserscs serious 
aiu-nbou lioni a public health pcrspeeiive and iU 
disinbution .a AusiuIili shoul.l therefore be tnoiiiiored. lis 
kavac cinild also tv encountered in loieusit eases m Ihe 
,\dehude rcaton; but hecaiiic ihcu moijiholo^v bears 
superheat siuiilanly loibal o| some olhci common tWIItIM 
breedmy species of |i|(twllv". ihey COuUI be mistaken loi 
I hem. Since the rale ol development off Iflt'tftiVt'phtJitt Ul a 
.t.'.u'ii diflers si^nifieanlly from lliese olhet 
species^' . misidcnlilicaiion ol Lhis intpoitant |l\ could 
le,id to seiious eiiors in estimates ol the time since death ol 
human corpxcH. 

I ihank Uis I'), A Huekhottse. K. K. N.mts and A K hud 
lor Iheit useful comment on the iiiauuscnpi The aiuhoi 
also tluinks l>r Nonv. lor Ins belpbit advice and assistance 
in ariiiutjui!.' die loan ol specimens. 

I.i/fiulmulu, /. ( l'.<"M |;h allies iKiehmoud 
I'ublisluiiy i 'o lad Sloupln, 

(irei-nhiTU, It. t IQZJt ' flies ,hk\ vol. 2. Iin»b .>*> 
and Incase ltan>mi.-;sion" tl'ru.Leton L'niscrsii) l^.ss 

I ihriiiiis. ,|. ('. (|794) "LntomoloiTaa s>sientatiLa 
euiendala el a.uja Secilidum el.i-^o, oidmes- ivmr., 

• p.' t..-. ad|cetis synoiiimis. Ions nk.ersaliniulni , 
de-uir-iionibu./ lomelir-Ci, IVoli, llafuiaer 

/u.upi, l\ ( l%5j"WyW> in Man and Animals m ihe Old 
W.nld " tLUtiierwortbs. I.ondoni, 

1uiiauetlt». S. VI. R, ( ampus, M, I., ( ., ||;irsi. i . \]„ 
ISurulli, f;. M. X Ishihala. G, K, i hMi lies Murobn.l 
15, I7<M74 

H;irris. \. II. & Down. H. A. < I l M*o Am. J- "l-op \k- ( | 
2n. 7Scj_si)(). 

l.imu. M. I.. V. S. X In/., I-:. i I'M I i Ada Jiiof 1'amna. 20, 
f.l Si 


x Monzon, R. B., Sanchez, A. R., Tadaiman, B. M., Najos, 
O. A., Valencia, E. G., He Rueda, R. R. & Ventura, .1. V. 

M. (199!) Southeast Asian J. Trop. Med. Public Health 22. 

"Spradbery, J. P. (1991) bt A Manual for the Diagnosis of 
Screw-worm Fly" (CS1RO. Canberra). 
"Herman, T. J. B. (1990) Weta 13, 12-13. 
1 Braack, L. E. O. ( 1991 ) Onderstepoort J. of Vet. Res. 58. 

1 Guimaraes, J. H., do Prado, A. P. & Linhares, A. X. 
(1979) Rev. Bras. Entomol. 22, 53-60. 

Kurahashi, H., Wells, J. D. & Ogino, K. ( 1994) Jap. J. 
Entomol. 62. 860. 

'-'Wells, J. D. ( 1991 ) J. Med. Entomol. 28, 471-473. 
1 Kurahashi, H. (1984) Dispersal of Filth Flies through 
Natural and Human Agencies: Origin and Immigration of a 
Synanlhropic Form of Chrysomya megacepliala pp. 37-63 
In Laird, M. (Ed.) "Commerce and the Spread of Pests and 
Disease Vectors"* (Pracger Publishers, New York). 

'Kurahashi, H. (1982) Monosr. Biol. 42, 689-698. 

1 7 Wallman, .1. F. & Adams, M. ( 1 997 ) Aust. .1. Zool. 45. 337-356. 

*Dear,J. P. (1986) Fauna N. Z. 8, 1-86. 
'"Wells, J. D. & Kurahashi, H. (1996) Med. Entomol. 
Zool. 47, 131-138. 

1 Wijesundara, D. P. (1957) Ceylon J. Sci. 25, 169-185. 

'O'Flynn, M. A. ( 1983) J. Aust. ent. Soc. 22, 137-148. 

.1. F. WALLMAN. Department of Zoology, The University of Adelaide Aust. 5005. 


Brief Communication 


Cyclostrongylus medioannulatus Johnston & Mawson, 1940 was originally described 
by Johnston & Mawson 1 from three females found in the stomach of Macropus thetis 
(sic) now Thylogale thetis collected from the Burnett River district in Queensland. 
The authors commented that the worms differed from other species of 
Cyclostrongylus Johnston & Mawson, 1939 only in having a narrow supporting ring 
around the buccal cavity. Mawson 2 revised the genus and considered its relationships 
with related genera. She found the type species of Cyclostrongylus which had been 
erected in 1939 3 , to be identical with that of the type species of Oesophagonastes 
(Johnston & Mawson, 1942) and placed Oesophagonastes in synonymy with 

ttttn.Mtt !>t>nt <'l tlw Rnvtif StH Ivty ifi& Anst, ( \Wf\ 121(4). 165, 


C \i h\iiwi^\tn.\ iiht_lu>uitunkttti\ Johnston & Mnw.soii. 
l')4d was originally described hy JohnsUm & Muwmhi 1 
from three IcMintlf, found ill Ihe stomach ol \lti< V7J/ZR4 tltftl,\ 
(ml-) now ihvhr^ih ihttis col leu led Irotii the Burnett River 
district in Queensland. Tlw authors commenled thai uV 
worms differed from oiler species uf <~_\>-lt>\nw>i. , \iu.\ 
lolinsiiHi & Mawson. |£>3V only in having .1 tinnow 
supnorttne litBJ around Ihe buccal cavity. Mawsoiv re\ 't%eit 
tin -jcilus aild cuilsideied Us relationships With related 

genera, She found the typo species nl CypJifswigyttti 
which had been erected in l*>'W m bffuleiukul with rJttn ol 

(he type species of < h'\itf>lhi^i>rtn\tf'\ (Johnston A: Mawson 
l l HJ> and placed (U's>>i>lnii,'i>ini\ii\ m synonymy with 
t !■( I<>\tn'tu>\hf\, Mawson' listed the valid species t)| 
( \tlo\honuYltt\ a- C- »(ilf->hi(h' Johnson $ Mawson. fV^y 
(type speviesi Micuiriiii! in Wallah/it htpsltW. C. HCfllarrfi 
Johnston & Mawson. I'M 1 ) occurrinc: in \tat ntftus 

mfvitriwm. < kiirUma (Muwwut ltf?51 occurring in Mi 

m^Ltm and flif m)tfi;ri.\t'u-\. V. /e/'/m ' MaWsOH. 10fr5 * 
oil urriny in M- 1/0/ w//m and ( ptirntfi John -Ion A 

via\\:.«'M. I'H'j tjccurriijg irj A/. jjarnm> 

( H tllC OtllCI SpCUCS plcVioiJsly USSlLJliL'd U' 

( vti<K\tnint;vlu\. C <li\.snt>ili\ Johnston & Mawson. ] i )\ i > 
occurring 111 W, f>in>(or was referral Iv 

\hi< H'pitslH'H'JxIoitlts f Vaiiiajuiti. I96T 1 tflld C citlomli 
Johnston & Mawson. IW occurring in Af w«f/w now A7 
\>\\>iinu-if\ in a qam genu* later vlencrifted .is Altwosumm 

Mawson. IM70 : Stibse.(|iic itlv C T iittflanh wus rcdcsenbei.! 

bs WttlttihttH'ttni ftulfantPl 

lii kite iafte oTC itic<litHinnul<uu t \ Mawsoii'' h'lind Iti.u ilic 
'cpotfuWU kilx-llrd ;U- Hk lypi,- was a IVnuiltJ Rfigttpfuirytft 

iw\thtli\ (M.-niiin'-. \^lt>i, obviously placed iherff in error 
Siiiec die orieiiial inaluiial could liol he louud she deetji'ed 
Ihe >peci.> ;i \{u>i vVv inttiutwufa 

ViLiWMHi' also revised the ^etiti.s Mtn'mfh^tronvvlits 
Yorke A: Mapleslone, |^2/i fcTflU eiveled three new yenen 

including Piri>*n j ct$rKtm$*lii& Ptipov&siiwiftYtus was 

i_luiiacleri>etl hy ait uesitphaeux Willi a eVlitkll it al tolplis H 
nai rim isthniiis and ending in a bulh; and a hueeal eapxiile 
whuh is ihicke^i .u midlen^ili ami liiis an anicrior holder 

Without projections, fnc speetc- ntcUided the type speeies 
P. walluhiut (Joliii-sLini & Mawson, l L l_^Mi, (.ceLHtiilf in ,V7. 
nilogrisvtis and W. bfpflttiK P. ('curso/ti oecurrin^: in 
PcrrVMUlt! tiiinafis. M t n\><Hii and A/. ntl<>yjiuus .tnd P. 
irnui DCLurriny; in M. inmt. BevcndiiL dien jincndcd die 
genetic JiiigHOHih, cMended the hosi range oi P. (Utitsoni 1o 
include Mm fnft,itin<t,ui.\ and cte.scrihod tv, o new Speck's, P. 
itHh ntpixPis oceinnnt' ill \L i>ii</tii/t'ns, Af rufits and M. 
ttthtt\ttt\, and /' th\i<>xnl<' occurring in Tkvlti$at$ 
SliXiHatiru, I hnuiii and Pnn)^a(c pcrscphutu HcveiidLV 
QtTmrneilted ituil /' thvhmtth 1 occurred commonly in /. 

sfixmdfuti 111 mtitliciu Quccilsljiid. ami Ifi P< In-^nh' 
pmeptami Iroui eenlial Queensland hut hud uui heen 
found in /; ,uti!i>ttitnti or I. fkcHh in soudiern Queensland. 
SpuU v! tii listed C ttuJiiuinnuhitu^ as .1 species nl 

Recently a slide illl" -fi(iO) labelled C\il<>Mn>ttuii<> 
HH'ithHittfmhUtts in R M Vla^sun's handwrilin|_', was Inuiid 
ill ihe Siuuh Australian Musetun. Adelaide. The slide 
consists ol pieces ol iwti Icinak' wtmns inottliled ill resin 
Lint) has allucbcd 10 il anolhct hand wnUen n*Mc by Miiwmih 
indicating that the specimens :uv iwn rtl the tally three 

females ever ei'llcclcd- The eluuaeLers uf the oesophiiiiils. 
buccal capsule, Icmale tail and Dvfcjeeloj thai can be 
distinyuislu-il in these two temales are consistent with the 
characters ol PnfV)v t t\trnnu\his buriher ihe buccal capsule 
has an annular thickening around the middle as is found in 

P lh\hn>tilc The oesophagus (0.7 1 nun is slmrtet 111 C 
nn'iliiHiunni<iHi\ than in /' ihvla^ih' {\\^A)}^\ mni- The 
measurements of the posterior cud ol C. mitifftttmuIuilHi 
are tmt eonmueiil with LboseorA' tf\\'h)^ult; vulva lit tail lip 
VS compared to 0,7? 0,80 and tailO.M compared U»0 4/S 
W.M nun respectively. Ihe vaein.) vera ol (. 
nitilnmttt)nl(itn\. however, is the same length tr$ (hut ot P. 
thvUwtit'. C\i-liKsirt>nu\ftt\ mct/iouiitmliittis. therefore, is 
clearly a species of Pain/Vd\!H'iti;\lit\ and is most siiuilai to 
/' ihsl'^tilr Additional maleiial- inclndinjj male 
specimen-*, in needed before an exacl determination can be 
made. No olliei speeiiiicns nl PojunnMnnmyln* lmwc\ci 
have been louud in T, ifwtis i\> dale 

Johnston, T.ll. iV Mawson. l'.M. 1 OMOi fnic. lann Soc 
NSW 65, 46X470. 

Mawson, P.M. ( W7t Tunis. R. Soc. S. Ausl 101. 1^ 20 
Johnston, T.I I. & Mawson. KM. I \H (,, '» I'i-l 1 .inn, Sue 
NSWf>4, "M4 53o. 
'lU-veridKi-. 1. 1 I0S3) Ausl. J. Z00L Suppl. Ser. 91. I-SS. 

Mawson. PM.It')77)'l"rans, R, S<.e S AuM. 101. S|-f>2 

"Heveririjie, 1, tl'JKbl [Still. Mus, Natn. \t\M, Nal.. Pans 4 
Set S 1 A. 2^ 257 265, 

SpraO, U.M., CTevoudue, I.. & Walter. K.I.. | IWI | Rec 

S. Ausl. Mus. MonoLT- Ser. I. 1-105. 

L.R. SMAtLS. Depaitment uf Kiologv, bacullv i»l Applied Seieoee {'entrat Oiiecnslaml I Ini^ersily Rockhamplim (Jld 
471 C. 





T. C. R. WHITE, BSc, BSc(For), PhD 



M. A. J. WILLIAMS, BA(Hons), MA, PhD, DSc 




Assistant Editor: 
N. F. ALLEY, BA(Hons), MA, PhD 


Programme Secretary: 
S. BARKER, BSc(Hons), PhD 

Minutes Secretary. 

Membership Secretary: 

Members of Council: 

P. KOLESIK, BSc, PhD A. F. BIRD, BSc, MSc, PhD. DSc 


R. D. SHARRAD, BSc(Hons), PhD, DipT(Sec) 

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