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


Transactions of the 
Royal Society of South 
Australia 


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


bandicoots (Marsupialia: Peramelidae) - - - - - - 
Zeidler, W. A new species of freshwater amphipod, Austrochiltonia dalhousiensis 
sp. nov., (Crustacea: Amphipoda: Haale) from Dalhousie 
Springs, South Australia - - - - - - - - 
Bird, A. F. & McClure, S. G. Composition of the stylets of the aera Macrobiotus 
cf. pseudohufelandi - - - - - - - 
Bird, A. F. & McClure, S. G. Studies of the eggs of Macrobiotus cf. pace tnaie tis 
(Tardigrada) from wheat fields in South Australia - - - 
Kolesik, P. Two new species of Asphondylia (Diptera: Cecidomyiidae) from 
Halosarcia spp. (Chenopodiaceae) in South Australia -  - - ~ - 
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 
Brief Communications: 
Jago, J. B. & Haines, P. W. Poorly preserved trilobites and brachiopods from the 
Kanmantoo Group, Fleurieu Peninsula - - - - - - 
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 - - - 


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


75 


79 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 121, PART 1 


TRANSACTIONS OF THE 
ROYAL SOCIETY OF SOUTH AUSTRALIA INC. 


CONTENTS, VOL. 121, 1997 


PARTS I & 2, 30 MAY, 1997 


Smales, L. R. A revision of the Echinonematinae (Nematoda: Seuratidae) from 
bandicoots (Marsupialia: Peramelidae) - = - 2 ~ “ 
Zeidler, W. A new species of freshwater amphipod, Austrochiltonia dalhousiensis 


sp. nov., (Crustacea! Amphipoda: Fopields) from Dalhousie 
Springs, South Australia - - - - 
Bird, A. F. & McClure, 8. G. Composition of the stylets of the tardigrade, Macrobiotus 
cf. pseudohufelandi - - - - - - - - 
Bird, A. F. & McClure, S. G. Studies of the eggs of Macrobiotus cf. pseudohufelandi 
(Tardigrada) from wheat fields in South Australia - - - 
Kolesik, P. Two new species of Asphondylia (Diptera: Cecidomyiidae) from 
Halosarcia spp. (Chenopodiaceae) in South Australia - ~ - 
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 


Brief Communications: 

Jago, J. B. & Haines, P. W. Poorly preserved trilobites and bende from the 
Kanmantoo Group, Fleurieu Peninsula - = - - 

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 - - - 


75 


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 - - - - - - - 

Vaucher, C. & Beveridge, I. New species of Potorvlepis Spasskii (Cestoda : 
Hymenolepididae) parasitic in dasyurid marsupials from New 
Guinea - “ : = 5 ? ; - - ~ + 

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 - - 


Watson, G. F. & Gerhardt, H. C. The breeding biology and advertisement call of 
Litoria splendida Tyler, Davies & Martin - - - - - 
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 - - - - - - - - - 
Gullan, P. J., Cranston, P. S, & Cook, L. G. The response of gall-inducing scale insects 
(Hemiptera : Eriococeidae: Apiomerpha Riibsaamen) to the fire 
history of mallee eucalypts in Danggali Conservation Park, South 
Australia - - - - - - - - - - - 
Olsen, A. M. An intensive monitoring study of two wetlands of the River Murray 
in South Australia; physico-chemical parameters and cyanobacteria 
concentrations. - - - - 2 . E - ? - 
Kolesik, P., Whittemore, R. & Stace, H. M. Asphondylia anthocercidis, a new species 
of Cecidomyiidae (Diptera) inducing fruit galls on Anthocercis 
littorea (Solanaceae) in Western Australia - - - - - 


Brief Communications: 


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


Smales, L. R- The status of Cyclostrongylus medioannulatus Johnston & Mawson, 
1940 - - - - - > = 4 . : < 


Invert to Transacnans af the Rayal Saciety of South Australia, Vol, 421, purts 3 & 4. 28 November, 1997 


95 


103 


119 


137 


147 


163 


165 


A REVISION OF THE ECHINONEMATINAE 
(NEMATODA: SEURATIDAE) FROM BANDICOOTS 
(MARSUPIALIA:PERAMELIDAE) 


By LESLEY R. SMALES* 


Summary 


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, 
1997. 

The name Echinonema being preoccupied the genus here designated Linstowinema 
(nom. nov.) 1s 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. noy. is established for E. cinctum sensu 
Yorke & Maplestone, 1926 and sensu Chabaud, Seureau, Beveridge, Bain & Durette- 
Desset, 1980. Linstowinema inglisi comb. nov. 1s 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. 


Trasdetions Af the Ravel Sector cf S. Ame (P99T DAB 1-27 


A REVISION OF THE ECHINONEMATINAE (NEMATODA:SEURATIDAE) FROM 
BANDICOOTS (MARSUPIALIA:PERAMELIDAE) 


by Lestrfy R. SMALES* 


Summary 


SMaAtrs. dR (1907) A revision of The Echinonematinae CSetratocdaSeuniidee) frony bandicones 
(Marsupiilie Perumehdaes. rans. Ro Sve. &. Ang VEC, 1-27. 30 May, 1997. 

The name &eiWienenia being preoccupied the genus here desiited Lins roveinenied (ner, nov) bs edeseribed, 
The type species L. cinciypr comb. ney. is synoayyious wath Eo prertdfeneléy (ate Chabaud, Seureau, Beveridge. 
Luin & Durelle-Desser, 1980 but nor with Ay eurecn seri Inglis, P9267, senaae Chbaud, Scurenu, Beveridee, 
Ram & Durette-Desset. l9RO. Lo werrinedoni sp. nuy. is established for &. oimenin sevisu Yorke & Muplesione, 
1926 und sev Chabuud. Seureau. Beveridge. Buin & Duretie-Desset. 1980, Litisfeviiene ingtist comb. nay, 
1s aYnonyMous with EL cient seas Inglis, 1967 and A. dedist seis Chubaud, Senreuu, Beveridec Bain & 
Diretle-Dessel, (SO umd ts redeserhed and four new species Lo fafeny spo nov, Eo feweuaitielise sp. nov, Le 
maplestane) sp. nov and 4. permimelis sp. nov, are described. All seven species can be differentiated as [ollows: 
fo crnetai 14-18 rows oF body hooks wilh unduluting edges, cesaphagus teeminacine atthe level of the sth tlth 
cows 4 ivahrinweni. 9-13 cows of body hooks with Gndidating edges, vesophagus terminating wt the Jevel of the 
Oh Wth row: 2. Jngliay, 1O-b rows of body hooks with undukiting edges, oesuphugus terminating at the level 
ob me Suh-Yih rows L. dares, YP fows of body hooks wih undulating cdes, otsephagis terminating 
posteriorly ta the Tbth row: L. daspreaniense, 1a1S rows hooks without undoliting cdes, oesophagus 
termimatiny at the level of the #th-LOth ew: 2 aaplestoel 12-13 rows of body hooks without undulating edges, 
LEXOpPhAZus termining posteriorly to Path tow ol heoks, 2, perdmeliy. 11-12 cows oF body hooks: without 
Unduliting cdees. oesophages termtinaung at level of the }2th row. A key ta the species ie given, Casiranarent 
AC OCs Lh Avadenobesilus, Lmaciaurus, Péruneles nasi and Bogut, Coidulio’ and 1. tasneditionae 
only ind obesitas, L. fates only inh nieeronens, boned and mec, bclesntis, dirty ak PB 
nevi, Loomupiestonen ind, digernrey and Poorer aod b peramelis onty in Ft howeaimytte 


Kei Wonns: Giafeinene, Newntoda, Eehinonenidnie, baade, Peraieles, handicuos Marsupiitia. 


Introduction 


Whet) Linstow (18984) described a thorny headed 
hematode occurring in uv bandicoat host, he called it 
Hoplocephatus cincius, He subsequently found thar 
Hoplocephalus was preoccupied. so he renanied the 
conus Eehinenem laer in the sume year (Linstow 
ISO8h) Eehineneaned has continued to be used for the 
genus anol the present. Kehineneme also is 
preoccupied, however, Having been used previcusly 
fol a penis of sponges by Carter in I88t) b now 
propose the name Lisroninema norm, nov 

The bandivoor hese orginally was identitied as 
Perneles abesilus, Linstow (1898a) deseribed and 
firured a pematode with seventeen rows of hooks on 
the cuticular dilation of the oesophagul region of the 
hody and Gesophuaus terminating on a level with rhe 
Oth row of hooks Yorke & Muaplestone (1926) 
desenbed a nematode with 12-13 rows of bocy 
hooks and a relatively short oesophagus which they 
identiiied us “AL einer”. They neither give 
meusurements nor provided a figure to show the 
relationship between the posterior end af the 
Hesaphacus and the rows of hody hooks af their 


Depertogor a) Hatayy, Cente! Oueonstornd Cniversiny 
Raekleanpren lel 4702 


specimens. Yorke & Maplestone (126) assigned the 
type und, at that lime only speeies. 1. cient 
(Linstaw, 1898). te the spirurid family Rictulariidae 
Later Johnston & Mawson (1939) reported L. 
cinoma Tron a native cat, Dasvucks viverriiies, near 
Sydney and re-evaluated the available informiuvornon 
the host species, They coapeluded that Linstow's 
original descripuion wits based on muterial collected 
from Lseccton abesulus (Shaw. 1797) fron the Upper 
Burnett River in Queensland while Yorke & 
Maplestone’s redescription was bused on material 
from a bandicoat, posstbly fo maecreuriy (Gould, 
1842). collected in the vicinity ef Townsville. 
Johnston & Mawson (1939) fierther concluded that 
both Linstow (7898a) and Yorke & Maplestone 
(1924) were describing material from the sume hast, 
namely. 7. mteerauray. Sinee 1 ebesuliy does not 
occur in the Burnett River region (Braithwatte 1955) 
but 4 meeronrus does (Gordon 1955), this would 
seen to be a reasonable conclusion, 

The first confirmed eccord of Liston inema frou 
the bundicoot genus Peraineley Geotlroy. 1803: ty by 
Johnston & Mawson (1940) Fron 2 nasite Cealiroy, 
1804 collected near Sydney. These wathors noted 
differences. in the male tailor the specimens thev 
examined, Irom earher deseriptions by Linstow 
(18988) und Yorke & Maplestone (1926) but, 
nevertheless, assigned the specimens to Le cineniv, 


: Le. SMALLS 


OF the differences noled by Johnston & Mawson 
(1940). |he number ol papillae on the rule tail ts 
problemanc as neither they. Linstow 1 1898a) oi 
Yorke & Muplestone (1926) described the raniber 
and placement of papillae found on other specimens 
of Lrnsteowinemu, The expansion of the male badly 
annul the cloacal region described and figured by 
Johnston & Mawson (1940) ts alse) a Signitieant 
stctive, Although hot fentioned by Linstow 
(I898a). Yorke & Muplestone (1926) or Chabuud vr 
ot (1980) in their deseriptions of G2 eivernian. it ts 
computable to the nivale tail deseribed by Chabaud ev 
ul (LYS) for Echinenema sp. (fey occuring in # 
niatire Trem: ar unknawn touuhby, 

No further work was done on the genus until Inglis 
(1967) re-examined the relationships of the 
superfamily: Seuratoidey. He redeseribed L. e/neniay 
from material collected trom £ abestlus near Perth 
and placed the sole genus within a new subfamily. 
Ihe Eehinonemiatinge. lneated vwathir the seuratord 
family Schneidernematidae on the basis of the form 
of the moull opening. distribuljon of cephalic 
pipilae, long spicules and short guhernaculurn, 
Subseyuently. the allinties of the genus were 
clunfied by Quentin (1970) and the Echinonermistinie 
Included within the Seuratidac. 

Chabinud ef el, (1980) re-eximined all he avalible 
malerial. established two new venen) Seareokind 
Chabuud, Seureau. Beveridge. Bain & ure 
Desset, 1980 and Jielechind Chabaud, Serireaiy 
Beveridge, Buin & Duretle-Desset, [980 for wors 
from dasyurte marsupials and cedelined Echimeneeia 
tow Linstenvinena)y und £. einetune (te) Linstow. 
PSUS nee Inglis, 1967. In their desenpoon of LL. 
cineium they holed difheulnves in interpreting 
Linstow’s original Myure of 1R98a, bul decided that 
an oesophagus 18 mm dong agreed with their 
definition of a “long” oesophagus, that is, one 
terminating al the level of the most posterior body 
hooks. Linstow's (1898a) figure clearly shows. the 
oesophagus lerminanhy at the leyelol the th of 17 
rows OF body hooks. Chabaud ef al. (1980) alse 
described three new species, namely FE. edinenlss 
(sic) trom a dasvurid, 2. navdronalts (ye) from 
obesuluy and Bo inglixi (ied = Eo ined seitsu 
luglis, 967.also from /, obésulay, These authors also 
indivated thal there were possibly addityonal species 
from the bandicool genus Peryimedes but they had 
insulficient material for detailed deseriptions. 

The bandicouts (subfiunily Peramelinge) ate 
rabbit-sized aminiveruus ofarsupiils with dau 
puinted heads unit compuce bodies. They forage by 
digging conteal holes with their short forelimbs and 
explote these holes with their pointed snouts 
(Gordon & Hulbert 1989), Perdmeles jieesata, the 
lome nosed bumdicom, 1s found alone te east cons! 
ol Australia, from rainforest in the rovth thposussty 


wetland and dry weodhand to areas walb little groupid 
cover in the south (Stoddart 1995). Irs distr7bution 
averlaps wilh A. drecieens. the turthern brow 
hundieaots which is found of the east coast north of 
the Hawkesbury River and veross the Morihenn 
Territory Lo the north ef Western Australia, in areas 
of tow ground cover weludios grassland. woodland 
and open forest (Gordon 1995). To the south, the 
distribution of P wayute overlaps thatol 1 vhesulies. 
the southern brown handicool which is found weress 
southern Austraha in Western Austria. South 
Australia and Victoria. sourhern coastal New South 

Vales. and Tasmania and prefers sandy soils with 
scrubby vevelition or low ground cover phit are 
burnt out fram Gime to ime (Braithwaite 1995), The 
cusiern barred handicoor, PR ennit Gray. Lo38. nw 
restricted to Tasmania, where vs disteibutfen 
overlaps with that of J, ebesulin, and a few peliet 
colonies in southern western Victoria, prefers open 
wrussland, bur may ulse forage in serub und 
heathland (Seebeck 1995) The western barred 
bandieool 2 bouceanville Quoy & Gaimurd, |s24. 
now exisung ently on Bernier and Dorre Islands 
Shurk Bay, Western Austiuia was previuusly found 
uvross mueh of the southern hallof Australia (Friend 
& Burbidge 1995), The only other bandigoat species 
sill extant, Loanredduy, the golden bindicoor, now 
survives only on Barron and Middle [slands off the 
coast bf Western Australia, the north western 
Kimberley region und sub-humid paris of the 
Northern Territory. having been previously recorded 
fram a much wider range of habitats (MeKenzie e 
i. 1995). 

In this study, all the available records nel rmtertal 
collected from peramelids, imeluding matey 
dissected [rom hosis beld in museum collections us 
Well ws live caught bandicours, have been examined. 
This hus provided sufficient material to reassess. the 
laxonomic Characters uvanable to use for species 
discriminaljon. redesignate and redescribe (he type 
species LE. einetum (Linstow. (898) comb, poy, = £: 
merdiengi« (vie) Chaband, Seureau, Beyenddye, 
Bain & Durette-Dessel, YSU. redeseribe und name 
fovorrargtae sp. nov. = L. carerune sensi Yorke & 
Muplestone, 1926; seni Chabuud. Seureuu, 
Beveridge, Bain & Durette-Desser, 1980 and 4, 
ingdiyé Chabaud, Seurcau, Beveridge: Buin & 
Duretle-Dessel, T982 and distingucsh the (oer 
additional new species front bandiceots that ane 
deseribed below, 

Materials and Methods 

Material and disseetion neeords from 719 

Dandheoots were exananed, This matercal was 


derived from three sources. Firstly, the gisine 
intestinal tracts af $6 bandicools, catlected belween 


LINSTOWINEMA FROM BANDICOOTS 


Darwin 
OL. macrourus 


Mitchell Plateau a“ 


———p, 


bousainville 


Perth 


Fig. | 


Melbourne 


= 


Cairns 


Townsuille 
Ponstel a 


Rockhampton 


AP eunnii PoaSttTe ah 


Hobart 


Present and former distributions of Australian bandicoots Gifter Gordon & Hulbert 1989), The symbols indieate the 


localities where bandicoots were collected between 1989 und 1996. Iscoden auras, L macrourus, @ Loobestlis i 


Peraneles bousiiiville, Po git, & Po nasiite, 


1905 and LOSS and deposited in either the South 


Australian Museum (SAMA) or The Museuin of 


Victor (VM), were examined. These animals had 
probably been fixed in 5-10% formalin before being 
stored in 7O% ethanol, The nematodes dissected 
from these hosts were stored in 70% ethanol. 
Secondly. 79 animals were either collected us fresh 
roud kills or trapped alive. in spring-loaded wire box 
Wraps baited wath peanut butter or a peanut butter, 
honey and oats mixture, between 1989 and 1996. 
The trapped animals were killed by intraperitoneal 
inoculation of euthanasia solution pentobarbiione 
sodium (Nembutal @). The digestive tract of cach 
animal was examined under a dissecting microscope 
and any nematodes found were washed in normul 
saline, fixed in glacial acetic acid or hot or cold 10% 
formalin then stored in 70% ethanol. Alb the 


available specimens of Linsfowineme held in the 
Queensland Museum (QM). the Australian 
Helminthological collection of the SAMA (ALC). 
the Western Australian Museum (WAM), the CSIRO 
Division of Wildlife and Ecology (CSIRO) and The 
Natural History Museum, London (BM(NH)) were 
ulso examined. The preservation history of material 
from the former institutions is largely unknown but 
probably it was fixed in ethanol or formalin. Material 
from the CSIRO collection was fixed th hot lO% 
formalin. All material is now stored in 70% ethanol. 

Specimens were examined from all the extant 
bandicoot species (number af bandicoots in 
parentheses) from $1 localities across Australi 
fsooden auratus (5), 7 macrourus (1), 1 obestultes 
(85), Perumeles bougainville (13), Po guanii (6), P. 
nasuta (14), handivoot, no species given (9). Host 


4 1... SMALES 


TABLE 1. Distribitian af bauadicaat species oxmmined fot Linstowinema species by State ec lecritary. Where iis specific 
Jacedin Hew bean given dale divseetion tecara dhe location is listed aw Austretia. 
Abibreviatinas Ay Auvtrelica, WAS Western Austria NT Northern Territory SA) denith Australi incduediny 
Kunseroo dvi und Pranklin Wleond> Q. Queensland: NSW New South Waless Vo Vietertay 2 Tasmenyre. 


Species A WA NT 


Dsencndenit cttrarns a - 3 
Zo macrourus a 2 13 
1, bbesulus 2 19 
Peraneles baugainville 13 - : 
PL ounnir - - 
PL nasaret | - - 
bandicoat | 

2| ri | lo 


Location 
SA Q v T 
- AU) \4 7 - 
30) y i5 13 
- _ - 2 4 
- 7 4 2 - 
fl 8. n - - 
30 65 27 V7 16 


(istributions and the lowations of the 79 dtnimuls 
enllected since 1989 are given in Big. 1. Details ofall 
the localities where specimens were collected are 
listed in the deseriptions of species given below. 
Latitudes and Jongitudes are provided for all 
loculities that are listed in the Australian Gazeteer, 
The location, by stite, of all hosts species examined, 
is given in Table |, 

Specimens were examined after clearing im 
lactophenol or beeehwood creosote. Measurements 
were Wade with the aid of an ocular micrometer or 
drawing tube and map measurer, Meusnrements ire 
iver WT pas a rdnge from LO specimens followed 
by the inean in parentheses. unless otherwise stated, 
Alb the new material bas been deposited in the AHC. 


Comments on taxonomic characters 


Prior to L980 only one species was recognized in 
the genus Linstvavieme ie. Lehinonema enretine. 
Then Chabaud ef af (1980) described four species, 
three from perumelid hosts und one from the 
dausvurid, Deasvarns hallucatis Gould, 1842, The 
species aceurring in the dasyurid was distinguished 
from the others by having the first row of cephalic 
hooks longer than the second, The three species fron 
bandicoots were differentiated from each other on 
the basis of (he arrangement of hooks ahd spines on 
ihe body, the relulive sizes and positions of papillae 
op the clonal region. the relationship between the 
ventral spires aid pre-eloucal papillae, the extent of 
sind cuticular bosses surroundiig the cloud, Whe 
number of papillae on the tail oF the male and the 
length of the oesophagus relative 1 the hooks on the 
dilated cuticular partof the anterior body associated 
With the region of the oesophagus, 

Although Chabaud ef a (1980) recorded all the 
sensory organs on the mule tail tipas pairs of papillae 


tngls (1967) had noted pitirs of papillae and a pair ol 
phasmids. Thus Inglis (1967) reported 4 pairs of 
papillae and a parol phasinds on the tip of the tal 
of Eehinonema einer (sie) and Chabaud et al 
(1980) reported 4 pairs of pupilie for the sume 
species, Spicule morphology is uniform across the 
venus, differing onty iy total length and proportion of 
body length, 

Chabaud er ef. (1980) deseribed the oesephagus as 
enther “long. extending to the level of the last row al 
the body hooks or “short, terminating within the 
dilated cuticular region, Detailed examimilion ol 
specimens for this study has showe that alihough the 
lermdnation of the oesophagus relative to the 
surrounding hooks is consistent within each species, 
the aetual length of the oesophagus and its 
relationship to the body hooks may be difficult to 
determine, Speemens that are not camipletely 
straightened prior to or during fixation are 
problematic. [tis difficult to tell the extent te which 
the vesophigus may have contracted mie the neek 
reson and the cuticle surrounding (he oesophagus 
thay also be contracted, Furthermore, the range of 
lengihs of the oesephagus within populations ola 
species ean also he quite varnible. depending on tie 
sizes and maturity of the worms being measured. 

Both spines and hooks originate in the cuticle, but 
spines in this context, are defined ay being sal to 
Hiny und rootless, whereas hooks const of a thorn 
und a root anchoring the thorn within the cuticle, The 
smitllest hooks may be only slightly hanger than the 
largest spines, The relative lengths of the three rows 
oF hooks on the cephalic bulh ts a consistent 
character, but the dimensions af the hooks mity vary 
markedly between individuals of the same species 
from the same individual host, Therefore hook 
(limensions are noluselul for discriminating between 
species, The dimensions of the cephalic bulb are also 


LINSTOWINEMA VROM BANDICOOTS A 


unrelitble because of the potential variation caused 
by the extent ot relaxation of the specimens prior to 
fixation and the method of fixation. Other consisient 
chirueters at the species level, however, ure the 
number oF rows of hooks on the dthited anterior body 
and the morphology of these hooks. Ln seme species 
the roots of the hooks have undulating lateral edues 
giving them a “frilly” appearance: in others, the 
edves of the hooks are more or less plain, 


Key to the species of Linstowinema 


I, [st pow of cephalic hooks longer than 2nd row: 
PAUSES OP MUSYUPTUS oc ceercreernecen Leedmenielst 
And row of eephalie hooks longer than Pst row: 
purasitus OF bandheouts oe ee AY 


2, Body hooks without undulating edges... 43) 
Poly Hooks willy Uniden CULES ce (OF 


~ 


Ocsophugus terminutes posterior to hooks on 
cuticular dilation of oesephageal region: male 
will pairs of papillae, | pair ol phasmids on tail 
Ovsophigus fermimiles ih OF uNLeTIOr Lo pasteriar 
row OF body hooks on cutieular dikiion oF 
oesophageal reeion; male with 3) pairs ar 
pupilae, | pair of phasmids OF bil Upc et) 


Lo Mile with 8-10. demale with (0-12 body hooks: 
tnale with © pairs clowed! papillae all same size, 
body spines extend dong 800 of dorsal surface 
terminate at level of most anterior pair of lateral 
pre-vloweal papiihie on ventral surlaee 0. farens 
Male with (1-134. female wath 12-13 body hooks: 
male with 6 pairs of cloaeal papillae, | pair 
lateral ad-cloacal papillae lareer than other 5 
pairs: body spines extend alone 75% of dorsal 
surface ferminate markedly anterior 16 clouval 
papillae on ventral surface 0 teplestenel 


5. Male with alalike expansmons of body 
surroumding elowes with @ pats of cloweal 
papillae. 3 pairs of fateval cloacal papillae larger 
than 3 pains oF ventral clowcal papiliie. spicule 
leneth about 1/9 of body length, female wath tail 
longer than OQ. 0 cece be MANU te nse 
Male wilhoul alelike expansions of body, with 7 
pairs clowcal papillae all same size, spreule 
leneth about Yio of body length; femule with tal 
shorter Un S00 va. perainelis 


6. Male with 13-16. female with 14-18 rows of 
hody hooks; male with ala-like expansions of 
body surrounding cloaca, body spines terminiute 
atlevel of mostanterior pair of lareral pre-cloacal 
papillae on ventral SUmlaee cee L. einen 


Male with | 2. feniaie wilh FLL rows of body 
hooks; male without ali-like expansions oF body 
surrounding cloaca, body spines terminate 
inarkedly anterior to cloweal papillae on ventral 
SUIMPECE: scipernccuccupeesyesesece es: seereeecee crepes en reese 7) 


7, Male with oesophagus shorter than 1570, termule 
With vesophagus shorter than L830. male with 6 
pairs of clodeal papillae. purr of lateral ad-cloucal 
papiliae larger (haw other S pairs, 4 pairs of 
papillae, 1 pair of phusinids on Gal tip, spieule 
length ubour is of hody length: female with tail 
MMOMCT TAU OAD ccc seretecrensceeenese L, inglist 
Male with oesophagus longer than 1570, female 
Wilh gesophagus longer than |S60; rate with 6 
pairs of cloucal papillae all sunve size, 4 pars of 
papillae. 1 pai of phasmids on tail tip. spicule 
lenvth about '/20 of body lengthy female with Kul 
Jonwer thin YSso Lo Warelietartt 


Systematics 


Order Aseurididy 

Super fhinily Searalores 

Family Seuratidae 

Sublimily Echingvemarinae 

Genus Lintowinesire Won, Way. 

Type species: LIST MEHL CHICTHA (Lap stow, L968 | 
comb, nov, 

Synonyons: Joplocepidus coictis Tinstow, USUs, 
Kehinonema mteriediovalis Chabad. Seurein, 
Beveridgs, Bain & Dureve-Desset, L980 nee 
Keinouena cinetunm send Yorke & Maplestone, 
1920: Inelis, 1967; Chabaud, Seureau, Bain & 
Durette-Desset. 19X80 fin part), 


Liistowinema (\oln, Vey.) 


Genente HARMONIES 

Antenior end with cephalic bulb hearing 3 rows af 
I4-1 files of large hooks. Month opening Gtangulir 
i outline, without lips or lip-like struecures, with + 
pairs double cephalic papillae. | pair amphics, inner 
circle of sense organs on edge of mouth (see Inglis, 
(967. Figs 6. 7). Neck with 2-1] rows of very ting: 
spines. S-T8 rows wf 14-16 hooks surround an 
anterior culieultr dilution, assoeiuted with 
oesophageal region. Body with nurerous rows uf 
spines, miuhber of files of spines inereasing 
progressively towards mid body. decreasing Lawards 
posterior, continuing ta caudal ip of female, 
terminating at abou! '/3 of length dorsally, anterior to 
cloawwa ventrally on male. Short oesophagus simple, 
club shuped. surrounded by nerve cing at level of 
cephalic bulb. Deirids simple. conical. atlevel of [st 
row of cervieal books. Spicules lone. equal. 
identical, small gubernaculum present. Vulva at nvid- 


LINSTOWINEMA FROM BANDICOOTS 7 


region of body; monodelphic ovejector directed 
anteriorly, Parasites of Australian dasyurid and 
peramelid marsupials, 


Linstowinema cinctum (Linstow, 1898) comb. nov, 
(FIGS 2-14) 


Synonyms: Hoeplocephalus cinctus Linstow, 1898a: 
pp. 469-471, Figs 3-11.) Echinonemia meridionalis 
Chabaud, Seureau, Beveridge, Bain & Durette-Desset, 
1980: pp. 436-438, Figs 4, 5A, D; Spratt, Beveridge & 
Walter, 1991: p. 26. Echinenema cincta Linstow, 
1898b: p, 672: Johnson & Mawson, 1940: pp. 473- 
474, Fig 25: nee Yorke & Maplestone, 1926; nee 
Inglis, 1967: wee Chabaud, Seureau, Beveridge, Bain 


& Durette-Desset, 1980. 9 Echinonema vincta 
Mackerras, Mackerras & Sandars, 1953; p. 62, 


Echinonema sp. 2. Chabaud, Seureau, Beveridge, 
Bain & Durette-Desset, 1980: p. 438. Fig. SC, F: 
Spratt, Beveridge & Walter, 1991; pp, 23. 24 (in purt), 


Material exainined 

From /soodon obesilus New South Wales: 18, 
fragment Lismore, (28° 49'S, 153° 16'E), April, 
1965, AHC 4413. 44, Timbillica State Forest, 
(37° 19'S, 149" 43'B), 15.xii1.1978, CSIRO N733; 
l4d4, 1922, Sidlings Swamp North, ‘Timbillica 
State Forest, (37° 17'S, 149° 45°F), 19.vii. 1994, 
17.111,1994, 20.vii.1994, 23.11.1994, CSIRO N4213, 
N4074, N4228, N4075; 854d, 8822, Sidlings 
Swamp South, Timbillica State Forest, (37° 18'S, 
149° 45'R), 14.vi1.1994, 20.vi1.1994, CSIRO N4212, 
N4230. 

From Victorias 26d, 149 2%, Melbourne, (37° 
47'S, 136° 5Y’E), 91x.1991, AHC 30292; 229, 
Monash University, (37° 47'S, 136° 59’E), no date, 
AHC 30293, 30294; 26d, 229, Gorge Forest 
Road, (38° 21'S, 141° 36’E), Sept. 1962, AHC 
30296; 26d, 11° 2, 7 fragments, no locality, no 
date, AHC 30295, 30298. 

From South Australia: 1d, 89 2, 6 anterior ends, 
Waitpinga, (32° 36'S, [38° 32'E), no date, AHC 
4460; 33d 722, Scott Creek, (35° 04S, 138° 
42'E), 5.x.1992, AHC 30291; 3¢d6, 399, 
Myponga, (35° 23'S, 138° 28'E), May 1966, AHC 
4446; Kangaroo Island, South Australia: 1d. 29 2, 
Cape Willoughby, (35° 51'S 138° 08'E) 13.x.1990, 
AHC 30301; 29 @, Seal Bay, (36° 00'S, 137° 20'E), 
41.1987, AHC 30304; 2d d, 22 2, Binowie, (37° 


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

From Perameles gunati Tasmania: 646 4, 142 9,4 
fragments, Grove, (42° 59'S, 147° 07'E), AHC 
30025; 334, 42°, Kingston, (42° 59'S, 147° 
18’E), L1.vii.1992, AHC 30056, 30057, 30058. 

From Perameles nasuta: Queensland: 2° 9, 
Wongabel State Forest, (17° 20'S, 145° 30'E) 
9.vii. 1982, CSIRO N1753; 12, Mt Nebo Road, (27° 
23'S, 152° 47°F) tl.vini.1993, AHC 30316. New 
South Wales: 1c, Epping, (33° 46'S, 151° 05'E), 
1 L.viii.1993, AHC 30316; 1& Epping, (33° 46'S, 
151° 05’E). 14.vii. 1933, QM GL 12048; 344, 
29, Sydney, (33° 50'S, 151° 15’), no collection 
data, AHC 1820; 19, Nadgee State Forest, (37° 
26'S, 149° 54'E), 13.11.1978, CSIRO N493. 


Description 

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 13- 
IS rows of 14 (male) (Fig. 2) or 16 (female) files of 
body hooks; Ist and last rows smallest, 4th-7th rows 
largest; roots of hooks with undulating 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 */; of dorsal surface, 
terminating about 400 anterior to cloaca, almost 
reaching level of anterior pair of caudal papillae, on 
ventral surface of male body (Fig. 7), Ocsophagus 
simple, club shaped, terminating about level with 8th 
- 11th row of hooks, within the anterior cuticular 
dilation; '/s to '/j) body length (Fig. 2). Nerve ring 
surrounding oesophagus within cephalic bulb; 
secretory-excretory pore in neck; deirids conical, at 
level of Ist 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 Ist 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, 935-1150 
(1035) long, about '/i\4 body length. Gubernaculum 
short. simple, subtriangular 60-63 (n=5) long (Fig. 9). 
Nine pairs caudal papillae: 3 pairs ventral and 
immediately pre-, ad- and post-cloacal respectively, 


Figs 2-14. Linstowinema cinctum (Linstow. 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. Gubernaculum (lateral view). 10, Body hooks (lateral view). 11. Male tail (ventral view). 12. Male tail (lateral view), 
13. Body hooks (lateral view). 14. Female tail (lateral view), Scale bars = 500 pm 2; 100 pm 3, 4. 5; 50 pm 6, 8, LL, 12; 


25 um 7,9, LO, 13: 250 pm 14. 


x L. BR. SMALES 


| pair literal ad-eloacal, 2 pairs Jateral pre-eloacql: 
ill same size, 4 pais papillae, parr phasmids well 
posterior to cloaca. near tail tip (Fig, 1h), Cloacal 
resign with small cutieulur bosses; ahelike 
exprbsion OF body unterior and posterior to clone 
(iw, 11), Tail 310-450 (365) long (Fig. £2), 

Pemale: Length 16-22 (20) mun, wadtth 80021035 
(RKO). Cephalic bulb 455-380 (505) lone hy 455-600 
(500) wide: cephalic Hooks Ist row PRO-235 (210), 
Mel row 220-265 (245). 3rd maw 130-780 (155) long. 
Ocsophugus 1785-2125 (P9007) Jones eutieulyr 
dilation bearing 14-18 rows bedy beoks. Deirids 
W500 780 (G25), nerve cing 440-520 (470), seeretory- 
ckeretopy pee 520, S85 (n=2) from anterior end, 
Vulva 7140-10370 (S830) Prom aiterior end (Fug. 8), 
Vavinw aboul 300 long (i=1). Tail 985-1120 (1050) 
line t Figs 6. 14). Eves ovoid. 45-54 (48) log by 36- 
$1 (43) wide 


Type duane 
Perameles nase Geollroy, (X04 


| ye ten “liny 
syeliey, Australia 


Sire an host 
Small intestine 


(ype specimen 
Neatype AHC 1%20 


Remarks 


The present Ineation oF the material sleserbed by 
Linstow as Heydneephelns then renamed Lelvuey 
dent is inknown, Yorke & Maplestone (1926) and 
Chabad ef af (L980) neither gave a lecdtion tor the 
lype niuerial nor indicated whether they hie 
examined if, Linstow was working in Gottingen in 
18908 but neither Mefinenema nor Maple lids ts 
fisted under specimens held in the Zeologishes 
Muscuin) der tlumbolt. Universitit, Berlin Ths 
Museum does, however, list holdings al other type 
specimens fran Linstaw, The specimens are not hela 
We (he parisite collections ef the BM UNH), the 
Inlernational Instinie of Parasitology, St Albans. or 
the US National Museunr Parasile Collection in 
Beltsville. 

The type host was given by Lamstow (1898a) as 
Perameles abexulus. one of iwo bandienat species 
collected by Richard Semon. These species were 
identitied hy Romer (1901), using the catagae of 
Thomas (1888) in the British Museurn, us Peveneler 
ohosulius, from Burnett River and Po pnete pure trom 
Couklown, with inedsurements being given for / 
meerura Perameles macrara is formdly listed as a 
synonym of fvacdon maerauins (Mahoney & Ride 


L988) but B obesilis is not. This is surprising 
hecuuse the taxongmie stitus of Po vbesalias was 
discussed by Maekerras & Maekerras (1960) who 
intheated that Perameles obese = Didelphix 
obesul = Tseddon abesulis, but that since the 
novtbern Laiutof disutbution af fseodon ubesalas is 
near Sydney, bumdicoals recorded iis A ebevuliy Gor 
Oneensland should be referred to us 2 meerouras, 
the species oecurring [rot north Queenstand te 
northern New South Wales 

Yorke & Maplestone (1996) Hat the type host as 
Peraineles abextla, dolinsten & Miwson (1930) 
decided that sinee Yorke & Maplestone tcl drawn cut 
original figure their material came From Townsville 
i) forthe Queensland and stared that Linatow's 
material from Jyaudon ebesday came tron Upper 
Burner River also in Queensland On that basis, 
Chubund een), (1980) voneluded that the host ot bath 
the Linstow atefiel and the Yorke & Maplestane 
material wis Dveeden meerouris, lhe northern brown 
and nit 4 abesilas: the southern brow buntigaut 

This does not, however, explant why Romer (M901) 
ia his denutication of the bandicoots eullecred: by 
Seynon Lists thent as two separate speeles rather thay 
us Perumeles macrira now bvoodon macrourus, The 
cHlalpeue of Thomas (LA88) however. does Tist 2 
nasaia. so the Upper Burnen River bundicoots were 
probably 2. macrourus 

Chabaud ef ah (L980) noted That Les ion. the 
measurement given for te length of the oesophagus 
by Linstow (18980). was sinker to measurements ot 
specimens oxamincd by then that had heen collected 
fron) 2 mecrouras fron northern New Sotth Wakes 
(hroweh to nerthern Queensland. tn particular these 
specimens had the pesophagus lerminating wt uboul 
the level ol the posteriorend of the cuticular dilavon. 
4 character they deseribed as a “long oesophagus’. 
Yorke & Maplestone (1926) did not indicate the 
lengil of (he oesophagus relative lo the rows of body 
hooks onthe cuticular dilation but they described 12 
or 3 “eireles” OF 4 ip 16 rows of hooks, Two 
specimens eolleered by Nivoll in L915, currently 
held in the BM(NELY. which eouldl possibly be the 
fuiterial described by Yorke & Muplestone (1926) 
ahd vedeseribed below have the “long 
oesophagus” deseribed by Chabaud er al, (LOS) 
The specimens described hy Linstow (E898) ine 
drawn and libelled as haying (7 “circles” of hooks 
With Lhe Gesophages terminating awe the level of the 
Oth row of hooks. These churucters are consistent. 
net with LE einer seas Chabad eral (1980) but 
with specimens deseribedl by Chabaud er a/ (1980) 
as A. meridiaralis, occurring ta the southern browe 
bandicoot /xeaden obesuliy collected in South 
Australia, 

In a re-examination oF the material deseribed hy 
Chahaud ef al, (M980) as EL meridionativ., tovether 


alsa ' 


LINSTOWINEMA FROM BANDICOOTS 4 


wilh speciinens collected fron long-nosed and 
hirred handicoots For this study. i) was found that all 
speciinens bad TetS rows ef body books aad) the 
ockoOphawos terminuted at the level of the Baie tl th 
row of hooks. Measurements of the oesophasts 
ramped front 1445 19 2041) for males amt 1785 to 
2125) lor females, also consistent with the 
Measurements given by Linstow CUYosa). 

Johnston & Mawson (940) deseribed three 


forniles aid two ttales obtamied froin the intestine of 


the long oosed bandicaol Fo vex collected in 
Sydney and attibuted these to 2. cine They 
described ind figured differenves in the male tail. 
minely an expansion of the bady surroinding the 
Cloaca. similar to, bul not transparent, as are caudil 
ali Chabaud eral (1980) figured the posterior end 
ol aimale which they designated Fehinemema sp.2 
from @ nasi registered in the ALC as (S20. They 
commented (hat their specimens were comparable 
wrth (hose described by Johnston & Miuwsen (1940), 
Neither sroup deseribed the anterior ends of the 
watts they cxaummed. The only material registered 
in he SAMA which might he the original Johnston 
de Miwsen specumehs is AHC [X20 The 
iorphology of the anterior. ends of these wornss,. |3- 
[8 body hooks, the oesophagus termining level 
will) the Ol) TLE row of books. is consistent with & 
mevidionalis seusit Chabaud, Searedu, Beveridge, 
Bain & Durelte-Desset, 1980, 

Chabaud erat (T9880) dit nol comment on the 
presence or ubsenee of an expansion te the Clowes on 
(hen Speconens bot contrast A. meridiomaliy with 
Cohinonemea sp, Disee Chahaud etal Vasu. py, 438, 
bie A) whiting thal Eefiinememe ap. 2. was 
comparable with the Tohnston & Mawsen (1240) 
desenpuun. 

Are exuniibition ob all the avartible specimens of 
Fe neridjenally (sie and Echinoneme sp. 2 (ie) Was 
fuled to shoy, aniy stntticunt dillerences beiweea 
them Sieh cifferenves us do eist can be anribited 
te the Rice chat Chabad en at L980) were doula 
Wil a mixed infeetion oF two species. namely L 
ciation and tL. weareiierane (described below) 
oveliceng Ta the material gollecred fron Wealtpinge 
ail Myponga, The male 2. cfietenn ask (ars 
popubiion were al the small end of the sive range 
und oesophageal length varies with work leneth and 
HHALLUDIy 

The déserption by Linstaw (PSUS of Ze cdots 
is congruent witht Hie revised deseripnog of muteyral 
desienaled L. merteivnlia by Chabiued of et. (TOR). 
The deseriplion by Yorke & Muplestone (1926) of 
specimens from h mecrinis und identified as &, 
cine. ts ebagrucnt with & carmen sensu Chabwud, 
Seurean. Beveridge, Buin & Durette-Desset, LOSO 
hee Linstow, PS9S, Theretore, | desienite a specimen 
fron ANC 1820 a the nearype of Lo efecnen 


Liiytowinemea cinch (Lanstow, 189Sh most 
closely resembles LZ. inylist (redeserihed below) in 
that the oesuphugus is celutively short in relation to 
the namber of rows of hooks. ending within the 
citicular dilation. The males of both species have 3 
patry of caudal pupilae and spicules ‘ya - “/\= body 
length. Linsrmawinena cieenue con he distinguished 
frony 2. aiglive uy having 13-16 Gale) and [4-15 
(female) body houks compared with 1-12 and 12-14 
in L. frefisi. Although the ovsophugus ts cehibively 
short itis, however. longer than in 2. ing/isi, being fy 
body Jeggth in nmile Lo evenun compared with Wy, 
bady Jength in 4. delist. The pair ol lateral ael- 
cloacal papillae is bo Jarger than the other pains oF 
venteal pupillie in 2. eynedue but ws larger an 2. 
iuelis? (see Taglis WOT Pie, 9) The ale-like 
expansion of the body surrounding the cloaca af L, 
cliendn ys not found on 2. welsh (see Inglis 1Y67 
Fig. 9). The body spines of L. eireiamn onty cover Ya 
ol the dorsal body surface bat cover |'7in of Lhe dorsal 
body surfiec ot 2, ineli’. Eggs of Lo emcaun ditfer 
from those of all other species in that they ure ovoid 
rather (hin almost spherical. 

The miternial (rom 7, oPeyalas from South Australia 
described by Chabaud ef ah (1980) as 2 
meridionaliy was ound to be wniixed infection all. 
cincnint, Pemales with up to 18 rows of hedy hooks, 
odsaphagus terminaring at about the level of the 9th- 
Lith row)and 4 werrorgivn? Gemales with up to 3 
rows of body hooks, the oesophagus termoiating 
about the level of the LOth -l3th row) deserihed 
helow. A conyparison OF the meusurentents piven by 
Chabaud ep af, (1980) ad {he speaiinens caamined 
for (his study reveal that the pnales (hey measured 
were smaller in size. the oesophagus, spicules: are 
ful were Shorter than in the specimens examined for 
this Study. ‘The females, however, were within the 
same size ringe ds for this study aud the comparative 
meusurements wre more consistent, 

The nuiterial dissceted from the scuthern brown 
hundicoot vollected it Seow Creek melided some 
females up to 46 mn Jong. These were larger than 
The specimens from Saul Australie studyed by 
Chabatider af (1980), (ip to 27h lowe), trap 
eustern Dirred bandicoots tram Tasmania (ap to 
721nn long), und from long-nosed bandienols. from 
Tasmania. (30-32 mun fone) Other variations 
observed hetween feniale specimens collected in 
different hosts wid Jocalinjes in the present study and 
thase of Chabaud eb af Ch980) included the 
oesophagus fonvest in southern brown bundieaols 
Tram Seatt Creek (1853-23980), and shoriest in 
southern barred banediewots from Myponga and 
Waitpinga (F100): the tail Jongest in eastern brown 
bundivoots from ‘Tasmania (986-1122) and shortest 
in soothern brown bindicoors fram Seat Creek 
(S84-98O), Gd the vulva slightly mere Pastertar 


10 L, k. SMALES 


(10,0380-14.450) in southern brown bandiceots from 
Seorr Creek than in eastern barred bandivoots fron 
Tasmanne (7140-10370) or southern brown 
handicnots From Wailpinga (10,800), The eggs from 
the Seott Creek specimens were smuller than those 
from Waitpinga or Tasmania, beng 33 by 36. 42 by 
45, and 43 by 48 respectively. These differences 
could be either yvandhons befweer populations 
within the species. or the resull of contraction during: 
frxvation, 

Linsrowinema cinenun, originally deseribed as 
ovcurring iP. ehesuta tyre) Trony Queensland ts now 


recorded as also occurring inf ehevuluy, Pe nesuler 


and 2. guani, The geographical range new includes 
Vietoria, New South Wales. South Ausuibie 
meluding Kainigeoo Ishind, and Tasmania as well as 
northern and southertr Queensland. The record af f 
einem in L ebesulus from Lismore, New South 
Wales, ATIC 4413 1s a dubious bost record because 
the southern brown bundicoot ts net found newly as 
fur north as Lismore (Braithwaite 1995), In all 
probability the host was L wnererenrs. 


Linstowinema warringtoni sp nev. 
(PIGS 15-24) 


Synonyms: Lefironrene cineta sense Yotke & 
Mauplestone. 1926: pp. 347-348 nee Linstow, 1895; 
nee Trelis, 19672 yest Munday & Green. 1972: p. 1) 
(in party, sels Chabaud Seurcau, Beveridge. Buin & 
DuretleDessel, (980: p, 435-436 (in party, sensu 
Sprat, Beveridge & Walter, TU9T p. 25) Cin part, 
helinonema emer sensi Johnston & Mawson, 
199) yy. a3 


Material creniuned 

Prom dred aurains: ba 8, ne collection data 
ALC 30319. 

From (sondern meray Queenstands |. 17. 
Rollinestone. (19° 03'S, ld6° 24" Ra 24.1915. BM 
(NH) $450) 12.6.163-}665 Mossman to Daruotree 
Road. (16° 15'S, 457 19'B), 2.8 1991 AHO 30278: 
2284, WV SY, Mossrrath, (16° 28'S, 149° 93'E 4, 
Wit b9S8, Sal T85s,  Fand¥5h bait, PYAs, 
12 PUSS. WAL ss. U4) 1958. OM GLI4351 
GE 1436), GL43a63, GL.b6d. GLIAS65_ CL 14366. 
GL IS370, GL14372, GL14374, GL14377. GL.144s1 
GL)4383; 264, 429, Caims to Mossman Read, 
(16° 58'S, 4S" 46H), 2a 1991 AHO 30274, 244, 
Gillies Highway near Lake Barring, (17° 15°S, 145° 
ARE) Ww 19901 AHO 30279 Goo, They, 
Atherton. (177 16'S. 145° 29"). 25,y, 1982. CSIRO 
Nigh 8s TES 2. Yorgubures to Atierton Road. 
(17> 1a’S. 145° 35", 29x 19YL AME 20278: Te of, 
1} 2, Miviwinng. (17° 24'S. bbs 55 °b, 25.16 1957, 
OM GI1d368, ste, (892 2, Diddgee (17° 29'S 
146° 00°R), QOGi,T8S& OM GE12653, GL4307; 


Res. 79 2, Millaa Millaa to Lonislail Read. (17> 
W'S. 14S" SPE) Shaw 1991, AHC 30276: Tod 2. 
133-29) Innisbail, (177 32'S, 146" ONT). 16d 1959. 
5.x 1953, 2ONTI9S6, 27-956, Ovi 1956. 
19.yi 957. TG 1960, ALC 4528. OM GLI4457, 
GL14360, GLI4376, GLAS378, GL14379, GLI4382; 
746 4,39 9. Atherion to Ravenshoe Road, (17° 46'S, 
145° Q9'E), 3ELx 1901, AHO 30277; 16, 4%. 
Ingham, (18" 39'S. 146° 10"B), 31-x.199t, ALC 
240; 25.0. 6% Y, Palmerston Aighway, 29,9. 1959. 
QM GL 14369277 °°, 60k south of Proserpine. (20° 
14'S, 152° 357E) AHC 30266: 3© 4, Roekhumpton to 
Yeppoon Roud (23° US'S, 150° 44'E), Lo-viti 1990, 
Syii 1992. AHC 30271, 30267, 322. Yeppoon te 
Emu Park Road. (23° 10'S. 150° 46'R), 16i1x.1990 
AHC 30270; 4a ¢, 15% 2. Rockhimptow to Emu 
Park Road, (23° 13'S, 150° 30°F) 17 411.1992, AHIC 
W272; 1, 24%. Rockhampton to Keppel Sands 
Road. (23° 20'S, 150° 48'R). Tait. 1992) AC 
A268: 69 9. Mt Glorious, (27° 21'S. 152° 54’E) 
27.6. 1955. OM GL.L4R59: 692d, L204 F, Ashereve, 
(27° 27'S, 153°, 02H), 274.1956. OM GI14347, 
1214 4. 20729. Paddington. Brisbane, (27° 28'S, 
153° OV BR), 25.ni11 1955, 21,x%,1955, 14x, 1995, AHC 
4371 QM GLI4350. GLI437l: 2945, Site Pp, 
Brisbane, (27° 24'S. 153° OIE). 28.48.1954: OM 
GL M4340, GLAda80: Ta a, P82. Mogeall, (27" 
20'S, 152° 54’). 12.4.1967, CSIRO NIST, 2s, 
705 Mt Nebo, (27° 3378. [52° STR), Tx. 1954, 
23,y. 1994. OM GLI4359, AHC 30317. 3081S: 
Sef 139, Broukfield. (27> 30'S, 1S7° S5"b). 
200.1973, My, 1967. AHO 19367, CSIRO NISI: 
(876,210 9, Rockles Crossing, Moy. 1867, CSIRO 
NIS52: New South Wales: 1. oa other ith, APIO 
4462: Nu locality given: 97 4. 171 GS, SL 193s, 
Wax lG54, Mx 19Sd Bax ts. QW 45e. 
(2.795%, OM GL(4345. GLI4346. GL14352, 
GL14393, GLI4354, GLI4355. GL 14362, 

Pram /seadon obesitus New South Wiles: Te, 
Sidlings Swarnp North. Timbillica Stale Forest. (47 
W'S. 149° 45’), TO) 1994 CSIRO N4228, 
Vielurias Tbe. JO 2 no eollectiow data, ALE 
446|. ABC 30207: 66 4.39 9 Hulls Gap (377 08'S. 
(42° S1'hy ne date, AHO 30207) 94 49.32, Moura 
Reservoir, Grampians, (37° 148, 142° 30°), no dale. 
ANIC 30299. Sout Austrailia: 32 2. Mypemaa, (35 
23'S. 13K° TS'R). no dae AHC +A. 395 | 
anterior end. Wuaitpinga. (35° A'S, 138° 42'B I nb 
date, AHO 4460. Kanvaroo Ishind, South Australie 
OF. Lot. Vivonne Buy, (25° 59°S, 137" 13°F), 
Lv.JO8S, AHC 30302; le 2, 39% 2, Parndana, (35 
ATS. IAT? LSE 350986. ATIC 30307; 14d 4, 
287%. Binnewie, (37° 47S. (36° 5S98'R), Soviit, 1987; 
ALC 30403, 

Prom Perankcles nasi Queénshunds bt 38S. 
Mareeba, (17° GO'S IAS” 26") 24a LOU). ALC 
30036. 


LINSTOWINEMA FROM BANDICOOTS i] 


4 
* 
4 
¥ 
* 
¥ 
¥ 
# 


Figs 15-24. Linstawimema warringtoni sp. nov. 15, Anterior end, (lateral view), 16. Cephalic end female, optical section at 
level of first row of cephalic hooks (en face view). 17. Female. hand cut transverse section through body hooks on 
cuticular dilation, 18. Body hooks (dorsal view), 19. Body books (lateral view), 20. Male tail (ventral view). 21. Female 
tail (lateral view). 22. Vagina (lateral view). 23. Gubernaculum (ventral view). 24. Female tail tip (lateral view). Scule 
bars = 200 por 15, 21: 100 pm 16, 17, PS. 19. 20, 22. 24: 50 pm 23. 


12 LR, SMALES 


Deseription 

Cephalic bulb with 3 rows af 14 (male) ar 16 
(enale) files of hooks (Pig. 16), 2nd row longest, 
3rd row shortest; neck with a-7 rows tiny spines: 
cuticular dihition of oesophageal region bearing 9-13 
rows OF J4 (mule) or L6 (lemale) files of body hooks 
(Pig, 17), Ist and last rows smallest, 6th - Sth rows 
largest, roots of hooks with undulating edges (Figs 
18, 19): remuinderof body with up to 48 (male) or 54 
(female) small spines at cach aonulation, over whole 
body af females extending over V/s of dorsal surface, 
terminating about S00, anterior to cloaca, not 
reuching Jevel of thost anterior pair of eaudal 
papillae, om ventral surface of mule body (Mig, 20). 
Ovsophagus, ‘io - Yin body length. simple, elub- 
shaped; terminating ul level of 9th- 13th row of body 
Hooks (Fig. (5) neat posterior end of anterior 
culicular dilation, Nerve ring surrounding. 
vesophagus within vephalie bulb; secretory- 
excretory pore On neeks deride conical, atlevel ot tsi 
row of body hooks. 

Males Length (5-20 (17.5) min, width 450-750 
(505), Cephalic bulb 270-340 (295) long by 270-320 
(205) wide; cephalic hooks Ist row 140-170 (155), 
2nd row 160-190 (175). 3rd row 100-130-(1 10) lang 
Oesophagus 1575-1925 (1743) lone. colieular 
dilation bearing 9-11 rows of body hooks. deirids 
490-550) (530); seerelory - excretory pore 360 (n=1; 
nerve ring 310 (n=l) fron: anterior end, Spicules 
similar. equal, without alae. G90-LO90 (850) long, 
about '/a body length, Gubernaculum short, simple, 
sub trangiilar, 50°75 (n=5) lone (Pig, 23), Teo pares 
viidol papillae: 3 pairs ventral and iinmediately pre-, 
ad- and posreloawal respeelively. | pair daleral ud- 
Cloaeul. 2 pains fateru! pre-eloucal all suime size: 4 
pales papillie, pair phosmids posterior to cloaca near 
toil Vip (Fig. 20), Cloaeal region with small eutieu liar 
bosses, dlidike expansinns ihseal Mil 430-430 
(90) long, 

Vemiale: Length 3241) (48) mm. widih 720-990) 
(edi) Cephalic hulls 450. 390 (470) long by 350-400 
(280) wide, henring 32 rews al hooks, Ist rew 1 70- 
JHC 85), 2nd tow PHO-230 (95), 3rd pow P20. 150 
(730) Jong, Qesophaiis TS70-2500 (2260) bans. 
vuteulie diladoo bearing |1-13 rows of bouly books. 
Deuirids S8O700 (O67): svererory-exerelory pore 
A7O-AOO) (390): nerve rings 200 360) (A301 drain 
uiferior end Vulva 12.5-05.6 (14.2) nim tron 
anterior end (Fig, 22). Tail 100-1900 (1265) long, 
(Figs 2), 24), Vagina about 300 Jong (n=l), Fags 
ilimost spherical 3044 (38) by 43-52 (40), 


Erynralagy 

This species is maned after Wirrnigton Yorke who 
with POA, Miplestane carried aut much af the early 
work on this ecnus, 


Type deeaulity 
Townsville, Queensland, Australi 


Type rest 
dyoedom macrourus (Ciguld, P42) 


Site in host 
Small intestine 


Type specimen 
Neutype BM (NM) 1950. 12.6. 163 


Remarks 


The iwo worms, |. b¥. BM (NID 1950, 
12.6.165-166 trom Perameles vbesula collected by 
Nicollin 1915 in Queenshind dre the only specimens 
registered i) Austalian or United Kingdon: parasite 
collections which could be the material deseribed by 
Yorke & Maptestone (1926). On examinution these 
worms were found to have the TL (imate) and ts 
(female) body hooks deseribed by Yorke & 
Maplestone (1926) for Behivenemea efor and te 
have the “long” oesophagus aiid all the other 
charaelers aliributed to 2 cineca by Chabad eat, 
(1980). J therefore designe the male speeiicn ul 
BM (N11) 1950, 12.6.165-166 us the notype, 

All the specimens ientiticd in this study as 6, 
warrington? conformed to the description siven by 
Chabaud ep al, (LORD) of Feiner. Adusrenvineniet 
warrington’ sp, nov, differs from. all other species i 
the genus in having Up to 9-1) rews of hooks (iale) 
or t1-12) (emile) on the oesophageal cunedta 
dilation amt a “long” oesophagus, thi as, the 
vesuphigus laemindtes: abo ee mew the fim tow ot 
body hooks. Male 4. weeengtoh can he 
distinguished fron L, civetiir by the combination of 
characters al (he posterior end. bey the terminating 
venteal body spies dir net extend te the most 
wotertor pare ol pupilhic in 2. wearing? buries orf 
Crean, whe disianve bepyeen spines yind clea is 
S00 pin not OO as tL. edaedan, the Tintited extent 
Of vuticular bosses surrounding the clowen in 
werrdigion’ compared with Le cinetan, bh, 
warrington? does net have an ulilike expansin 
surreundinyg (he clowea bul 2 cited does, and four 
pans of papillae vot three un the tal tip) The spietles 
of Lo werriitent foo body length ane shorter phan 
those of L. cinetane Vio body lenwih, “The opps al b. 
werrmeten; ire almost spherical hut tose ol 7. 
Curelae ure ovoid, 

Chabad ef af, (1960) in ther deseription of &, 
cineruny (sie) indicated (hab they had studied 
bumerous specimens from a range of localities 
including Woolwonga and Darwin in the Northern 
Territory. A re-cxuinioition of the imitertal from che 
Northern Territory has shown that ibrepresents a new 


LINSTOWINEMA FROM BANDICOOTS 13 


species of Linstowinema, L. latens, described below. 
The measurements and figures of specimens reported 
by Chabaud ef al, (1980) however, are congruent 
with L. warringtoni rather those of L. latens, 

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

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 1. abesulus ts 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. cinctum. The geographic 
distribution of L. warrington’ theretore 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 of /. obesulus trom the 
southern states needs to be examined before 
ecographic distributions can be fully mapped. 
Populations of bandicoots in NSW, Vic. and SA, now 
have patchy distributions over a reduced range 
(Braithwaite 1995) and attempts to collect additional 
specimens of /. obesuluy for parasitological 
examination have been unsuccessful to date. Further 
work on the southern geographic distribution of L. 
warrington’ will be problematic, as bandicoots 
become more difficult to collect. 

The specimens of L. warringtoni found in a single 


P nasuta suggest either a natural low prevalence of 
infection, or an occasional, incidental infection of 


this host. 


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


Synonym: Echinonema cinctum sensu Chabaud, 
Seureau, Beveridge, Bain & Durette-Desset, 1980: 
pp. 435-436 (in part); sensu Spratt, Beveridge & 
Walter, 1991: p. 25 (in part). 


Material examined 

Type material: Holotype cd, allotype &, from 
lsoodon macrourus, Walsh Point, (15° 08'S, 125° 
46'E), Mitchell Plateau, Western Australia, 
23. vil. 1982; AHC 30322, 30323, 43d, 6592, 
paratypes AHC 13028, WAM 110-83, 116-83. 


Other material: From Isoodon mucrourus Western 
Australia: 1¢, 49 9, Mt Hart, Kimberley Ranges, 


(16° 48'S, 124° 55’E), 22.x,1993, 24.x.1993, AHC 
30289, 30290. Northern Territory: 1634, 189°, 
Darwin, (12° 27'S, 130° 50'E), no date. 19.vi, 1993, 
27.vi.1995, 28.vi.1995, AHC 4703, 30287, CSIRO 
N4413, N4414; 764d, 72 2, near Byers Rd, Stuart 
Highway turnoff, (12° 30'S, 130° 50'E), 18.vi.1993, 
AHC 30284, 30285; 7d d, 102 2, Adelaide River, 
Arnhem Highway, (12° 28'S, 131° 14°E), 
20.vi.1993, AHC 30288: 13d d, 122 2, Bees Creck, 


off Stuart Highway, (12° 35'S. 131° 04'B), 
17.vi.1993, AHC 30286; [1d d, 89 2. Jabiluka, 


August 1979, AHC 6421: 3° 9, Woolwonga, (12° 
45'S, 132° 39'E), 19.x.1972, CSIRO N159. 
Queensland: 6d ¢, 112 2, Atherton (17° 15'S, 145° 
29'E), 25.v.1928, CSIRO NI610: 11d d, 142 9, 
14km north of Atherton (17° 15'S 145° 29'E), 
15.41.1982, N1532: 1944, 129 9%, Yungaburra to 
Atherton Rd, (17° 15'S 145° 30'E), 29.x.1991, 
L.xi. 1991, AHC 30278, 30281; 1d, Gillies Highway 
near Lake Barrine (17° 16'S, 145° 35'E), 29.x.1991, 
AHC 30279; 64d. 82, Gillies Highway near 
Yungaburra, (17° 16'S, 145° 35’), 29.x. 1991, AHC 
30283; 2344. 1169 2, Mareeba to Kuranda Rd 
(17° 00'S, 145° 26'B), 2.xi.1991, AHC 30282. 


Description 

Cephalic bulb with 3 rows of 14 (male) or 16 
(female) files of hooks, 2nd row longest, 3rd row 
shortest (Figs 26, 27); neck with 2-9 rows tiny 
spines; cuticular dilation of oesophageal region 
bearing I4 (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 of hody with up to 44 (male) or 60 
(female) small spines at each annulation, over whole 
body of female; extending over °/jy 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 '/\y - 1/1) (male) ‘is = '/ie (female) body 
leneth, simple, club-shaped. terminating posterior to 
the oesophageal cuticular dilation (Fig. 25). Nerve 
ring surrounding oesophagus within cephalic bulb, 
seeretory-excretory pore in neck, deirids conical. at 
level of first row of body hooks. 

Male: (measurements of specimens from Western 
Australia, followed by measurements of 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, Ist row 150-180 (152), 105-165 
(140), 2nd row 160-190 (168), 125-195 (155), 3rd 
row 100-110 (108), 72-117 (94) Jong (Fig. 26). 
Oesophagus 1200-1825 (1555), 1325-1990 (1715) 
long, cuticular dilation bearing 8-10 rows body 
hooks. Deirids 540-690 (610), 390-650 (530); 


L. R. SMALES 


AA’ 


“ean 


Figs 25-36, Linsrowinema latens sp. noy. 25. Anterior end. (lateral yiew). 26. Cephalic bulb (lateral view). 27. Cephalic end 
female, optical section at level of first row of hooks (en fuce view). 28. Vagina (lateral view). 29. Body hooks (lateral 
view), 30. Body hook (dorsal view), 31. Female tail (lateral view). 32. Male posterior body spines (ventral view), 33, 
Mate tail (ventral view), 34. Female tail tip (lateral view), 35. Male tail (lateral view). 36. Gubernaculum (ventral view), 
Scale bars = 200 pm 25, 26: 50 ym 27: 100 pm 28, 31, 33, 35; 25 um 29, 30, 32. 34. 36. 


LINSTOWINEMA FROM BANDICOOTS (5 


secretory-excrerory pore not scen, 230-495 (305): 
nerve ring 300 (n=1), 225-365 (275) from anterior 
end, Spicules cqiil similar, without alae. 830-1090 
(995), SAN-1100 (1005) Tong, about Yin to “is body 
leogih, Gubermaculum short. simple, sub-triangular. 
O1-68 Ure3). 66-79 (n=) long (ig. 36). LO pairs 
miudal papillies 3 pairs ventral and immediately pre-. 
fd and post-clodcal respeetively, | pair lateral ade 
chougal 2 pours Tateral pre-eloacil, all sume sizes 4 
pairs papillae, pair phasmids. alaike expansions 
ibsent posterior to Clowes. near dul ip (Rig, 33). 
Clowcul region with siiall cuticular bosses. ala-like 
expansions absent, “hal 250-340 (31), 255-320 
(305) long (ip, 35), 

Pemate, Length 20-38 (30), 25-41 (30) mim. widih 
40) 750 1590). 665-1175 (840). Cephalic bulb 300- 
380 (350), AOS TO (ASS) hone by 350-500 (385). 
S7TS-5A0 (430) wide, bearing 3 rows ol frooks, lst 
row LA0-T90 (175), P3S- (90 (155) 2nd row 170-190 
CIRO) 40-195 (170). 3rd row TO0-1 200) EO, 80-1 35 
(100) long. Oesophasus 1450-2075 (1790), 1S05- 
3430) (J970) long: cuticuhir dilation hearing 9-17 
holy hooks, Detrids 600-700 (660), 325-770 (405): 
seerelory-excrelory pore nol seen, 295-590) (420); 
nerve fing not seen, 205-405 (3455) (n=5) from 
anterior end (Pig. 28). Vulva 2-147 (Ee) ¥2- 
17.2 (11.9) fin froin anterior end. Vagina about) 15 
(n=2). Eves almost spherical 36-00 (50), 33-42 (35) 
by 34699 (34), Til THOT 210 (980), 920-1615 (1160) 
home (Fins 31. 34). 


Enyinaaleus 

The species nanie is derived tron the Latin fareny. 
mening Hidden. since ib wis nel found wher (he 
materi wits first examined, 


Type hese 
Jyoudor macrourus (Gould, e425 


{ype lowelity 
Mitchell Plateau, Western Australia, Australia 


Nite tee dake 
Small iitestine 


(ype specimens 
Holotype mule. ALC 30322. allotype fentile. ATIC 
AQS23, parutypes ATIC 13028 


Reimarks 


Linstowpreme fares sp. nove resembles L. 
WrPowonaise oceurriig inh daerouriy, in bein 
of siomihie sizes Qnales S21 mim in Lo deateny 
compured: with 15-20 nim dong in Lo werringieni), 


having the oesophagus of similar length (1575-1925 
in Lo hens vounpared with 1200-1825 in L, 
warring males) and fodr pais of papillae Gn the 
inale Gul, Phe agsaphiaguis i 2. laren. however, 
terminates posteriorly to the hooks surrounding the 
oesophageal region whereas that of Zo wereiagron 
terniinittes at about the level of the Yih ~ 130h row of 
hooks. L. dates hus 8-10 Gauley or 9-12 (emale) 
rows Of hooks, roots without undulating cdees, while 
Lo werringtent bas 91) (male) or 11-13 (female) 
rows of hooks. wilh toots having cinching edges 
oo the dikited cutie region, The body spines on 
the dorsal surface of male 0. fates extend further 
Jowurds the posterior end, Giboul aor the body), 
than ond. warriveron’ (about “Vs ol the body) 
Ventrally the body spies extend to the sume level as 
the inost amerior pair of cundual papillae oud, laren, 
but de pot on Lo warhead The male tail is longer 
aL. waertigtent (A30-430) than in Le etsy (250- 
340) The vagina of ho haens (115) ts shoner than 
thiol. terraced (300), 

Specimens were found ind prevonras Tron 
wortheryy Western Australia, the Northern “berritory 
vod northern Queensland, The population of 
nerthern hrawa bandicoots in Western Australia is 
salted from that of the Northern Territory and 
Queenslind (Gordon 1995), bib measurements of 
worms from hosts occurrii fh the Koinberley, 
Western Austrati. are consistent with those (rom 
Darwin. Northern Territory. The only niorphological 
difference observed between these pupulations was 
that the first row Of cephalic hooks of the Kimberley 
specimens Was alvest as long as Te second ri, 
ISQ-180, compared with 160-196 in males, bul in the 
Darwin specimens the difference in length between 
the Iwo rows of hooks was more murked. 105-163, 
compared with 126-195, This difference is got 
considered to be significant and us the Queenstind 
worms were similar to those from the Norther 
‘Verritory. the material from all three localities ts 
coustdered to be vonspecilic. 

The northern brown handicoots colleeted From 
horthern Queensland were infected wath 7, 
Warn S lrosts, Lo laters. 4 hosts. ot bow 
specues, 3 hosts. Chabaud ef af, (1980) wleatitred all 
the material they examined from the Northern 
Territory and northert) Queensland as 2 edverane. 
now 2. Wwercinglonk Was not possible to determine 
from their paper which, Wo any, specimens fron 
northern Queensland, presently lowed in the OM, 
SAMA or CSIRO collections, they examined. The 
specimens they examined trom Woolwonga and 
Darwin in the Northern Territory lave been re- 
examined for this study, and ave all datens, The 
Queensland material exumnbed by Chabaud ey ef. 
(M980) could hive been either 2. Werrington 1, 
fatens, or both, 


L.R. SMALES 


4| 


40 


LINSTOWINEMA ROM BANDICOOTS 7 


Linstowinema inglixi (Chabaud. Seurcuu, 
Beveridge, Bain & Duretie-Desset, 1980) 
comb, mov 

(FIGS 37-47) 


Synonyms: Lehinonemed eine sensu Unglis. 1967: 
py. 122. (28, 131-134, Figs 8-10, Eehinonenne aeglist 
Chobaud, Seureau. Beveridge. Bain & Durette- 
Desset 980: pp. 437438. Spratt, Beveridge & 
Waller, (994) p. 26. 


Material examined 

From Jyoodan obesulus Western Austrailia: 447. 
39 9 Murdoch, (ale 37'S. LIS* STR). 27vA8Sh, 
AHO 84901; 402-4, 10649, Perth. (34° 57'. 115! 
Sth, Jan. 1993, 95,1993, 311.1993, 1995. AMC 
40257, 20258. 30259, 30260, 30262. 20263, 30264. 
30005: Lhe ot. 44-29. Wartle Grove (3202'S, Elo" 
QO'E), 3.y, 1906. BM (NH) 1967. 616-626. WAM 25- 
7; We 2. TYP, Glow Forest. (Oovitt.6@. ATIC 
2u7A0, BM (NH) 1967, 473-523: 4¢ a. 109 9. 
Horesulule, LSI 1978, AHO 8885S; 40d 2. 108 
Jorrahdile, (AZ) 30'S. 16° O7'L, 26.x.1993, ATIC 
30261-44489 4, Albany, (357 00'S, 117° 52°B), 
oxi F944. TONE 9d CSIRO N4242_. N4243: 
Wad. 322%, Manjimup. (35° 15'S, 1160 09"). 
°7.vi. 1903. 16.6x.1993, AHC 30295, 30256: 625 4, 
IAS] 3, no locality given, 22.68.1977, AHO Sd8s 


Descriplton 

Cephalic bulb with 4 rows of 14 tale) (Figs 38. 
49) or 16 (female) files of hooks 2nd row longest, 3rd 
row shortest, neck with 5-11 rows tiny spines: 
cuticular dilation of oesophageal region bearing |) 
i4 rows, 14 (mide) or 16 (emule) files ot body 
hooks. [st and last rows smallest, 7th - 9th rows 
hooks Jargest (Pigs 3, 7), roots of hooks with 
lindulating edges (Figs 40. 41). Remainder of body 
with up to 35 (inale), or 45 Gemaley small spines at 
euch annulation, over whole body of female, 
extending to 400 yim from tail tip Gn dorsal surlace 
and ubout 300-500 anterior to cloaca, not extending 
to anterior pair of caudal papillae, on ventral surtuce 
of ynule body (Pig. 47), Qesophagus '/11 (mile) to 
H/ja (female) bedy length, simple. club-shaped. 
lerminiting level with Bth-9th row of hooks. Nerve 
ring surrounding oesophagus within cephalic bulb. 
seeretury-exeretory: pore in neck. deirids comical. at 
level of Ist raw of body hooks. 


Male: Length (1-L8 (15) mm. width 325-625 
(540), Cephalic bulb 266-325 (272) long by 247-350 
(309) wide; cephalic hooks, Ist row 140-180 (155). 
Ind row 160-200 (180), 3rd row 115-140 (125) long 
(Pigs 3, 7). Oesophagus 1120-1565 (1400) longs 
culicular dikition bearing 10-12 rows body hooks, 
Deirids 410-650 (520); seerctory-caxcretory pore nol 
seen: nerve ring 412 (H=1) from) anterior end, 
Spicules similar. equal, without alice 700-7200 (985) 
long. about ‘y= body length. Guberniculum short, 
sub: tritngular. 90-58 (n=5) long. (Pig. 45). Nine pairs 
caudal papillae: 4 pairs ventral and inimediately pre-. 
ad- and prost-clowcal: | large pair lateral, ad-cloaeal. 
2 pairs dateral. pre-cloacal; 3 pairs papillae, pair 
phasmids well posterior to elowea, near bul lip (Fig, 
47). Cloucal region with small cuticular bosses, ala- 
like extensions of body absent. Tail 150-350 (205) 
lony (Pig. 460). 

Female: Length 15-30 (24) nim, width 500-935 
(700). Cephalic bulb 312-357 (334) long by 293-422 
(334) wide; cephali¢ hooks, Ist row 155-195 (180). 
2nd row TR0-235 (205), ard row | 15-150 (125) long. 
Ocsophagus I385-1835 (1655) logs cuticular 
dilation bearing 12-14 vows body hooks. Deirids 
485.570 (S45): secretory-exeretory pore 390 (n=l) 
frony anterior ends herve ring, sol seen. Vulva 9 &- 
(0.2 mm (n=2) (Pig. 4a) from anterior end. Vaya 
about 350 (n=L). yg almost spherical 40-48 (441) 
by 36-45 (40), Tail 610-920 (730) long (Pigs 42, 44) 


Type hose 
Jyvodon obesuluy (Shaw, 1797) 


Tyne foceltty 
Waitle Grove, pear Perth, Western Australia. 
Australia 


Site i hast 
Small intestine 


Type specimen 
Nestype BMINH) 1967 616 
Remarks 


Material from /. e@hesifuy tron Wattle Grove. 
collected on 5.v.1966 und identified by Inglis ts 


Figs 37-47. Linafeavinemer taglist (Chabad. Senreau. Beveridge. Bain & Duretic-Desset, 1980), 37, Anteriun end, (lareral 
view), 38. Cephalic end (ea face view), 3 Cephalic end nile, optical section at level of first raw of hooks ter face view) 
440, Body hook (dorsal view), +1 Body hook (lateral wiew), 42. Female tail (lateral view). 13. Vieina (lteral view). 44 
Female tail tip (literal view). 45. Gubermaculum ¢yentral view). 46, Male tail (htteral view) 47, Male fil (ventral view) 
Seale bars = 200 yon 47, 42250 pm 38, 39.43, 44, 100 pin 40.41, do. 47. 25 pm +5. 


Is 1. ROSMALTS 


deposited in Wie WAM and BM (NE). This appears 
lo be the material deseribed as 2. cineuun by Inglis 
(1967), T therefore designite a specimen trom BM 
(NH) 1967. 616-626 as the neatype. 

Linnie dteliyi — (Chabauidl — Seureaut, 
Bevenidee, Bai & Duretic-Desset, 980) can be 
distinguished front L, werrtaetoud and Lo lens in 
having (he oesophagus terminating within’ the 
wileror coliculer dilution. Linstowineme inglisi has 
1-12 (rate) and 124 (emule) rows of hody hooks 
compared with 81 and lbela rows for o., 
weeringwo) and SAO and 8-12 rows far do lererey. 
The spicules of £. felis, (Wis body length) ure 
relitively longer than for Lo weevingtond (oy body 
length) and 4. deren (iy body length), The male tuil 
al Lo jnglisi also differs fom both Lowarringions ane 
Hh larevis i being shorter 150-350 (265) compared 
With AXD-130 (390) ond 250-340 (407) respeetively 
vod in having only 3 pairs of papillae rather than 4, 
The female tal is also shorter (10-920 07300) in 
L. justivicompored with L. werriaatoi and Le hares 
(1000-1900) (1265)) and 7916S (L070). 
respectively, 

The pairof lateral pupitlic level with the cloacal 
ppentig is more prominent than the other pairs of 
pupillue surrounding aod anterior to the chouce of Z 
inglisi, Wi (es pespeect Lo dawlist resembles 2. 
edimendst trom dasyurid: nursapials but 2. daelisy 
differs trom &, edmeaney in having the 2nd. not the 
Ist row of eephulie hooks the longest, Lovstawinen 
cdinands? oceuts i Dayyerny falluveins trom ale 
Northern derrmtory while 2. daelivi oecurs ti 7, 
vheyuliy from southern Western Australia. 

Linstowinernes (iglisi can be distintished from L, 
cena, Which alse beours in abesulits and has the 
Oesophagns termining within the cuticular dilation 


OF the oesophageal region at about the 9th row al 


hooks. by (he number of rows of body hooks, 10-12 
(mle) amd 12-14 female) compared with 13-16 and 
Id-I8 in Lh. eimenin. The differenees between J, 
clon anid O. ediai are discussed i) detail under /. 
CTCL, 

The measurenents of L. dagtiod trom this study are 
waiiuent with those give by Toglis (1967). Any 
differences belween the Iwo sets Ol imusurenents 
are because Inglis (1967) measured smaller woriis, 
329-11 39 Fur the males compured with [1-18 inthis 
study and 11-184 for the females conipared wilh 
IS-AU in this siidy: Chabatid ep al. (1980) described 


L. Inglis) mules as having spines covering only 2 ol 
the body dorsally. Ao careful examination of 
specimens for this study, however bas shown tbat ibe 
dorsal spines, allhough tiny. extend alone about 2% 
vl hady, that is. further towards the tail tip than de 
the ventral spines. 

Linsewinemd inelisi has been found only in? 
Obesulas fron the south of Western Austeatia, 


Linsfrowineme taymantonsé sp.nov 
(PIGS dh-61) 


Synonyins) Echinoneme cinetuin sensu Munidiy & 
Green, M972: p21 Cin part, Aedinonemed tirelivi 
wi Sprit. Beveridge & Waller, 1991; p. 26 (in 
purty. Lelinonemea spt Chabad Seureau. 
Beveridge, Bai) & Duretie-Desset LOO: p. 453: 
Spratt, Beveridge & Waller. 199}: ps, 26. 


Material examined 

Type material: Lolotype oo, allotype '. tram 
Iynodon obesulios, Kingston (4h2" 50°78, JAN Rob, 
Tasnminits Thi (992, AMC 30820 40R71 
Parutypes [209 EES WARE AOS10, 30411 


Oren material: Fron lyovdemr obeadlus South 
Australia, Kangaroo Isdands bite, 499. Viviane 
Bay, (35° 59'S. 137" (312), no date. Lav LOSS. ABC 
4458, 30302: 133 2. 2492. Hundred of Grosse, 
Tune 1983. ATIC 30305: S28 172), Kacatta, (a5° 
59'S, 136° 56"), Sept, 1983, ALC 20300; [3.42 7- 
IS 9, Pulndanw (3S 47'S, 137° LOR U2iv, oud 
S.X1 1986, ANC 30300, 30307; bet. 12) Seal Bay, 
(36° 00'S, 1378 QO. 4atUs87s AH! 320304 
Tusmania: $3 % 1529  Beuconsfield, (41% 12'S, 
46" 49'E), 1991 AME 30315: 34 4, Glengarry, 
1" 21'S, [46° 52°B), 1992, ATIC 30304 J. 
Hobart, (42" 53°S, }47° JOR) 25. yi 1982. CSIRO 
NI674;, Fad. 8. Gow Runge. (42° 50°S, (47! 
PRED, Suivi 1992. AH 303808; Bsa, 139 2, 
Margate tip, (43° 02'S. 147° 16°E). (Oj 1993, AC 
30312: Loh. Upper Dromedary. ne date given, AHC 
4530, 1244, Ye 9) hoe collection data, ATC 
40313, 30344, 


Deseriplion 

Cephalic bulb with J rows ol [4 Ginaley av to 
(female) (Figs 49, 50) files of hooks. 2nd raw lonsest, 
id now shortest (Fig. 49). neek with Ss rows liny 


High 44-01, Aiaienydaenn femranieive sp. now. 48. Anterior end, optical section (literal yew). 10 Cyphalic bull cuter) 
view), 90, Cophili¢e end feniile. aptical scetion al level at Hist row of hooks (em fee view). 56. Fentale unl (ateral view). 
52. Mile posterior body spines (ventral views 53, Body louks Cateral views, 54, Bony hooks (dorsal views S35. Vagina 
Hater view), 56, Male til (ventuil view). 57. Gubernaculunt (vental view), 58. Kenale (ail tip Uateral view), 54 Male 
Tai) Up vente view), 60. Male tail tip (ventral view), 61 Gubernactdun Uateral view). Scale burs = 300 yim 44. S12 200) 
Jr OOO, TOO pn SO, SO pan 92, 938, 54, 55. 5A: 75 pom SF. 58,50: 12 fH OL 


aly) I. RO SMALLS 


spines; Cuhecular dikition of oesophagenl region 
bearing 13-15 rows af L4 (male) or 16 (female) files 
ol body hooks Tstauel last rows smallest. 9th Oth 
rows lurgest: moots of hooks without undulating 
edpes (Pigs 34.54 neotainder of body wath up to 4h 
(mule) or 66 (emale) small spines at each 
annulation, over Whole body of female, extending 
over “/) Of dorsal surface, terminating about 350 
unterior to cloaca, not reaching level of most anterior 
mae Gt caudil papilhie. on ventral surface of nile 
beady (Fig. 56). Oesephugus about Mois body 
leneth, simple, Clobeshaped, (cemimanng at level of 
Sth-TOth row OF body hooks (Fig. 48). Nerve ring 
sunounding cesophagis within cephalic bulb: 
seUreiory-e8crelury pore in neck: deinds conical at 
leweh or bat pow ot hedy hooks. 

Mile, Length [le T4 (12.6) jam, wilt 453-715 
(605). Cephalic bulb 215-295 (255) long hy 270-390 
(225) wide; bearip 3 rows of hooks, Ist raw L4U- 
195 (170), Ind row 175-225 (200), 3rd row LOa-130 
(120) long thie, 49), Gesophagis 1105-1580 (1380) 
Jong; cubeular ditaboo bearing 12-14 rows of boty 
hanks; deirids 355-480 (430). secretory-excrelory 
pore 300-440 (365) (n=O); nerve cing 240-300 (2801 
(n=4) from anterior ene. Spicules snvilar equal. 
ayittioal alive 720-1030 C890) tors, about /)) body 
length. Gubermiculim short, simple, sublriangalar, 
S0-55 (n=3) Jong (Pigs. 57. 61). Ten paies caudal 
papillae: 4 puirs ventrul and immediately pre- . ad- 
and post-eloical respectively, Lo pair heer! ad 
Slowed, 2 pairs hicral pre-cloweal (Pig, 50): 3 hiteral 
pairs larger (hig, 56); 3 pairs papillae, pair phasmiads 
posterior to cloaca. near tail tip (Piz. 39), Cloueal 
region with small cuuucular bosses. ula-like 
expansion of body surrounding cloaca, Tail 235-325 
(285) long (Fig. 60), 

Female: Lengthy 19-22 (21) moi, width 8TS-1155 
1965). Cephalic bulb 195-325 (255) lang by 470-305 
(460) wide; bewing 3 rows of hooks, Loi row 190- 
JNO (205), Ind rew 220265 (245). 3rd row 125-170 
(150) long. Oesophugus 1445-1990 (615) lone. 
coliculur dilaGon beanie [4-75 rows body hooks, 
Deirids 355-450 (405); sceretury-excrelory pore 375 
jn=1) [root anterior cna: nerve ring pot seen. Vulva 
3525-7600 (63500) (n=3) long (Fig. 55), Vayina 175 
(n=1) long. "Tail 715-935 (810) long (Pies SL, Sa). 
Leyes almost sphericil 36-48 (42) by 33-45 (48). 


Remarks 


Linstawineme wisnniieuse sp. nov. resembles 1. 
inelise and L, etmeram. alsa eecurring iL eheslas 
id hawviog three pairs af pupiiae on the male tual snd 
the Gesophagus terminating within the cuuculiu 
dihition af the vesophageal region, at about the sth - 
Lith (ow of hooks. Listawearenica Lasmeainiense can 
he differeritiited font 2. reelisr in having $2-13 


(male) and 14-15 Gemaley raws ot body hooks 
Without Undulating edges, compared with Le} 2 
(male) and 12-14 (female) rows of body: hooks wath 
undulating edges and fram 2. cine whieh has 13- 
16 (nme) and Les (lemme) rows af body Hooks 
With undulating edues Linger fisinaiiedse 
huirther differs from Z. veglive baying all tiree pats 
Of papillae literal and anterior to the clowea, larger 
than those surrounding the elouca. ane in having the 
vloacal region with ula-like extensions of the body. 
Linsrowinema inelist has only one pair ol larger 
lateral ad- cloacal papillae and does pot have the ale 
like extensions, Linglewihesur cinetun, which does 
have vhelike extensions of the body, has ventral houdy 
spines extending (othe level oF the mosianterior parr 
of latent] caudal papillae bur ZL. lesiaiieise does 
nol. Liwlewinema cinetam hax wi spe patty ot 
papillae literal ynd anterior to the ¢loaca the sans: 
sive bul Lo tesmiunienve tus the three hater pairs 
larger The guberniculan ts more U-shaped in 
vental view in L, trsmatiense than th, iulisior f. 
ernein, Pemile Le tasmuniense Haye a shorter far 
TISA35 (S10) than Feiner 986-1122 (1050). The 
besophugus is whoue To body length in ob. 
fasmanienye males compared with Us in Le cient 
andl T/ib ny de dewdist The vulva is elosey to the 
anterior end im be tewmanense, about OO mm 
compared with 1O mn int 2. taedise and 12 mm in 
cei, The vagina of Lo teasmaniense. VS. ts 
shorter thin in Lo emenon 300 and Lo tiglive 350, 

Chabagd et ah ()980) fignred the (ail ofa male 
specimen, registered as AHIC 4530, tran A efesutinys 
Upper Dromedary. Tasmania, There is new only ane 
dimuged male specimen in the bottle avaiable lor 
Comparison, bub such characters us van be seen, Und 
(he drawiigs of Chabaud ef ad (L980) (Fig. S BLE, 
p. 438), are consistent witht L. faymaniense, 

Cinstowaiemae teesmaniease appears fo have vv 
googruphic range whieh extends weross Tasminma 
tmd Kangaroo Islund. Three hosts trom Kungaron 
Island were infected with two species at 
Linstowinend. one With Lo efter and be 
lannnunie@nye, und Iwo wilh Le wyweringdent and LL. 
tasmaniense. This stiggests past links hetween 
Kangaroo [sland hosts. dadvlind Australian hosts 
and Tasmania hosts. 


Bivinelouy 

The species ts named according ty a Tabet found ant 
AHC 1820, "EA. cinedam tasmaniensiy’, here 
determined to be ZL. civetum. which had appurenity 
been writen by Chabuud era when preparing their 
paper ol 1980. 


Type lavatiry 
Kingston. Tasmania. Austria 


LINSTOWINEMA FROM BANDICOOTS 2) 


Type host 
lsoodon obesulus (Shaw, 1797) 


Site in host 
Small intestine 


Type specimens 


Holotype male, AHC 30320, allotype female, AHC 


30321, paratypes AHC 30310 


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


Synonym: Echinonema  cinetum 
Beveridge & Walters 1991: p. 22. 


SENSU 


Spratt. 


Material examined 
T\pe material: Holotype d, AHC 30023, allotype 
°, AHC 30097, from Perameles bougainville. 


Other material: From Perameles bougainville 1, 
19. 1 anterior end, | posterior end, no collection 
data, AHC 30055, 30054. 30053: 1d. L. 3 anterior 
ends, 2 posterior ends, no collection data. AHC 
4522, 23 5d. 1%. captive, University Adelaide, no 
date, AHC 13928. 


Description 

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 


63 


Figs 62-66. Linyfowinema peraneliy sp. noy. 62, Anterior end (lateral yiew). 63. Body hooks (lateral view), 64, Body 
hooks (dorsal view). 65. Female tail (lateral view). 66. Male tail (ventral view). Scale hars = 200 jim 62: 25 pm 63, 64: 


100 jim 65, 66, 


a2 L. RE SMALES 


liny spines; cuticular dilition of oesophageal resin 
bear LO-12 rows, 14 Gmale) or 16 (femule) Files of 
boely hooks, first and last rows smallest. bth-8th rows 
lamest, roots of books without undulating edges 
(Pigs 63, 64); remainder of body with a row of small 
spines at each curieulan annulition (auinbers of 
spines nol counted), over whole body of female; 
extending over 7) dorsal surhiee. terminating about 
300 unierior to elowea ventrully. reaching level of 
Wost uitenor pum ool cuudal pupillac, an verral 
surfave OF male bady (Pig. 66), Qesophagus simple 
Clubeshuaped, about Yine'/o body length, lermmunating al 
Vth lth row of hooks (Fig. 62) Sevretory- 
exerelun pare. derrids und nerve cing nut seen. 

Males in=2), Length 9 9 mim, wilth 475, 475. 
Cephalic bulb 235, 325 long by 234, 267 wide, 
cephihic hooks st pow 123, 129 2nd vow 117 150. 
Yl row 73, “3 tong Oesophagis FHS. 970 long, 
vuticuhe dilanon beuing b)-12 rows hody looks. 
Spreules stimilar, eqaal. without alice, 950, 1050 long, 
about 4) hedy length, Gubermeolin short, simple, 
subetyaedtir TT pairs candal papillae: 3° pairs 
veniral dnd dromediately pre-. ad-aid post-cloaval 
respeclively. | pair lateral ad-elowcal. 4 pairs Tatenal 
pre cloucak 7unterior pairs all about sume sive: 3 
pairs papillac, pair phasmids well pasterion lo clowes, 
nea) Gal Up (big. 66) Cloacal region with small 
riticular bosses, alt-like expansion wbsenr, Til 210. 
20 long. 

Ponale: (v2) Length |) inne. width 520, Cephalic 
haoks Ist row 2600, 145, 2nd row 140. 260, 3rd row 
78, 140. Oesophuyus (190, 1250) longs cuticular 
dilation hearing If-12 rows bouy Hooks. Tait 460 
louy (Fay. GS) Vulva not seen. kees more on less 
sphericul, 5t-S7 hy 60-63, 


Remurks 


Alihouph ther was only aosmall done ol 
Specrniens tind they were al in paar condigon, 
significant differcuces between these specimens und 
Oher species Of Lrwlaveiinemed Could be foe, 
Althouel) elements such as the vulva cold tol be 
seen, (he worrns appeared fo be mature. having 
fertilized eyes feamero. Since & homgativite is now 
extingbon meintand Austratia aud Tully protected an 
the istinds ti Shark Bay Western Australia, it is 
unlikely Cal any inure speeinens will becente 
avtilable, Accordingly this somewhat Taeemplete 
Jeseiplion is presented ae the best possible of the 
species under (he crreumstunces, AdaMeandieuy 
peramelly Sp, nov, most closely resembles 1 
warrington’ and the speeies thar have been 
(istiniished a the discussion Ob L. wanvingtend, ty 
Unit there are no more than [1-12 (male) and 1O-12 
Houle rows af body hooks compared with 9-11 
Give) and Lh) 3 Gemaled and the oesophagus ts 


relalrvely lone, extending (othe Tlth or 12th row nt 
hody hooks. The awa species van be differentited hy 
sive. Linswinenet wareingiont Wales, 15-20 wie, 
und Jemules. 32-41 pm, ate much larger worms. than 
L perameliy, Y and Tt im, respectively, Lin 
Mewinwne peruned’s has plain edges on the roots of 
is body hooks bue 4. werriigroné fas undulaune 
edges. Phe numbers anil dnranvement of papillae on 
the posterior end af the male are alse different, Lid- 
SOwinend percwielis ts the only species of Lin 
Mewinema wilh Tour pairs of pupillae Taleral anid 
anterior tothe clones ull other spectes bave three. 
The spivules of 2. penumeliy (950-1030, Wa body 
length) ane relatively longer haa those of 
merrier (680.1090, 1/20 boy: demethy, Ltr 
sme perwmelis has three pairs of papillae mean the 
caudal op hur Lowesriigread has five. The lai eld. 
peramehs. (210, 240 (male, 460 Wena, is slorter 
thun that of 2. anaiefigtand (330-430 gid 1OaO 
1900), The exss ob L, peranelis (51-57 by 60-63) are 
hirger than those oF 2. wurriagiont (3-44 by 3348), 

Liasnavineme perumely issiinilar ted. fredise anu 
4. eet. in having three pairs of caudal pupae, 
Iican he differentiated front these (wo specs i 
having (relatively lod oesophagus, terminating: sl 
the level of the postenor rows ol liaoke van the 
cuticular dilation of the pesophipgea! rewion. and net 
WKH toad in hiwing up te 12 rows of body looks 
without undulating edges compared with Lo olelian 
Up to be anh Le crrera, up ter TS cows of bouly 
hooks with undulating estes, 

Linstewinena peraniel’s rescanbles L. rsnmeniense 
Wy diiving, Four pairs of caudal papillae eur the 
cuudal lip and body hooks without wiadilatine cdees, 
Kt differs from L, resmenienve tn having the 
oesophians terminating at the end of the eumeuli 
dilation and not within it, up ta C2 tows ol buds 
Hooks rather than [5 and larger eggs, 51-37 hy 60-63, 
compared with 36-45 by 33-45_ 

Althotigh the third tow of cephalig hooks js 
relatively much smaller than the first and second 
hows in Lh. perameliy Vii mn other species uf 
Liiiowinema, the cophalic hook siaes vary wreath 
berween jndivicuads und se this muy web he a 
consistent chiaraeter, 

The cight B. howedinuille, dissected to prnvide / 
peramelis tor this study Uhree ot whieh were 
ifeeted), Were all rewistered in the SAMA in 1936, 
hut wo other collection date were given in the 
museum revister. The collection data for AHC 452% 
and 13928 are equally sparse. the information Gn the 
labels giving Goly the laculity as possibly Soul 
Austad und captive mi the Zoology Department ot 
ihe Liniversity of Adeluide. Examination of the AHC 
records jiedicates what five additional bundicoots were 
dissected for helnioiis, (wo ol whieh were mfected 
with 4. pecamelis, This suggests that the prevalence 


LINSTOWINEMA FROM BANDICOOTS 21 


al infection in Po beongeinitle by Lo permuneliy was 
ubout 48%, 


Epyiiolawy 

The specilic name ts taken from the label ot ALC 
4522. the material originally registered as 
Fohiianemie cine perameles, here determined ts 
bef peranielis. 


Type lacality 
Unknown, Australiu 


Type hese 
Perameles bouvarwville Quoy & Gitimard, 124 


Site tH dest 
Stall intestine 


Type specuuens 
Holotype mule, AHC 30023. allotype fenide, ABC 
40007 


Linstowinema maplestonei sp.nov. 
(FIGS 67-78) 


Material examined 

Ipe material: Ubolotype fd. allotype 9'. fan 
Perameles hasta Dinger Creek. (17> 26'S, 146 
OO'T), Queensland, Ty.60. ATIC 30094, 30095: 
Paratypes 40. 1 9. 2 anterior ends. 1) posterior 
end OM 1446341, ALI 19763. 


Oren jnatental From — Perameley — mrsite 
Queensbind; 2/7 5, no locality given. Jax.57, OM GL 
M4S7 1d. 29 S214 fragment. Fmishial (17) 42'S. 
146° OLR), 15.81.59. OM GL 4as56: 20 0. 2 
southern Queensland, no diate, AHC 1726, 

bio fyeadon meeronrus Queensland 2° 5 , 
Mossman, (10° 28'S, 142° 23°). 41-1958, OM GT. 
4303/2. 296. 299  Innistail, (17° 42'S, 146° 
QUE) 22.yi,. 79 ALIC 1726: 12, Brisbane, (27° 28'S, 
IS} OVE) no date, AHC 1738:54d 4,79. 1 antertor 
end, Paddington, (27° 28'S. 153° OLB) Aug, 195s, 
AHC 4371 from bandicom, io collection data, 3 3, 
ALLO 19667, 


Desc ripiis 

Cephalie bulb with three rows of Ib (vale) (Fig, 
69) or 16 (female) files of large hooks, And pow 
laraest 3rd row sipallest (Fig. 68): neck with 5-8 rows 
ol tiny spines: cuhicular dilation of oesophageal 
rewion bearing 11-13 rows of 14 (male por 16 female) 
files-of body books, firstuod best 2 rows stallest. 7th 
- Oth tows hurgest: roots of hooks without undulations 
(Pigs 72. 73), reinainder of body with a row of up to 
42 unaley or SO (female) small spines at eae 
wnndlation (bis. 71), aver whole body of female. 


extending over ‘4 dorsal surlace. termioaimy about 
400 anterior to cloaca On ventral surface of (ale 
body. Oesophagus simple. club shaped uboul iii 
body Jength jerminuting posterior to oesophageul 
cuticular dilation (Fig. 67), Nerve ring surroundiny 
oesophagus within cephalie bulb; secretary -excretory 
pore in neck, deirids conical, at level of [st row of 
body hooks. 

Male; Length 13-23 (17.5) mm. width 475-560 
(550). Cephulie bulb 285-350 (345) long by 260-310 
(290) wide: cephalic hooks Ist row EE7-150 (130), 
And row 140175 (160). 3rd row 90-130 (120) long. 
Oesophagus [3602210 ()804) Tong, cuticular 
dilation bearing 11-13 rows body hoaks. Deirids 405- 
S30 (APS). nerve ring 255-300 (285). sevretory- 
excretory pore 325-405 (360) [rom anterior end, 
Spicules equal. similar without ale 820-7 100 (1030) 
long. about Yo body length. () pairs cauclal papihwe: 
3 pairs ventral and iminediately pre ude and post- 
cloacal respectively, | large purr literal ad-clodeal, 2 
pairs lateral pre-cloucal 4 pairs papillae, pair 
Phasmids well posterior ty clowea, near ual lip (Figs 
74, 75), Clowcal region with small cultcular bosses. 
alu-fike expansions absent. Gubernaculun stort, 
simple. sub-triangulir in ventral view, 60 (n=l) long 
(Fig. 76) Tail 305-440 (375) long (Fig, 7X), 

Feniale; Length 18-35 (28) mm, width 560-765 
(070), Cephatre bulb 325-425 (362) long by 274-425 
(330) wide: vepbabe looks Ist row 150-176 (1604, 
2nd row 170-225 (190), 3rd row 115-145 (125) Jong, 
Oesophigus lolS-3740 (2272) longs cuticular 
dikttion bearing 12-13 rows body hooks, Deirids 405- 
Sas (495) nerve ring 260-390 (3350), secretory- 
exeretory pore (n=2) 405. 470 trom wnterior ena. 
Vulva nob seen. Tail 565-970 (790), Eggs not 
measured, 


Remarks 


fiastowinenta maplestone’ spo now. resembles 7 
warrington’ in having up tof fows of body Nooks. a 
rekilively Jong oesophagus and five pairs of caudul 
papillae on the male. TW differs from 2. weeringraad i 
having: FE-l4 (male) and 12-13 Gemale) compured 
with ® 11 (mated and 11-13 Cfenmiled rows of body 
books, The vesophagus of Lo mneaplestanes, extends 
beyond the cubeulur dilution of the Gesaphageal 
regan Of the hody whereas in L, warriteeront it 
termingtles at the levelof (he 9th - Tarh rows of body 
hooks. The hody hooks of L. maplestoved ave withour 
undulating cclves and thoseal Lo werrieren are with 
undulating edges; Mule body spies extend along Va 
of the hody dorsally and well above the lateral caudal 
papillae ventrally on Lo viaplesraner bur on L, 
Wario they extendatoig */s of the boudy dorsally 
undalmost to the lateral caudal pupitlae ventrally. All 
6 pars of pre-, post- and ad-cloacal pupillac are the 


LINSTOWINEMA FROM BANDICOOTS a3 


same sive on Le awarriteron’ bul L, anaplestenet las 
larger lileral al-cloucal papillae, The spicules of 2. 
maplestoner (/\ hody length) are relatively: longer 
(hit those oF Lo werraguiné (fn body length), The 
arrangement Of the papillae surrounding and unleror 
WO the elowca of 1, mreplestered inust closely 
resembles that for Lo img/id ie. wilh the haberal ad- 
clowcal parrthe lamest and the ventral body spines net 
extending to the level oF the cuudal papillie, 
Laistoyiineiie maplestone? differs. tram L. inedlisi im 
haying the oesophagos extend posterior to the body. 
hooks without undulating edges rather Wii an 
besbphagis which termites within the cubieular 
(ikiHon ait about the level af rows 4-9 of the boily 
hooks will undulating edges, Male 2. fy/isy hive 
only three pairs Of papillae on Gre caudal tip but /.. 
maplesione? have four, Male Lo taplestened on 
uverive huve longer tats (405-440 (A7S) than 2. 
inelish (150-350) (205)), the body spines al L, 
maplestone’ extend along oF the beady dorsally 
compared with almost the whole body (4a) on ZL. 
inelent. 

Linvtowinena meaplestone’ can he distinguished 
from £. litfeas. which also hus the Gesophagus 
extending posterior lo the body books, body hooks 
wilh undulating edges and four pairs of papillic on 
the caudal tip. dh having t)-13 Gmaley) and 12-134 
(lemale) rows of Nooks compared with 8-10 (mule) 
and 912 (emake) rows ef body hooks ind. datens, 
Male Lo mapdéestaned have body spines extending 
Wong “of the body dorsally and well phoave the 
Kuteral couch papillie ventrally compared with / 
lateus. which hits body spines extending alan ia ot 
the body dorsally und level with the inmost anternar 
Lateral caudal papillae Vventtally, The literal ded 
clodeul pair Of papillae os largest On Le mraplestene! 
compared Willi wil 4 pairs of lateral elovedtl papi tlie 
heme the same ste oo L. tates. Memute 2, 
aplestane. Wuve on average shorter daily 65-970 
(790)) tha 2. lereiis (790. TB ES CLOT). 

Linsionwineni ineplesiamet Gan be drflerentucded 
rom Forel in having bI-ta 0 (miley, T2-13 
(emailer body hooks without undibaring edges 
vompured With }A-L6 (malen I-TS (female) hoy 
hooks with unduhiti cages: a ling oesophagus 
depminating posterior (6 (he pows af body houks, 10 
A shor GesOphiigie lernmutine at the devel of te 8 
Li rows ol hooks. Liason nuples(ones tas Tour 
pues af caudal papillae at the tail tip. awhile / 
crete has Three, the pair ot Lateral audbcloacal 


Figs G7 7S 


papillae is the largest on 2. deplestonter but all six 
pas of cloaci) papillie qe the same sive on L 
cine, Linslownrenre nreplesteme? does: not have avy 
alitike expansion surrounding the clowea bur le 
Cinetim dues: the yentul body spines oof 2. cieria 
extend dlmost tothe level ab the inost anterior lateral 
pairol pro-cloucal pagatlae but nL. aaydestener they 
do nots the female tail of 2. wuplestene’ (3035-970) 
(790)0 os shorter than thar of Le einer (986-1122 
(1050). 

The other species of Lyostewimmena, lL perunntis 
wd 4. davaniense. which Nave body hooks without 
undulating edges have three pairs OF caudal papillae 
on the ual tip. These can be further differentiated 
from Lo dplesrone’ by the number ef rows of body 
hooks and the relative lengths of the oesophagus, 
Linsiowinema purametis has 10-)2 rows of buds 
books with the oesophagus terminating al the level of 
the L2ih rows... feayvmenmienve has (3-15 rows of body 
hooks with the oesophagus terminating at the level of 
the &th = (Oth row anid £. meplesroned his [l13 tows 
of body hooks with the pesophasus terminating: 
posteriorly to the (3th row. Lantenvinema peranelis 
hus four pairs OF kvteralad- and pre- cloacal papillae, 
compared with three pairs on L. aaplestoned and 
relatively shorter spicules ('/o of body lengthy 
compared with (jn) Linwtawinemer teasnaniense (ibs 
three large pairs of lateral cloaeal papillae compared 
with the one lange ad-cloueal pair of L. mreplestarrer. 

Beeuuse af the small number of female specimens 
wiluble, none was dissected, so allliourh enes Were 
seem, iL was diffieull to delermine which were mature 
and therefore suitable for measuring. As a result. no 
INGUSUreMenks were made. 

Linsnuinenid meiplestonel oeeurs in Poet and 
J mecroures (ron) Queenshuid and New South Wiles 
alhauvh (he examinition of mere hosts is required 
before the full extent of the geographic range wl (his 
Species cun be deterinined. In twa 1 retinitis. b. 
maplestaner occured ait iiwed mmfechons with J. 
warrt elon. 


Etyinndogy 

The species is nunmed ufler RAL Maplestone, whe 
together with W. Yorke. cured uut pioneering work 
on othe nematodes of Australian marsupials. 
Eelinenenne eplestone? Was Used on in tindited 
museum libel by Chibaud ef af (P98O) ton 
Liayioawineme specimens (rom Zo macrourus that were 
subsequently determined tobe Lo werringrean 


Lawton jee maplestened sp. nov. O72 Adnenor cid, optical section (litera view) 68 Cephalic bull Hater 


view) OY Cephitic end male. optical sectional tevel OF rst naw at hanks be ice SOW L720, Fenaite tal lateral view 1 
1 Male, posterior houy spines (ventral View). 72. Body hook (later Wiew). 73. Bady haok (darsal yew). 7 Male gal 
Lip (ventral views 79. Mile dail (ventral view). 74. Gubermiculini (vented) view) 77. Flonule if ep cheteeal view TR. 
Mule till Hatera) view, Suale burs =S00 Li O72 200 pans. TOL S50 pin 69. 7577. 25 pm FD TITY Ta, Tee HOO i 7 


ab LR SMALES 


Type liast 
Peranteles Waste 


Type locality 
Dinner Creek (17° 26'S. 146° OO). Queensland. 
Australia 


Site it feast 
Small intestine 


‘Type speemnens 
Holotype tale, 
OMA0N95 


QOM30094, (lotype female 


Discussiun 


Linshovinenae warrington’ appears ta be the 
domimint species of Livacewimenn, in eastern 
Austrian bindicoots. being found) from oorthern 
Queensland throweh to South Australia, including 
Kawigaroo tshiid, He ocedrs ib all extiit species: of 
Avoedon as well us one individual of Po acyl, 
indicating a low prevalence ii (his latter host species, 
Althourh ©. faery wes found) in northern 
Queensland populations oF the wurthemn brown 
bandicool, together with L. warringteni, it was the 
only species oecurrimg in northern brown bandicools 
fromthe Noriher Territory and (he north of Western 
Austialia, while L. fragilis? wis the only species 
ovcurtiiyg in southert, browa bandicouts m rhe south 
of Western Australia. 


Only one species, L, peramielis oecurtue a Ave or 


13. 7. horganvilte examined, has been found 
exclusively in Perales spp. and three species 1 
nasmanenye, L. larens and &. inwlist exclusively in 
hsondon spp. £. dates Oecurs in fe macrourus, L 


jasmiemicnse aid L, iedit oceur itl obesutis, OF 


the other three species. 2. dapleatoner has a north 
easter distribution occurring in 7 yee rouris and P. 
nosuid. Lo Cincvieea more south-custern distribution 
pecurring nL. ehesdlus. Po nesete and Boa, 
Further collections of material from south-custerti 
Austidia are needed before any typothesis on the 
distimbulion of species of Liasrowienur cun be 
Weveloped. tt does appear thar bandicoots (/ 
hess) Ot Kangaroo Iskimd, bur ot Tasmania, 
miy have been derived from stock in which all three 
species (2. weiinyrond. Lo einetiin, and b 
lusmentemse) Were prevent The continuing 


detrimental elects of Ruropean serlement Taye 


resulted imu putehy distribution off obesitas over a 
reduced yunge (Braithwaite 1995) This decline inthe 
host population may have affected the distribution 
and prevalence Of species ol Linstearinema on the 
iidinkind. The current prevalence of the three species 
of Linstawwiena in banweoots on Kangaroo Ishind 


may reflect past prevalences of these species i 
bandigoots on the mainland. 

The whility to trap bandicoots yaries with species 
their age. species and Ipeulity (Gordon & Hither 
1989). Peranedes nasune is apparently more difficult 
lo trap tha species of /yoadun (Menkhorst & 
Seebeck 1995). This nay be the reason for the small 
number of Peraeles compated with Ssecdon 
collected in this study (see Table |) and in the 
amolntof material deposited in musciim collections 
froin each host genus. A farther coniplicuting lactor 
is the possible difference in prevalence of infection 
with Linsiowinena berveen the two, The records ol 
the SAMA and QM indicate that 90 2. nase have 
been examined for belnmth parasites and. of these. 
only eight were infected with LL. taaplestanel, six 
with Lo elven and one with Lo weannietens. 
Similarly, of SEA gem examined, only tour were 
infected with Lo cinenue and one with Ff 
werrtiwton’. dos unlikely chat Linvtowinena could 
have been Overlooked during dissection as (he worn 
are Taree nd cine be readily detected in the simill 
intestine. These low prevatences of infection contrast 
with Une prevalences found for species ol deeaden 
disseeted in this scudy. OF 72 hancicoots examines, 
53 were infected with Linstowimenk spp, 

A working hypothesis would be that species af 
Lintomineme are dominant in the helminth 
communilies OF fseovdow but net in those ol 
Perameles. Wi some areas. species of bandicoot haye 
overlapping geographic ranves, do acronvuy and PB 
Hasite in the north-east, (oohbesi/is and P pasiited in 
(he soulheeast and 7 obesity atid Po gee 
‘Tasmania, “Their habiti! preferences within each 
geographic region are different, and although they 
Inay nol be ii striek symypalry. opportunities foe 
Incidental infeedion and bost switching would Gx ise 
Observations trom this stidy suggest hit species at 
Linstowinenae nay switch trom 7. wheal be P 
gunn and PO eesues Veron Eo abesidige loa P 
honviwitte ond perhaps fron / onusutia ta | 
mrechnens, Cho mht aecaunit for rhe Geeurenee af 
Lingiwvinemea i Perameles. 

Additional collections of material fon Percmieles 
spp. across Australi and especially 7 ebesedtay trom 
south eastern Australia are needed to lest thes 
hypothesis, 


Acknowledaments 


Thanks are due to the lollowing collectors whe 
provided miatterial for (his study; [, Beyerndec, 
R. Beavis, T. Th Cribb, T. Dennis, M, Driessan, 
T. Foley. To Priend. Re Purer, Bo Gosslink 
P. Hayeoek. Po Hayward. O. Hiekiman, Bo Mundsy. 
R. Noking R, Norman, D. Obendorf, b. Owes 
D_ Spratt, the late M_ J. Mackerras, P. Presidente and 


LINSTOWINEMA FROM BANDICOOTS 27 


the late A. J. Bearup. Thanks are due to-C. Kemper of 
the South Australian Museum, J. M. Dixon of the 
National Museum of Victona and T. Friend of the 
Department of Conservation and Land Management. 


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


References 


Brarhwarre, R, W. (1995) Southern Brown Bundivoot pp, 
W7O077 fa Strahan, Re (Rd) “The Miarnmals of 
Austrailia” (Reed Books, Chatswood). 

Chiabauo. A G,, SbukiaL, C.. Beyeripct, 1. BAIN, OL & 
Diunerve-Drsser M.-C, (1980) Sur les Nematodes 
Eehinonemiatinae. Amy. Parasol, lum, comp. 55, 427- 
da, 

Carink, OJ. CES81) Supplementary report on speeimens 
dredeed up from the Gull of Manaar together with others 
trom the seu in the vicinity of the Basse Rocks und from 
Buss's Strails respectrvely, presented to the Liverpool 
Free Museum by Capt, Th Cawne Warren. Aan, Afar, 
Nat. [livt. 7, 3601-385. 

Prinp, J, AL & Burpipdh, ASA, (1995) Western Barred 
Bandicoot pp. T7180) fe Striban, Re kaha Phe 
Miainimiils of Australia” (Reed Books, Chatswood). 

Gorpon. G, (1995) Northern Brown Bandicoot pp, | 74- 
175 shied, 

& Unninien. A. TE (1989) Perumelidae pp. 603-624 
fn Walton, D2 Wo & Richardson, B. J. (Rds) “Fauna of 
Austealia’ Vol 1B Maminlia (Australian Government 
Publishing Serviee, Canberra). 

Incuis, WG, (M967) ‘The relationships of the nematode 
superfamily Seuratoidea, J, Mefminthel, 41, 115-136, 
JouNstos, TFL & Mawsos, PM. (1939) Sundry 
nemiitodes From eastern Austealisin Marsupials. Trevis R. 

Sac §. Anst, 63, 204-209, 

R940) New and koown nematodes from 
Australian marsupials, Prac, Linn, See. NSW 65, 468- 
476, 

& (1952) Some nematodes trom 
Australtae birds and mamoials. Tren Ro Sue, 8. Aust. 78, 
30-37, 

Linstow, O. Vor (1898a) Nemathelminthen von -lerrn 
Richard Semon in Austrilien sesimmell, Denkschr vid: 
neathw Gesellseh. dene 8, 467-472. 

()989h) Nemathelminthen. you Herm R. Semon in 
Australien gesummelt,  4eelogisehes Central Blan 
Letpziv 5, 672 


MeKenvih. No1 Morris, KR. D.& Dickman, CR, 11995) 
Colden Bandicoot pp, 172-173 dn Strahan, R. (Rd.) “The 
Manimals of Australia” (Reed Rooks. Chatswood). 

MackErRAS, L Mo & Mackrreas, M. J (1960) Tuxonomy 
of the vommon short-nosed marsupial bundicoot of 
eastern Queensland. last do Ser, 23, 51-53. 

_ & Saspars, D. F. (1954) Punasies of the 
bandicoat, Proc R. Sec, Qld 63, 61-63 

Mantowgy. J, A. & Ribk. W. DL, (1988) Peramelidie pp. 
36-42 Jn Walton, D, W. (Lid.) “Zoological Catalogue of 
Australia” 5S Mammalia (Australian Government 
Publishing Service. Canberra), 

Menwnnorsr, PW. & Srapeen. J. A, (1995) Long-nosed 
Bandicoot pp. 77-78 fa Seebeck, J. bE (Ld.) “Mammals 
of Victorit Distribution. ecology aud conservation™ 
(Oxford University Press. Melbourne), 

Munday, Bo LL. & Grken, R. G. (1972) Patisites, of 
Tasmanian native aed feral fun Pert Uh blelminthes. 
Rec, Queen Vie, Muy. 44, 1-15, 

Qurntin, J.C. (1970) Sur le cycle evolinil de Searanin 
cvadarachense Desportes. 1947 et ses uffinites avee ecus 
des Nématodes Subulures (Ascuridia) ef Rictulires 
(Spirurikiy). Ann. Parasitel, non. conip, 45, 605-628, 

Romer, & (1901) Monotremuta und Marsupialta. Derdsclie 
med-naitew Gesellyeh, dena, 8. 153-160, 

Srepecn, J. G. (1905) Eustern Barred Bandicoot pp, bs82- 
183 /n Strahan, Re (Rd.) “The Mammals of Australia” 
(Reed Books, Chatswood), 

Sprarh DL MM. Beveruogi. 1 & WaAuTiR. Eb, (hoor) A 
cutulogue of Australasian monotremes and oursupials 
and their recorded helminth parasites. Ree. 8, Aust. Muy, 
Manor ser t, 1-105. 

STODDART. B.( 1995) Long-nosed bandicaot pp. Us4-1K5 fv 
Struhan, Ro (id) “The Mammals of Australie” (Reed 
Books. Chatswood), 

Yorke, W. & Marcestonk. PA, (1926) "The nematode 
parasites Of vertebrates’ (Churchill, London), 


A NEW SPECIES OF FRESHWATER AMPHIPOD, 
AUSTROCHILTONIA DALHOUSIENSIS SP. NOV., 
(CRUSTACEA: AMPHIPODA: HYALELLIDAE) FROM 
DALHOUSIE SPRINGS, SOUTH AUSTRALIA 


By W. ZEIDLER* 


Summary 


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. 


Tenscetions of the Rovet Suciely uf S Aust (1997), 12161), 29-42 


A NEW SPECIES OF FRESHWATER AMPHIPOD, AUSTROCHILTONIA 
DALHOUSIENSIS SP. NOV., (CRUSTACEA: AMPHIPODA: HYALELLIDAE) FROM 
DALHOUSIE SPRINGS, SOUTH AUSTRALIA 


by W. ZEIDLER* 


Summary 


Zio, W. (L997) A new species of freshwiter amphipod Auvrechiltonna dalhousrensiy sp, ney. (Crustiven: 
Amphipoda: Hyaleilidaey fon Dalhousie Springs, South Australia. Frans. A. Soc So Aust T2101), 29-42. 30 


May, 1997, 


A treshwiter amphipod Ansochienia duliousienas sp, noy, is deseribed and Ulusteated, Iis-endemie tua 
lew urfesian springs amongst he Dalhousie Sprigs comuples In the north of South Austealia. Morphologically 
itis very Similar to other species of Ansireeitonia found in mound springs near Lake Eyre South but 
preliminary cleetrophoretic analysis of allozymes suipports the revagnition of a distinet species. IL imost closely 
texernables \ aresdraliy (Sayce. 1901) 1h thal uropod 3 js lwo-urlieutale, bul differs in woumber of minor features, 


Which collectively distinguish 1 fromm ils congeners, 


Key Worps: dasieeditiena dallomyiensis sp. noy.. 


(Uxonomy. 


Introduction 


Aniphipod species of the genus Austrchitontesure 
among (he most common crustaceans Found in (he 
permanent freshwaters of soulhern Australia ranging 
from New South Wales to Western Australia and 
including ‘Tusmania. More recently Austrochilyn le 
Nasaudso been found in the inlaid water's of artesian 
Springs ii South Australia (Zeidler 1989) and at 
“Edgbaston” north-east of Aramac, Queenslond 
(personal colleehion. May 1988), 

When | re-estublished the genus Ausrochiltonia 
(Zeidler L988) it was my mlention to proceed with an 
Australian revision of the genus beginning with the 
description of species found in the mound sprigs 
Wear Luke Eyre South and at Dalhousie Springs im 
Northern South Australia, Since then | have exainmed 
a large number ol specimens from wide-ranging 
habitats in Southern Australia and have found them 
all tobe very similiar morphologically ond difficult to 
distinguish from the only previatisly-described 
species, A. anseralin (Sayee. LOO1) and Al subtests 
(Sayee, 1902). Williams (1962) revised the 
s¥slematics Of these bWwo species based om type 
inalerial and a range of specimens from New South 
Wales. Victoria, “Tasmania and Rotimest Istand. 
Western Australian and likewise found that, 
morphologically, specihe differences are minimal 


However, a preliminary andlysis af allozymes of 


specimens from the South Adstrahan mond springs 
usin electrophoresis, idiodtes that Austrochilionia 
in inost likely a yery speciose genus. Given its 


Soul Atiatailion Maseurie Sarthe Terie Aelia 8. Aust 
SUM) 


new species, Amphipod. artesian sponys, Australia, 


potentiul enormity, the project was abandoned duc to 
lack of resources. 

The species found at Dalhousie Springs is most 
similario A, ausirafiy Sayce. 1901 in that uropod 3 is 
two-arliculate. Tt has a very restricted distribution. 
oecurring at only three of about &0 active springs ti 
the region (Zeidler 1989). “Two of these springs are 
quite durge, with large outflows of warin water 
(P40°C) bul Auvtreciitenia is found only in the 
distant overflow where the waler is colder and close 
to ambient temperature. However. one isolated 
specimen Was colleeted from the edge of the pool of 
the main spring, which has w water temperature ot 
about 35°C. The other spring is a small relatively 
vold 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 Cyperus laevivars 1. 1771 
and sometimes alse the reed Phragmites australis 
(Cuv., 1AebL). 

The restricted and isolated distribution of this 
species of Austrachilionia makes i) vulnerable to 
habitat distarbance even (hough Dalhonsie Springs is 
within Wihira National Park, The purpose ob this 
paper is to establish the taxon so that park managers 
und visitors can appreeite tS significance and 
potential vulnerability, 


Materials ond Methods 


The Dulhousie Springs coniples (Pry. 1) consists 
oF about SOQ uclive springs all of which were sampled 
in 1985 (Zeidler & Ponder 1989) hut Ausrochilnania 
was found in only three springs (Fig. 2). The springs 
we coded following Zeidler & Ponder (1989, Fig. 2). 


a0 W. ZEIDLER 


Animals were collected from amongst sedges and 
reeds with a small hand sieve or picked off plant 
debris with forceps. A total of 424 specimens (230 
90.174 oo, 20 juveniles) was collected and 
exumined. 

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


: . 


1 LE SONS H 
: \ 
| | / 
{ Northern Territory | Queensland 
' \ j 
i} 
\ \ h 
\ | j 
kL = - -—=- Se — ——yPrdelle 
\ Dalhousie Springs* 4 
| F 
i 
| 
; 
1 py Lediardatia if 
! i 
\ ° | 
| oO i 
) v 
\ iD RYH 
| Leaver Helps AW 
\ Matinee rv 
| South Australia ; | 
\ at) j 
! j 
| / | 
i \ 
\ a) | 
1 \ I 
ae j 
rig \ 
é \ 
gies nugeane | 
i) i 
: | 
i 
| 
. i 
i Borcelside | 
i ' 
WoT | 
~ \ 
\ | 
1 
\ I 
1 
| 
! 
| 
Saule \ 
a 


Fig. |. Lovation of Dalhousie Springs. South Australia, 
From Zeidler }O91, 


Table | and data on nearby springs are also available 
(Smith 1989). 

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

Material reported here is deposited in the South 
Australian. Museum, Adelaide (SAMA) and. the 
Ausiralian Museum, Sydney (AM), All specimens 
are preserved in 75% ethanol or 2% 
formaldehyde/propylene-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 
mouthparts, uropods and telson are mounted in poly- 
vinyl lactophenol on a microscope slide, 

Specimen length is measured along a. lateral 
parabolic line drawn from the anterior extremity of 
the head through the mid-line of the body to the 
posterior limit of the telson using a pair of dividers 
and seule, 

The thoracic limbs are referred to as guathopod | 
and 2 followed by pereopods 3-7. Size comparisons 
of gnathopods exclude the coxa and dactylus, and of 
the pereopods, the coxa, with articles being 
measured along the mid-line, 

The following abbreviations are used in the text 
and figures. Al. A2 = first & second antenna, G1, G2 
= first & second gnathopod: LL = lower lip: Md = 
mandible: Mxt, Mx2 = first. & second maailla: Mxp 
= maxilliped; O2-5 = oostegites fram pereopods 2-5; 
P3-7 = pereopods 3-7: PI = first pleopoda: T = 
telson; UL-3 = uropods 1-3: UL = upper lip; r= used 
as sufhx to indicate that appendage was taken from 
right hand side of the animal. 


TALE b. Temperate measurements and phystoachemical data (from Smith 1989) far springs fram which aniphipads were 
vallected at time of collection (except far Cal - data from [983 expedition) 


Field Chemistry 


Spring Temp. Temp. 
Aw Wiiter Temp. Cond. TDS pH bo 
Cc *C “C ASHE mgt! ppm 
siemens 
Cal - channel to main pool - - 43 1490 865 73 3.8 
Cul - main pool x) 37 34 1780 1o00 7.9 67 
Cal- main discharge channel = 25 36 33.5 2050 1150 7.7 47 
Cd? - SW edge of pool 15 yu 32 1550 850 7.9 11.3 
Cd2 - at or near swamp 15 11 ts 1650 - TAS 7.6 
Gb 13 16 20 THO 4850 7d 45 


A NEW SPECIES OF AUSTROCHILTONIA 4 


st ANS | 


Ls 


.0 


=I e | = St ‘ 5 ’ . 3 afl y , Hf 
‘Dalhousie (ruins)} 
wo 


« 


Fiz. 2 Dalhousie Springs complex showing springs (coded) from which Austrochiliania dalhousiensis sp, nov. was col- 
lected. Collection sites for Cal and Cd2 are arrowed. Other major springs are shown as dots. Swamps [rom springs and 
ebeck heds are stippled (light stippling indicates ephemeral stream beds and heavier suppling areas oF permanent” walter). 


32 W. 7EIDLER 


Systematics 
Austrochiltonia dalkousiensis sp. nov. 

(PIGS 3-9) 
Austoclifimoa sp. Zeidler 1989 83-K4, fig, 12.) B. 
1YOLS 185 
Holton &, Dulhousie Springs SA, from amongst 
reeds and sedees along eastern edge of swamp 
crealecl by outflow from main spring (Cal), 3.3 kin 
doth of edge of old airstrip. 26°23'07" S 135°30'26" 
hi. b2vi1985, W, Zeidler & K.L. Gowlett-Rolmes. 
SAMA C565]. 
Alfplypes Oviverous & SAMA C5652, Collected 
with holotype: 
Paratypes: AM P48840, 10 2-9 100" oT sume data 
as holotype. SAMA C5653, 24 29 (one 
oviserous), 17 cf sane data as holotype. SAMA 
C5054, 37 GY (three ovigerous), 11 oo, same 
data as holotype except 14-vi. 985. SAMA C5655, 
24.9 DV OO Lsame data us holotype but ).) kim 
further north, 26°22°26" S L35°30°2A" EL 


Giher material examined: Al From Dalhousie 
Springs urea (Vig. 2). AM P4a841, 1 2 (damaged), 
sprit Cul, 26°25700" 8 135°29'53" BE. trom edge ol 
main pool, WF, Pander & BD. Winn, 3.vi.1985, 
SAMA ©5656, 2) GY (three ovigerous). 31 a, 
spring Cd2, fron -edges of swamp formed by 
outflow, approximately 0.9 km NW ob souree. 
262433" S 13597845" E,W. Zeidler & K, lL. 
Gowlett-Holmies. 6,11 1985, SAMA C5657. 14 
PO, 2 Aa, same data as previous lol except 
ori (O85. AM P48847, 10 2 2 (one ovigerous). 6 
Oo" spring Cb1, from edges of swamp, 2604112" 
S 135°29726" EWE Ponder & D. Winn, 5, vi, L985. 
SAMA C5658, 9) 2 & (three Oviverous!, 61 O'S, 
20 juveniles, Sume cla as previous lol except 
colleeted W. Zeidler & K.L Gowleti-Holmes. 


Descriprion of holotype male (Figs 3-6) 

Length 3,8 nin, Head wbout as Jong as deep. length 
Ulitoast equivalent! lo first two pereoniles, Antenna | 
about 3x length of bead; pedunculur article | length 
15x width, uricles 2 and 3 subequal in length about 
0.75% length of article 1: flagellum slightly longer 
than Lax peduncle, of ning articles with one ventral 
aesthetise dt base of each of last four articles. 
Antena 2 about O7x Jeapth of Al wath 
characterise ghind cone ab base; peduncular article 
L slightly wider thin long, article 2 wilth about 0,7 


lenyih. 2 as long as article | and 0.78 length of 


ardele 4: Flagellum slightly longer than pedunele, of 
vight articles, 

Upper lip shahtly wider than Jonp. upieally 
rounded, bearing numerous short sete apieally. 
Lower lip With vestigial inner lobes; outer lobes 
sudowale Wil sctose distal abd (aver miuruias, 

Mancdibles without paps lel with inepsor ol sis 


teeth, acina mobilié of five teeth, spine row of three 
feathered spines und trituralive molar; rh with 
incisor of five teeth, lacina mobilis: of three teeth, 
spine row of two feathered spines and laiturauve 
molar with one long feathered seta. 

Maxilla | without palp, notched at palp's qorial 
posiion: outer plite with mnpe comb-hke spraes 
upically: inner plate very narrow with two feathered 
spines apicily, 

Maxilla 2; outer plate about !5y length of inner 
plate. setal row pesticted To apedt inner plate with 
one laree seta medially about 1.4 from apex. setul 
row apically and medially. almost to large seta. 

Maxilliped; inner plate Jarge, sub-rectanpulir, 
reaching end of merus, maximum width about 3« 
length of outer margin, wilh three apigal spine teeth, 
ihe inner one smaller. four plumose selae on inner 
marin and several apically: outer plate ovate, 
reaching midway alone inner margin of carpis, 
about. as wide dis inner plate. apical margin with three 
selue. Inher margin with several setae for distal hall, 
palp large, 4-articulale; merus proximally nurrovy, 
sub-triangular, outer margin about 2x length of inner 
margin with two selec on inner distal angle; curpais 
slightly broader than long. slightly expanded distally, 
distal Iwo-thirds of inner margin with row of setae, 
hwo sefae on outer distal angle and also pear inner 
distal angle: propodus slightly narrower and shorter 
thin carpus, distal margin with several strong series 
curved daetylus with strony ungurs. 

Coxul gills sausage shaped. present from G2 to Pb 

Gnathopod |; coxa shghdy longer than inaximun 
width, proximal width about (7x distal width, 
uNnicrion Margin concave, postenor Margin straight, 
distal murgin eyenly rounded with several evenly 
spaced sete: canes lridngular with large 
posterodistal tobe. with anterior nargin almost 2. 
length of postemor marine maximum width about 
1.5% that of anterior twreit, posterior iigia with 
¢lose-sel row of pine stout. pectinale spines: 
propodus subsrectungulap about 14a length ot 
curpus, slightly wider distally, widih O,6x lengit, 
posterodistal corner with two stout spines on elther 
side of ductylus. cluster of Jong setae on anteradistal 
corner, row of seven long sete medially. mixture Of 
tong and short setac near distal margins dactylus 
Shehtly shorter than width af propodus Htlnie neally 
wwarost palm. Gnathopod 2 length 1.6% Unit of Gi]. 
cosul gill length 2s wath, like shorler than cosa: 
coxa rectangular slightly longer than wide, abow 
O.88 length of basis. disud margin evenly rounded 
with several evealy-spaced Setic. merus with right 
angled bend: eurpos similar to Gl but withuut 
pecunate Apines; propodus slightly shorter than 
basis, length anterior margin LOX saaximum width, 
posteroproxrmad corner forming rounded lobe, palin 
oblique wilh numerous spines of varying lengihs on 


A NEW SPECIES OF AUSTROCHILTONIA 33 


Fig. 3, Austrochiltonia dalhousiensis sp. noy., holotype Go. Scale bars = 1,0mim (whole animal), 0.2 mm (A,U.T). 


a4 W. ZEIDLER 


Wz 


Wey 


a 
SS 


Fiz. 4. Austrochiltonia dalhousiensis sp. nov., holotype co. Scale bar = 0.1 mm. 


A NEW SPECIES OF AUSTROCHILTONIA 35 


Fig. 5. Austrochiltonia dalhousiensis sp. noy., holotype o. Scale bar = 0.2 mm. 


36 W, ZEIDLER 


Fig. 6. Austrochiltonia dalhousiensis sp. nov., holotype 6". Scale bar = 0.2 mm. 


A NEW SPECIES OF AUSTROCTULTONIA 


either side of cutting edge followed by shallow 
wroove lor tip ol dactylus: dactylas claw-ltke, as long 
as anlerior margin of propodus. 

Pereopod 3 with part of propadus and dactylus 
inessing On right: coxal gill length almost 2x wrath, 
about 0.75x length of coxa: coxa like that of G2 but 
slightly larwer. shehtly shorter than basis; merus 5x 
uy long as basis, anterodistal comer produced, carpuy 
78x length of merus. Pereopod 4 similar to P34: 
slightly longer than G2: cox with distinc 
posteroproximal excavation, maximuny width 
slightly more than length, shehuly longer thai bists, 
propodus slightly longer than merus; dactylus stout. 
length slightly less than 0.5x of propodus. Pereopod 
S slightly Jonger thu P4, coxal gill length ubout 2x 
witb, shghtly longer than basis; coxa width about 
5x that of busts, anterior lobe slighty more than 
05x length of basis, posterior lobe uboul 0.8% lengur 
of basis: basis slightly longer than wide with typical 
expanded posterior margin and posterodisial lobe 
veaching to about midway of ischjum: merus with 
posterodistal comer produced. length about (78 (hal 
of basis; carpus slightly shorter than mierus: 
propouus length bax that of carpus, ductylus stout. 
0,5, length of propodus, Pereopod 6 length | 3x that 
of PS: vost! gill length about 2x width. about (75x 
length of basis; coxa almost as wide as basis, anterior 
lobe (3% Jength of basis. posterior lobe (5% length 
Of basiss renmining articles bike those of PS bat basis 
with siedight posteroproximal shoulder and carpus 
atihthy longer thay mers. Pereopod 7 longest, 
slightly exceeding PO, like PO but coxa semi-cinvalar 
ane lachkumge coxal gull widui ty length. about Od 
leneth oF basis: posterodistal lobe of basis more 
capanded reaching lo about midway of mesus. 


Pleopuds all onmoditied (vot as in Clete. 


Urepod [ whout |.5x length of (2; peduncle wiih 
spine on inner and outer distal comer, three large and 
one smaller spine On dorsal onter manga, one small 
spe On Inger margin: outer mumus SlighUy shorter 
than inner, length 0.7% that of peduncle, with twe 
yocutan and three terminal spimes: omer ranms with 
two sivalland three larver terminal spines and three 
medially. Ureped 2: peduncle with spine on inner 
andl ooter distal corner ind additional one on dorsal 
marpin; toner ramus |x length of outer ramus.and 
[3x that of peduncle, live liege spines clustered 
terminally and three spaced evenly medially: inner 
ramus wiih 20 spines of varying sizes gradually 
closer together lowiuds Up. Uropud 4 (vo-urienlate. 
imanially niece than O.Sk length ob rebyen: ranits 
0.58 length of peduncle with three long sefae and one 
short setu revminally. 

Felson votire, subrectanpulin, shyltly wider than 
Jon. distal imurgin slighdly coneave with two snail 
Sete ut each vorner 


~ 
a) 


Deseription of allotwpe female (Figs 7-9) 

Length 3.8 mm. ovigerous with 23 eggs in brood: 
pouch, same as male except for the fullawing, 

Antenna 2, Nagellum of seven articles. 

Gnathopod |; coxu relatively narrower and longer 
than for mule, width distally 0.8.x length: posterion 
margin of carpus with vlose-set row of LO peetinate 
spines: propodus relatively oarrawer than for male, 
slightly longer than carpus, Gnathopod 2 length 1.2% 
that of Gl: coxal onl eelatively smadler, less than 
0.5x leneth of cox voxa with posterior margin 
produced to pout miccdilly, ratxamium width 0.5% 
length, as long as basis: remaining articles like those 
of Gt only relatively more slender. Pereopod 4 
length about b.3% that ol G2; coxa similar in shape lu 
that of. G2, Pereopod 4 slightly shorter than P3; coxa 
without distinct proximal excavation, almost as ware 
as Jong. Pereopad 5 only marginally longer than P4; 
coxa width about 1.7% that of busts. merus, cirpus 
and propodus relatively shorter than for mite, 
Pereopod 6: basis with posterior miuruin rounded 
proximally: merus, carpus und propodus relatively 
shoner than for male, Pereopod 7 slightly shonet 
than Po: basis relatively narrower, ind merus, carpus 
an! propodiis suecessively slightly shorter than for 
male, 

Oostegites on Coxe 5, all with curled margins 
and numerous small hooks, together lorming ueht 
marsipium. First heartshaped, length 1.6. 
marion width. about O.7x lent of C2) second 
wapezoid, length almosr 0.5 that of P3, maximum 
width almost O.5x length; think oval-shaped of 
similar size to second: fourth sub-reeiangular with 
oblique distal margin. length anteriorly almost (55 
(hat of PS. maximum width almost equal to length ol 
posteriur murern, 

Uropad | length 1.64 tht of U2. pedimele with 
five large and one stall Spine on Guler margin, miner 
margin With tWwer small spines proximally in addition 
{o lurge spine on distal corner; outer ramus as long as 
inner, length O.8 that of peduncle. with two lage 
and two smiller spines terminally and (wo medially; 
inner runius with three lirge dnd Iwo smaller spines 
terminally aed two medially. Uropod 2 peduncle 
with two large spines on oliter murgin, onter ramts 
slightly shorter than mner length tax that ol 
peduncle, one large and two smaller spines 
renuinally. three large spines medially; timer conus 
with wo terminal spines. cluster of fear near tip and 
another two tedially 

Telson with group of three small setae at wach 
corer, 


Evils 
token from the type locality in reeogmition of the 
restocted distribution of the species. 


3K W. ZEIDLER 


Fig. 7. Austrochiltonia dalhousiensis sp. noy., allotype Q. Scale bars = 0.2 mm. 


SOLS Ws ut 
me-peee | 


\W 


A NEW SPECIES OF AUSTROCHILTONIA 39 


Fig, 8. Austrochiltonia dalhousiensis sp, aoy., allotype 2. Oostegites on P3-5 not illustrated. Scale bar = 0.2 mm. 


40 W. ZEIDLER 


Fig. 9. Austrochiltonia dalhousiensis sp. noy. Oostegites from allotype 2. G1-P4 from paratype @ , 4.8 mm, from 
SAMA C5653. Scale bar = 0.2 mm. 


A NEW SPECIES OF AUSTROCHILIONIA +| 


Variation 


Apart from minor variations due to size. paratype 
and other material @xaimined is very Similar ro either 
the holotype or allotype, The maximum recorded 
size of males is 5.2 mm and that of females 6.5 min 
bul Host specimens eXxamiied are around 4-0 Min 
long. Minor differences between specimens 
generally were noted us follows. The number of 
flagellar articles of Al varies fromm eight fo (en wath 
one small specimen having seven: A2 has from six to 
nine Flugellar articles but most specimens have only 
seven or eight. The number of gesthetses on AL ts 
remarkubly conmstunt will) only some larger 
specimens having an extra one Clive), ‘The nuanber of 
peetinate sete on the carpus OF GL varies from seven 
tonne Hiimales and ciaht to ten in females (similurly 
lor G2 of teniales). In the allarype the coxuc of G2- 
P+ ure of unusual shape, dilfering fro fyles and 
qoreovieerous females (Pig. 9) in thal the poster 
margin is produced ty a point medially und Pa is 
without a proximal excavation. In the hololwpe the 
basis of Po bus a relatively straight posteroproximal 
shoulder bur ii newly all other specimens examined 
the posterior margio is evenly rounded, Pereopod 7 ts 
usually longer than P6 but in the allotype i is slightly 
shorter, probably because of the relatively shorter 
propodus Which js normally longer (han the carprs 
The spinavion of 1 & 2 varies slightly with larger 
specimens Tavine one or two ext spies ob the 
peduncle and rami. Uropod 3 is usually two 
articulite and only one Specimen (a female trom 
05654) had US with one article and then only on the 
jivht-hand side, Oostegites of females vary 
considerably in size but are expanded, as illustrated 
forthe allotype, in ovigerous specimens. 

The possibility (hat speciation may have occurred! 
between springs withoul any obvious morphulowical 
changes was considered ahd specimens for allozyme 
electrophoretic unalysix were collected fron all three 
loculives. A preliminary anilysis of this matertul 
using methods outlined hy Richardson ef ef. (1986) 
indivated fixed venelic differences of 10% or Tess 
(for Tt foci). thus supporting the morphological 
evidence of one species with hte yanation. Giyen 


these oresully, i more dehulled analysis was 
corstdered) (unecessary. 
Diseussiont 


The new species desenibed here closely resembles 
A, austratiy to What U3 is dwoeurticulite. However, a 
cuiiber of minor features collectively reudily 
distinzursh i from this species audats only other 
convenen AL syblenis, The nin distilpiishing 
features ure as follow. Remmles reach a larger sie 
(han les and the species is wenctally vot as tare as 
A. oiuntadiy (ales up te LO. gain, females up to 8,3 


mm) or A. subiennis (ides up to 1,0 min, lemales 
up to.6.4 mm). Antenna | hax Jewer vesthetuses (5-7 
in A, aitstralis). Both antennae have fewer flagellar 
articles (AT up to 17. A2 up te LL tn AL eistralis). 
The coxae of A. defhousiensiy sp. nov. are relatively 
wider and the excavanion on coxa 4 is nol us decp as 
in A, atstealis. or AQ subtennis. In both A ainvtrtlis 
and A. subteques, the lareral maven of the excavation 
of coxa 4 js at oghtangles to the posterior margin 
whereas in A. daliousiensis the aagte of the coxal 
excavation is much greater than 90°, In oviperous 
females of A. dalhousivrnss the coxac of G2-P4 have 
the posterior margin produced to a point medially 
und coxa 4 is without a chuwueterisue exeuvarion 
There are fewer peelinute spines on the exepus of Gt 
(iiales) and G! & 2 Vemales) than in AL australis 
(usually >>l0). For Gl (males) and G) & 2 
(females) the carpus ts. Slightly shorter than the 
propodus whereas the reverse ys. truc for A, duspilis 
Other miso: differences between the new and other 
species no doubt exit but were not evident an the 
present study, 

Austrochilionia dealionstatsix is aly similar to 
Phrearschilroata unuplihalma Aeidler, V9, a 
phreatie species whieh also has a limited distriburion 
at Dajhousie Springs (Zeer L991). cypeciatly in 
that ovigerous femules of A. deliuisiensiy have coxa 
4 without an exeavation, a feature chardeterishe of 2 
qnophthalaa, Given the isolated habit of 
Dalhousie Springs, one would suspeet thut these two 
species would have Common aneesiy., Howeve), 
since electrophoretic analysis hay shown thal they 
diflerat about 80% of fhe 2} loct examined, (his does 
nol uppeur tu be the case, 

The closest relatives geoeraphically, apart fron? 
dmophititima, ace species ol Agatrchiltenia Fouad 10 
the moand Springs newr Lake Eyre Sooth. Although 
A. dalhousiensis is morphologieally very similar to 
these apecies. eleetrophoretic analysis has shown thut 
it differs from thent at 73-80% of the 2b low 
exuynnmed. Clearly a more detailed morphological 
and genetic study of the genus is requited to 
defermine relationships. 

A single, damyzed female of Ay cdelhoninivnyts wis 
Jon in the pool of spring Cat (AM Pass4id hts 
record apmay be due to contaminated collecting 
equipment as the Watley Ceniperature at that locality: ts 
37°C and freshwater amphipals preter cooler waters 
(Barnard & Barnard l9RA) Th therelore seeiis 
unlikely (hat A. de/Amsiensiy Occurs matupally por tn 
pool of Cal but is possible occurrence al this 
logulity warrants further ivvestigalivn. 

The factors determining the distrthatwou ol this 
species ure unknown, Us restricted distbution at 
Dalhousic Sprmgs ts puzelmy us riany appaccntly 
suitable habitats eaist im which this species was tet 
ound, Although restacted in its aistrbutran, the 
species is pchilively abundintat all of the collector 
sles, 


42 W. ZEIDLER 


Like P. anophthalma, the presence of this species at 
Dalhousie Springs on the edge of the Simpson Desert 
suggests that it ts a remnant of a once more widespread 
fauna during a time when central Australia was much 
wetter than it is today (Krieg 1989), 


Acknowledgments 
lam inost grateful to Dr W. F. Ponder (AM) for his 


assistance in organising the 1985 expedition to 
Dulhousie Springs. He is also thanked for his 


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

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


References 


Barware. J. L, & BakNARD, C. M, (1983) “Freshwater 
Amphipoda of the World. 1. Evolutionary Patterns” 
(layfield Associates. Mt Vernon. Virginia). 

Kaeo, GW. (1989) Geology pp. 19-26 Jn Zeidler, W. d& 
Ponder, W. F. (Eds) “Natural History of Dalhousie 
Springs” (SA Museum, Adelaide). 

RICHARDSON, 3. J.. BAVERSTOCK, P_R. & ADAMS, M, (1986) 
“Alloyyme Electrophoresis” (Acuderme Press. Sydney ). 
Sayer, O. A. (1901) Description of some new Victorian 
fresh-water Amphipoda. Proc, R. Sow. Viet. 13, 225-242. 

(1902) Description of some new Victorian lesh- 
waler Ainphipoda, No. 2. Lhid, 18, 47-58 

Sairy, PC. (1989) Hydrogeology pp. 27-39 In Zeidler. W. 
& Ponder, W. Fo (Eds) “Natural History of Dalliousic 
Springs” (SA Museum, Adelaide). 

Sati. M. J, & WILLIAMS, W, D. (1983) Reproduction 
cycles ino some freshwater amphipods ino southern 
Australia. Mem. Aust. Mus. V8. 183-194. 


WILLIAMS, W, D, (1962) The Australian) freshwater 
amphipods 1, The genus Anstrechilronia (Crustacea, 
Amphipoda. Hyalellidae). Aus. /. Mar Freshw. Res. 13, 
189-216. 

ZVIDLER, W. (1988) A redescriphon of Afracsiltonia capen- 
wis CK, H. Barnard, 1916) with a review of the genera ol 
the family Cemidae (Crustacea, Amphipoda). Ani. 8. 
Afi. Mus, 98, 105-119, 

(1989) Crustacea pp. 79-87 Ja Zetdler, W. & Ponder, 
W. F. (Eds) “Natural History of Dalhousie Springs” (SA 
Museum, Adelaide). 

(1991) A new genus and species of phreatic amphi- 
pod (Crustacea, Amphipoda) belonging in the ‘Chiltonia 
generic group, from Dalhousie Springs, South Australia 
Trans R. See. §. Aust, 115, 177-187. 

__& Powprr, WF (1989) Preface pp. ix-xi /7) AZeuller, 
W, & Ponder, W. F. (Eds) “Natural History of Dalhousie 
Springs” (SA Muscum, Adelaide), 


TRANSACTIONS OF THE 


| ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 121, PART 2 


COMPOSITION OF THE STYLETS OF THE TARDIGRADE, 
MACROBIOTUS CF.PSEUDOHUFELANDI 


By ALAN F. BIRD* & STUART G. MCCLURE 


Summary 


Bird, A. F. & 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, 
1997. 

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. 


Transecttoas of tre Reval Seotery af 8 Anat (997), 122), 43-50, 


COMPOSITION OF THE STYLETS OF THE TARDIGRADE, 
MACROBIOTUS CF. PSEUDOHUFELANDI 


by ALAN F. Bikp* & STuarT G, MeCiure 


Summary 


Biko, A & Meci ure, SG. (1997) Composition of the stylets of the bordigmade, Muncohiotis cl 
preudohufelandi. Thins, KR. See. 8. Aust, 121(2), 43-50. 30 May, 1997, 

The chenucal composition of the two stylets of tardigrades has received little attention in the past. probubly 
because they are so Jabile. We have studied these structures in the tardigrade. Maereliuiis el psendohifelundi, 
und have shown, using histochemiesl techniques and energy dispersive X-ray analysis. that these structires are 


composed of calcium carbonate 


The response of M. ef. preadofufeland? to the environmentul stimuli of anoxia, differences in pLh und 
leniperature change demoustate that dis tardigrade can teabsorb its stylets under stress ane reform them when 
the Stress is removed, ‘he possible evolutionary links between tardigrades sind molluses that Have clierts 


composed Of calcium carbonate are discussed. 


Key Woks: Tardigrade, stylets, Macrahiotis cf pyendohufeland!, SEM, energy dispersive X-ray andlysis, 


histochemisiry, salivary whinds. 


Introduction 


Members of the Phylum Tardigrads (water bears) 
all feed using a pair of stylets Whose composition is 
such that they dissolve and break down on the death 
of the tardigrade and in some commonly-used lixa 
lives. Thus. they are frequently nor included in the 
camer lucida daiwings or photographs of the hucco- 
pharyngeal apparatus commonly used for taxonomic 
idemification. The general composition, shape and 
purpose af the twin stylets seems io be similir 
throughout the phylum and so the chemical compo- 
situ al stylets af one species probably holds for all 
Becuuse of theie transient nature, dissolving on deuth 
or fixation, turdigrade stylets have aot been anu 
lyzed, Th has been sugvested thi they ure calcareous 
in nature (Kaestner L968: Kristensen 1976; Wenck 
}O14. Kristensen unpub. both cited by Nielsen 1906) 
and that they resemble nematode stylets in forn) and 
fiunehion (Rigen 1962 cited by Kinchin 1994), The 
stylets of Macrebrolus cl psendohufelandi are 
curved, sabre-shaped structures about 400 pm in 
length that exhibit marked birelringence under poliar- 
ized light (Bird 1996), 

In this paper we examine the dissolution of the 
stylets in Various media under the polarizing micre= 
scope. Weir shut properties Under braght field 
microscopy and their elemental composiion using 
energy dispersive X-ray analysis in the scanning 
electron microscope, We alse examine the tetorma- 
bon of the stylets aller dissoluuon und disetiss these 
structures in the hehe of tardigrude evolution, 


2 7havbord Row) Micha s. Aust Ste2 
SSTRO bat and Willer, MH 2 Clon Osea 8 yust Sti 


Materials and Methods 


Locality, sot wee and extraction 

The tardigrudes were recovered from soil classified 
asa solonized brown earth from an experimental plot 
ona firm at Avon, South Australia (Bird 1996), After 
thorough mixing, 50 2 aliquots of the soil were 
pliced in a misting apparatus and processed as pre- 
viously deseribed (Bird 1996), The turdipricdes so 
collected were pieked out using a dental No. 3 nerve 
hroach. Walter which hid passed through the soil wis 
vollected from the misting upparatus and filtered 
through w 0.2 pin membime filter, The tardigrades 
were placed in a shallow layer oF this water ina ster 
ile Petri dish. Under these conditions, tardigrades 
remuined viable fora week or more without Feeding, 


Shlel dissetuiion 

The break down of the stylels was observed, on 
slides with vovershps sealed with nat varnish. i 
solutions with pH ranging: from 4-8 for different 
lengths of time (from the sturt of the experiment (a 
several days) and at teniperatures ranging From 15 - 
30° C Observations were made with polarized light 
and differential interference contrast (Nomarski) 
aptics using a Vanex Olympus ABT research 
Microscope 


Sryler reformation 

Specimens were mounted in distilled water in 
scaled slides with pieees OF No. 1 hiss coverslips 
acling as spacers fo prevent crushing. These speci- 
mens were kept under observation. usiiig Nomarski 
optics. until the stylets could no longer be observed 
(usually about 3h), The vovership was then gently 


Al Ab BIRD & $.G, MeCLURE 


removed under a dissecting oicroseape and the 
motionless tardigrades phiced, together with some 
small nematodes to stimulate feeding activity, in shal- 
low distilled water ina Petri dish. After about | 2h the 
lardiaades, which bad resumed activity. were placed 
in sealed slides and re-examined under Nomarski 
optics, 


Prerzy dispersive X-ray analysis of the seylets under 
the scanning electron microscope (SEM) 

Because of the difficulties of obtaining intact 
stylets (rom tardigrades on sealed slides or after the 
use of acid fixatives such as PA 42), as used for the 
fixation of pemutades (Hooper 1986), a special lech- 
nique was udupted for these studies as follows. The 
fardigrades were transferred to a small drop of water 
which was then swamped with at excess of 6% 
gluteraldehyde in Sorensen’s buffer at pH 7,34, The 
lardigrades were then gently crushed ander a coyer- 
slip to permit rapid entry of the fixative without eaus- 
ing undue damage to them, After 2 h in the fixative 
the turdigrades were removed, washed four limes in 
0.2 un membrane-filtered distilled water und placed 
ina sihall drop of the membrane-filtered distilled 
waler on the surface of a polished carbon stub. The 
lurdignides were manoeuvered close together and a 
second polished carbon stub was lowered on to the 
first so that the tardigrades were caught between the 
polished curbon surfaces, They were left in this posi- 
tion for 24 hto dry and then the uppermost stub was 
lifted direetly upwards, thus pulling the rardigrades 
apart and exposing their inner organs, including the 
buceo-pharyngeal region. This technique was also 
used on fresh. unfixed muterial. The stubs were kept 
iv a sealed dry container until examined and pho- 
togruphed in uw Cambridge § 250 Mk 3 SEM opera- 
ted at 20. KY using Ilford 120 roll film (FPS Plus), 

Energy dispersive X-ray (EDX) analysis was by 
means of a Link EDX system attached to the SEM. 
BDX analysis of characteristic lines with energies 
less [hun approximately O48 keV is pot possible with 
this delecior because of the absorption of the low 
energy X-rays by the deteetor’s beryllium window. 
Consequently, the direct identification of elements 
with alomic numbers less than that of sedium is not 
possible, 


Histochemistry: af the sedivary glanes daring sivler 
dissolution 

The unthraquincie dye alizarin red S’, sed as a 
stain for calcium. was made upas a 1% solution (50 
me 5 ink!) either ima buffer solution at pH 4.0 or in 
distilled water (pil 5.0), The tardigrades were placed 
in a drop of the alizarin red S solution and gently 


Aldevh Chemical Compiny 


Fig, |. Head of living specimen of Maerebioniy cl preudo 
dufeland? immediately after beg placed in distilled 
water on a Sealed slide. Note the lwo eye spots in lower 
part of photograph and the two curved: stylets. Girrow 
heads), Norurski optics, Seale bur= 10 pm 

Fig. 2. Sume speeimen as in Piz. | but photographed just 
ailer the sivlets hud dissolved 166 rora later, Arrow heads 
tidicale the original location of the stylets. Nonrarsks 
opties. Seale bar = 10 pm. 

Fig. 3, The same as Fig. 2 but viewed under bright field 
optics. Arrow heads indicate the original Jocation of the 
stylets. This photograph illustrates the buccvo-pharyngeal 
structures used in the taxonomy of this genus. Seale hur 
= (0 um. 


TARDIGRADE STYLETS as 


squished. without causing gross damage to then, by 
placing a ghiss coverslip over the cop and with- 
drawing the stain sulation from under the coverslip 
with picees of Mer paper, The edges of the coverslip 
were then sealed with nail vartiish and the speeimens 
observed and photogriphed wider beight feldl optics. 


Results 


Disselatian af ite stylets 

The time tuken for diysolution ot the stylets uy cli 
tilled water. when westute of anoxia fas been induced 
by sealing the tardigrades under a coverslip on a 
side, is teriperatare dependent, Thus, a 30° 0) total 
stylet dissolution occurred within three hours whee 
us att 15° C the stylets were only partially dissolved 
afler this ume. 

The appearance oF the heud region ab Ww eb. 
prendohufeldid?, viewed under Noiiarski opues. as 
shown prion ty commencement of stylet dissolution 
we25° C (Vig. (und aller its completion three heurs 
later (lag. 2). The head region ts alse shown utter 
three hours. phatogriphed under nerinal bright feld 
Hplies (Fig hy, 

The sequence of sliges in the dissolution of the 
sty lens is ilustrated pos! cloudy ina seres ef pho 
lamicrieciphs taken under the polarvang miera- 
scope (Fig. 4\-H). Styler dissolution ty gridit aver 
the first (Wo hours (Pig. 4+ As) but aeeculerites over 
the third hour (Pig. 4 BH) so that warked ohunees 
were cletecled at approximately |S-iirmute intervals 
fron two hours onwards, 

Stylet dissolution Gecuered: more rapidly we ue 
solutions. than oy neutral ov alkaline solutions. took 
place within 50-90 tin ig O,0SM ACL and occurred 
immediately in. IM ACL and se is Similitr to Caco, 
inthis vespeet, 

The hirefringence exhibited by the touseles of the 
irivadiate pluiryox of Mech, pyewedolufelimet dil nor 
disappear over a period of three hours (Pig. +) ane 
the brighiness of these muscles, although mot as 


Warked as thut ol the siylets at the conmencemeat af 


the experiment remained cemstant througheut, 


Reformation of the spylets 

Turdivrales appear to huye the capacity to reform 
their stylets under favourable conditions in) what 
appears to be a reversal of the dissolution whieh 
ocvurs When they are placed in unfavourable situa 
tions, Speeimens that had been imduced to dissolve 
their stylets ri scaled slides and had hecome motran- 
less Were Observed to regan (heir Muscular activity 
and reproduce stylets. over w 12 h period, when 
placed in shallow distilled water inn uasealed Petri 
dish. in the company of several smull nematodes, 
This retormation did not always lead to precise 
realiznmtient of the stylets and the effieieney of this 


reformation and correct realignment may be depen 
dent on the state of health of the tardigrade at the 
commencement of the experiment. 


Enerey dispersive X-ray anitlysis ef the stylety 

Esxumination vader the SEM of the polished carhon 
surfaces bolding the dried. squished and disrupred 
lardivvades, cleatly shows whole stylets. or parts 
thereol (Fig, 5), By means of this technique, itas pos- 
sible to obluin suitable PDX analyses of the element 
tal composition OF the stents. Lecan beseen from the 
DX spectrin (Pig 6) thatthe sivlets ace rich in cal 
cium. wilh maximum X-ray robedsily Counts al erer 
sies matching the Cu pat 5.690 keV and the Ca Pal 
4.012 KEV chinueroristic Nery lines. 


Hiyiochennyty of the salivary glenes divine siyfer 
dissolution 

When the jardigrades were venily squashed in a 
Ge salutian ob alc red Sut pH 3-0. rhe salivary 
glands surrounding the stvlers @raduully became: u 
deep red (Pigs 78) This reaction was bottle tnarked 
and Joculized, provided that the body of the turdi- 
wrade had been erushed co allow rapid penetration oF 
the stain. These vhinds. which are thought to be 
responsible for (he secretion uwod dissolution of the 
stylels (Rinchin 1994), ane apparently rich in-cul 
cium as indigated by their peachion to alizacin real S. 


Discussion 


Liformation on the nature of the tardigrade Siylets 
iS Sparse und hits received Tittle or no-satlention iA the 
two most recent general acequnls of (hese organisms, 
notably a most informative book on their general 
biuluey by Kinchin (1904) and a collection al papers 
published in the Zoological Journal of the Lrnnean 
Soviely i 1996 and edited by Mefines & Normiin. 

Tardigrade sivle(s were (hough! lotuye seine Sune 
ilarity with neniatode stylets by Riggin (1962 ) (cited 
hy Kinchin 1994). However. ibis clear that tardignade 
stylets (Bird (996) differ markedly from nemiutode 
stylets (Bird & Bird 1997) both in composition und 
structure, Thus, FA 4) }. a mixture of formaldehyde 
and avelic acid and & common lxalive for nema 
tudes. causes rupid dissolution of the tuntigrade 
atylets beauuse ol its low phl and the bigh tempera- 
ture used i the procedure, This fixative, however, 
does nol cause dissoluhon of nematode sbylets, 
Hoyers mediam. a moisture of guar arabic, chloral 
hydrate yond ghyeerol (Kinehin 1994) does nol dis- 
solve tardignide stylets and his the added advantage 
of functioning as a combined fixative and embedding 
medium. ininany illustrations of the bucea-pharyn- 
geal region of tardigrades, the stylets are not shown, 
presumably because they dissolved when fixed, 

The small size Ol tardigrade stylets (40 pin 1 


AUR 


TARDIGRADE STYLETS 47 


Vig. 5. Anterior region of dried and fractured Macrobiotuy cf. pseudohufelandi (see Materials and Methods) viewed under 
the SEM showing parts of the two stylets (arrow heads). The site of EDX analysis is circled. Scale bar = 10 pm. 


12000 
10000 


Xray B000 


Intensity 


6000 
(Counts) 


4000 


2000 


.00 2.00 4.00 6.00 | 8.00 
Energy (keV) 


Fig. 6. EDX spectrum of the stylet shown in Fig. 5, 


Fig. 4. Series of photomicrographs viewed under the polarizing microscope illustrating the stylets of a living specimen of 
Macrobiotus cf. pseudohufelandi exhibiting diminishing birefringence as they gradually dissolve in distilled water on a 
sealed slide as Shown in Fig. | - Figs 2 & 3. A. At commencement. B. After 56 min. C. After 96 min. D. After 113 min. 
LE. After 120 min. F. After 136 min. G. After 150 min. H. After 165 min. Arrows indicate parts of stylets. Note that the 
muscles in the pharynx do not lose their birefringence as the stylets dissolve. Scale bar = 10 am. 


48 4.F BIRD & S.G, McCLURE 


length in Macrabiociy el preudolufeleard() and their 
lable nature, have made difieull determine theli 
composition. We fitye overcome this hy udupting it 
luchmique used to separate the layers tral uematode's 
cuticle (Bird & Deutsch 1957) whieh takes ante 
accuomil the adhesive qualities oF animal (issues fo the 
surfaces oo which they have been dried, In this 
instance. the Iwo surfaces were polished eurbon rods 
mounted on SEM stubs its described in Materials ane 
Methods. In some instiinces, this technique led Wa the 
exposure Of either (he whole or lure enough pices 
ol stylet to be recognizable (Fig, 5), These were then 
eusily photupruphed und subjected to chergy disper 
sive Neray analysis measurements 70 the SEM. These 
measirements clearly show that the elemental com 
pasion OF the stylets consists miiily of culenin 
felenents with omic numbers less (han that of sedi 

unture not detectable). "The rapid dissolution of the 
stylet in dilute acidsyand their unarked hrrelrireenuce 
tinder polivived light. indicite ahat they dre con 
posed of caletui curborate, ‘Thus, te copposa or 
ol durdigrade stylets differs fromm that of nematode 
stylet whichoare proicin in nariire, 

Hach tirdignide stylet Hes in the linen of a sali 
vary dha (Kinehin 1994). Kristensen (1976) cetted 
by Nielsen (996) has stveested that separate lobes ot 
(he salivinry whands are responsible for the Formation 
of the styles und their supports in the tondigradte, 
Ratillipes. Whether or not this applies to Mt ef. 
pacjalolufelanill pemiins to be determined, We have 
nor observed the Termation ov dissolution of the 
Sivlet supports oy Wis tindirade, iicating thal their 
chemist) compasioon is diflenent fom that of the 
wivlew. We have co inflienimbni on ihe origin and 
Cheiieal Compaspiion or the stylet Supports jy i, et 
parndolifelands 

Ou experinvnts indice that the seylets of Ae ef 
paren iifelandi are secreted and reubstithed by the 
salavy ekinds, We have shower that (Pies 78), 
wheo the Gindigrades ure gently squashed an a dilupe 
SHlivon of the stain alrzurin red Seal pit wl S20 ja 
sedleds the cell surrounding eagh stylet stairs as (Ire 
siylers break down, indicating (hat ealeturir is lifer 
tedhund diffuses throughour these two cells. Alizarin 
nl S is meoguived usa histuehemical rouge far ihe 
detoelony of caleium (Conn 1977) changing in 
colour from yellow toyed to purple over the pH 
tanve 3. 7-5.2, 


The gradual break down of the stylets ia slightly 
acidic aid anoxie covironinent, as demonstrated i) 
this paper, when photogruphed under polarized light 
(Fig. 4), closely resembles their step by step forma- 
tion us depicted in drawings by Marets (1929) (vited 
wn redrawn by Kinchin (994) showing stuges in 
their formation, Although stylets of caleiuin earboye 
ate ure urtusual in the animal kingdom und appear to 
be both unique and common (oall kudigrades, some- 
what similar structures are found in other graups at 
animals, Por instance, many of the terrestrial pub 
mone gastropods shoot durts of caleiuin curhomule 
into cach other prior te copulation. This behaviour is 
thoughr to actas a stimuiiie tothe sexo er The 
dart of Heliv dsypersa is about & - LO niin in lenge 
(Tompa 1982), at least ten times the si7e ot un entire 
tardivrade and some 200 Gyies larger (ha the 40 pe 
stylet of Macrobietiy cl) prendohufelandi, The cir, 
whieh takes ahout five days to fori. iy seecered Hr a 
dart sae that ts part af the reproductive system. The 
composition of (hese darts, wher subjected to energy 
lispersive X-ray analysis, is identical to the tardies 
gerade stylets with both Cy Kee uml Ca KE peaks 
(Lhint 1979). Phe evoluttonury srumheanee of these 
findings is Ohseure. However, they de lend same sup 
port to fhe phylogenctio position of the Nirdigryda 
proposed, as st result oF studies gia the 185 ribopornal 
gene sequence, by Moon & Kint (1990) These 
uuthurs shue “the tardigrade chide appear as an 
dependent lineave from the seimutode chide’ ana 
the culewreous niuire of the tardigrade stylet as 
Opposed to the protemmecous malice of the heitiatode 
stylet. lends siipport to this Hypothesis. Moon d& Kirn 
(1996) Jurther suggest Unat the turdligride clude is st 
sister group Of the pretostome cucoelomiale wssent 
blage (hat emnented belore the imolluses. animelids 
arthropods und sipunculids evolved. This would sug- 
gest independent evoluwon of the Culeareous str 
tires ja tarcdigrides and imialins. 

Somme deasere of The wags i Ou kiawledgee of the 
origins and relaGorships of the Hurdistada is bidicut: 
ed by the fact That hirdigrides are not mentioned ty a 
book or the origins and relationships among lower 
invertebrates. edited by Morris et val (1985), 
However sume priiitive Cambrniy fossils share 
some characteristics with both Onyehophora anid 
Tomigrada, ineluding poorly articulutect limbs ene 
ine in chiws (lobopodia, (eriiial ious abel the 


bis, 7, Whole specimen of Meerebrores ef, pseudehufetcd: That bas been crushed lo permit the entry of alizarn pou Scat 
pltS.0, The photograph was liken under bright fell ophes about 40 nia ater We conmnencement of stamiag. Showiny 
vye spots (small arrows) andl claws (iene ares). The area around the tidignide comlains material that fas been exid 
ed from the ruptarce animal and the two salivary glands Mar lie just anterion to the wuscular planyins are heavily stained, 


Seale bur = LOO ja 


Hig ¥ Rillitged portion of hig 7 showing eve spots (small arrows), the stamned salivary glinds (Sey che Wuscular pharyny 
(py amd material that has heen exuded by rupiuring to permit Gntey oF (he stiin Cire arawsy, Seale bar= 10 pm, 


TARDIGRADE STYLETS 49 


@” 


50 A.F. BIRD & S, G. McCLURE 


lust 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 
Onychophora 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 “Panarthropoda”, and that the ony- 
chophorans diverged first. It is clear that the origins 
und lineage of the Tardigrada remain obscure and 


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


Acknowledgments 


We 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) CSIRO for 
accommodation and facilities and to the Australian 
Biological Resources Study for providing equipment 
that made this study possible. 


References 


Biro. A. F. (1996) Studies on the soil-inhabiting tardigrade, 
Maecrobiotus cf. pseudohufelandi, from South Australia, 
Traits, R. Soc. 8, Aust. 120, 147-154. 

& Birp, J. (1991) “The Structure of Nematodes” 
(Academic Press. San Diego). 

& Deusen, K. (1957) The structure of the cuticle 
of Ascaris lambricoides var. suis. Parasitology 47, 319- 
32S, 

Coxn, H. J. (1977) 
Wilkins. Baltimore). 

Hoover, D. J. (1986) Handling, fixing, staining and mount- 
ing nematodes pp. 59-80 Ja Southey, J.P. (Ed_) 
“Laboratory methods for work with plant and soil nema- 
todes” (HMSO, London). 

Honr, 8. (1979) The structure and composition of the love 
dart (gypsobelum) in Helix pomatia. Tissue & Cell 11, 
51-61. 

KAgSINER, A. (1968) “Invertebrate Zoology” Vol, TH (John 
Wiley & Sons Inc., New York). 


“Biological Stains” (Williams & 


Kinciin, I. M. (1994) “The Biology of Tardigrades™ 
(Portland Press, London). 

KRISTENSEN, R. M. (1976) On the fine structure of 
Batillipes noerrevangi Kristensen 1976.1. Tegument and 
moulting eyele. Zool. Anz. 197, 129-150. 

Mctwnnes, 8. J. & Noxman, D. B. Eds (1996) “Tardigrade 
Biology” Zool, J. Linn. Soe. 116, 1-243 (Academic 
Press, London). 

Moon, 8, Y. & Kim, W. (1996) Phylogenetic position of the 
Tardigrada based on the 18 S ribosomal RNA gene 
sequences. [hid. 61-69. 

Morris, S. C., Gtordk, J. D.. Gipson, R. & PLArT, TL M, 
Eds (1985) “The Origins and Relationships of Lower 
Invertebrates” (Clarendon Press, Oxford). 

NIELSEN, C. (1996) “Animal Evolution ; interrelationships 
of the living phyla” (Oxford University Press. Oxford). 
Tompa, A. (1982) X-ray radiographic examination of dirt 
formation in Helix aspersa. Neth, J, Zool. 32, 63-71. 


STUDIES OF THE EGGS OF MACROBIOTUS CF. 
PSEUDOHUFELANDI (TARDIGRADA) FROM WHEAT FIELDS 
IN SOUTH AUSTRALIA 


By ALAN F. BIRD*® & STUART G. MCCLURE 


Summary 


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 exuyia, 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. 


Treasaerous a the Rayal SocieiyafS, Mave, (L997), E212). S157 


STUDIES OF THE EGGS OF MACROBIOTUS CF, PSEUDOHUFELANDI 
(TARDIGRADA) FROM WHEAT FIELDS IN SOUTH AUSTRALIA 


by ALAN FE. Bikn*® & STUART G. MCCLURE! 


Summary 


Binbo A, Bod Mec) ure $.0 (1997) Studies of the eges af Mocrbiotaeel. prevdolipfetind: VVardignada) fren 
Wheat Helds in Sour Australie Prams, A. See SAMs 12102) 5 1-57.30 May. 1997. 

The lindinnie, Mecoblotas eb pyendofmfetends, and lsc were isolated Hoi soil Prom a Wheat telat 
Avon in South Australia during the winter ol 1996. The surfice of Ihe eges was examined ender the seanning 
vleeiron microscope and was slieawin lo be iehty ornamented and redculate with numerous “verted gobler 
Shaped” projections, The dentate margins af the hends of these projections eansist of clusters oF coraldike 
globules. Statistical aniilysis of the sive of the projections reveuled (wo significantly dillerent types of eges in 
The sainples, EXaimples oF ormamented eges were Lound in discdrded exuyin, a condition whieh is Wor inadecart 
WITH recently proposed hypothesis on tandienile ese evellition. Observations on feedinu beh ioue, population 


density und exe layin habits are presented, 


Kby Woks: Macrae el, prendahujeland) iiteroseapy, tandimndes. eves. cenplology. eyaliinin sail 
1 HY e. 


Introduetion 


Tivdigrades, also known as waiter bears ar mess 
piglets (Rinchin 1994), belong to a diserete phy lim 
ul vosmopoliiin distribution Tron diverse habitats 
ieluciig Waring. fresh water and semieterresuial 
cuvironiments. “The tardignides responsible for the 
was described jn this paper were identified as 
Mincrobiolus ef, preudohufeland: Iharos 1966 by S, 
Choxton (Aird 1996) and are semi-lerrestrivl, having 
heen isokited from sandy foun soil aia wheat fel ar 
Avon, South Australia. 

The tardigrade exe shell is ay tse taxonomic 
wile Co speeles identification, particuhurly in genera 
such as Minrbions where the shell is ornamented, 
Meacrohions et psendelufelund?, which are unty 
about SOO pm long hy [50 yim wide when fully 
sown, lay comparatively laree virculir eggs which 
hive brphly ommunented reliculited shell surlieces 
with numerous “inverted goblet-shaped” projections 
(Bird 1996), 

Ii tus been shown by Bertolini & Rebevehi (1993) 
What differences in exe shell morphology an 
Macrobiarus hufelindi, previously thought to be due 
fo Variability within this species, fall into seven iis- 
finel types that ure related to different animal hor 
photypes, Using ege shell morphology. along with 
other chiunteters. these workers have deseribed a num 
ber of new species from the M. fuife/andi group. bggs 
that belong to this eroup have pitted or reticulated 
shells with protruding processes shaped Jike inverted 
goblets, chulices. thread spools or “cooling lowers”, 


eu 


2 Playtord oie Mitcham S. Aust, 3002 
CSO taimd und Water, PMB 2 Glen Osmond S Aust 506d, 


Berlolanies af (1996) have suited that ornamented 
eps ire generally laid ree in soil or water ana 
smooth shelled egus ure laid in the moulted cuticle 
(exuviuiiy) These Workers inelide the tantly 
Muacrobiotidie, to whieh Mo ef. pweredeluefelumell 
belongs. in those families that hay tree, ornamented 
cees. Benalani ef al. (1996) Iawe put Torwand an 
hypothesis ta which they explait the evolution ot 
lurdigrade epes, 

In this pauper we examine the structure of fhe eve 
shell of At ef psendelvgetmed? und measure the 
processes provuding from the surlice of the ce 
shell. We also discuss its egg living habits in relation 
ly the Wypothesis of Bertolami et ad (L996) allel con) 
ment on feeding behaviour and population density. 


Materials und Methods 


The lardigeades were collected on ES July [996 
fron sandy loam Soil and from (he same Tneulity at 
Avon, South Australia clatitude 34° 14S. longiude 
138° 19° EB) us those collected previously. using the 
sumpling technique described by Bird (1996), The 
soi) samples were collected in mid-winter so that the 
sites were wet and the kirdigrades were feeding, 
reproducing und depositing czas. 

Vhe tardigrades und other metofauna, consisting 
predominantly of nematodes. were isolated from tits 
soil over a period of three divys using a misting appa- 
ritus (Yeates & Bird 1994). Tardigrades and nema- 
todes Were counted and tardigrade eggs were picked 
oul using a dental No. a nerve browch and examined. 
alive, in distilled water under a coverslip, with the 
light microscope begs to be examined under the 
seaming electron micrascope (SEM) were fixed in 


ta 
nm 


A. F. BIRD & 8. G. McCLURE 


EGGS OF MACROBIOTUS Ch. PSEUDOHUFELANDI 


6% gluteraldehyde in Sprensen's phosphate buffer at 
pH 7.2 and at 5° Cand kept ui this Gxative for-sey- 
eral days. They were then washed three Himes in dis- 
tilled water and sonicated in an Elna T420 sonivator 
al adrequency of 35 kHz for W) see or unt shown by 
Microscopic observation to be free at debris, 

The eygs were freeze-dried by placing them 
between membrane lillers which were frozen rapidly 
by placing them tna slurry of freon cooled by liquid 
nitrogen. These filters with the eves were quhickly 
(ransterred to a [reeze driee and freeze-dried at - 70" 
C, The dried eges were picked up with double-sided 
tape whieh was ullached lo un SEM stub and couled 
with 30 nin of wold. This material was examined und 
photographed m u Cumbridge § 250 Mk 3 SEM 
operated yt 20 KV using PP4 Plus Tord roll filn. 


Results 


Numbers and feeding 

Atthe tine of collection the nutio of tardigrades Lo 
nematodes in SO gy of soil was 842297. Feeding on 
Hhenmatodes was also observed during the course of 
(this investigation. and the nematodes held by the 
tardigrades usually did not move although once a 
nematode broke Iree and moved uway. In one 
instanee, a tardigrade wus observed to be arching its 
back in the manner of a scorpion, curing feeding 
uclVily 


Buy levine ane eyes 

Leys thal were about 90-100 pm in diimeter in (he 
living unfixed stare (Pigs 1 2) and about 60-70 pun io 
the fised and dehydrited sue (Pig. 3), were Tad 
cither within exuvia (east cuticles) (Figs | 2) or free 
(Pigs Ai Figures 2. 3, 4. 5 clearly show that (hese 
eves buve a morphology of lhe MW frafelaimlé group 
with reticulited shells and charucteristicully 
uplurncd-chiilice-sNuped — protanding — provesses. 


i 


i 


Although ihe putrern of the reticulavions remains the 
same with the apertures on the reticulate surface of 
the cee shell being about 0.25 yar in chameter (Fig, 
6), the shape of the protruding processes falls inio 
two distinct groups. Type tis shorter than Type IL is 
narrow at the base and has a wide distal head (Fig. 4). 
Conversely, Type His taller wider at the base and 
hus a narrower distal head than Type T (Pra. 5). For 
10 processes af cach type. the differences in the 
mens ure statistically significumt with 95% confi- 
dence (Table 1) 

When the processes were examined under the 
higher magnification of the SEM (Fig. ©), the dentate 
cog-shaped margins of the head were shown fo con- 
sist OF clusters of globules with a structure resem 
bling madreporarian corals in uppearinee and were 
approsmmately 0.5 pin in diameter. The mudreporarian 
globules and reticulated shell surfaces wre simalar in 
both the shell types desenbed above. 


Diseussion 


Bertolani ep ai. (1996) have proposed an hypothe- 
sis fa expliin the evolution of tardigrade egys. 
According to these workers, the eggs of tardigrades 
have evolyed us a result of two events, the Urst bemg 
the wequisition of Ornamentation and the second the 
use of the shed exuviun as the site for exe laying. 
wilh the subsequent loss of ornamentauion, Thus, the 
ornamented eggs of the Macrobiotidae are thought te 
be laid free, Ole observations that the ornamented 
eees Of MO cl, psendolufelandi can either be band m 
exuvia (Pigs |. 2) or free (Pig. 3) do not appear to he 
th aceord with this hypothesis. The exuyia conning 
ortamented eggs were transparent and devoid of 
body contents and chu not appear to be females that 
had died before completing cep keying. A possible 
explanation Vor the laying of the e#gs in ihe exuyiiun 
by the Woch, pyendohufelundi fromthe solar Aven, 


Tahu) Le Meaviiements af the nee ipes vl prmceses prendiding fron the ene shell sifaces oe) Macrobioms of 


peuctohiufelanada, 


Type No. Heigl (int) 

Mean + 5D Ritiye 
i in a4 5 AAS 
Il 1 (ib (4+ 57-74 
10.95] = 2.26 247 + 


Basal width (yim) 


Distal Width (lim) 


Menit +58D — Ringe Moan + ALD Range 
43 0.2 3.048 5.3 2 3A? 
5.3 2 405-4 2.1 ul Uy See 

WAAR 16,07 


SD = Stundund Deviahon, | = Students (test 


Fie | Boe of Macroblons ch pyendelifelonads im cast cunele (esuviun) Arrows Tdicare posmon ofcust lees and ehiws. 
Bright field optics, scale hur = FOO yin, 
Nig, 2) Same specimen wt tgher magnification showing shell projectors (Pp vovered by cast cugele (OC), Bright Held opdes 


Scale bur = 20 pan, 


54 A. F. BIRD & S$, G. McCLURE 


EGGS OF MACKROBIOTUS CE PSEUDONUFELANDI 


is that this may be an adaptation to the hol, dry: sum- 
ners experienced there. 

Our studies on the tardigrades and nematodes in 
soils froin Wheat fields at Avon in oidwinter when 
the soils were wel and (he populations of the mer: 
fauna could be expected to beat their peak. show that 
tardigrades make up) a SubstanGal component, 
although they are not as numerous us nematodes. The 
ratio-ot 84 tardigrades to 297 nematodes per 50 y soil 
found in these eXperiments varies at other sites where 
the tardigracde numbers per 50 ye soil may: be less ane 
nematode nutnbers greater (Bird 1996). However. i 
is Clewr hat the tardigrade presenve al Avon is wide- 
spread, 

The lardigrades isolated from soi at Avan feed on 
bematodes and can survive hot dry summers m an 
anhydrobiotic state (Bird 1996 ). Feeding on nemia- 
todes was ulso observed durmg the course of this 
mvestigation and the nematodes held) hy the tardi- 
erodes did not usually appear to be movitig. although 
they were coiled and therefore probably not dead. 
suygesoig tous that they might have heen paralyzed 
by some type of injeeted narcoue. Since there is pa 
Information on tardigrade diversity and geographical 
distribution in South Australia. frether studies are 
warranted. particularly on their feediug habits, is 
they may hive w role inthe biocontrol of the parasitic 
nematodes which have heen shown to oceur in the 
wheat felds at Avon (Yeates & Bird 1994), 

The line structure of the egy shell iy the genus 
Macrobioms is of greal iixonomice inportance, Prom 
our Observations on the ultrastructure of the exp 
shells of the Avon turdierades, wv would seem that 
there may be two populuions GF Ma pseudolufe- 


LA 


> 


lanai in the Avon soil or there may be two different 
Species, neither of which completely resembles those 
described so fur for (he Aufelaned: group (Bertolani & 
Rehecchi 1993: Biscroy 1996). The reticulated sur- 
fave of the shell und the structure of the glabules on 
the heads of the projections are Similar in the two 
torms of egus deseribed above but the diameter of 
the apertures on the reticulited surface of the ege 
shell is much less than that shown in the eggs of 
other members of the /iv/claned? eroup described by 
Bertoluni & Rebeechi (1993), Kinchin (1994) and 
Bixerov (1996). To our knowledge. the globules on 
the heads of the projections have not been described 
belore and might prove. together with the reticulated 
surfuee. fo be usetal taxonomic eriteria al the fade 
lend? group is further divided on the basis of egg 
ormumentahon. 

Clearly further studies ure required on the taxon 
omy of these lurdigrades und on their distribution in 
the semi-vrid aerivultural areas of South Australie 
dnd olfice Sika! regions of the Australian continent, 


Acknowledgments 


‘The senion-autior is graterul for cogrant front the 
Australian Biologigal Resourges Study which pro- 
vided tuvilities that made this research possible anc 
10 CSIRO Land and Water Adelaide: for aeeomntoda 
tion and equipment Dr WR. Miller, Department of 
Biology Southwestern College Winfield Kansas 
USA and Dr W.I,. Nicholas. Department of Botany 
und Zoology ANU Canberra ACT are thanked lor 
constractve eriticism of the manuseript 


References 


BeRror ANT, RB. & Reaecont £. (1992) A revision ol the 
Maeralyonrs Jrifelannli oroup (Vardigracta, 
Macrobiotidae), will same observations on the Gxogon 
we chunicters of ctitardigrides. fon, Sormtay 22. 127- 
192, 

& Chaston, SK. 1996) Phylogenetic siz 
nifivance of epee shell variiion in tardigrades. Aol. 
Lin Soe. V6, | AY aR- 

Hien A. F (1996) Studies on the soilanhabiting taurdigrade, 
Macrobioms of prendahafelmd: trour South Australia 
Trams. Ro Soe S. Ausé, 120, 147-054. 


Riserov. Vo 1(1996) ‘Lordietades ot the Thinyr peniisuli 
with descriptions. of lwo new species. Aival I La Sea 
Mh. 215-257, 


Kawcnin, TOM, (194) “Phe Biology oof Tareligrades” 
(Portland Press. London), 


Yeares. GO. Wo& Birep. ALT (1994) Some observations on 
the influence of agriculiueal practices on the nematode 
fiumac of some South Australian soils bandane Apple 
Nematol. 17, 133-145, 


Fie 4, Scanning eleetron mieragruph of whole cegs mt Maerobferiy ef, psendoinfelind’ showing the inverted: goblet 
shaped prajections of Types F(T 1) and 112). Seale bar = 20 pin. 

Fig 4 Scanning eleetron micrograph of part of the egg shell surfice of a Type | eg showing projections with) hartower 
buses-wnd hurger dentate cow-shaped heads thunthose.ol Type WH, Note similarly rehendaled surfaces of ery: shells, Scale 


hours 3 on 


EGGS OF MACROBIOTUS CF. PSEUDOHUFELANDI 57 


Fig. 5. Scanning electron micrograph of part of the egg shell surface of a Type Il 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 um. 

Fig. 6. Scanning electron micrograph of the head of one of the Type II projections at higher magnification showing the 
madreporarian 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 = | pm. 


TWO NEW SPECIES OF ASPHONDYLIA 
(DIPTERA: CECIDOMYITIDAE) FROM HALOSARCIA SPP. 
(CHENOPODIACEAE) IN SOUTH AUSTRALIA 


By PETER KOLESIK* 


Summary 


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 Royal Society of S. Aust. (1997), 121(2), 59-66. 


TWO NEW SPECIES OF ASPHONDYLIA (DIPTERA: CECIDOMYIIDAE) FROM 
HALOSARCIA SPP. (CHENOPODIACEAE) IN SOUTH AUSTRALIA 


by PETER KOLEsIk* 


Summary 


Koesik, 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. noy. 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 Worbs: Diptera, Cecidomytidae, Adelaide, Strzelecki Desert, South Australia, 


Introduction Halosarcia indica subsp. leiostachya is a small, 
decumbent to erect shrub widespread along the coast 
Halosarcia 1s a plant genus comprising 23 species — and around inland salt lakes of mainland Australia 
commonly called samphires. The genus is endemic — (Wilson 1984). It is a common plant in the Strzelecki 
to Australia except for AH. indica (Willd.) Wilson 
which also occurs in Malaysia and other countries 
bordering the Indian Ocean (Wilson 1986). Two 
species of Halosarcia 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. pergranulata (Black) 
Wilson subsp. pergraniulata (Fig. 1) and A. erici- 
Jormis sp. noy. forms spherical, spiky galls on branch 
segments of H. indica subsp. /eiostachya (Benth.) 
Wilson (Fig. 2). 


Halosarcia pergranulata subsp. pergranulata is a ‘amar’ 8 
shrub about 0.5 m high which grows in southern 
Australia (except Tasmania) associated with coast- Fig. 1. Gall of Asphondylia inflata sp, nov. on Halosarcia 
lines, estuaries, swamps and margins of inland lakes pereranulata (Black) Wilson subsp, pergranulata. Scale 
(Wilson 1984), The plant forms a substantial part of bar = 10 mm. 


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 
of the mangrove swamps. Areas that are regularly 
inundated by tides are typically dominated by 
Sclerostegia arbuscula and Sarcocornia quinqueflo- 
ra, while areas that are only occasionally flooded are 
dominated by Maireana  oppositifolia and 
Halosarcia spp. In May 1996, a large number of 
galls caused by A. inflata sp. noy. was found on H. 
pergranulata subsp, pergranulata at Port Adelaide, 
about 400 m south of the Torrens Island bridge. 


Fig. 2. Gall of Asphondylia ericiformis sp, nov. on 


© Department of Horticulture, Viticulture and Oenology Faculty of : ‘ 
Agricultural and Natural Resource Sciences, University of Halosarcia indica subsp. leiostachya Wilson, Scale bar 


Adelaide PMB1 Glen Osmond S. Aust. 5064. = 10mm. 


P. KOLESIK 


60 


NEW GALL MIDGES FROM HALOSARCTA al 


Desert where it grows tna variety of habitats, includ- 
ing salt lake margins, open elay plains and gibber 
plains, Trois one of the dominant plants around 
Lyndhurst, where. in bebruary 1996, all examined 
shrubs exhibited a low (o moderate infestation by the 
gull midge Alericifarnity sp, ney, 


Material and Methods 


Galls were sampled trom Halosareia tndicu subsp. 
leiastuchye at Lyndhurst (15.16, L996) and Helosercte 
pergrdnuldi subsp. pereranulaca at Port Adelaide 
(5,y, 1996), The galls collected on both occasions 
were processed im one of two ways. A small nimber 
Was cur open and the larvae preserved in 70% 
ethanol A larger number of galls was kept in plastie 
bags ane the lirvae were reared to adults. Pupution 
look plaice within the galls. Pluste bags were exgurt- 
inca daily and emerged adulis preserved. together 
Will tein pupil skis, tn 706% ethanol, Canada bal- 
Sunk amounts of type speciinens for microscopic 
c\amination were prepared according to the teeh- 
mque outlined by Kelesik (l995a) ALL rmeasure- 
ments refer lo Lype series. The type series and other 
mate rekined in 70% ethanol are deposited in the 
South Australian Museum, Adelaide [SAMA and 
the Australian National Inscet Collection, Canberra 
[ANIC]. 


Genus Ayphandyite Loew, 1850 


Loew. 1650; Dipterologische Beitriige. 1850: 21 and 
37 (us subgenus Of Cecidopinia Meigen, 1803) 
Type species: Cecidumiia serothanny Loew. 13s. 
Jace 38 (des. Karseh. IS77): Revision der 

Gallmiieken: 15), 


Asphondvia is a worldwide genus thal currently 
Comprises some 260 deseribed species (Gagne 
1904) Tt cantiins species that have a ventro-distal 
spine on the first tarsomere, the ovipositor with) lurve 
hasal lobes, female fMlagellomeres 9 - 12 progress- 
ively shortened. the ganovosite with aw ventro-apucil 
lobe anda dorsally situated gonostylus that is about 
as Wide as long and hears two basally merged teeth. 


Asphondylia inflata sp. nov. 
(FIGS |, 395, 7-9, 12-14, 1-18, 21. 24-27. 31, 44) 


Halotype: &, Port Adelaide. South Australia | 34°50" 
S. 138°30' Ef. cinerged 6.v.1996, P. Kolesik,, reared 
from braneh gall on HW. pergranudata (Black) Wilson 
subsp, pereranulini, gall collected 5.,]996, 12 | 283 
|SAMA|. 


Paratypess 1a. 382, 2 pupae. 1 pupal shar 
[SAMA], be. 324. 1 pupa. | pupal skin [ANIC], 
all same dati but emerged 5.9.-13.vi.196: 7 lurve 
[SAMA], collected with holotype. 


Orher material: 29 2, 13 popil skins (SAMAJ, col 
fected with bolowpe, 


Mule (Pigs a-5, 7-9. 12, 14) 

Colour: selerotized parts of body dark brown, non- 
selerotized purty of abdomen orange. 

Head: Antenna: seape broadest distally. length 1.3 
x breadifi ar distal end, 1G - 1-8 % length pedicel, 
pecicel about ay broads long. first Tagellomere 2,0) 
~ 2.2% length of scape. flagellomeres evenly eylin- 
drical. eireunfila dense, equally distribured along 
scemients. Eye ficets close together. hexugunalid, cye 
blidge 6-7 facets long, Frons.with (2 - 18 setae per 
side, Lubella reduced in size. fused. Talerally wath 3 
G setae. setulose. Masillary pulps 3 segmented, 
length of third segment. us well as total Jeng. vari 
able 

Thorax: Wing length 3.5 mm (range 3.4 ~ 3.5). 
width |b mm (14 - 15) Se cell pigmented proxt 
mally. Anepistermunr and anepimerou covered with 
scales. Claws of all legs subequal in size, similar in 
shape, as Tong as empocis, 

Abdomen: Genitalia: zonocosties short. with long 
veolo-apieal lobes gonostylus with 2 unequal apical 
frecth. ventral about 2 x length of dorsi: aedeimzus 
elongate and narrow. 


Fermele (Figs (4, lo-bS, 21. 24) 

Frons with 9 - TT setae per side: Twellth tagelly- 
mere sometimes fused with eleventh. Cireumliti 
comprising two longitudinal bunds connected by two 
short transverse bands.. Wine length 3.4 mun (2.7 - 
37). wittth 1.3 min (11 - 14). Seventh ubdominul 
sternite 22 x (2.0 - 2:3) Jength of sixth. Genitalia: 
ovipositor 2.2 x (1.9 - 24) length of seventh sternite, 
cere) glabrous, Other characters-as i inale, 


Bins 3-140 4 Head of mule Asphomdylia inflata sp. acy un frontal View. 4. Genilatie of mule A yplieid\ a ciflatet sp. now, 
in dorsal view. 5. Gonostylis ol male Awpliwidvbie difatd sp wy in posterior view, 6. Gonostylus of male Asphonidy ta 
erieifarniin sp. (ov. Ti posterion View, 7. Last three Magellameres al malé Asphotdlia inflira sp.nov Bo Sixth Mazel 
lomere of pile Asphoiedviia fafiita sp. ney. 0. Male Asphendyia aafliid sp. nay, Last tirsomere with chiw and empodi-~ 
wi, LOd TL. Maoathiey palpus Of tale Asplandviierertedoriuy sp, doy. 12, First tursamere of male Asphondvlia miflo- 


He sp. NOY 


13. Wing of male dyphondyiie vefer sp, now, Seale bars = LOO wor 4.4. 7-122 50 pint 5. @. S00 us 13, 


P. KOLESIK 


NEW GALL MIDGES FROM HALOSARCIA 3 


Papa (Pigs 25-27, 31) 

Colour: abdomen orange. remaining parts dark 
brown Total length 3.0 mim (2.8 - 3.3). Antennal 
horns serrate medially. P83 pin (al - 191) long 


Upper and lower frontal horas simple. Two pairs of 


papillie on lower face. one of each pair with w seta 
Prothoracie spiracle broad at base. narrow on distil 
half curved beyond tracheal Opening at mid-length, 
Abdominal segments 2-8 with bwo pairs dorsal 
papillae. two pairs pleural papillae, and one pair yen- 
tral papithie. all papillte setose, Abdominal dorsal 
spines simple, prominent pairon last segment curved 
faterially, 


Lasse dasteee larva (Vig. 34) 

Coluur: orange. Integuiment covered with dense 
spietiiae. Length 3.0mm. Head capsule strongly pig- 
mented, postero-hitend extenmsiods shatter thar 
length of head capsule. Spatubie with two tong, pom 
ed anrerioe teeth. shaft narrowed pear mighdle. 
Widened again posteriorly, surrounded iiteriorly ind 
Jurerally by extensive pigmented, glabrous urea 
Papilhur pattern generally ay lor Asphondylte (Mahi 
1955) exeepr only 2 lateral papillae on each side of 
sputulicaind ho terminal papillae visible au the avail- 
able specimen 


Lovnitoey: 
Phe name “hake” pst Latin adjeerive Tor i Fkated 
referring: 6 the appearince of the galled beanel, 


Galland hinlogy 

Brweh seaments of Milosareia poreraniulitte 
subsp, pererunilate infested by this gall midge are 2 
- A times larger than normal in-votume, greyish green 
und Nard in eontrast to the vivid. green colour and 
sof texture of uninfested brunch segments (ig. 1) 
Bach wall has one to three chunthers, with one lurve 
in wich chamber, The chamber wall is fined with 
bard. pale-green, 0.25 - 0.33 mint thick tissue, 

Pupation takes plaice inside the gall A] cieular 
brown area appears on the top of the gall before the 
pupa cuts a opening wilh its antenoal horns, Ov 5 
May. 1946, at Port Adelaide. the gulls appeared very, 
conimon in the host plant popalition covering sever 
al hundreds of me. 


Asphondylia ericiformis sp, Woy, 
(FIGS 2.6, 40, 11, 15. 14. 20, 22, 23, 28-30, 
32, 33, 35) 


Holotype: 2. Lyndhurst. South Australia {30°17' 8. 
138921" Bj, 204.1996. PB Kolesik, reared front 
brinch gall on Malevarere indica subsp, leresracliya 
(Benth.) Wilson, gall collected 15.411,1996, 121284 
|ISAMA], 


Porapes: V2 20) pupa. lb pupal skin [SAMA], 
IFS, bE pupae 7 pupal skin PANEC]. all suame dita 
but emerged 25-27,1),1996: 3 larvae [SAMA]. 2 Tar- 
var |ANIC], collected with holotype, 


Other material 7 pupae. + pupal skins [SAMA], col 
lected with holotype, 


Male (Figs 6, 10, 11) 

Frons with 6 - & selue per side, Wine length ab 
mm (2.1 - 4b). width bmn (9-16. Bye bridge 
6-9 facets long, Ventral tooth on vonostylis as long 
ats dorsal, Otherwise as in AL laflane 


Female (Figs (5, 1Y. 20, 22. 23) 

Prous with 4-8 sete per side. Wing length 3 
ny (2.3 3.9), width 1.2 nam (09 Tab. Seventh 
abdominal sternite 2.305 (2.0 - LA) length of sixth, 
Ovipositor 19% C14 - 2.0) length of seventl sternite, 
cere) tibrous, With line oF teeth dorsally. Otherwise 
ere, Hflata, 


Mipa (Figs 28-30, 32) 

fot length 4b a (4t - oh 7 Antennal horns 
198 pre (1S4- 214) Jone. Upper frontal born simple, 
Hooother Hore presen. Dorsal spines of last abdomi- 
Hal segment bout same length, strarehe, 


Lusr instar larvae (Wigs 43,345) 

Total lengil 3.2 nin (2.4- 4.8). Spatula with long, 
pomled anterior eth, shaft stort broad, parallel 
sided. Tiree Literal pupitlae on eieh side of (horace 
segments. fo terminal papilkie visible on tevailubly 
specimens, Otherwise as an ch. iffede, 


hivinalogy 

The name “erieifecnos” isa composed Latin adieu 
tive formed from “erieius” (hedgehog) and “Tormnis’. 
referring to the the hedgehop-shuped gull, 


Gall und binlegy 
This species transforms brinch segments ol 


hias 14-24. 14, Female doydiendvia iillare sp. now, Busal lobes on pyipesitor in dorsal view. 15, Female Aspliondyliin eris 
olferinis ap. nov, Basal lobes of eyipositor in dorsal view, 16, Antenna al femiule Avplroweylica aiflata sp. voy. 17 & 14, 
Masillary palpus of female Asphondylic inflata-sp, nev. 19 & 20, Maxillary palpus of female Asylonedylia erroifarnits 
sponoy, 2), Sixth (hiwelomere af femiile Ayphendyvlie taflate sp. nov, 22. Female Asphondv lia evicifarmis sp. nov. bn 
of abdomen in laleral view, 23, Bemale Asphondy fie ericiformis sp. nov. End of ovipositor in lareral View. 24, Female 
Ayphondytia inflate sp. nov. End of ovipositor in duteral view. Seale bars = 100 pin 14-21-24, 24: S00 pm 22 


64 


P. KOLESIK 


NEW GALL MIDGES FROM HALOSARCIA 45 


Halosarcia indiey subsp. leioytachye into spherical, 
spiky. monothalamous galls, each oceupied by one 
larva (Mig. 2). Outer diameter of gall @ - 12 mm- 
inner diumeter 2.0 - 25 mm, Chamber wall lincil 
with hard, brawn, 0,25 - 0.33 mim thick tissue. 

Pupation dukes place inside the gall, On 15 
February, 1996.at Lyndhurst. (0 examined shrubs of 
the host pluat bore a total of about 200 gully of the 
new gall midge species, The galls contiined larvae or 
pupite. 

Remarks 


Four species Ol Aspliondylia have been prev ioushy 
knowe to occu i Australia (Gagné 1989: Kolesik 
1995b). Asphondvlia dedenueue. a South Australian 
species commMan inthe Adelaide Hills, ntalforny term - 
inal branch stems und primary leat veins of Dedonuea 
Vixcase (Sapindaceae) (Rolesik 1995b). ‘Twa species, 
A dogwi and A. rubicinde, were described in the pre- 
vious century from adults caught in fightin Sydney, 
New South Wales. and their bialogy as unknown 
(Skuse PRSS, L890) The fourth, AL filli, was 
described froma feniles and pupae bred from an 
unknown plant in Darwin, Northern Territory 
(Edwards 1916). These last hice species, which were 
deserihed superficially and can not be conipured on 
then’ deseriptions, are not Considered in the present 
paper, but | plana review of Australian Asplonedytte 
spp dita hater stage. 

Morpholowical similarities helween the (vo new 
species and the fact that their réspeetive lost 
phuts belore to The saine wepilis, sugwest ac lose 
rehitivoship. They form a distincive group that 
excludes AL coderede Asplimedyvifa dodeitucue 
differs from the two few speciesin the following 
respects. Adulis hove prominent labetla and scapes 
is long as brow al the distal ends, The male hus u 
shoraeyentro-apreal lahe on the gomocoxite, srall 
lobes udjuwent ta iweth on the gonmostylus and ue 
wedeagus Wich shorter than the gonocusites. The 
seventh abdumrmal steiiie in the fenyale is three 
Himes longer than the sixth. The pupae has nie 
frontal bors. (be antennal horns ave trivnwelae and 
seimaled: Lamerdiy amb the prothapacie spiacte 7s 
Noe Considerably beodder atthe base The ared sur 


rounding the spatula inthe hurva 1s not przmented. 
Adults of the hwo new species diller from euch 
other most prominently in the shape of the gono- 
stylus and the end of the ovipositor, The yonu- 
stylus hears teeth of unequal length and the end of 
the ovipositor lacks external processes in A. iafla- 
tt. In contrast, the gonostylus of A. erieffermis 
bears teeth of equal length and the end of the 
oVvipositor is serrated. Mare differences are evi- 
dent in the earlier developmental stages. Uhe pupa 
ol A. inflate has both upper and frontal horns pre- 
sent and a prominent pair of dorsal spines on the 
last seginent is curved laterally: ind. ericifaraiy 
the pupa has the upper horn only and all dorsul 
spines on the last abdominal seginenr are equally 
stony and straight, The lacvae differ in the shape 
of the spatula and the sumber of lateral papillae 
two per side in A. daflata but three per side ii A. 
eriomiis. The wo new species resemble cach 
other in the shape of the antennal horns und pro 
thoracie spitucles in pupac. tbe antennal seaments 
in adults. the ventro-upicul lobes on the gomoeox- 
ites In midles and the relative lengths of the sixth 
und seventh wbdonimeal stertites mi females. The 
elongation of the yventro-apical lobe on the gone 
coxite 14 Unique to these bWo species und disuiog- 
wishes them from the other Ayala vlia spp. 


Acknowledgements 


Tamogratetul to PG Wilson, Western Australian 
Herbarium Comino for the wentification of Halayarore 
jidicu subsp. feisrarhya. Ro J. Chinnvek, Seuth 
Austrauhan Herburiuin Adeliide tor the identification 
Ol Halayetrcd pergranulie subsp. pergranntere, 
Tt, B Reardon who led a Sooth Avseatian Museun 
collecting trip during which Asplorndylie ericifearnis 
sp roy was discovered, A. Stark. Walle Germany for 
providing copies of Kasel’s aod Loew's papersaind 
J 0) Gray, Department of Hortieulture, Viti 
culture and Oengloey University of Adeltide and 
Ro} Gayne, Sysiemahe Entomology Laboriwory 
USBA Washinglom DC for ihrem coments on ay 
early dralt of the manuseript 


Pips 25-48. 25 Aqphaniviio dative ap naw Antenne parent pupa in ventral views 26. Ayydanediir inflate sp. new, Anteriot 
park el pupa rater stew. 27, Aphids tie (u/ledé sp, now, Prothoragie spiiaghe of pupa, 2S, daplonedyiar eeloiorays 
ap Hs, ARLEFLOL pare a parpa Or ventral yas. 24, lytic enietovins sp. roy Anterior parhol pupal in literal viesy 
40, Aguiar lo iciloruits spony, Prothoraeicspiracle of pupa, 47 Pupa ol lsphendy tig taflala sp now. Last ybdtonn 
ind seanene iT dorsal view. 72. Pupaol Ataenedviie cercdor ssp now, Least abdoniina segment in dorsal view. 43 
Larva ol Asplondylin eretiowiy spy, now Last (wo abdominal) segments in dorsal view. 442 Agterion part of tive ot 
Ayphornmdyiir ioleatiesp, toy, tn wut view, 35, Plead quid fist thorauie segment of larva ol lap hemidvdien vei ipernity op) 


noy, meventral view, Seale burs = TOO ya 


66 P. KOLESIK 


References 


Epwarps, F. W. (1916) Two new Australian Diptera. Ayn, 
Mag, Nat. Hist, 103 (8th Series, Vol, 18), 498-502. 

Gaane. R. J. (1989) Family Cecidomyiidae pp. 152-163 /n 
Evenhuis, N, L. (Ed.) “Catalog of the Diptera of the 
Australasian and Oceanian Regions” (Bishop Museum 
Press and E.J. Brill, Honolulu). 


(1994) “The Gall Midges of the Neotropical Region” 
(Cornell University Press, Ithaca New York), 
KaArscH, F. A. F. (1877) “Revision der Gallmiicken” (E.C. 
Brunn, Miinster i. W.). 
Ko esik, P. (1995a) A new species of Eocincticornia Felt 
(Diptera: Cecidomytidae) on Eucalyptus fasciculosa in 
South Australia. /. Ausir. ent, Soc, 34, 147-152. 


(1995b) Asphondylia dodonaeae, a new species of 


Cecidomyiidae (Diptera) damaging leaves and branches 
of hop-bush, Dodonaea viscosa (Sapindaceac) in 
Australia. Trans. Ro Soc. §. Aust. 119, 171-176, 


(1890) 


Loew, H. (1850) “Dipterologische Beitrige. Vierter Theil. 
Die Gallmiicken. Zu der Offentlichen Priifung der 
Schiller” (Das Konigliche — Friedrich-Wilhelm- 
Gymnasium zu Posen, Posen). 

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

Skuse, PF. A, A. (1888) Diptera of Australia, Part 1. Proce. 
Linn, Sov. N.S.W, (2nd Series) 3, 17-145. 

Diptera of Australia, Nematocera 
Supplement 1, /hid. 5, 373-412. 

WILSON, P. G. (1984) Family Chenopodiaceae pp, 81-317 
In George, A.S. (Ed.) “Flora of Australia.” Vol. 4 
(Australian Government Publishing Service, Canberra). 

(1986) Family Chenopodiaceae (Dysphaniaceae) pp. 
236-311 J Jessop, J. P. & Toelken, H. R. (Eds) “Flora ol 
South Australia”. Part | (South Australian Government 
Printing Division, Adelaide). 


A LATE EARLY CAMBRIAN TRILOBITE FAUNULE 
FROM THE GNALTA GROUP, MT WRIGHT, NSW 


By J. B. JAGOo*, LIN TIAN-RUIf,, G. DAVIDSONE, 
B. P. J. STEVENSY & C. BENTLEY* 


Summary 


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. 


Tramsac rons af the Raval Soetwly of S, Aunt (1997), T28(2), 67-74 


A LATE EARLY CAMBRIAN TRILOBITE FAUNULE FROM THE GNALTA GROUP, 
MT WRIGITT, NSW 


by J, Be JAGO*, LIN TIAN-RUU, G. DAVIDSON, B. PJ. STRVENS! & C. BENTLPY® 


Summary 


IAGO, 7. BL. LIN TARE, DAvIDSOM. GL STEVENS, BL POA Brainy, ©. (1997) A Late Early Chombrin 
(nobite Taunule from the Goalit Group, Mt Wrinht, NSW, Fras, Ro See S, Agst, 1202) 67 74, 30 May, 1997, 

Irilubites from a new Jogilily wilhin a siltstone of the Cyimbrie Vale Formation, western New South Wales, 
we deseribed Here as Aeedliohia el. ciguiensiy Din 1878 and Ayuwaypis cerustey (Opik 1975) The species 
deseribed us Sireniay cerates Opik und Exiinga blobare Pocock from nearby localities are included jn asingle 
redefined species. 7 coraytes, The gener Pyeudi tema Chu & Zhou mn Lier al (1974) wad Ate Opik 
ITS are phicecd! ia synonymy with Asay The funy described here is of hile barhy Cumbrian (hale Bolomun) 


ape, 


Kiy Worps: Cambrian. Trilobita, Aastedia. New South Wales. /Avneapis, Cymbrie Vale Formation. 


Introduction 


Davison’ discovered a small area of fossiliferous 
siltstone of the Cymbric Vale Formation in the Mt 
Wright urea, western New South Wales (Pig. 1). 
Davidson identified the trilobites as Avreren cerayles 
Opik and Myiiagia bilobala Pocock of Opik 
(19756). More samples were colleeted reeeutly and 
the results of the examinition OF the fossils are given 
below. 

‘The Barly to carly Middle Carmbrian Gnalte Group 
comprises three formations, from bottom to top: (he 
Mount Wright Yoleanies, the Cymbrie Vale 
hormution and the Coonian Pormation, The distrih- 
ution af these formations us shown in Fig. | is ater 
Warris ind Rose (1968), 

The Gralla Croup crops oul along a broad valley 
west of Cymbrie Vale homestead and ts mostly con- 
Jined by the Mt Wright Fault on the east and the 


Lawrence Poult on the west A smaller urea of 


Cymbrc Vale Formation crops out in the core of an 
anticline west ot the Lawrence Fault, The stricture 
of the Gniltt Group is very imperfectly Known as a 


Deputmenel Append Gieolupy. Sehowbot haineernnge 
Tlityorsiy of Sant Atistedlie The bevels So Atist 3095 
Department ol tgerth Sciences. Nanjing Uaiversiny 22 Hankew 
Bowl Nun 22008 People's Repablie of China, 

Geology Dopartinicit. Gliversity al isnt GPO Box 252 
Hobart Tas TOOL 

| Cealogieal Survey of New South Wiles, 32 Sriphide Street 
Brokon HAL NSW 2880, 

Dowibsoa, GO (MOR A contibniron tothe geology at the Mi 
Wright area, BSe Clons) thesis, Austin ational Unversity 
(unpub) 

Wokiis. Bu (1967) The sfiiligriphy unit pakicontoloyy of 
Horthwestera Mew South Wales. PhD thesis. Sydney University 
(itu 


result Of geological complexity (faults and: folds), 
poor outcrop in much of the area and a lack of hed 
ding in the abound voleunies of the Mt Wrelit 
Voleunies, 

The siltstone to fine sandstone outerop from which 
the fossils were collected (Pig. 1) is tsoluted trom 
Oller outeraps by u vover of soil and cobbles repre- 
senting a Cainozoie lag deposit, Henee there ts nu 
outcrop continuity with any identified Gnali Group 
formation. The outerop also contains po diagnostic 
rock type. Phe structure of the Caialla Group is sulft- 
crently complex to render it impossible to vonfident- 
ly plive the ourerop ina formation, Davidson placed 
this and nearby oulerops im the Cyimbric Vale 
Formation, while Krise (1982) placed the pearby 
oulerops in the Coonigan Formation, As shown 
helow, the species deseribed by Opik (1975h) from 
the Cyinbric Vale Formation as Lyteiingian bilatrte 
and Strenay cerasfes are found wt the locality being 
considered here, Na such fossils ate known fan 
either the Mt Wright Voleanics or the Coonan 
Formation, Henge, the outerap is probably Cymbrie 
Vale Rormation. 


Previous Work 


‘The teem Gialtt Group was mtroduced by Warris” 
who ussigned ao Early to early Middle Cambrian age 
to the group. Prom limestone in the Mt Wright 
Voleames, Warns identified algae and archacocy 
athids, including Tidlumoeyatiiy tracheatiy Taylor, 
From the Cynmibrice Vale Formation he identified 
Fetiingia biloba, Calodiseus sp. and Pagzerides sp, 


and assigned a mitddle or hue Barly Cambrian age, 


He identified the trilobites Redliehia idenea, 
Nyseridura saints. Pageria signtficais. 
FPeranopsis narmata. Orvetovephality sp. and 


J B.JAGO, LIN TIAN-RUL G, DAVIDSON, B. Pd, STEVENS & C, BENTLEY 


an Ow 


Vay 


5 kilometres 


ais 


REFERENCE 


Ganazdic seuinierits, Willies Puplialy Go Ustune, sv ate 
Wyuatennity alluvin. 


Deyoran- Garlornlurous Rayanddle Punmatiin acifl 
Weetlerno sandstones, 


Mile - vate Davenian Snake Cave Sandetcne 
sarlatone, menor congloinemte 


wn a us ine 


= | Pally - Middle Ordeyician Rowena Fornitation, 
Salton, quanwte, sitslone, 


Dorvpyge sp. fromthe Coonigan Formation, indieat- 
ing an early Middle Cambrian age. 

In jerms of the Cymbric Vale Formuion, Pleteher 
(1964) first recorded archaeoeyatha from lenticular 
limestones near the base of the writ. Kertise (1982) 
noted four arehdeouyath Hinestone-bearing lenses: he 
considered two to be of Aldabanian or Lenan age and 
two to be of Lenan age, The uppermost! Cymbric Vile 
Formation contiuins well-bedded sandstones amd cur 
honutes with abundant fossils. Opik (1975b) record 
ed the Wilobites Divesuy alt, wraiidosius, Testeliipedal 
hilohata, Strenax verustey, 8. fletcheri, Serveadiscus 
deedalus. Meniscuchis tienen, Pinvenesttes 


fregumt und Pagetie, the monoplicephoran Scenely 


retieulta Ahe inarticulate brachiopods Borsfardia el. 
cdelutd, Livenlella and Nenbolus, eoerinoids ini 
sponge spicules, 


Age and Correlation 


As shown below we consider [hat the species 
described by Opils (LY75H) as Stremaey cenisfes anal 
Lying bilabrea tron nearby toualities should 
both be ineluded within w single redefined species 
Heispis cerastes. These: localities Opik eonsidered 
ju be equivalent to the Siemshtyk gol Horizon oF the 
Altay-Sayan region which, as noted by Zhuravlew 
und Oravestock (M94) can he roughly correlated 
with the Botonjun of Siberia. The archieoeyathia 
from the Cymbric Vale Formation whieh oecur 
below (he trilobites described by Opik (1975b) were 
correlated with the informal archaeoeyathid unit the 
Sviigecnema favus beds by “Aburaviey und 
Gravestack (1994) which wus considered fo he 
equivalent to the middle (o Lite Botoman by these 
aulhors, These authors considered that the 
Syrinvecnema fevus beds fall within the trilobite 
based Pararaie janue Zone of Jell (in Bengtson ep al. 
1990). dell suggested that the Po jamae Zone can be 


Lal Garten Crifly Ghlidian Nootumbiilia Randslonae 
aafiletorn Getiolomerats, shale, limertore, 


correhded with the Botoniui and probably with the 
jute Botoman (see also brief discussion by tage 


MOOTWINGFE 
GROUP 
I i : 


Pp 


GWALTA GRA 


e) 


7 =| Miditta anthem Conner F atnalior 
a Wnestons shale 


Cambrian Eyre it Yale Pay nation 
= Sele Paiee (terperiniitte: gelertnyes 


Sd Wie} 


Tar) Cambie Mb Wright Maluanios, 
Piormmciets snd del Volearies. rrierar alate. lenestanes 


< ARLES PAINT PURI 

fee ee Gai]! Ronit Ged. Te cuns TipMicHU ete ibbAbacelt 
Stl Peet HAS) Selene GIS, Het geo Sel iiss, 

PDR EA Letter | ee? 


ol bale Vrotermoyhe Early Garnbriiny’) Mitt fede, 
WOLD Stutle sthditinS Causal litnSti ties aan 


[sane Hihyenvetin betes 
Sryjron bait veth t kicks ol ant aal lliatite 


Tal oy) Lips ded Shi 

Trinh . Hui auleas 
--£ yey ° Walerloreu 
——P——-  Aplutyic ee fasulLowhly 


1996), As noted above, the exact stratigraphic posi- 
tion of the present fuuna is a Hille unelear bul it prob- 
ably occurs high inthe Cymbric Vale Formation und 
Stratmraphically close fo the faunus deserthed by 
Opik (lY7Sh). The Coonipan Formation which over- 
lies the Cymbric Vale Formation has a very carly 
Middle Cambrian ave (Opik [975a, Jell (975; 
Shergold ef al, T98S), this providing an upper limil 
fo the age of the Tauna currently being desembed, 
Palmer and Rowell (1995) note that Heuuspia is 


Vig. 1. Geolowy of the Mt Wright siren shawing lovality 
of fossils described herein, Geolagy from Warris_ Rose 
(1968 Davidson! Kruse (1982), BL Stevens and 
A. Criwloul Cimpub. field data) 


EARLY CAMBRIAN TRILOBITES FROM WESTERN NSW nu 


elosely related to the Siberian genera Bergeriniel/us 
and Bergeromieaspiy which churaclerise the Botonjn 
of Stheria. 

In China Reedlichie cigviensis Gveurs tn the tate 
Lurly Cambrian Poldealenus Zone af the Yargtze 
Gorge This is approximutely equivalent to part of 
the Toyontin of the Siberns Platform, 

The above discussion suggests that the faunule 
deserihed herein is of late Barly Cumbriin age und 
probably of Late Botomuan age. 


Systematic Descriptions 


‘Temmnvlogy essentially Follows Darrington es al 
(1959), Specimen miimbers refer to the palacontalag- 
ical colleertow of the South Austria Museum 
(SAMP). 


Family Redlichtidue Poulsen 1927 
Sublimily  Redhehime Poulsen 1927 
Genus Reedlehia Cossman 1902 


Redlichia ct. ciguiensix Lin V978 
(Tig. 2A) 


W978 Redlichia cieuieasis Lin a Zhou & Lin 1978, 
p. 14S. PL 20, Fi. 8. 


Material 
Gne incomplete cramadium, SAMP35 344, 


Deseripliamt 

Criuntdium subrectinyular, about 35 mm long. 
Glahellw (including ocvipial ring) about 0.9 length 
OF crunidium. Ghibella tapers evenly forward to 
rounded glabellue anterior. Narrow shallow axial fur 
rows. Three pairs of lateral glabellar furrows; |p, 
moderately deep. directed slightly to posterior and 
meet at centre OF elahella to form ain evenly posteri- 
orly arched furrow: 2p pair, shallow, direeted slight- 
ly to posterior, 3p pair, represented by faint depressions 
on glabellar margins, Occipital furrow deepest later- 
ally: shallow medially where itas urched slightly to 
umeries, Anterior border furrow of moderate depth: 
anterior border very gently convex, and of uniform 
Jenvth (Sag), Very short (sagz_) preglabellur field. 
Small anterior areas of fiXigenae gently Conver; long 
wide palpeboul lobes extend from just lo anterior ol 
Sp furrows to level wilh occipital furrow. Narrow, 
distinct palpebral furrow shallows pasteriarly. 
Pulpebral areas of fixigenue very gently convex. 
Width of palpebral areas of fixivenie at their widest 
is about Gre-third that at Glabella. Presculir seetions 
Of fasial suture markedly divergent, 


Discussion 
A single incomplefe internal mould of cranidiumn 
makes a definite specific identification difficwlt 


However. 1s sive, outline, slender glibella, the posi- 
hon of the postenor of the palpebral lobes. the nature 
of the preglabellar area and the shape of (he preocu- 
lar seefion of the facial suture suggest Reelichia 
signiensiy Lin (ie Zhou & Lin 1978, p. 145. Pl. 20, 
Fig, $). However, it Jiffey's (a having a more tapered 
glabella, a forward arching at the centre oF the oceip- 
ital ting. and a more sharply rounded glabellur ante 
ion 


Superlamily Lllipsocephaloiden Matthew L887 
Family Ichangiidae Zhu LO80 
Genus Astiaypiy Chung 1957 


Ayyayprs Chang (9572p, 45. Lu ed al, 96S. p. 85; 
Zhang ef al, JOSQ. p. 244; Sell in Benytson yr al. 
1990. p. 310. Palmer aud Rowell. 1995, p. 16; Nedin. 
L995. p, 36. 

Lstaingia Pocock lU64. p. 462: Opik 1975h, p. 10 
Peudichanwia Chu & “Zhou iy Lavet af, WTA, p, 93: 
“bute Zhane eral, TY p, 230. 

Syenax Opik 1975 p13. 

lotvietlee Zhang & Zhu in Zhang ef ul, LYS, p. 247. 
Type species: Lusiafops vitesis Chang, 1953, p. 
12h. Pl. 2, Figs 1-16. 


Disettssron 

Jell (in Bengtson et af. 1990) placed Lsrafnera 
Pocock 1964 und Zhadella Zhang & Zhu (in Zharrg 
eluh I980)in synonymy with Asyaspisy Chang 1957, 
a move supported by Palmer & Rowell (1995). 
although the fitter authors expressed some doubt as 
ta the suprageneric position Ol Asuaspis In addition, 
Jel sugested that Strenar Opik 1975. might be 
regurded as a junior synonym of Pyeudichangid Chu 
& Zhou fn Lo er af, 1974 fron southwestern China. 
The present authors sopport Jell in placing Srremety in 
synonymy With Pretadicfangre bur alsy consider that 
Prendichangiir ty a junior synonym of Asaespis, thos 
also placing Saeaay in synonynry with Aswaspiy. 

The following species ot Pyeudiehangia as figured 
in Zhang eral 1980, P. daniianensvis (Chats) (PL 76, 
Figs 13. 14; PL 77. 1-3). PB ronwejfensis Zhu (PL 77, 
Figs 6-8). shuvessiy Zhang & Zhe (Ph 134, Fig. 
2) and Pyeuelicfhangia (21 inwalite Zou. PL 77, Pigs 
910, may all belong ina single species. we. Ayuasyrs 
domiaoeusis (Chang). 2haviella hubtensis Zlang & 
Zhu (see Zhang ev al, M80, PL 134. Pig. 3) also 
appeurs fo belong in H. demmianensis us far as. can be 
determined from the available figure although the 
glabelki of ckiiqeerses extends Curther forwards 
than dior the type species ol Ayuaspiy, A. vinenyiy, 
and the anterior sections of the facil sutures. of 
sinensiv are more divergent than those of daniteoen- 
wis. We would cegard these as specifie tather Un 
genenc differences, 


7) J. B, JAGO, LIN TIAN-RUL, G. DAVIDSON, B. P. J. STEV 


| a : 


ENS & C. BENTLEY 


Fig. 2. A. Redlichia ef. ciquicnsiy Lin 1978, SAMP35344, cranidium, internal mould, x 1.5. B-L. Hsuaspis cerasyes (Opik 
1975). B. SAMP35329_ cranidium, external mould. x2. C, SAMP35340, cranidium, internal mould, x5. 
D. SAMP35335, cranidium, internal mould, x2. E, SAMP35331, cranidium, external mould, x2. F. SAMP35343, crani- 
dium, internal mould, x2. G. SAMP35337, cranidium, internal mould, x2, H. Top. SAMP35327, cranidium, external 
mould, x2. Bottom. SAMP35328, cranidium, external mould, x2. 1. SAMP35342. cranidium, internal mould, »2. 
J. SAMP35333, pygidium, external mould, x6. K. SAMP35332, cranidium, external mould. x6, L. SAMP35341, par- 
tial thorax, external mould, x2. 


EARLY CAMBRIAN TRILOBITES FROM WESTERN NSW ra 


Jel in Bengison er af, 1990) quenced the taxcian- 
ic position of Estatigia bifobata trons western New 
South, Wales as described by Opik (1975b). 1h our 
view the specimens deseribed by Opih (19735b) doy 
Nol belong in Aywespis brloheta (Pocock 1964) 
hecause the vlabella oF Opik's Speenmens caxtends 
further to the anterior than does that of TL dflahena as 


deserthed by Pocock (L964). In addition, in three of 


the foureranidia figured by Opik. there is a marked 
forwards expansion of the glabelhke wheres si the 
pe inaterial of Ho bilibata figured by Poeork 
(1964) the glabella is either tapering forwards or has 
a slight waist, In our view the speciinens desenbed 
and fizwred hy Opik (l975h. po UL PL ot, Bigs 1-7) 
belong i the same species us those described by 
Opik (197Sb, po be PL 2) Figs 1-6) as Sirenen 
eefistes. Apart from the specimens fgured hereny. 
Numerous other specimens are available ad there is 
a coynplete gradation from specimens such as those 
figured here as Figs 2B.E whieh are quite similar tu 
the eranidium ilustated by Opik (lO7Sb, PL 2. Kaw. 
T) as the Holotype ob Siren cerastes get ck sp. Noy 
lospecnnens such as (hose figured herein as Pies 2G, 
3C which are indistingdishable from those figurecl as 
Evtcinge biloba by Opis (WY75h. PL. 1). 

Hpik (L975, p. 16, PL 3, Pigs 1, 2 teu. Fig §) 
erected a subgenus of Srremay, el Syren 
(Semarscus) bused oo four crunidia with the oaly 
Hlustemed cranidium having a lengrh of only 3.8 nim. 
Hoos clearlyun ioinalure speennen which should net 
he the basis of a new taxon 

Jef (1990) erected Ayueapis eceipilasping which) 
is churacterised by a relatively long preelabellur Held 
crossed by a preglabellur median Hdwe and the pres- 
ence of a short slender oecipial spine. It is possible 
that the specimens described by Palmer & Rowell 
(1995) trom the Central Teansantaretie Mountains as 
Hsnaspis Cf Te bitohune (Pocock) belong in 
necimtospiia, becuse the Anturcue specimens hive 
a glabella of similir length to S72 accipitesyina as 
well as a osimihir preghibelkur fedian ridge. 
However. the Antarctic Specimens have an occipital 
node rather than an occipital spine. 


Hynaxpis ceraytes (Opik 1975) 
(Pig, 2B-L, Fig. 3) 


1975 Stree cerasias Opik W975b, p. EPL 2, bises 
1-6, leat. Mig + 
1975 Exrainwid biloba Povoek. Opik 1975b. p_ 11, 
PL 1, Figs 1-7. 


Marevial 

Almost twenty crumdia, one partial free cheek 
(SAMPS5434), un ingoniplete thorax (SAMP35341) 
ancl an incomplete pygidium (SAMP35333). 


Dienst 

Species oF (/yvaxpis with distinedly expundedt anter- 
joy part of glabella: length of ghibella (excluding 
occipital ring) ubout O.75-0.8 that of cramadium 
fexcludipe oecipiul rine): occipital ring bears a 
sprog which in some speeimens is a siiall nodes i 
others ibis long und slender. 


Description 

Gently convex cramidivm with wichth slightly 
grester than length pag.) Deep wide uxtal furrows 
shallow anteriorly. Gently conver elubella of leneth 
fekcluding occipital jie) about O 7548 thar of 
cranidium, Cabell ticreuses tn width io antenor 
with sheht waist qear tp furrows: wlibella widest at 
eve ridges, broadly rounded yhabellar unteriar, hour 
pairs Jatera) glabellir fureiiws; Up fureows made rite 
jy deep and each extend abopr 1/3 of distinee aeross 
glabell and directed sliitly fo postertur, 2p tarrows 
shiutlow. extend about 1/3 distuice aenoys glabella. 
and directed very Slightly lo posterior, Ap furrows 
quite shallow and directed almost simight aeross 
vlubella; 4p furrows oeeur as small indistinct pits, 
Occipital ring longest medially; it bears a spine the 
length of which varies considerubly rom specimen 
Lo specimen. In Semi specimens a smull nade pre- 
sent; in others there is u lone slender spine which 
muy hive a leneth about W735 that of ghibellia 
(excluding oecipital ring) in muture specimens (Pig, 
2D) and somewhat longer io immature specimens 
(2B\P). 

Occipital furrow shallows medially, Very short 
(sau,} venily conves preatabellie held, Anterior bor 
dey vently convey with similar tength (sue) be 
prewlubelkir field. Poorly-developed jalectrum pre- 
sent tr some specimens (Pre. 2G). Shallow anterior 
border furrow. Gently conver palpebral areas ob fy- 
igenue have width ubout sume us that ol vlabella, 
Prominent eentro-pasteriorly placed palpebral lobes 
have a length (exsug.) about 0.55 that of glabella. 
Postetior of palpebral lobes meet the broad postero- 
lateral border furrows, Well-developed eye ridges 
slightly narrower (han palpebral lobes, Shallow 
palpebral Surrows, Narrow posterolateral borders, 
Preocular sections of facial suture diverge forwaris 
atubout 30" to the lransverse, Short postacular sec- 
bons of ficial suture. Narrow postecoliteral border 

The single parbal librigenae fis a Wide border 
which extends mto an ineomplele genal spine. It Tits 
wreliculate ornamentation. 

Thorax with at Jeast nine segments. Convex axis 
has willl about 0.3 that of segment. Centrally places 
nodes on 4th and Sth segments of ivailuble thoras. 
Shallow pleural furrows; short bread pleural spines. 
The only available pysiditun as poorly preserved. [tis 
Small and transversely etliprieal. Axis has a width 
about 0.4 (hat ot pysidiuim, Axis extends almost to 


72 J.B JAGO, LIN TIAN-RUI, G, DAVIDSON, B, PJ, STEVENS & C, BENTLEY 


Fiz. 3, Uyuspis cerires (Opik 1975). A. Left, SAMP35325, cranidium, internal moul. x3. Right. SAMP35326, eranid- 
ium, ioternal mould, x3. B. SAMP35338, evanidium, internal mould. x3. C. SAMP35330, eranidiain, internal mould. 
2. D. SAMP35339, cranidium, internal mould, x2. Bb. SAMP35323, eranidium, internal mould, x3, F SAMP35336 
cranidium, internal mould. x2. G, SAMP3S3344, librigena. external mould. x2. 


LARLY CAMBRIAN TRILOBITES FROM WESTERN NSW 7" 


posterior margin. Axis comprises Wwe uaal tings plus 
terminal axial piece. Two pleural furrows and one 
imMerpleural furcaw present. Two border spines can be 
seer in avatlible specimen. 


Morphogenesis 

The simallest ciaiditind is that figured as 2K. In 
ihis specnnen the length oF the glabell (exclading 
oceipiial ring) is 0.7 that of cramdium, tr larger 
specimens such as those figured in 3B,D the glabel- 
la has a length of O.8-0.85 that of eranidiam, The rel- 
auve sizeof the oegipilal spine decreases ~ in siull- 
er specimens (2B.B) the spine has a relatively thick~ 
er base than tv nore mature specimens (3B.D), 


HNC UANT OU 

The considerable variation in the shape of the 
whibella, the path oF the fei suttire. the shape itd 
length of the becipital ring, and the length of the 
vecipiall spine. of the speeuncns described and: tig 
ured here as AL cerastes raises the qhestian as to the 
validity of plieiig all these specimens within a single 
species, However, us noted above, there appears 1 be 
a-coniplete morphologeal yariation present in the 
uvailable specimens ind henve we feel the erection of 
more than one species can not be justified. A/siesypis 
cerasieys us desevibed herein differs tram UL 
daandoensix in having a relatively shorter glabellay the 
glabella of AL, dantldwensiy esther reaches or almost 
reaches the anterior border furrow, whereas that ol 1, 
cerasfes stops short of the border, However, its worth 
noling (hal wilh respect to the length al the occipital 
spine, JE damioensis, as Tyured in Zhang ef al 
(1980. PL 76. Higs 13, 14: PL 77. Figs 1-3) shows con- 
siderable variauior in length of occipital spine as dues 
A. ceraytes. The anterior of (he ghibella of H, cerisres 
is more expanded Unan is that ol A, diamtevenvis. 


Tsuaspix necipimsping at sell (1990) tas a small 
well-developed oecipital spine. The spine of H. 
cerastes arises From further back on the oecipital rng 
than thal of JE eceipiospinag: the spines on sone 
specimens of 4d. cerayfey ure much longer than the 
biggest spines on /7, eccipitosping. In Ayers the 
ratio of the length of the glabella (excluding oceipi 
tal ring) to the length of the cranidiam (excluding 
oceipilil ring) 1s O7S0.805 in A, ecelpitosping it 
ranges from 0.65-0.70. 

The glabella of AL stesiy extends further ter 
wards than does that oF (7. cerastey. The unterfor part 
of the dlabella of 7. cerdsres is more expanded thin 
H, sinensis, H. yinensiy bears only a sthall pedi 
node rather than a prominent spine as mn many spec- 
imens ol AL ceresies. 

The anterior purt of the glubella oF Z/. ceraytey ts 
more expanded than that of J. bitebate: the glubella 
of //, verastes t§ velanvely longer than that ofA. bile 
bette, The glabetla of (ef T. bilubitia as described 
hy Palmer & Rowell (1995) 0s shorter than that of A, 
rerester. 


Acknowledgments 


Sample collection was carried out under a licence 
Irom the New South Wales National Parks and 
Wildlife Service. This paper is published with the 
perinission of the Director-Geoeral, New South 
Wiles Department of Mineral Resources, Lin Tiun- 
Rut was supported by a visiting researcher's grant 
from the University of South Australia, Dr PA, Jell 
constructively reviewed an earlier version of this 
puper. Mr JS. Bau gave valuable technical assis- 
lance. Dr C. Jenkins supervised G. Davidson's hon- 
ours project 


References 


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AJ & Rowancak. BOS. (900) Lark Cambrian fossils 
from Sou Austratat, Meni. Assoc. Auxtretlas 
Patueuntal, 9, | 364 

Foereniba. BLO) (1964) New Tilo shells tron the 
Lower Ondovicnin ind middle Palaeozore rovks of New 
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HargRINGTON, HLS etal (L959) Arthropoda l pp. F560 hr 
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13-21 September 1996, Meld Trp Guide une Abstracts. 


Jett, PA I975) Australing Middle Cumbrian codiseaids 
wilh review of the superfamily. Paloemndographice A 


150. [17 


— (1990) Trilobite pp. 257-322 Jn Bengison, §,, 
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Krviss, PD. (hYTHT New Archaeoevathi front the Burly 
Cuinbrian al the Mt Wrucht area, New South Wales, 
AMlcheringa 2. 27-47, 


(1982) Archucoeyathian bipstaiiweaphy of the 
Gniulae Group ait Mt Wright New South Wales 
Fealeeoutographicd A UT7. | 212 


74 J.B. JAGO, LIN TIAN-RUI, G. DAVIDSON, B. P. J. STEVENS & C. BENTLEY 


Lu YANHOU, CHANG WENTANG, CINBN YIYUAN, CHU 
CHAOLING, Lin, H. L., ZHOU, Z. Y., QIAN, Y., ZHANG, S. 
G.& Yuan, J. L. (1974) Cambrian trilobites pp. 82-107 


Jn “Handbook of stratigraphy and palaeontology of 


southwest China” (Science Press, Beijing). [Chinese. 
English summary]. 

Nurpin, C, (1995) The Emu Bay Shale, a Lower Cambrian 
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Mem, Assoc, Australas, Palacontol, 18, 31-40. 

Opi, A.A. (1975a) Templetonian and Ordian xystridurid 
trilobites of Australia, Bur. Min. Res., Geol. Geoplrys., 
Aust., Bull, 121, 1-84. 

(1975b) Cymbric Vale fauna of New South Wales 
and Early Cambrian biostratigraphy. hid. 159, 1-78. 

Parmer, A. R, & RowskLe, A. J. (1995) Early Cambrian 
Trilobites from the Shackleton Limestone of the Central 
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1-28. 

Pocock, K. J. (1964) Eytaingia, a new trilobite genus from 
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Rose, G. (1968) Broken Hill, 1:250,000 Geological Series, 
Sheet SH54-14, Preliminary Edition. (Department of 
Mines, NSW). 


SHERGOLD. J. H., SAGO, J, B., Cooper, R.A. & LAurIE, JR. 
(1985) The Cambrian System in Australia, Antarctica 
and New Zealand. Jat, Union Geol. Sci. Publ. 19, |-85. 


ZUANG WENTANG, Lu YAN Hou, ZHU ZHAOLING, QIAN YL. 
Lin, H., Zou Zatyt, ZHANG SANGUT & YUAN JINLIANG 
(1980) Cambrian tilobite faunas of southwestern China. 
Palaeontol, sin, ser. B 16, 1-497 [Chinese with English 
summary]. 


ZHOU, T. M. & LIN, T. R. (1978) Early Cambrian trilobites 
pp. 143-154 7/n “Sinan to Permian stratigraphy and 
palaeontology of castern Yangtze Gorges” (Geologicul 
Publishing House, Beijing). [Chinese]. 


ZUURAVLEY, A. Yu & GrRavestocK, D. 1. (1994) 
Archacocyaths from Yorke Peninsula. South Australia 
and = archaeocyathan Early Cambrian zonation, 
Alcheringa 18, 1-54. 


POORLY PRESERVED TRILOBITES AND BRACHIOPODS 
FROM THE KANMANTOO GROUP, FLEURIEU PENINSULA 


BRIEF COMMUNICATION 


Summary 


The Kanmantoo Group (Fig. 1) 1s 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’, the uppermost formation of which, the Heatherdale Shale, contains a poorly 
preserved fauna of possible mid to late Botoman age’. 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’. 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’, i.e. late Early Cambrian or early Middle Cambrian 
on recent Cambrian time scales*®. The stratigraphy of the Kanmantoo Group is 
described in detail by Daily & Milnes”. 


Dansacnons af te Raval Seciew af’ S. Aust 1997), PRU 7 


BRIEF COMMUNICATION 


5 


POORLY PRESERVED TRILOBITES AND BRACHIOPODS FROM THE 


KANMANTOO GROUP, FLEURIEU PENT 


The Koomintoo Group (bine Fy) is a thiek (R-TO) key 
succession al predominantly ehistic metinedimentury rocks 
which crop oot avan urenate bell from near Australi Plains 


in the northeastern Mt Lofty Ranges to the western end af 


Kanvaroo Uslund On Fleurieu Peninswla, the Kanimnuntoo 
Group rests unconforimably on the Normianyille Group), the 
upperfoost formahoa af which. the Heulherdale Stale 
contitins a pourly preserved fiw of possible mid to lite 
Boroman age’, Phe highest well-dited aun fom the 
Nonnunville Group are aehieocyuthids frou the Pork Tree 
Limestone atSellick PHI whieh indice in age close to the 
Atdubatiin-Botoman boundary. These are at least 400) 1 
below (he top ol the Heatherdate Shale. The top of the 
Kuumainloo Croup is hot exposed bet lawer parts af the 
croup are iitruded by the Rathjen Gneiss whieh hus an age 
oF Alé+ + Mat ne. dane Hark Cambrian or earhy Misdle 
Combruia recent Cybern time The 
alnitigraphy of the Kainminton Group is described in detail 
hy Daily & Milnes’ 

The Cambrian sequences as exposed on Fleurieu 
Peninstila. Kangaroo Istand aad) Yorke Peninsula show 
considerible Tateral aud vertical lithologies variation’. 
Currently, the lanited blosianivraphie inforinanc makes it 
dilficull to correlate accurately trom one region to. another 
jm hence derermine the Gining and sequence of Bork 
stmupraphie and tectomie events 

One of the problems tn fhis regard has been the lack of 
biostrutgruphicully uselul body fossils in (he Ryuommantot 
Group. The only stich fossils reperted to dite are the 
imarnculie brachiopod Lingiledée from about 10 ta above 
the base ob the baswl unit ob the Kanmiantoo Group, the 
Carrickuliiga Plead Formation, a Carrickalinga (head!) and 
‘possible hyolithid fron acar the top of the Curriekalinga 
Head Formation near Blowhole Creek!!, Trace fossils have 
been reported from the Backstuirs Passage Formation al 
Accommoedaitlon Hill near Prue! 

This qule reports the presence of Vurther inarticulate 
brachiopods (Fig, 2D), from about 100m above the base 
ob the Carnckuliga Head Formation al Cumickulinga Head. 
They are sivalaind poorly preserved, but appearto represent 
uff lexist (Wo Species. 

In quarry near Parawa (Pig. 4). 01 627,551 (Torrens Vale, 
1S04100 topomaphie map) on Cullovonua Creek Road, the 
vuthors have found the firsh known (rilobiles from the 
Kinmiantoo Group, The specimens come froma laminated 
iehisiltstone. probably part of the Tunkalilly Formation but 
possibly upper Tapinappi Formation, To dite we have 
found dhoot 20 specimens, allot whieh are almost coniplete, 
thus qidivaling low enerey depasitionul conditions As 


on scales 


SULA 


shown in Fe TAB.LC the speciinens are poorly preserved, 
leclanically deformed gn of no blostratigraphic use. 
dlihough, within the fimits of preservation, they seen to 
represent au single species, However, (herr presenve pndicules 
the possibility oof the eventual discovery of 
hiostraticnplieally riselil iilohites front the: Keamgatoo 
Group 


Middleton Sandstone 


Petrel Cove Formation 


Balquhidder Formation 


Group 


Tunkalilla Formation @ 


Tapanappa Formation 


Talisker Calc-siltstone 


Kanmantoo 


Backstairs Passage Formation 


Carrickalinga Head Fm (4) 


Heatherdale Shale 


Fork Tree Limestone 


Sellick Hill Formation 


Wangkonda Formation 


2 
=) 
2 
1) 
2 
> 
e 
G 
E 
tw 
5 
= 


Mount Terrible Formation 


Fig |. Stratigraphic outline oof Normanville anil 
Kaniewloo Groups, Fleurieu Peninsula, showing Ue 
levels of the fossils figured herein, 


MW 


Pip, 2, ACC, “Trilobites from) quarry fear Porawal AL 
Rubber cust of caxternal mould, SAMPS35445, 33. B- 
Internal mould, SAMP35446, x4. CL) Rubber cast of 
external mould, SAMP3S347, x3, DEL tnarticutate 
brachiopods from Carrickulinga Head, DBD. Internal 
moulds. SAMP35348a and SAMP3S448b0 x13, ob 
fuurtially extoliated specimen, SAMP35349, x10. 
Specimen numbers refer to the palaeontological 
collection of the South Australian Museum, 


AUSTRALIA 
a 


Adelaide 


Fleurieu U 
Peninsula area 


Gulf St 
Vincent 


Carrickalinga 


Southem 
Parawaarea Ocean 


2a see inset 
Outcrop of 
Dies Kangarcd Kanmantoo Group 


Island 
0 10 20 30km 138° E 


Tunkalilla Beach 


36°S Southern Ocean 


Fig. 3, Locality map. 


‘Jago, J.B. (1994) Aust. J. Earth Sci. 41, 445-453. 

Jago, J.B. (1996) Some comments on the Cambrian time- 
scale in relation to recent radiometric dating of Australian 
Cambrian rocks pp. 105-106 /n Linan, E., Gamez Vintaned, 
J.A. & Gozolo, R. “II Field Conference of the Cambrian Stage 
Subdivisions Working Groups” Field Trip Guide and Abstracts. 
‘Thuravlev, A. Yu & Gravestock, D.I. (1994) Alcheringa 18, 
1-64. 

‘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). 
°Tucker, R.D. & McKerrow, W.S. (1995) Can. J. Earth Sci. 
32, 368-379. 


Tel. 


Daily, B. & Milnes, A.R. (1971) Trans. R. Soc. S. Aust. 95, 
199-214. 


‘Daily, B. & Milnes, A.R. (1973) Ibid. 97, 213-251. 


°Gravestock, D.I. (1995) Geol. Surv. S. Aust. Bull. 54, Vol. 2, 
3-61. 


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. 


Daily, B., Firman, J.B., Forbes, B.G. & Lindsay, J.M. 
(1976) Geology pp. 5-42 In Twidale, 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. 


SIGHTINGS AND STRANDINGS OF THE PYGMY RIGHT 
WHALE CAPEREA MARGINATA NEAR PORT LINCOLN, 
SOUTH AUSTRALIA AND A REVIEW OF OTHER 
AUSTRALASIAN SIGHTINGS 


BRIEF COMMUNICATION 


Summary 


The pygmy right whale Caperea marginata is a rarely sighted species known primarily 
from strandings. The latter occur frequently in South Australia, especially along the 
north coast of Kangaroo Island and near Port Lincoln’. Migratory patterns are not 
known although it has been suggested that this species moves inshore in spring and 
summer™*. Sekiguchi et al.° 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**. There appears to be a broad mating, calving and weaning period 
between June and February’. 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. 


Transactions of the Rayal Society ef S. Aust. (1997), 121(2), 79-82, 


BRIEF COMMUNICATION 


SIGHTINGS AND STRANDINGS OF THE PYGMY RIGHT WHALE CAPEREA 


MARGINATA N 


The pygmy right whale Caperea marginata is a rarely 
sighted species known primarily from strandings. The latter 
occur frequently in South Australia, especially along the 
north coast of Kangaroo Island and near Port Lincalna!? 
Migritory patterns are not known although it has been 
suggested thal this species moves inshore in spring. and 
summer’. Sekiguchi ey a/> postulated that such a move- 
ment off South Africa may coincide with an increase in the 


abundance of copepods, one of the presumed main prey of 


Co marginata. Spring and summer are also the seasons 
when juveniles most (requently strand’4. There appears to 
be a broad mating, calying and weaning period betweer 
June and February’. This paper describes two recent sight- 
ings of live animals off Lincoln National Park, about 10 km 
east-south-east of Port Lincoln, South Australia and sum- 
mirises past sizhtings and strandings in that area. Other 
known sightings in the Australisian region are alse 
reviewed, 

While sailing a 6-m yacht in Spalding Cove, off Lincoln 
National Park, two of the authors. J. D. and B, F.. observed 
lwo pyginy night whales, an adult about 6 m long, accom- 
panied by a calf about 2 m long. Paired blowholes (Fig. 1) 
confirmed that the adult was a baleen whale. Species iden- 
tification was bused on the curved jaw-line, medium-grey 
colour, a falcate dorsal fin placed well back on the body and 
the adults broad back (Figs 1-3). The animals were seen 
between 1030 and 1130 h on 4 January and 1300 and 1400 
h the following day. On both occasions they were deep in 
the cove al the fur south-western side, about 300 m from 
shore (Fig. 2). Water depth (as determined by a depth- 


Fig. | - Adult Caperca marginata in Spalding Cove, South 
Australia, 4/5 January 1996. Arrows show the paired 
blowholes and indistinct white bar behind the head 
Note also the broad back. Photo: J. Dutton. 


(AR PORT LINCOLN, SOUTH AUSTRALIA AND A REVIEW OF 
OTHER AUSTRALASIAN SIGHTINGS 


sounder) was about Sm, water visibility 8 m, surface water 
temperature 20°C and the sea was calm. Spalding Cove ts a 
shallow, sloping bay with a sandy bolton and extensive sea- 
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 m from the boat. About 2-3 min later 
and 50 m away a large animal, the presumed adult female. 
surfaced and blew. This was followed by the blow of the 
calf near the adult. The calf swam slowly around the boat 
coming close to the bow and turning off. much as dolphins 
often do, It swam with an undulating motion, surfacing fre- 
quently for air (30-40 see or as long as 2 min). At this stage 
the boat was under motor at a speed of 2-3 knots (3.5-5.5 
kph) and the calf showed no sign of being disturbed by this. 
The calf alternated between swiniming next to the boat and 
the nearby adult about 20-30 mi away and sometimes swam 
above the adult, a common position for dependent calves of 
the southern right whale Cubalaena australis (C. Kemper 


Fig. 2 - Juvenile Caperea marginata in Spalding Cove, 
South Australia, 4/5 January 1996. The arrow marks the 
prominent, falcate dorsal fin set well back on the body, 
Photo: J. Dutton. 


40. 


pers. obs. The calls grey back und sometimes its head 
showed as WH suifieed und blew. Later ecamination of pho- 
tos showed (hal the almost white ventral colour extended 
well up the side of the hody and that a distinetive dark at 
cea pately tilerrupted this just behind the head (Pig 3). A 
similar colour pattern has been observed ony South Atricad 
juvenile C. margitata® and at recently stranded neonate in 
New Zealand (van Helden pers. comm.). 

The adult swam jn lirge circles in the general vicinity of 
Ihe bowl, somerimes voming very clase 16 i and at fimes 
turning on its side. [showed no signof being disturbed by 
jhe presence of the boat. The adult surfaced to breathe much 
less often than the call. ubout every 3 mi on more. Later 
exumination oF photos showed that the adult was a medium 
erey colour with a pale band or eheyron just behind the 
head (Pig, 1), The belly was lighter (han the hack. 
Matsiioka also deserihed and iMustrated light chevrons on 
the backs ofa large group of adult Cyanine observed 
use (Table 2), 

The afiiials were in the same part af the cove during the 
allenngon of the next day when JD) and BOR returned to 
sail there, A net fisher operiting in Spalding Cove during 
hilt laniiay and tiich of February 1996 reported seeing a 


smull Whale there on sevent oceysions. His deseription al 


jhe animal does not allow species identificution but it may 
hive been CL mearuiniie. possibly one ob the same ania 
ubserved in eirly January by J.D. and B. Ti 

The other recent South Australian sighting was made dur- 
ing the aternoon al 7 July 1996 by RM, who observed, 
with the aid af binoculars, a small whale from the shore 
new Cape Donington lighthouse, Lincoln National Park 
(Big, 4. Hi was about 50 m from shore and 60 m from the 
observer, The weather Was sunny and the sea calm, with a 
light southerly to south-westerly breeze of 1-5 koots (2-0 
kph), The animal was moving slowly towards deeper water, 
i a south-south-casterly direction. At times it swam just 


— 
-_ 


below the surface with its mouth open, [twas nol possible 
to tell if the animal was teeding and no obvious signs of 
plankton swarms were visible ta the Observer Several low, 
thin blows were seen, The animal’s colour was dark grey on 
the back wad Tight grey underneath, The features which sug- 
gested thal this was a C marginaia were: 1) lighter patches 
between the mouth ind the Hippers. 2) whitish baleen plates 
which darkened towards the outer edges, and 3) a small, 
curved dorsal fin about -/) of the way along the back 

However, without observing the bowed jaw-line, the identi 

ficahon as Coren could jot be vonsidered confirmed 
since minke whites (Baldenaprent aeureraxtrai) share 
several of the above-mentioned features. Estitated: bey 
length of the animal ROM. observed was 45-40 mi. sue. 
gesting thal if was a post-weaning juvenile’, 

Nine stinunings of Co aaginata have been recorded i 
the Port Lineoln area from before 1948 to 1993 (Table 1) 
and, as suggested im 1964 by Hale! many more have prob 
ably aecurced without being reported. ALL the reportud 
strandings have been from Port Lincoln Proper and in, or al 
the entrance to. Spaulding Cove (Fig. 4). Both are shallow, 
scugriss-coyvered, sandy/mudllat bays with large (dal 
movements, All strandings involved single animals. 
abhough i some cuxes There may have heen a connection 
between certain events occurring within a short time ol 
cuch other, For example, u 3.38 fa juvenile was found 
recently dead on 2 February 1989 in Spalding Cove nd on 
| March 1989 a decomposed adult (unknown sex) was 
found on Bigkers Islind at (he entrance (othe cove (strand 
ing nos 6, 7. Fig. 4), On 6 April 1993 iin adult deme, with 
no evidence ol lactation, washed up tear on the south side 
of Port Lincaln Proper and six days liter an extremely emia 
cited 3.15 m juvenile stranded alive in Spalding Cove 
(stranding nos X, 9 Fig. 4+). Hts possible that both cases 
involved mother/ealf pairs, OF the five adults that have 
stranded, three have been fenmiles and two of unknown sex. 


Vig. 4 Juvenile Caperee inetrginata in Spalding Cove. South Australia, 4/5 January 1996, The line drawing helps to dis 
tinguish the bowed jawline. which is slightly distorted by a wave. The arrow marks a lateral colour pattern of white just 
behind the head and a-dark patch posterior to: this, Photos J. Dutton, 


— : 
,y Pl dustim H 
pe | | Ane M 4 b , 
| va - beh " 
oe 
DO .t ) 
Fs ~ 
| ales 3 
a F a Le } 
, f  ertrt ESANHREN Pa 
a, ro WHORE ® g mss 
. OO _ a 
f ~ f 
~~ } 
= oe 


fig. 4 - Map of Port Lincoln region showing positions of 
sightings (squares) und strandings (circles) of Caperce 
mergindta, Numbers reler to strandings in Tuble | and 
Sightings in Table 2. 


These stranding results suggest (hal (he region is Frequently 
used by females und calves. although such iformation can 
be misleading if there is. as in this ease, much human aetiv- 
iy inthe region and fherefore possibility of discovery 
CUPCUISSUS 

Reported sightings of ©. werginata, summarised in Table 
2. are hor eoninion tn the Australasian region. To: our 
knowledge, the Spalding Cove sighting of Jumuary 1996 is 
the first time a cow and calf have been sighted and pho- 
lographed anywhere ip the seutherm hensphere. Nat 
included in Table 2 are two unconfirmed records found in 
the Australian Native Conservation Agency sightings data 
hase. one of three animals off Montague Ishind, NSW in 
June [992 (record no. 876) and the other from new Bernier 
Istand. WA in dune 1993 (io 1558). Since there was no sup- 
porting deseription ol the aninals to allow positive identiti- 
eulion and © angie can be confused with the minke 


Tanbh b 


st 


while, B aentorasirata, the records have not been included 
here. 

Many of the sightings listed in Tuble 2 and several Iran 
South Africa! were made inshore, suggesting thal ©) war 
ginara inhabits coastal waters. at least for some part of its 
life or annual eyele. Unpublished stranding data show that 
dependent young und recently weaned juyeniles (3.0-3,3 m) 
are more common ilong the central South Australian and 
westem Victormn coast (Remper vapubo, Buur of the sitht- 
ings listed in Table 2 involved dependent young or animals 
that, from ther sive. would appear lo tive been recently 
weaned. Three were from the Port Lincoln aren and ane wis 
from Portland tn western Vietoria. Larre. proteetend! bays 
with shallow, sandy bottoms and extensive seagrass beds 
may be importunt calving and weaning areus far CL ma 
vim. Some exanples are Porthind) Bay. Nepean Bay and 
(he Boston Bay region. Several strandings ol C2 ateargimater 
have been recorded in the Nepean Bay arcu! 

The sighting made in Cockburn Sound in }990- by B, and 
D, Parker (B. Parker pers. comm.) merits a specnl nove 
because ibrecords some holable behaviour as well as prob- 
able feeding by the 5-m animal they observed over a period 
of about one hour, The aninal was swimming very fas, 
Joaving oowake of water, and nodding its head poticeubly, 
The whale came to investigite the em bout, scraping itsell 
the first Gime aust the battond of the beat The second 
lime if approuched (he bout very quickly, almost in at 
charge. lifted the boat out at the water aod ainose caused 
The vecupants ta he thrown out! 

The authors wish ty thank B, Parker and D, Coughran tor 
supplying information on the sucht mm Cockburn Sound 
gad the Anstralian Nature Conservation Ageney for searely 
ing their records for Capered sightings, ©, Kemper thanks 
all those who have provided information on pyemy right 
whale strandings and sightings records, especially South 
Australian National Parks and Wildlife officers and 
Department ol Primary Industries South Australia 
(Fisheries) officers, J. Thurmer suggested and prepured the 
line drawing for Mig. 3, 


Revordy of Caperea mannake strandings iy tle Part Lincoln area. Stranding number (Fig, Ref.) croasy-metched 


lo Kino 4, “estimered length based on skeletal measurements. Museum no, M = specimen in South Australian 
Museu, Ss = he specimen in South Austeudlian Muse. 


Date Location ree Sex ore 
<1948 SW Port Lincoln \ 3738 
Proper 

26121955 Tulku ? M 3.05 
1681960 Tulku 3 r dull 
4.1ON4 Spalding Cove 4 - 6.05" 
IS.S.1985 Port Lincoln Proper 3 Ib 6,20 
2.2 1089 Spalding Caye 6 M 3,38 
1.3, 1989 Vickers Iskind North = 7 - 3,31* 
64, 1993 Thin SE Horse Rock 8 F 6.08 


42k SE 
Stamford Hill 


YA pous ¥ 
12.4,1992 y MAIS 


Miisquir Comments 

ho, 

M5753 

Mol It) seen ulive before stranding 
50009 decomposed, not collected 
MI4580 decomposed 

M14465 washed up deud 

MI5024+ very fresh 

M15374 —syery decomposed 

M1I7362 washed up dead 

S0085 alive. very emaciated. returned to sea 


82 


TABLE 2. Sightings, including captures hut not sightings inumediately followed by strandings, of Caperea marginata in the 
Australasian region. Latitude and longinide given in dearees and minutes. Rel. is reliablitiy of tdentification (1 = 
certain, 2 = probable, 3 = doubtful). A =udult, S = subudult, J = juvenile. 


Date Location Lat. ! Habitat No. Rel. Size Comments Rel, 
Long. 
1.1874 north end 46 508 - - \ S captured amongst 8 
Stewart 1, NZ 168 00 B black fish , 
911959 Bruny Island. Tas 43 178 bay, in - | S killed by fishers 9 
147 18 EF 2-3m water 
7.1960 Port Lincoln, SA - protected bay 2 2 A.J ne description of LO 
animals 
4.1980 ~ 50 om SE 38 205 open oceun 5 | observed from I 
Cape Howe. NSW 150 20 E olf shell ship 
4.) 985 Svela Seamount, 43508 open ocean, many 2/3 - feeding. no 12 
120 nm SE Tas 15022 BE over sea mount description of 
animals 
28.11.1986. Portland, Vie. 38218 sahdy, 1 | S shipping harbour 13 
5.2, 1987 141 36E — protected bay possibly feeding 
1.1489 Spalding Cove. 34475 shallow, 2 e A.J Vig. 4 ref. no, 2 this 
SA 13558 FE protected hay. stucly 
sengerasses 
19.10.1990 Cockburn Sound, = 32 105 offshore | A photos surgest B. Parker 
WA IIS44E feeding behaviour — pers. 
comm, 
26.11.1992 420 nm S Cape 41375 pen sea ~80) | A three groups near 6 
Leeuwin WA 11S 38 E euch other 
4.1.1996 Spalding Cove, 34478 shallow. 2 | A.J stayed in area at this 
SA 13558 E — protected bay. least two days, study 
SCUTUASses Fig. ref. no. 3 
77.1996 Port Lincoln, SA 34.44 8 edge of 1 2 J Fig. 4 rel. no. 4 this 
13600E Spencer Gulf study 
References 


‘Kemper, C. M. & Ling, J. K. (1991) Trans. R. Soe. 3. ‘Kemper, C. M. /hid.accepted), 

Aust. 115, 37-S2. ‘Hector, J. (1875) Trans. Proce. N. %. Inst. 7, 251-265, 
Pavey, C. R. (1992) Aust. Mammal, 15, 1-6. CCiile: BROT Ry-Phe, Pais ane * $2,780. 
‘Davies, J. L. & Guiler, E. R. (1957) Proc, Zool. Sov i tuiler, E.R. (1978) Pap, Proc. R. Soc. Tasm, 112. TSi- 
Lond, 129, 579-589. 213. 

‘Ross, G. J. Bo. Best, PB. & Donnelly, B. G. (1975) J “Hale, H. M. (64) Ree. S.A. Mus. 14, 334-419. 


Fish. Res. Board Can, 32, 1005-1017. "'Cawthorne, M. W. (1981) Rep. Int. Whal. Conn. 31, 
“Sekiguchi, K., Best, P. B. & Bozena, Z. K. (1992) Mar, SC/32/Progress Report N, Z.. 201, 

Mam, Sci. 8. 288-293. ‘ . ' ; so a oy 

‘Matsuoka, K., Fujise, Y. & Pastene L. (1996) (bid, 12, Blaber, 8. J. M. (1986) Emu 86, 239-24, 
594-597. “Arnold, A. (1987) Aust. Nal. Hist. 22. 266-270. 


CATHERINE KEMPER, South Australian Museum, North Terrace Adelaide S. Aust. 5000, JOHN DUTTON, 23 Mitcham 
Aye. Lower Mitcham S. Aust. 5062, BRIAN FOSTER, 51 Britainnia Road Nairne 8, Aust. 5252 and ROBERT McGUIRE, 
83 Veran Terrace Port Lincoln S. Aust. 5606. 


VOL. 121, PARTS 3 & 4 


28 NOVEM 


BER, 1997 


Transactions of the 


Royal Society of South 


Australia 


Incorporated 


Contents. 


Brief Commu. 


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 - - - 

Vaucher, C. & Beveridge, I. New species of Potorolepis Spasskii (Cestoda : 
Hymenolepididae) parasitic in pag hat soe from 
New Guinea - - - - - 

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

Watson, G. F. & Gerhardt, H. C. The breading Rivioes anti qavetiscenent cat 
of Litoria splendida Tyler, Davies & Martin - = 

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

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

Olsen, A. M. An intensive monitoring study of two wetlands of the River 
Murray in South Australia; 2 eee a 
and cyanobacteria concentrations - —- - 

Kolesik, P., Whittemore, R. & Stace, H. M. Asphondylia pes idis, anew 
species of Cecidomyiidae (Diptera) inducing fruit galls on 
Anthocercis littorea (Solanaceae) in Western Australia -  - 

nications: 

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

Smales, L. R. The status of Cyclostrongylus medioannulatus Johnston & 
Mawson, 1940 - - - - - - - - - 


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


83 


95 


137 


147 


157 


165 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 121, PART 3 


THE AGE OF THE POORAKA FORMATION AND ITS 
IMPLICATIONS, WITH SOME PRELIMINARY RESULTS 
FROM LUMINESCENCE DATING 


By R. P. BOURMAN*, P. MARTINAITIST, J. R. PRESCOTT? & A. P. BELPERIO£ 


Summary 


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 (1.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 fluyial 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. 


Fresnel if de Raval Saetery ol S dase (1997), PEGA) BA OL 


THE AGE Ob THE POORAKA FORMATION AND ITS IMPLICATIONS, WITH SOME 
PRELIMINARY RESULTS FROM LUMINESCENCE DATING 


by RL PR BourmMan , P. Martinarns', J. R, Prescorr & A. PB Brrperny! 


Summary 


Bouin, ROP. MARINAS, By Prescorl J, Ry & BbipeRion A, P (L997) The age of the Poorake Pormanon 
and its plications, with some preliniinaey resulis from luminescence dating, Trai AL Sey Ao Att D2ECA), 83 


4, 28 November, 1997, 


SmMighiphic relianships, supporwed hy liminescenve dling. sia@@est thatthe Poorake Rocnation spas alae 
vrewer (ime interval than previously recognised on the basis of ridiocarboo dating ant stratigraphic analysis af 
discrete sedentary sections. extends back is fir as the Last Tnterghieiil Re-evaluarion of the niuhecar bor 
ues That idicute an titerstudhtl age (ie, Oxygen (sotope Stige 3:45 10 30 ka BR) forthe sediments is required, 
Miernittively, a considerable time interval for deposition af the Pooraka Formation would necessitate that the 
wit he ditehronois weross the tindseape An age extending biek to the Last Niterglacial (Oxygen fsorope 
Substige 50.6, 125 ka BP) would provide the appropriate palieo-climates and paliea-enviconiients for (luyisl 
sedimentation, The revised age has impligaions for kindseape evolution, aehiwolowical and palaeomugnetic 
prospectiog as Wella the antiquity of thy Diprajeden inthe Adelitide area. 


Ky Wokbs: Poorake Portndtion, Plestucene stratiraphy, Last Witerglieial, Wimineseence dutine 


Introduction 


Linge areas of the Adelie Plans are underlain hy 
the Pooraka Pormation (Firman (67, (969, Callen 
eral, M95: Sheard & Bowman 1996) w reddish: 
brown coloured Pleistocene ahivial deposi with 
weakly developed calcareous pedogenic horivens 
that underlies river terraces and alluvial fans, Plies 
Unit is also Widespread beyond the Adgliide Plains. 
extending On to the Fleurieu Peninsula ane inte the 
mid-narth wl the state, where it Minky the Plinders 
whl Giwler Ranges. 

The red-coloured sediments that comprise the 
Widespread Pooraka Pormation have been aseribed 
different names by workers over time and indifferent 
areas, bor exaniple, they were originally referred to 
as the “nanimaliterots drift by Tate (1879) becuse 
skelotul remains of the extinet. giant marsupial, 
Vipratoden apitiin, were recoyered from thenr in 
areas Cothe west al the city of Adelaide, Ward (1966) 
relerred) to the sediments as the Christies Beneh 
Formation on the Noarlunga and Willunga sub- 
busing, Twrlule (1968) tumed them the Klemzig 
Sand during his investigation of the terraces of the 


Sehool or Bnviromnenal and Keeredion Mawar, (iene vl 
Eicineeriiye aid Lanwroniient: Ubpwersity ot Southey Avatedics 
Warrenty Ral Tlie bawels So Apst, S005 

Dypatareat ed Pliysies vil Mathematical Miysies. Pl biivrgity 
cb Nolo lide nist, SOS 
Vonvierly Mines iid Eneniy doxgurees Soh Australian PO) Hos 
19) Lastwrmd 5, Agst 3004, Cuerenthy Minti Gold, ta 
Cohalatore Ne Peron S. Atist Sue 4 

Bev ADA RP OPYOU) Ladforn Siadies near Vietor Phorbour Bay 
(Homes thesis. The Cntogrsaey ou Adehttules ciapiity 


River ‘Torrens, and Bourn (1968, 19694) referred 
to them as the Adie Clay where they Mank the 
Rivers Hindiaesh and Taman in the Vietor Harbor 
aren (Localities shown on Pig. 1), The red-coloured 
Alu bears consistent stratigraphic relutignships 
food younger, greyeblivk alluviuin | Waldeita 
Formation of Ward (1966), Wilkerville Sand of 
‘Twidiile (1968) und the Breckan Sand of Boutros 
(1968) ) whieh forms Tower terraees and Moodplains 
set within valleys carved out ol the red allivinina of 
the Pooraka Formation, Bstuuine shells collected 
from within the Walder Pormation in the lower 
reaches of the Onkaparinga River (Bourman 1972) 
retumed a ridigearbon uve of 4.580 + 160 veurs BAP. 
(Bourman 1979). During this Middle Holocene time 
the lower, dearecoustil reaches OF any streams were 
shillow, shellered estuaries as revealed by the 
presence of fossiiferous narine deposits at depth up 
valley, This interpretition ts Supported by evidence 
Worn a locality seyerul Kilometres rom the coast on 
the lower Onkaparinga River where ain uboriginal 
kitehen midden Containing estuarine shells citec at 
S820 + 90 yours B.PL(N.B, ‘Tindile pers, conom. in 
Twidale ey a 1967) is sired ona well dramed sand 
dline site at 20 mast ane adjaeentta the former, more 
eNLOMAIVE SULLY, 


Ages useribed io the Pooraka Formation 
and its equivalents 


The Pootuka Formaion tas generally been 
ascribed to the Late Pleistocene, with nest 
nuroerical ages, based on radioearben dating. falling 
within the ange nf S0.000-20,000 veurs (see review 


K+ 


R. BP. BOURMAN, P. MARTINAITIS, J. R. PRESCOTT & A. P. BELPERIO 


9 


AUSTRALIA 


SOUTH 
AUSTRALIA 


oe 


AB 


ADELAIDE \. 


d 


Beach /* 


7 


Hill 


Pym. 
“Ue p> . 

Weymouth 
Hill 


f 


Cape 
Jervis 


Fleuriel 


( 


Waitp 
Bea 


tei 


“~ Tunkalllla 
mi Beach 


We RIVE 


Hallett 
Cove 


Embayrnent / 


ft 
ln Selly 
Sellicks \. o~ 


Myponga gay 


XN 


gawler River ee a 


/ 
/ 
f 


Oe 
pata SLA a 


Mt- 
Pleasant 


A: 


3 
. 
v 
rT 542 


MAL 
Gawler 


: / 
~ ADELAIDE 
|_| / 


Mt, 
726 Loatty 
a 


£ 


7 
os 
rf 


vf Noarlunga 
Embayment 
An nse 


ie 


fe: 
Rone 
en 


arind® 


Willunga 


~ 


Mt 


a 

oN Currency C4 
Ty See SS 

vA Hindmarsh 

7 Is. > ae 


eB 


i Victor 
} & Harbor 


inga” 
ch 


Pd 


Encounter Bay 


Study Sites 


OS ory Cc 


“ River Torrens, Walkerville 


Bie 
reek, Pooraka 


Ki Hindmarsh River., Victor Harbor 


20 30 


Scale in kilometres 


Fig. 1, Location map of sites. 


AGE OF POORAKA FORMATION 85 


in Callen ev af. 1995). There has not been universal 
agreement on to which part of the Late Pleistocene 
the sediments should be allocated. Twidale (1968) 
assigned the Klemzig Sand to the Late Pleistocene 
and demonstrated 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 fllstrath terrace cut into the Adare Clay, 
in the Hindmarsh River, was dated at approximately 
12,600 years BP (Gak - 2356) (Gill & Bourman 
1972). Firman (1969) and Daily ef al. (1976) also 
considered the Pooraka Formation to be of Late 
Pleistocene age, but younger than the Anadara- 
bearing Glanville Formation, which is now widely 
accepted to be of last interglacial age (¢. 125,000 yr 
BP; Murray-Wallace ef al. 1988; Murray-Wallace & 
Belperio 1991; Murray-Wallace 1995; Belperio ef al. 
1995), Confident separation between Late Pleistocene 
Pooraka Formation and earlier Pleistocene alluvial 
sediments is casily achieved in the coastal zone 
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 
(1966) assigned the Christies Beach Formation to the 
Last Interglacial as he considered that the surface on 
it was graded to the last interglacial shoreline (his 
Epimonasterian high sea level) at approximately + 3 
m above present sea level. However, at that time the 
Last Interglacial was thought to be considerably 
younger than the present 125,000 years BP. 

From the Dry Creek alluvial fan, Willams (1969) 
described reddish-brown clay overlying older grey- 
green and red mottled clay, now known to be the 
Keswick Clay (Sheard & Bowman 1987a,b; M. 
Sheard pers. comm. 1997), Williams (1969) also 
noted a calcareous red-brown earth developed within 


the sediments containing nodules and cylindroids of 
pedogenic 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 detrital in origin and thus was regarded as a 
rehable representation of the time of deposition. 
However, if the carbon were detrital, its age should 
predate the lime of sedimentation, which would 


mike the ''C date somewhat older than the time of 


deposition, The date was taken to indicate a last 
glacial (Wiirm) age for the sediments. Further 
radiocarbon dates supporting a last glacial (Wiirm) 


Guppy, DA, (1943) Geological reconnaissance of part of the 
Hundreds of Encounter Bay and Goolwa, BSe (Hons) thesis, The 
University of Adelaide (unpub.). 


age were derived from a study of 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. 


Stratigraphic observations 

Critical evidence concerning the age of the 
Pooraka Formation occurs at 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!) established that red-coloured 
alluvium forms fill-top terraces along the Inman and 
Hindmarsh Rivers and grades to a shoreline at ¢. + 6 
m ubove present sea level. The age of the shoreline is 
considered to be crucial with respect to the age of the 
terraces and the sediments which underhe them. 
Twelve species of shells have been identitied from 
this shoreline deposit (Guppy 19432) and it is 
significant that they contain the sub-fossil Anadara 
trapesia. Initially, Bourman (1968, 1969!) followed 
Sprigg (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 the above dates are compatible with 
those of Williams (1969, 1973) they are 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 of 
radiocarbon from modern activity. Thus, materials 
with an infinite age by radiocarbon dating techniques 
may yield an apparent age of 37 ka due to the 
incorporation of 1% MC with aw modern activity 
(Gupta & Polach 1985). Gill (1974) checked 
radiocarbon dates of 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 ef al. (1984). 

Not only do Anadera shells occur at the + 6 m 
shoreline at the coast at Victor Harbor, but extremely 
large Anadara shells were recovered trom a sewer 
trench c. 1.6 km upstream at a depth of 4m below the 
surface. within the Pooraka Formation equivalent 
unit and at the same absolute elevation of 6 mas al 
the shoreline (Fig. 2). A drilling programme (CSIRO 
Soils Division) further revealed the intimate 
association of Anadara shells with the Pooraka 
Formation equivalent unit, demonstrating that here 
the Anadara, last interglacial deposits (Glanville 


RG RP BOURMAK. FP MARTINATTIS. J. R. PRESCOTT & 4, Py BELPLERIO 


Formation) and the Pooruka Formation are a last interhienal age, 
intercalated coastal and terrestrial equivalents. The 


Anadara shells have been dated both by Uranium. Methods 
Thorium techniques (100.0006 (50,0000 yeuns BP) 
and by uming-acid racemisation studies (Kimber & The stratigraphic rehutionships between lust 


Milnes 1984). whieh provided results Consistent with interglacial molluses and the Pooraka Portion 


_ Anadara Shells («Glanville Formation} 


15 
WwW = 
7 — ~ 
a S - > “ 
Y— > wt =~ = 
Ee 
j 
Cc o+-———— ---— —— eS ——— 
= Soa Level 
3 . Pooraka Farmation —— 
ee 
ii 10 —— 
—— yo ~~—S—CS~;S Pormian Sediments 
1S a 
Xt —————— 
—— 
a2 
17] 
VOODIN Sen 


fio. 2. Sketch section showing the interfingeriny relationships af the Gkiille Formation aad the Pooruka Pornnaion i the 
lawer Hindmursh River at Victor Harbor. 


Fi 7 PHotosaph of 6m high ever bluff cut in Pooraka Pormution sediments downstreu of the Bridgeway Horelon Dry 
Creek, Powruke, Masinuin seetion exposed « 7m Red brown earth with assechkted calcium carbonate zones in upper 
purl of selon. AL the busc al the secvion The Pooraka Formation rests unconlormibly on ai older Plesitacene Unit tat 
Chisely resembles the Tartiipa Formation af Wank ted; bat whieh Sheard & Bown CM 87ah) consider is mure hkebs 
Ibe the Keswick Chay, 


AGE OF POORAKA FORMATION 87 


equivalent, near the coast, at Victor Harbor provided 
strong indications that the Pooraka Formation might 
extend back 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 for sample collection for luminescence 
dating was the Dry Creek (Pooraka) locality where 
Williams (1969) collected samples of detrital 
carbonised wood and carbonate for radiocarbon 
dating (Fig. 3). Unfortunately, the steep river bluff al 
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 haye suffered 
from human modification and landscaping of the 
former eroding river bluffs 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 mand 7.5 m (for Pooraka) and 4.5 m, 
5.5 m, 6.0 m, 7.5 m, 9.0 m and 9.3m (for 
Walkerville). The drilling site at Pooraka was located 
7.74 m above the base of Dry Creek and the second 
site was 11.68 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 PKIS from a depth of 3.5 m is close to the 
level from which Williams (1969) collected detrital 
carbon for 4C dating. 


Luminescence dating methods 

Three methods for luminescence dating (LD) of 
the sediments were used: selective bleach 
thermoluminescence (TL) of coarse-grain quartz 
(Prescott & Mojarrabi 1993) and green light 
stimulated-luminescence (GLSL) of both coarse 
grain quartz and of fine grain separates (Aitken 
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 ina generally colluvial 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, WV1S/7.5 
and WYV1S/9.3. Such runs are designed to assess 
whether the sample is likely to be datable and, if so, 


TABLE Lo Summary of collected data for Pooraka and Walkerville samples. 
SAMPLE PPMU PPM Th PPM U PPM Th = =PPMU Sct. PPM Th Sct. =“%@K XRS TK Set GUO 
DNA DNA 

PKIS/3,5 1540.6 7042 1484011 7.23405 1.744017 7.124029 1.08+0.03 0864002 6840.07 
PR2S/4.5 Ll+03 7841 1264011 731404 1.754017 7.684029  0.8940.03 0.82 +0.02 3.7 +0.06 
PK2S/4,8 Ll+05 6341 E1O+010 6.24404 1.344016 6.76+0.28  0.894+0.03  0.9140.03  4.4+0.07 
PKIS/7,5 12406 7142 1224010 691404 1.744017 9.694032 1.204004 1134003 1240.07 
WVISM.S LR 403 1241 LSP 4012 17405  1.8940.24 1254044 1854006 1.604004 8.8 40.07 
WVISIS.S 16406 1442 1934013 126406 2.334024 1054042 L8P+0.05 1324004 7.5 40.07 
WVIS/,0 19405 Wel 1864012 103406 2474025 1214048  15940.05 1544004 8.3 +0.07 
WVIS/TS 17405 142 1664012 1840.6 2.664026 IL1+047 1.71 40.05  149+0.04 15.7 + 0.07 
WVISMO T8405 IS4t LI7+012 134406 2.954029 1344050 2.194007 171 +0.04 18.0 40.07 
WVIS9.3 12405 1542 L88 +013 13.0406 2.954029 1344050 1.984006 1.714004 16.4 +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. ppin U Sct, ppm Th Sct & %K Sct were derived from the on-site gamma ray scintillometry data. 


88 R. P, BOURMAN. P. MARTINAITIS, J. R. PRESCOTT & A. P, BELPERIO 


to give a limited-accuracy estimate of the acquired 
luminescence and the sensitivity to radiation, and 
hence plan the schedule for a complete dating 
procedure. A pilot run consists of eight dises: half of 


these are bleached, after which two discs euch of 


bleached and unbleached are given a radiation dose 
of 60 Gy. Por all samples except PK1IS/3.5. the 
shallowest of the Pooraka samples, the TL was 
clearly saturated and no further work on them was 
justified. Although PKIS/3.S was approaching 
saturation, a full dating procedure was carned out tor 
both TL and GLSL. 

Such procedures give the Equivalent Dose, D.. a 
measure in grays of the energy absorbed by the 


3.0 


4 


9 


Counts x10 


1.0 


100 200 300 400 500 600 
Temperature (9C) 


Fig. 4a. Glow curves for sample PK 1LS/3.5, The figures next 
to each curve indicate the dose in Gy, 


Counts x10% 


0 10 20 30 40 50 60. 70 80 
Shine Time (in seconds) 


Vig. 4b. GLSL. shine-down curves for sample PKIS/3.5. 
The figures next to each curve indicate the dose in Gy, 


sample from radiation in the environment since il 
was last reset to zero. The age of the sample is found 
by dividing the equivalent dose by the dose rate in 
gray per kiloyear (Gy ka!). 


Equivalent doses, dose rates and ages 

TL glow curves ure shown In Figure 4a, shine- 
down curves in Figure 4b and corresponding dose 
curyes are shown in Figures Sa, b respectively, None 
of the curves is sealed. [It is evident that the dose 
curve of quartz (Fig. 5a) is close to saturation but that 
the growth curve for GLSL on fine grains (Fig. Sb) 
has a different shape and the curve continues to rise 
quasi-linearly for high doses. This is because the 


Counts «10° 


0 0.0 200 400 600 800 
Dose Gy 


Fig, Sa. Th growth curve for sample PKAS/3.9 for the [0 
interval at 305° C. For Figures Sa and b, the curves are 
fitted by the “Australian slide” method: the (natural + 
dose) points are shown by crosses: the (bleached + dose) 
points are shown by cireles. There is an apparent 
sensitivity change for the Tl bleached curve bul the 
scaling factor does nat differ significantly from unity. 


Counts x1o* 


9 200 400 600 800 
Dose Gy 


Fiz. 5b. GLSL growth curve for sample PRIS/3.5 
integrated over the first LOO s_ 


AGE OF POORAKA FORMATION Mel 


Panay 2, Bawls doses, dese ratey anlages far sample PR LS/35. 


COARSF GRAINED QUARTZ 


FINE GRAINS 


selective bleaeh Th GLSL GLSL IRSI 
1), Gy 240+ 58 JK & 44 24445 > 150 
dose mite TSAC/DNALRRS SCINT TSAC/DNAIARS SCINT 
Gy ku (90 #1, LG 1b 2544 (1,12 2542011 
Weighted mec LUT © 1.03 2.54 = 1).08 
ave kit 126 + 29 LOS + 22 llh+6 > 10 


undifferentiated fine grafts consist of a mixture of 
minerals in which quartz and feldspar ure donmnant, 
Thus although the quite component saturates, the 
feldspar component does not, Equivalent dose values 
from all three methods appear in the first tine of 


Vuble 2. Ih will be noted that the two methods of 


finding D, give different values; this is characteristic 
ol the methods, 

Two distinet methods for finding ihe dose rules 
were used (described i the appendix). the aim being 
iO get two independent values for euch sample, to 
inprove the statistical precision and ws cheek on 
(he presence of radioactive disequilibrium (Prescott 
& Hutton 1995). "The values obtained are ineluded in 
Table 2. As with D., dose rates differ for course: and 
fine grains. The values are in excellent mufial 
agreement and this shows that radiouctive 
disequilibrium is absent, 

The weighted mean dose rates are shown on ling 3: 
line 4 shows the ages derived from the D. using the 
age equation from the uppendix. The weighted mean, 
116+ 6 ku is dominated by the GLSL determination 
for Hie grains. 


Comment 

Allowing for stanstical Mtting uncertainties. all 
three ares are in good agreement. Bearing in mind 
(hal they are bused on different physical processes, 
allo? which assume that the luminescence sign was 
set to vero in the past it shows that this wis very 
likely the case and that the age being determined is 
indeed the tune of deposition, Th it is not so. the 
uppurent ages will be too large. 

As already inentioned, the dose growth curve of 
Figure Sa, which is lor quartz TL. shows: that the 
luminescence is close fo saturation. For this reason 
the estimate Of relative Gneertainty given by the 
ialysis is large. This is also true for GLSL, of quarts. 


For GLSL of fine grains, on the other hand, the 
presence of the Ieldspar component allows Dy to be 
found with significantly better precision, 

The best estimate of the the of deposition of 
sumple PKIS/3.5 is the weighted mewn age, 116 4 
6 ka. In facet. this is dominated by the GLSL 
determination on fine grams. [is probably best to 
reaard the quartz determinations by selective bleach 
und GLSE, ws being supporuve of the GLSL fine 
grain age. Al the two standard deviation level the age 
exvecds LO ka, Consistent with these numbers. the 
saturation of the luminescence of all the other 
samples shows that they we as old or older than (his, 
Therefore, although only one sample has yielded a 
numerical age, there js enough evidence ly establish 
a lower limit to the age of the formation and itis in 
support Of the geomorphological and sedi- 
menlolugical evidence, 

In addition to the three methods alreiuly desembed, 
wilta-red stimulated luminescence (IRSL) of fine 
wrains (Wintle 1994) was tned. IRSL uses only 
feldspars and is less susceptible to saturation, 
although if may be subject to Tongeterm fading. 
Although (he growth curve was similiar im shiape ta 
the GLSL curve of Figure Sa, the stiustical fitting 
procedures did net salisly our eritena Por an 
accepluble D.. beyond showing that it was greater 
than LOO ka. Whether this is due to the sample or the 
methodology remains to he determined. However, 
the result is sufficiently cocouraging to suggest that 
it may be possible to find ages from the lower levels 
i Pooraka und for Walkerville where quitrtz methods 
were unsuccesstul: 


Discussion 
The location of the date derived from the upper 
section of the Pooraka Formation at a depth of 3.5 m 
below {he surface 1s i complete aecordinee with the 


Ot) RT ROURMAN. POMARTINAPTIS. LR. PRESCOTT & A. PL REL PERIO 


numerical age oblaned oh 176 + 6 ka avilh the Last 
Inlerglacal ranging from approximately (32-108 ke 
tsee Chen ero. Yt; Lumbeck & Nukuda 1902) Zhu 
et dil (99S: Sarling er al 1995) Ejsentner ep af 
96). The GLSL technique olfers the possibility: of 
vbliining & Turninescence date for lower seetions al 
the Poorske Pormution thereby fucdiratiig the 
calculation of the rates of sedimentuion during the 
Taint Imerghiciah itmay be that sedimenhitton duping 
warmer, Weller, LIStTMberglaenu UimTes was quite rapid 
and this technique offers the opportunity to lest vis 
TV 

‘These resulls Sugeest a far greater age range for the 
Poorgkii Formation thin has previotishy been 
recoemsed, In the past, the gee of the Poorakit 
formation fas heen poorly consiained we abou! 20- 
50 ku BP pussibly heeause ol the now-known 
Hivikitions of padinearhon dating. Furthermore, 7 
Sfould be poled ifjab other methods such as Th, 
GLASS, and IRS were not available when many vt 
the madiues bon dates were cured but. 

Many observutions and cores indicate that, an 
places, the Poorthie Farmition appedrs to be younet 
(hun ihe Just inberetacial Cilunyille Poroatien, 
especially in’ subsiling areas. For example. 
Ludhrook (1970) nthe Port Adelaide itrea described 
Pooraka Formation overlying calented Glanville 
Formation, whieh th Wut overlay Hindmarsh Clay. 
Relperin and Rive (WY89) shawed thal ol 62 cores 


from) the Gillman area. Eb record the sequence ul 


Holueene overlying calercted Glunville lormation 
Without Poorika Formation, and 5 record Halocem: 
overlying Pooritku whieh overlies calercted Glanville 
Vormiation, The calereted Surfaces itre inlerpreteel as 
pedogenic features developed during subuactial 
exposure and wuld have dormed prior to the 
deposition of the overlying sediments of the Pooraka 
Fornmdton, In contrast, four of the cores reeurd at 
sequence Of Holocene/Pooruka/Glanyville Porniation. 
withour 2 well-developed culerete oi the Glanville 
Formation, and one revords intermixcig of Pooruku 
und Glinyille. These Just cores miry ihdieate coeval 
Canville and Pooraka sedimentation or litle time 
Break between the (Wa, 

Extensive work by Sheard & Bowriun (1996), 
which involved drilling cores to depths of FO mr it 
7H sites over the Adeliide Plains. intersected 
Pooruka Pormation in 80 cores. This work 
corroborites the Findings of Belperio & Rive (1989) 
for the voustal zone. More landward sites iniieated 
the sequences to be: Holoeene/Pooruka/Keswick 
Clay/Homdinarsh Clay and Holoverie/Poorakas 


Woon. Pi bat de Kose Pin Ply bid (18s) Appendix 3 
Ceatechiea) Investaatiens lu Holiival Shores ti Banildlonstone 
Thomiboioh Giles Canaan) Werke dun Terry Wilark Bepiart ty 
Urhii Projects Aviheriiy at SA. Gunputts,) 


Keawiek Cluy/Tertiary sands orAdehddean rocks, At 
the MM) sites intersecting Pouraka Purnddon 
sediments. ‘Sheurd d& Brym (1996) rarely 
eXpefienecd diffloulties a adenufying Pooraks 
Formation from the overlying and underlying 
materiily (M Shearl pers. comm. 1997), 

Recent drilling work by Woodhead er al (Yas) 
for Baulderstone Hormibrook in the Haldfist Shares 
(Glenele Patawalonga Redevelopment urea) has 
demonstriee that depostis of Poorake Formation 
2-4 om thick oeene well offshore frum the 
present eoust line. These deposits aire detavhei 
from more landward deposits either by vones 
Of fol-deposiuon or by carly Holocene erosion, 
Variuhle stratigraphic relationships are aipparent in 
this locality. Por exaniple. the tallow ing relariuinshipes 
Of Holovene/Pooruka/Glanvilles/Hindivirsh Chi. 
Holocene/Ghinvilley Hindinursh Chay ot Floloeenes 
Fulham Sand/Pooruka/Ghuryile/Minginarsh Clay! 
oveur within an S00 mM east-west section. 

Ie omuy be that the Pooruka Porniiion was 
Uepasiled dver i considerable period Of time This 
view 1 also supported by the oecurrence of 
palacosols within. Pooraka Pormation equivalent 
sediments is exposed it Sellicks Creck and Cobblers 
Creek. The luminescence data indicaie an ape 
approximately coincident with the Last Miler heh 
(125 ku BP), bul the resalts atany one place may be 
joflucneed by the terresirial/maripe interactions. For 
extmple. there is mut un achive Supply of alliwinin 
Othe coastline then marine deposition, exposure andl 
calerete formation will doittate, fh plices 
subsequently mintied by Pooraka or Holocene 
sediments. Th omay be fi only where there tsa 
sulfivient supply of terrestrial sediments to the 
coustline, sueh us where streams deboucl at the 
eoustine Th PelaGyely coystramed valleys (ee, 
Hindmarsh River at Vielar Harbor), that the 
Inrertinvering vhunacteristies af the two sediments 
‘an be demonstrated. Et fsalso possible that Puorakat 
Formation sediments Jo not reueh the position of the 
Glanville Bormation uniibaflerit has been culereted, 
It should be noted that touch of the Pooraku 
Formation bas been deposited infind, well beyond 
the elevation and plinimetric position of the last 
inlerlaciel shoreline, so thar in these situations (here 
Ison clear siraneraphie rehkiionship between the two 
uns 


hnplicationy vf the Mraruka Loriiation exetidiin 
hack te the Last datercleavtel 

Wo the Trterpretation Uhl the Poorika Pormutien 
extends back to the Last Interelacnal is correct, wpe 
evaluation of rhe radiowarbon aves thar indicate 
glacial or (nterstadial ages for the sediments 14 
required. Alternatively, the canflictuay results tiny 
suguest that there could be sediments, which 


AGE OF POOR AKA FORMATION 9] 


athoueh appearing similis Occupy a rune oF ages, 

Ati number of lowations, the Pouraka Formation 
his been noted to averlic the Glanville Bornution 
with « yariably developed calercle, Sugeesting 
surface exposure prior lo burial by younger Poortkit 
Fotmauon sediments, Elsewhere. such as in the 
Jower Hindnirsh Rivet. where there is uur interplay 
of coevail oustil anc terrestrial events. the 
strutipriphic relationships between We Pooraka and 
Glanville Formations may simply represent a hicies 
chunge and not ow geolagicul sucvession A 
considerable lime interval nay be requied for the 
deposition, OF the Poopake Pormition so that tts age 
conkl he diuchronie, with deposiion oecoriine 
during the rise in sea evel up to and beyond Lhat ef 
the hist imberchienl level, 

A lust intereletil awe privides appropriaie pulseu- 
chime and pulaeossea level conditions for 
exphiiniog the distribution oF the Pooruku Vormation, 
Close to the coast the unit rektes to a shoreline 
higher than the modern one, while inland the wetter. 
Woerelacial conditions would have hivourcel 
dwvradation of sediments w-opposed fo dpier ehicial 
conditions that would have (actlilaled dissecuen, 
This timing would alse ensure sufficient tine both 
for the build up of extensive deposits oF the Poerikit 
Portion aver large aveas ieluding much of the 
city of Adelaide. jind time during elicials and 
interstudials for erosion oF the Pooraka Parination to 
develop the extensive terruve system al the River 
Torrens (Twidile |968). 

A last intengiieial age for (he Pooraka Formation is 
al signiicunce Nrarchivalogicul prospecting. AL the 
Present Uine there ts debate concerning the untiyuiby 
of Humiins on the Australi continent. Substurrail 
chaes in Vesetition related to Aborginal burning 
pruchees faye been ilerpreted vis sesulling Yom 
human pmpach ural used bo infer the arrival of people 
in Australias lemme ics 140.000 yours (Rershaw 
eek WS. Kershuw 1993, 190d. 1995). These 
Claims have Beeb Questioned hy: varinus workers (eo 
Andersen Tad: Hope 1994; White 199d) Webb 
19S). Tile Pouraka Formation is of bist tiberetactal 
He ny present ad prospecting opportunity to best 
i Aboriginal colonisadon had occurred in southern 
Auistalia prive to 125.000 years BP ducing the Gime 
ofthe penullanate ehiekil law sea level, 

A minot palacomugnetic event, the so-culled 
“Blake Event, his heen nle tied ro the northern 
hemisphere irr loess sequences (Llpneho ed af, LY95) 
“eapprosinnitely [20 ka. Given that the generat age 
of the Pooraky Fornvarion ts almost certainty of last 
iMerelienl uve, and that the upper part ob the 
formation has been useribed vu GEST aye of blo = 6 
kit Uhere may be opportunities ta identity the Blake 
event in the southern hemisphere, 

A last Tnlerg ictal ave lor the Pooraka Formate 


has implications fev the ahquiyy ob the 2 peatertent 
in the Adelaide urea, Diprotodonlid remains. have 
been revurded well baek into Tertiary straui in the 
North Flinders-Cullabonna Phiins areas (Callen & 
Tedford 1976; Pledge & Tedford 1990; M. J. Sheard 
pers. comm. 1997.) but Hot se far, in the Adelaide 
vrea. Discoveries of Dipraloden remains have been 
mude i the Poorwku Formation (Twidale 1968) N, 
Pledge pers. comim, 1996) ab the Adelaide area und 
Winle the Diprarodon iray have predated and 
survived well beyond the age of this sedimentary 
unit, if was almost certainly roaming the swampy, 
iwprading Adelaide Phi some 125.000 years ago. 

The age has further implreations for landscape 
evolution as there appears ta have been a tajor 
erosional hiatus of some 120.000 years belween the 
deposition of the Pouraka Formation and the grey- 
black Wildeila Formation, which is oFimid-Holocene 
uve, There were ro sea levels hiuber than (he present 
in the iptervening period so that erosion would have 
dominated this interval of time. Alernitively, any 
sediments deposited during this (ime could: lve 
heen removed in bile stages of erosion. 


Canclusions 


The main conclusion from this work is that the 
Pooraka Formation must span a lar ereuter line 
period that previously recognised. probally 
extending as tar back as Lust Interglieiil dine te. 
125 ku, Th vppedrs that the tectonic ane 
environmental setuag. whieb influences the supply 
of terrestrial Sediments to the coasting, sale sireme 
Importance in interpreting the stlatighaphie 
relavonships between the Pooruka Formation ane the 
Glunville Formation. Mueh mure luminescence 
CIN Work Is Lequiied fo coystrai dr document the 
ie ranges in diffgrent setlings ae. in the Wietor 
Lhurbor Setong where the bWwo qe jotermived. tir 
contest to the Port Adelide Setting where a tia]or 
calerety pulaeosol separctles thei, We mi expect 
luminescenve ating: techniques to document wes 
from list interelicnl tines. possibhy through te 30- 
SO aL uf the quitrty erains are sufficienily reset. This 
offers The possibility of providing ae pore reliible 
dating tool than radineurbon techniques. whieh Sill 
Stiwele do prowile mean dates pasty 4d) ka. 

The red-brewe alluvial sediments, referred ba is 
Pooraka Formation and equivalents, are eopisidered 
to extenu back toot least dhe Last Inleralocil of ¢, 
125,000, vous BR This ts demonstrated by the 
Stratgfaphical relationships of the red-hrows 
sediments lo both younger and older sedinwits, the 
interdigitiing of dentonstrated last interghiectal 
miine depasits with then then grading tod + @ nM 
higher shoreline containing last interglacial marine 


ya R. PR BOURMAN, P, MARTINAITIS, J. R. PRESCOTT & A.) BRLPERIO 


fossils al Victor Farbor, and the fict that a higher sea 
level. together with associated warmer and wetter 
conditions would) favour ageradation. whereas 
colder, drier glacial conditions associated With lower 
seu levels would hive favoured erosion. AlLof these 
luclors Support a last interglacial aye. 

The luminescence data have demonstrated viability 
us un independent means of testing the hypothesis 
that the Pooruka Formation is of last interghieil age. 
Fluvial sediments present special problems for 
luminescence dating because the sediments being 
waler-deposited, may not hive been exposed to 
sunhieht for sufficiently long to zero the quart, 
grains, resulting in inherited saturation levels within 
the grains. A farther complication may be that in the 
Adehude area, the sediments may have not been 
transported sufficient distanees for the quartz grains 
to have been zeroed. Given these constraints, it is 
extremely gratilying Unt 1 was possible to achieve a 
rehuble lumineseence age for the Pooraka Formation 
at Dry Creek, offering the possibility of further 
advance in this area, 

An aim ot this projeck was (Oo establish the 
effectiveness Of luminescence as a citing fechmique 


for Quaternary: Mluviul sediments in order to resolve 
differing interpretations OF their ages and to (ucilitute 
correlation OF river terres jn different valley 
systems, Clearly, there is a peed for many further 
dates to be obtained from trver terrace and allawial 
fun deposits over wider-ranging afeas (o Verify the 
conclusions of the present work, However, the 
implications of un age foe the Pooraka Formation 
extending back to the Last Interglacial are so 
Sigmilicant that our preliminary results are presented 
here and provide the basis for further study. 


Acknowledgements 


Funding for the drilling of the Pooruka Founation 
und initial luminescence dating of the sediments wits 
provided by a University of South Australia Researeb 
Grant. The Th analyses were curried oul by 2. 
Marlinailis under (he supervision of Professor 
Prescott. We are grutehd ta Dr C. Murtiay-Wallace 
and Dr N, Alley for commenting on the text und te 
M, Bishop for assistinee with field work, We are 
upprecialive of the significant contributions of the 
referees Mr M, Sheard and De V, Gostin. 


Keferenves 


Atrbes, My ke (i904) Oprioul during: on nonespectilist 
feview. Ghat Geach. (Qual Set Rev) 1, 504-508, 
Anbrtson, At 1994) Comment on, Peter White's paper “Site 
H20 un the evidenee for early pecupation in Austialin’ 

Quel Mavtrnlavie V2, 0-31. 

Brrrecion aA. PB. SMinik B. Wo. Ponact. TL AL NP HOUR. 
COAG DeMasing. Dod PRescon dk Ro Pars bh Rd 
Gostin, VoOAL (TOR) Chronolowenl Studies of the 
Quaternary (iiring sediments of the northern Spencer 
Gull, South Answulian Man Geak @t, 265-296, 

& Ren Rl 
the Cillnuie development sie. Port Adeluice oxtuny 
South Aust, Dept Mines and Eneruy Report Book, 80/62, 

Vibes Warten, C, Wok Can, ET (19958) The 
Jost literehietil shoreline i seuthern  Ajatrulie 
Horphostralivraphic Vwairiations in a lemperite carbonate 
setting. Gueit, faternat 26, 7-19, 

Bovrvas, Be PO Cl968) Terrees ab the Time ane 
Hinciarsh Ravers Jing 7, 17-71 
— (N972) Some aspects of the Geamorphalogy af the 
Onkaparinga River, bil 9 LPP-143. 

(1979) Cieomorphological contibutions to couse 
nhikiwenment pp. BOB} J“ Procecdings Pocus ae our 
Southern Hlerikge” (Conservidtion Couneil of SA und 
Depuninent of Continuing Piueation, University of 
Adelaide) 

Bowman, Go& Hakvey, NL C1983) Radiocarbon deting 
firing shells in South Australin. Aden Aredieal. V7. 10% 
12, 


(VOR9) Stritiwraphic investigations af 


Canby, ROAM Trppoke. Leb lata Sew Chanaziie 
rock veils and depositional environments, Like Frome 
urea, South Austtalia, frei. A See 8. Aust HL | 25- 
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Appendix: Methodology for luminescence dating 


Quartz grains (90-120 um) were extracted as described in 
Huntley er al. (1993), Briefly, pure quartz of the right size 
was obtained by pretreating with HCI, followed by NaOH 
to break up clay aggregates, sieving, etching with 40% 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 HCl and NaOH extractions by settling 
from aqueous 0.01N NaOH, the 4-11 um fraction being 
retained. This was then deposited on aluminium discs from 
acetone suspension, about | mg per disc. Individual discs 
were 0.5 s short-shine normalised, 

As stated jn 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 & 
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+8) 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 ef al, 1993) and the data output is the equivalent 
dose D. expressed in grays. Such curves are known as “dose 
curves”. 

Two methods of dose rate determination were used: /n 
situ scintillometry (see e.g., Hutton & Prescott 1992) uses a 
sodium iodide scintillation crystal, 75 mm x 75 mm 
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 e7 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 for Th. These are the values shown in Table 


Table | 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!) 


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 fine grains extracted from the 
sample; and “dose rate” is determined from measurements 
in the field and/or laboratory. 


Transactions of the Royal Saciety of §. Aust. (1997), 121(3), 95-102. 


NEW SPECIES OF POTOROLEPIS SPASSKII (CESTODA : HYMENOLEPIDIDAE) 
PARASITIC IN DASYURID MARSUPIALS FROM NEW GUINEA 


by C, VAUCHER® & I. BEVERIDGE' 


Summary 


Vaucuer, C. & Beveripaer, L. (1997) New species of Pororolepis Spasskii (Cestoda: Hymenolepididae) parasitic 
in dasyurid marsupials from New Guinea. Trans. R. Sac. S. Aust, 121(3), 95-102, 28 November, 1997, 
Potorolepis aruensis 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 sae (0,27-0,42 mm in P arvensis, 0.14-0.25 mm in PB bradleyi) and arrangement 
of testes. Cestodes tentatively allocated to P aruensis were also found in Antechinus naso. Pororolepis woolleyae 
sp. nov., from the small intestine of Murexia longicaudata from Morobe Province, Papua-New Guinea, differs 
from all congeners in having longer rostellar hooks (163-182 um), The generic diagnosis is re-assessed as well 
as 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. 


Key Worps: Pororolepis, cestodes, Hymenolepididae, marsupials, Dasyuridae, New Guinea. 


Introduction 


Cestodes of the family Hymenolepididae Ariola, 
1899 are common parasites of birds, rodents and 
insectivores in most regions of the world (Czaplinski 
& Vaucher 1994). Vaucher ef al. (1984) 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 peramelid 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 Rodentolepis Spasskii, 1954, 
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 Spasskni (1994) as well as the host- 


Muséum d‘ Histoire Naturelle, Geneva Switzerland. 
Department of Veterinary Sciences, Universily of Melbourne 
Parkville Vie, 3052 


parasite relationships between sub-groups within 
Potorolepis and families of marsupial hosts, a 
relationship first suggested by Vaucher er al. (1984). 


Materials and Methods 


Cestodes from Myeoictis 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 (Pritchard & 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 ym in both longitudinal and 
transverse planes and stained with haematoxylin and 
eosin, 

Measurements are given in millimetres as the 
range, followed by the mean and the number of 
measurements made in parenthescs. Drawings were 
made using a drawing tube. 

All specimens studied have been deposited in the 
South Australian Museum, Adelaide (SAMA) or the 
Muséum d’ Histoire Naturelle, Geneva (MHNG). 

Host nomenclature follows Flannery (1995) and 


% C. VAUCHER & I. BEVERIDGE 


3, Me ey De ee ee 
2, Ply gy RSS Hes SOCAL PP MGV 


NEW CESTODES FROM DASYURID MARSUPIALS yy 


Sprall ef al (199). Tabulated morphological data 
were derived Irom Beveridge & Barker (1975). 
Vaucher erat (1984) and Jones & Anderson (1990), 
Host distributtan data were derived from Spratt er al. 
(1991). 


Potoralepis aruensis sp. wy. 
(PIGS 1-7) 


Tipexs Holotype fron srall intestine of Mivereriy 
melas wallace) Gray, 1858, Koboor ts., Aru goup (6° 
128 134° 32’E). Idan Jaya. 16-vi. 1993. coll, PA, 
Woolley, SAMA AHC 77877: paritypes, 23 whale 
mounls,-8 scolcees mounted in Berlese’s fluid. serial 
sections. SAMA AHC 27878-27905; 2 whole 
mounts MANG 23407 INVER: addilional specimens; 
numerous specimens 2. xi. 1992. SAMA AHC 
S0SX6-305K7, 3 specimens, 3. sil 1992. SAMA 
ATIC 30588: numerous specimens Twi }993, 
SAMA AHIC 30589, 


Material examined: Vrom  Myatens melas 
(Miijler.1840)) types. From Aarechinus — irever 
(lentink, HYPE): 5 speeniiens., Mt Kandi (7° 20'S. 
146° 41°). Papua-New Guinea, SAMA AHC 
27864-27876, SUSK4-30585. 


Deseriplion 

Based on types, Small cestodes up to 60 in length. 
Seotes globose. 0.52-0.65 (0.58.0 = 10) in diameter. 
Suckers sub-circular in Superficial views. unarmed: 
cup-shaped in section. with openings directed 
unteriarly, O.13-0.16 00, 15, 9= 10) & OL T5018 (O17. 
n= 10) Rostellum muscular, (180,26 (O21) x 
O.17-0.23 (0.21, n = 10): rostellar sae 0.26-0.38 
(0.30, n= 10) x 0.26-0.31 (0.28 n= 10). Hooks 16- 
IK (17. = [0), arranged th Single ring with broad, 
curved blades; slender handles prominent: blunt 
cuards often with irregular surface, core of hook 
blade hollow. Hooks (.128-0.147 (138. 0 = 10) 
jong. Neck vartable, 0.74-2.05 (1.26.0 = 10) lony. 
Segments craspedote: mature seginents much wider 
chan long, 0.06-0-17 (O10, 0 = 10) long x 0.93-1,.63 
(1.3). n= 10) wide: near gravid segments longer. but 
stivhtly narrower, 0.13-0.29 (0.22, n = 10) long. 
(.81-1.22 (104 mn = 10) wide. Genital pores 
Unihitteral Three testes arranged Imearly, one poral. 
two aporal: very little variation in testis distribuuon: 
single seement wilh + lestes: single segment wilh 2 
testes: single segment with 2 poral, | aparal testes, 
‘Testes oval. of similar size. (.10-0,17 (0,15. 1 = 10) 


long x O.08-0.1 1 (0.09, = LO) wide. Vasa efferenanit 
of antiporal testes run along dorsal margin of 
medulla (o elongate. pyriform external seminal 
vesicle (.06-0.16 (O10, 1 = 10) Tong x 0.04-0.09 
(O07. n = 10) wide. anterior and dersal to poral 
testes: distal region of external seminal vesicle 
slender, sittwous, enters clongate Gurus suc 0.27-0.42 
(0.35. 9 = 10) long x 0,04-0.05 (Q,0S. 4 = 1) wide 
Cirrus sac contains elongite internal seminal yesicle 
occupying two-thirds of volume of crerus sacs ne 
armature Seen On cirrus. Genital duets cioss 
osmoregulatory, canals dorsally, 

Ovary median, with 3-4 indistinct lobules, 0.06- 
O10 (0.08. n = 10) long. O10-0.19 (0.14. mn = 10) 
wide: vitellarium reniform, posterior to avury 0.03 
0.06 (0.04. 2 = 10) long ¥ 0,05-0,07 (0.06, n= 10) 
wide. Vagina posterior and ventral to cirrus site. 
dilating to form sacciform seminal receptacle dorsal 
(i ovary; seminal receptacle 0.14-0.24 (0.18 n= 10) 
lany x O.O8-0.L6 (O0b w = LO) wide. Uterus 
origingles as Uansyerse sac on ventmil dspect ot 
medulla. extends to osmoregulialary canals. 
developing small number of diverticula, never 
becoming reticulate. No segments found with fully- 
developed eggs materi. Ventral osmoreguliutory: 
canal 0,03-0,05 (0,04. m = 10) i) diameter, dorsal 
canal narrower 0.01-0.023 (0.02. 0 = TO) in diimetes. 
Longitudinal strobilur museulature arranged in two 
concentric rings! outer ring Composed of numerous 
small bundles with only [3 libres per bonule: 
bundles of inner ring larger with 5-10) fibres. per 
bundle. 


Potorolepis woalleyae sp. nev, 
(FIGS #-14) 


Tivpes. Holotype from small intestine of Mirrexia 
lotigicaadaw (SMegel. 1866). Mount Missim (7 
13'S. 146° 49'E), Morobe Province, Papua-New 
Guinea. voll. G. Gossek, 24.0.1084. SAMA ALHC 
27906: paratypes. 9 fragmented specimens. | scoles 
mounted in Berlese’s Aluid. serial sections. SAMA 
AKC 27907-27919, 30590; 2 specimens MUNG 
23408 INVEL 


Description 

Small cestodes, largest Imgment 35 long. Seales 
globose. 0.44-0.55 (0 50. n= 9) in diameter. Suckers 
Sub-circulin, unarmed, 0.15-0.21 (0.17. 9 = 10) tong 
x 0-70-4019 (0.15, n= 10) wide. Rosiellum inuseular 
0,13-0,20(0,17, 9 = 9) long x O.19-0,28 (0.23. 1 = 4) 


Figs 1-7. Badeeilepis arueusis sp. nov. Types. 1. Scales with custellim everted. 2 Seoles wath rostetliany wrhdrawn. 3, 
Resiellar took ta profile. 4. Rostellar hook, view [rom posterior surface showing entarved hook guard, S. Mature 
Sscemenmts. 0 Neareeriivitl seumenl 7. Troasverse listological section of riiture segment: dorsal aspect towards top ol 
pape. Soule bars = Q.Tim L225 7) 0.0bon 24. Legend By, dorsal osmoregulatory canal CS. external seminal vesicles 
SR seminned reeeptacte: | pestis; UL ters Y, ventral osmoregul ory canul 


98 


C. VAUCHER & I. BEVERIDGE 


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NEW CESTODES FROM DASYURID MARSUPIALS 99 


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 
(1.20, n = &) long. Segments craspedote; mature 
segments much wider than long, 0.05-0.06 (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 | 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 
ducts 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 = 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 yentral aspect of medulla, extends to 
osmoregulatory canals, developing few diverticula; 
never becoming reticulate. Eggs spherical, 0.032- 
0.045 (0.035, 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 
smull bundles with few fibres; inner ring composed 
of larger bundles with 10-20 fibres per bundle. 


Discussion 


Spasskii (1994) created the genus Potorolepis 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), PR aklei (Beveridge & 


Barker, 1975), P. bettongiue (Vaucher, Beveridge & 
Spratt, 1984), A bradleyi (Beveridge & Barker, 
1975), P. isoodontis (Vaucher, Beveridge & Spratt, 
1984) and P 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. 
Potorolepis peroryctis is morphologically similar to 
P. peramelidarum and P. isoodontis, also from 
bandicoots, and clearly belongs within the genus 
Patorolepis, 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 Potorolepis. 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 Potorolepis 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. 
bradleyi. 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 Icngths of P aruensis may 
overlap with those of P. bradleyi. Potorolepis 
aruensis Can be distinguished by mean hook number 
(Table 1) but also by the cirrus sac which is shorter 
in P. bradleyi (0.190 x 0.026 mm) than in P. aruensis 
(0.350 x 0.050 mm). In P. bradley, the central testis 
lies dorsal to the ovary (Beveridge & Barker 1975) 
rather than aporal to it as in P. aruensis. For these 
reasons, the specimens deseribed above from 
Myoictis melay are considered close to but distinct 
from P. bradleyi 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 eggs. This is 
surprising since the animals were ‘rausported to 
Melbourne following capture and maintgined in the 


Figs 8-14, Potorolepis woolleyae sp. nov. Types. 8. Scolex with rostellum everted. 9. Scolex with rostellum withdrawn, 10. 
Mature segments. 11, Gravid segment. 12-13. Rostellar hooks in profile. 14, Transverse histological section of mature. 
segment, dorsal aspect towards top of page. Scale bars = 0.lmm 8-9, 11; 0.2mm 10; 0.0lmm [2-13; 015mm 14. Legend: 


as for Figs 1-7, 


100 


C. VAUCTIER & I. BEVERIDGE 


Jaboratory unt] autopsy, providing adequate ime for 


cestodes to mature. 
The specimens 


trom Aateehinus 


wis are 


tentatively allocated to this species, They are poorly 
preserved and internal features are difficult to 
discern, Hook lengths are identical to specimens 
from Myaictiy, The number of huoks, 18-22 (20) (n 
= 5), is larger than in specimens from ML nels but 
the range overlaps. They may represent a distinct but 
very similar species, although the current evidence is 


equivocal, 


The cestodes trom Murexia lengicandata were 
quite severely contracted, limiting the morphological 


details which were 


visible 


in Whole mounts. 


Nevertheless. (hey represent a new species bused on 
the features of the rostellar hooks alone. Winle they 
resemble P. arvensis in terms of mean hook number, 
the size of hooks immediately distinguishes the 
material from all congeners. In having the three 
lestes arranged in a tnangular fashion, P. woollevue 
most closely resembles 2 uklei and PB. antechini, 
Apart trom adding to the hymenolepidid fauna 
known from mursupials in New Guinea. the new 


species described support the erection of the genus 
Potoralepixs by Spasskir (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 dorsally’. This is 
unequivocally the case in P aklei, P. brailleyi, P. 
aruensis, PL woolleyae and P. peroryetis and probably 
is similar in the remaining species of the genus, His 
generic definition (Spasskii 1994) also needs to be 
amended to allow for testes in cither a linear or 
triangular array and for cirri which are either armed 
or unarmed, Apart from these minor modifications. 
the generic definition provided by Spasskit (1994) 
appears to be reliable. 

The description of the new species also provides 
evidence in support of the suggestion first made by 
Vaucher ef ul. (1984) that cach family of marsupials 
Wis parasitised by a distinctive morphological group 
of hymenolepidid cestodes. Spratt e¢ al (1991) 
reported P. peramelidarum trom Antechinuys 
swainsont) which would represent a potential 
exception. However. the identification was tentative 


TABLE 1, Measurements and kev morphological features of species af Potorolepis fram marsupials. 


Species 


Species from dasyurid hosts 
Po aklei 


BF hlvadtev 
f antechini 
P aruensis 


P woollevae 


Species trom perumelid hosts 
P peramelidarum 


ft ispedontis 
BP perorvets 


Species fram potoroid hosts 
ES potornd 


P bettougiae 


Host(s) 


Amechinus sp. (indesertbed)* 
A. flavipes, Pseudantechinus 
bilarni, Sminthipsts lencepuas 
Anrechinus sp. (undescribed )* 
Antechinus swainsenit 


Myatets melas 


Murevia lanvicaidate 


Perameles Nana 
P gunnit 
Isoodon obesulus 


Iyoodon abexulus 


Perarvetes raffravanits 


Potorous twidaciylus 


Rettrongia gannardt 


Hook No, of Testis 
Length Hooks Distribution 
83-1) (OT) }-17 (13) tiangular 
103-128 (114) 1-15 (13) linear 
56-59 (58) 23-24 Inungular 
(28-147 (138) l6-I8 (17) Iinear 
163-182 (172) 15-19 (17) trrangular 
YA-101 (98) 35-38 linear 
71-2 (79) 43-39 sub-triangulin 
124-192 at) triangular 
98-103 (102) 29.33 linew or 
triangular 
T9-91 (SO) 24-27 linear 


formerly ientited as Amechinus sumarrit (see Strahan 1995) 


NEW CESTODES [ROM DASYURID MARSUPIALS Wl 


and based on incomplete specimens. This dubious 
record has therefore been elimimated from 
considerabon ntl mare material is collected and the 
host record confirmed, The species found in 
dasyurtds. Fo aklei, Pooantechini, Po aruensis, P 


hradleyt and Po weallevae, belong tou group of 


species with & small number of rostellar hooks (10- 
23) compared with 33-40 hooks tn Po tyeedentis, P. 
peramelidarum and Po pereryetis Yrom perameled 
hosts and 24-33 hooks in Po petoraé and B hettongice 
from potoroid hosts. The relative size of the ovary 
also separites the first Woogroups, We ratio of width 
of qwvary lo segment width being 9-15% in species 
trom dasyurid hosts compared with 26-39% in those 
from peramelids. In the first group the uterus 
contains relatively few egys. but itis clearly bilobed, 
More data are needed from hymenolepidids 
parasivsing peramelids and potoroids to confirm the 
ullity of this character. The observations of Jones & 
Anderson (1990) on a species froma New Guinean 


peramelid marsupial and the current desenpuons of 


new species from dasyurid marsupials from New 
Guinea provide additional support for the hypothesis 
advanved by Vaucher et al. (1984). The data alsa 
suggest that the hymenolepidids of dasyurids from 
Australia are similar ty those of New Gumea, as are 
the comparable cestodes of peramelid marsupials. 
Pinally. we agree with Spasski (1994) 1 allocating 
Hymenolepix cvercarteti Vaucher, Beveridge & 
Spratt. 1984 to the genus Kedentolepis, 


Key to the species of hymenolepidid cestodes 
occurring in Australasian marsupials based on 
rostellar hooks 


| Hooks small shorter than 30 yim Jeng, fraternoid in 
SHPO. eee vovenanl 
Hooks larger, Tength: ‘greater an in 30 yin “Tons. ht 
fralemmoid in shape - o scertasaxinc Oy srailverstreaiiey 


2. Wvoks number 20-30, 14-18 pm long 
Tere TETT ESE a ET ESE Sty Tt ee etre aT Tee TT Ridenralepis Hatta 
Hooks 17-22. 172 2 pm Jong... Redentalepis cercartent 


3. Fewer than 22 looks of if 22 hooks bragent, 
hooks >100 um long, 
More than 22 hooks or, if 22 hooks: presen. “hooks 
SU OWD gaty Neat sess sess ee assassin ea ee taasnaee ts 


4+. Hooks shorter than or equal to 100 pin 
Potovolepus akles 


Hooks longer than 100 pin... 


5. Hooks 163-182 pm Jang... 


Pintarolepis woallevae 
Hooks fess than (50 pam in length occ see eeeeee ere o 


6. Hooks 103-)28 pm long. [0-15 in number 
prorssasshhbeaastedteinsabtbnoane (cethtooeeassante ee ofa Paroralepis bradleye 
Hooks 128-147 i all tong. 16-22 in number 

cece POLOrOlEpiy ATUCHSTS 


JT. Fewer than 28 WOKS, ccc gig se perseser neeesey ceteeeepay ere 
More thitty 28 HOOKS ccc sens besten ere 9 


§. Hooks 56-59 jim long. 22-23 in gumber 
fo naire one bcobinsss bot .Patorolepis antechine 
Hooks 79-491 pire ton, 74 a7 id number 
eae ee ee, oe Patoralopis betngiae 


9, Hooks 124-192 ym lonp, 40 in number 
Poderalepis PEPOINCTIS 
Hooks. less than ‘120. yin long. ‘few ér than 40 in number 


a mu) 
10, Hooks shorter thi $3 WM. Pororalepis toodontis 
Hooks loner than §35 pmo 2 ee ee Wl 


Hooks number 29-34. 
Hooks number 35-38 ... 


at Potoralepis proton 
Patoralepis perametidarwn 


Acknowledgments 


We are grateful to Dr POA, Woolley who volleeted 
material used io this study and who also made 
specimens collected by others from New Guinea 
available to us. 


References 


Bevertoge. 1 & Barker. LK. (1975) Acuanid, capiliariid 
ind hymenolepidid parwsiles of the dasyurid marsupial, 
Mittechinus stiri Macleay. (8d. trom southeastern 
Australia. J. Meliiathal, 49, 211-227, 

CVAPUISSRE BO & Vancimr, ©. (199d) Family 
Hymenulepididae Anola. 1888 pp. 595-663 fn Khalil, b- 
F, Jones, A, & Bray, R.A. (Eds) “Reys to the Cestode 


Parasites Of Vertebrates” ICA Interminianal. 
Wallitigford) 
FLasnery, TOF 1995) “Maminals of New Giinea” (Reed 


Bas Chatswood ). 
Jonny. A, (1990) “Techniques Tor hand-sectioning thick- 
Viet Platyhelminths, Svat. Porusrtal, 15, 201-208, 


a & Asperson. TJ, C, (1990) Helminths at 
rodents and marsupils from Papuu-New. Guinea, with 
the description of two new species. Eelimastonu 
echymiperae usp. (Digenea Echinostomaticdae) 
und Manpiralepiy peranvctiy a ap (Cesteda 
bhymenolopididae ). Syst Pavonital. 15, 224-2497. 


Privciuaro. Mo HO & Keese. G. O | W. (19K2) “The 
Collechon and Preservation of Animal Parasites” 
(University of Nebraska Press. Lincoln), 

Srasskil. Al AL (1954) [Classification of the 


hymenolepidids of mammals.) Prdy Lab. tel mins 7, 
120-167 (in Russian), 


102 C. VAUCHER & 1. BEVERIDGE 


(1994) [On the systematic position of STRAHAN, R. (1995) “The Mammals of Australia” (Reed 
hymenolepidids from Australian — marsupials.} Books, Chatswood). 
Parasitologiya 28, 66-69 (In Russian). 

Spratt, D. M., BEVERIDGE, I. & WALTER, E. L. (1991) A VAUCHER, C., BEVERIDGE, I. & Spratt, D. M. (1984) 
catalogue of Australasian monotremes and marsupials Cestodes du genre Hymenolepis Weinland, 1858 (sensu 
and their recorded helminth parasites. Rec. S. Aust. Mus., lato) parasites de marsupiaux australiens et description 
Monogr. Ser. No. 1, 1-105. de cing espéces nouvelles. Rev. Suisse Zool. 91, 443-458. 


Transactions of the Royal Society of S, Aust, (1997), 121(3), 103-117. 


STRUCTURE OF THE ACOUSTIC SIGNALS OF CRINIA GLAUERTI (ANURA: 


MYOBATRACHIDAE) FROM SOUTH-WESTERN AUSTRALIA, AND COMPARISON 


WITH THOSE OF C. SIGNIFERA FROM SOUTH AUSTRALIA 


by Murray J. LITTLeEsJOHN® & JOHN R. WRIGHT® 


Summary 


Litr.eionn, M. J. & Wricut, J. R. (1997) Structure of the acoustic signals of Crinia glauerti (Anura: 
Myobatrachidae) from south-western Australia, and comparison with those of C. signifera trom South Australia. 
Trans R. Sac. S. Aust. 121(3), 103-117, 28 November, 1997. 

Advertisement calls of 51 males of the Australian myobatrachid frog, Crinia glauerti, from five sites, and 45 
mules of the presumed cognate species C. signifera from four sites, were analysed and compared. Patterns of 
geographical variation in the advertisement calls of C. glauertr were explored and a cline 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 cal] in the south. Because of possible similarity 
in function, the squelching call of C. glauerti was compared with the encounter call of C. signifera. 


Key Worps: Crinia glayerti, Crinia signifera, advertisement cal), encounter call, acoustic analyses, pulse 
structure, spectral composition, geographical variation, 


Introduction 


The commonly produced call (= advertisement 
call, sensu Litthejohn 1977, Wells 1977) of the 
endemic south-western Australian species Crinia 
glauerti (Loyeridge) 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 .,. 2’ This 
observation of inter-populational variation in the 
calls of C. glauerti 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 yalues were cited by Brown & 
Wilson (1956) from A.R. Main (in litt., based on 
andlyses 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 
(115°52' E, 31°59’ S), near the northern limit of 


* Deparment of Zoology, University of Melbourne Parkville Vic. 
3052, 

| Hawe, §. M. (1970) Calling behaviour and Territoriality in Males 
of Two species of Crinia (Anura: Leptodactylidae), BSe (Hons) 
thesis, Department of Zoology, University of Melbourne 
(unpub.}. 


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 
presented. 

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 ef al. 1958), 
Quantitative descriptions of the calls of C. signifera 
were also provided by Littlejohn (1964, 1970), 
Littlejohn & Martin (1965), Hawe', Littlejohn ef al. 
(1985) and Odendaal e? al. (1986). Interpopulational 
variation in advertisement calls of C. signifera was 
considered by Littlejohn (1959, 1964), and by 
Odendaal ef 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 (sensu Wells 1977) 


14 M. J. LITTLISOUS & J. BR, WRIGHT 


= territorial calls, Littlejohn ef al 1985) of ©. 
signifera) were identified through held) playback 
experiments by Hawe! and Littlejohn er al. (1985). 
The squelehing cally of C. glavertt may also have a 
territorial function but this hus yet be determined. 
In the absence of experimental documentation, the 
ternr “squelching call” (Main 1957) will be retained 
for the compressed series of pulses. 

Tape-recorded samples of advertisement calls of 
51 individuals were obtained at five localities across 
the geographic range of C. glauerti (Pig, 1, Table 1) 
io provide a measure of geographical variation. 
Mundaring Weir (Site |) is the type locality for C. 
ulauverti (Cogger ef al, 1983), Squelching calls were 


32° 


118° 


116° 
Fig. 1. Geograplic distribution (based on Tyler eral, 1994) 


of Crinia glaverti, and locations. of recording sites (see 
Table 1), 


TAREE 1. Denuls af recurring lavalites, 


produced in the recorded sequences OF 25 of these 
individuals, Sufficient squelching calls were 
obtained to provide a preliminary deseription of this 
type of signal for comparison with the conspecific 
advertisement call, and with the encounter calls of © 
signifera, and to-allow an estimation of geographical 
variation in the frequency of production relative Lo 
the advertisement call. Tape recordings of 
advertisement calls of 45 individuals from the closest 
populations of C. srgnifera were also obtained, 
namely from four localities in southern South 
Australia (Pig. 2. Table 1) at or near the western 
limits of the extensive geographic distribution of this 
species (See maps given by Brook 1983, 1984; Tyler 


33° 


37° 


140° 


=. 


Vig, 2. Geographic distrburon (based on Brook 1984) 0 
Crinta sivnifera in South Australia and locations of 
recording sites (see Table |). 


All sites are in Western Australia Cor Crinja elanverti aod in South Australia for OC) signifera: 


Species Sie Dates of recording Lacality Latitude Longitude 
(South) (East) 
©. vlanerti I 19, 23.vil. 89 Helena River, at base 
of Mundaring Weir 3157" Hla! 
2 O6.ViESG. 31 AV iSO 21 km south-south-west 
of Busselton 33°51 119! 
7 DAVIE. WITVIRG Diamond Tree railway siding: 34°20” 1 bode 
4 24..25.vii.86 3 kin cast of Nornalup 34°59° lin sa 
5 OY HOw. 89 14 kim east-north-east of Albany 44°59" Lise 
© sivnifera 6 TS8ovili BOD PS.Vii 49 9.3 km south of Wanilla 34°36! )a5°4u° 
7 03. O04 NY Yellowman Creek, 3.6 kit 
sonth-east of Melrose 32°51" yan 13" 
8 28, 29.yii. 90 9 kim north-north-west of 
Victor Harbour 35°29! 138°36! 
9 26.9190 4.4 km north-north-east 
of Millicent 37°31" ares 


CALLS OF CRINIA GLAUTERTIAND © SIGNILEKA rns 


1985: Cogger 1992). The deseription of the 
encounter call (Sterritorial call) of Co signifera is 
bused on accodnts of Hawe!l and Lithejolin et al. 
(1985, unpub,). 


Materials and Methods 


Recording af calls 

Tape recordings were obtained at the breeding sites 
(Figs t, 2. Table 1) with an open-reel recorder (Nagra 
IV-S) and a cardinid dynamic microphone (Beyer M 
88), Males of boty species gall from a variety of 
sites: om the banks adjacent to water, while sitting mn 
shallow water, or while floating and supported by 
emergent vegetation. The effective temperatures at 
the calling sites (surlace water, wet-bulb aie. 
depending on the calling position of the frog) were 
then measured with an eleetronie (thermistor) 
thermometer (Takura Digimula’ Model Doll, with 
sensor type SZL-64). The mean was used as. the 
elleclive Lemperature when an tndividudl was al the 
interluce between air und water. Where possible, the 
recorded’ mules of C. elawerti were colleeted. 
euthanased, preserved in Tylees fixative (Tyler 
1962), stored in 70% ethanol and lodged with the 
Western Australian Museum. (f the specimen was 
preserved. the measurement of smout-urostyle length 
was liter made with dial calipers (to 0.1 tin; 
rounded 10 0.5 mm), Otherwise, a short plastic ruler 
(15 em) was placed along the dorsal surface of the 
living specimen held tn a thin, clear plastic bag. and 
the Smout-urostyle fend determined to te nearest 
0.5 mm. 


Acoustic analysis of recordings 

For acoustic analysis, tape recordings were 
replayed on an Open-reel lape recorder (Revox B77 
MkIT or Sony TC.S10-2). Advertisement calls were 
dnalysed by using a digilal audio-spectrograph (Ray 
Elemetrics Model DSP-5500 Sona-Graph), Where 
there were high levels of low-lrequency noise, a 
passive filter (Allison 2B. high puss, cut-off set at 
512 Hz) was inserted between the output of the tape 
recorder and the input of the awudid-spectrogruph. 
Staustical procediires were carried out with 
SYSTAT, Version 5,03 (Systat Inc., Evanston). 


Characteristics af equipment used in recording and 
cUnaly8ix 

Tape transport speeds (1% cm 4!) of the recorders 
involved in recording and playback were periodically 
checked against a locally produced standard 1000 
KUlz calibration tape and a frequency counter (Heath 
Schlumberger SM-LISA or Good Will GFC-8010G) 
(Revox, Seny). or against the 50 Hy AC mains 
Hequeney by a buill-in stroboseope (Nagra); overall 
viriations in Lipe speed through recording und 


playback are estimated to be less than +0 5%. The 
frequency responses of all Clectronic vomponents 
used in recording and analysis dre presumed ta he 
close lo Linear within the narrow range of frequencies 
of interest (c. 2.0 - 5.0 KHZ), based on manufacturers” 
specifications. The relatively high simpling rates 
used in the digital analyses (=44 kHz) préelude the 
produchon Ob artiliels from aliasing. 


Surucnire and acoustic anribines of calls 

The calls of the lwo species are of Simple structure 
ahd each consists of a group of discrete damped 
oscillations (Figs 3-5). Por convenience. und 
following previous usage, euch of these oscillations 
is referred ta as a “pulse.” and the group (i.e. the 
pulse train) as a call (Figs 3, 4). The number of 
pulses ina call was delermined by direct inspection 
of the displayed wayeforny. The depth of amplitude 
modulanon of the last two pulses in a cull was 
sometimes. less. than 100%: in sueh causes, separate 
pulses were recognized if the depth of modulation 
exceeded aboul 75%, The duration (to nearest ms) of 
acall Was taken as the fnterval frem the peak of the 
first pulse to the peak of the list pulse (= “peak-peuk 
duration”). The pulses in the advertisement calls arc 
produced in a yuaseperodic tashion, and the pulse 
rale (as pos!) wats calculated over a complete call as 
(n - 1 pulses) x 1000/peak-peak durition in ins, 
Dominant frequencies were measured as the peaks in 
# power spectrum of the whole call, To describe the 
temporal and spectral properlies of pulses in 
advertisement calls, a tape recording of one call of 
each of three individuals of cach species (CL glanerit, 


io 


> wh 2.1, cu. uth. 0 
200 400 600 800 


milliseconds 


Fig. 3. Waveforms of advertisement cally. Lipper, C rive 
vlawerni= Reterence R439-7, Sire 1s effective emperature, 
wWel-bulb air = PLAN? CO. Lower, ©. sfeiiferd, Referenee 
R408-2. Site 6. elfevtive temperatures. wet-bulh wie = 
He O.. wailer= 12.5 €, 


106 M. J. LITTLEJOHN & J. R. WRIGHT 


Site 1; C. signifera, Site 6) was replayed into a digital 
sound card (Sound Blaster SB16, 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 


Time (ms) 


Relative amplitude (dB) 


[\WWrerannne 


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 (CF), 
by analogy with amplitude modulation in 
electronics. 

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. glauverti) or tenth (C. 


Time (ms) 


2 3 4 5 6 
Frequency (kHz) 
d. 
a@ 30 
2. 
o 
Gu 
z 
[-s 
FS 20 
a 
oO 
2 
2 10 
2 3 4 5 6 
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). 


Bo wn eo & 


onNn 


— 


CALLS OF CRINIA GLAUERTI AND C. SIGNIFERA 107 


milliseconds 


Fig. 5. Waveforms of squelching calls of Crinia glauerti. 
(a). R441-5, Site 5, effective temperature = 11,8° C. (b). 
R44\-1, Site 5, effective temperature = 13.1°C. (c). 
R404-2, Site 2, effective temperature = 9.6° C. (d). R441- 
4, Site 5, effective temperature = 13.1° C. (e). R407-6, 
Site 3, effective temperature = 11.1° 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 
millisecond, 


Numbers of advertisement calls of individuals 
analysed 

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 I4th (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. glawerti, 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 11.0° C, the nearest integer 
to the pooled mean for effective temperature (C. 
glauerti, mean = 11.1° C: C. signifera, mean = 10.8° 
C); these yalues were used in subsequent calculations. 
Where the slope for the combined samples for each 
species was non-significant, the raw data were used in 


HOS M.S, LITT LISOVN & JR, WRIGHT 


Taber 2, Influence of effective temperature on four attributes of the advertisement cally of Crima vlauentt (9 = 91) end C 


signiters (n= 45) 


Resulls of analyses by linear regression. Sce Table 4 for ranges of temperatures. 


Atribute of call Species Slope Significance of Coellicwnl of 
Slope (pt determination (ie) 

Duration C. shaven “0.070 <0. ),291 

C. sivnifera +0419 240 32 
Sumber of pulses C glauerti O.151 OS516 0.009 

C_ signiterdt +0,.290) (ddl 0.050 
Pulse rite C_ vlauerty HAAS <0. (249 

C. signifera r49o O.587 0,007 
Upper dominant frequeney C elanerti ~ 1), 107 0.008 (34 

C) sigmifera -156.566 <U.001 249 


(he subsequent analyses, 
Occurrence af squelching calls of ©, elauerti 

The presence of squelching valls was determined 
subjevtively. by teplaying the  tipe-recorded 
sequences of cach individual. In this analysis, three 
lypes of call were tecognised: advertisement calls, 
squelching calls, and [ransitional calls - as il was not 
possible [oO assign some to emher category. The 
squelehing valls of CL glauerti also consist of pulse 
(rains (Fig. 5). Becadse these calls are highly 
variable in temporal structure, and were produced 
only during the recorded sequences of about 50% of 
the individuals, they are only bricfly described ina 
subjective way, 


Results 

Steucnire of pulses in advertisement cally 

The polses in calls of both species are similar, each 
wilh a Sharp attack and a gradual (negative 
exponential) decay (Fig. 4), Estimated durations are 
4ms for C. elaverti and 4-7 ms for C. sienifera. The 
maximum amplitude is reached within three positive 
of fegative peaks of the carrier frequency. that is, in 
about | oms for both taxa, Examination of the 
expanded wave form of each pulse indicated the 
presence of a clear sinusoid (= fundamental/carricr) 
with a frequency that is close to the upper peak of the 
spectrum (Table 4). Accunlingly. only the ipper peak 
(= currier frequency) was used ino subsequent 
calendanons. The frequency bandwidth at 10 dB 


below the peak is ahout 1200 Tz for both species 
(Fig. 4), The envelope of the pulse is. amplitude 
modulated to a depth of about 30-60% (based on the 
first cycle) with the envelope modulating frequency 
(EMF) within a range of 694 lo 820 Hy for © 
glauerti, und of 505 to 885 Hz for C. signifera (Fix. 
4), These values are close to (he difference between 
the upper and lower peak of the spectrum for each 
individual (Fig. 4. Table 3). It is suggested that the 
Jower peak that is present in the spectrum of the 
advertisement cally of some individuals of cach 
species (Fig. 4) is a sideband of the carrier frequency 
(i.e. the upper dominant [requency), The level of the 
lower peak is about 4-9 dB below that of the upper 
peak (Tible 3). 


Structure of the vdvertisement calls 

Both species produce advertisement calls in long 
ind regular sequences. Por C. elaversi trom Site 1, 
calls were produced at jt mean rate of 26.5 calls min! 
(range = 22.9 - 28.8: n = 9) al a mean ellective 
temperature of 12.1" C (ringe = 11.4 - 12.8), For 
sivnifera from Site &, the mean rate was 124.7 calls 
min! (range = 41.) - 87.5 n = th) at & mean 
effective temperature of 11.4? € (range = 112 
11-7). Values for three primary attributes (duration, 
number of pulses and carrier frequency) and the one 
derived atuibute (pulse rate) for five samples of © 
glaverti and four samples of C. signifera. corrected 
for the éffect of temperature where appropriate, are 


Tanith 3. Specrral characteristics of pulses in advertisement cally of Crinit glauerti and C, signifers, 


ALL values are in Hz. See text for explananun. 


Species — Individual = Carner Envelope Lower 
frequency modulation spectral peak 
(CF) frequency (LSP) 
(EME) 

©, ohiverts ! 4065 R20 ARTO 
z 4167 694 3472 

4 4049 820 3266 

© sieniferit | 2494 505 2050 
2 2632 671 206% 

3 3413 885 2454 


Upper Dillerrenee Ditlerence — Difference 
Spectral peak = hetween between CF between EME 
(USP) — spectral peaks and USP and DSP 

(DSP = USP 
~ LSP) 
4048 OTR 17 (42 
Nad 662 33 32 
ADS 782 | 38 
2407 447 3 58 
2670) #02 ae qy 


V5 991 32 106 


CALLS OF CRINIA GLAUERTI AND C, SIGNIFERA 109 


Tanvr 4. Phasical characteristics of advertisement calls of Crinia glauerti and Co signifera, correeted ta an effective ten- 
perature af FLOOC, where appropriue (see Table 2) 

For each cell, he mean and standard deviation are given on the upper line, and range (io parentheses) on the lower dine. 
a 


Species Ste Sample Eflective Call duration Number of Pulse rate Carrier 
sive temperature (°C) (ms) pulses ips't frequency (HH) 
i 
€. vlimerti | 10 (2.17. 047 738, 87 9.70, 1.21 1,80, 1.04 4052. 229 
(114-128) (573-817) (7.5- 1bS) (9.4 - 16.3) (3584 - 4430) 
2 7) 10.14, 0.64 724.153 9.83, 1.50 ) 2.06, 0.56 4279, KS 
(9.2 - 11,0) (S82 - 982) (8.0 = 12.0) (I2.0- 13.6)  4a580- 4540) 
A 13 10.27, 1.03 705, 115 11.50, 1.88 4.94, 1.80 4068, 215 
(7.7 - 11,5) (547-935) (9.0 - 15.5) (IIS- 18.3) (3770-4430) 
+ s 10.56, 0.57 326. $2 10,06, 1.84 6.91, LAS 4031, 485 
(9.6 - 11.4) (458 - 655) (8.5 - 13.0) (142- 18.6) (3250-4570) 
5 1 12.27, 0.62 598, 85 11.23. 2.37 18.20, 2.90 4076, 292 
(11.0-13,1) (456 - 747) (8.0 - 16.0) (15,1 - 23.3) (3640 = 4450) 
Combined 51 W110, 1.18 604, 130 10.57. 1,92 4,93. 3.06 4098, 302 
sample (7.7 - (3.1) (4358 - 982) (7.5 ~ 164) (9.4 - 23.3) (3250 ~ 4570) 
C. siunifere 6 12 11.54. 0.54 272, 77 6.28. L54+ 19.08, 3.41 2790, 325 
(1024 - 12.2) (150 - 437) (4.0 - 10.3) (15.5 - 28.8) (2504 - 3568) 


7 13 10.64, 0.78 161. 70, S31. 141 29.05, 7.56 431. 41 
(9.2 - 11.2) (9} - 272) (4.0 - 9.40) (184 - 43.8) (2092 - 2871) 

8 10 1143. O18 182.55 4.87, 0.09 22.50), 4.53 2509, 87 
(1).2- 11.7) (10) = 237) (4.0 - 6.0) (175-297) (2471 - 2710) 

4) 10 9.25, 0.27 158, 20 4.37, 0.48 21.32, 1.91 2825, 169 
(8.9 - 9.8) (125-192) (4.0 - 5.0) (18.2 + 24.1) (2549 — 3103) 

Combined 45 10,75, 1,02 195, 77 §.26, L.34 23,48, 6,15 2645, 279 
sample (8.9 - 12.2) (91-437) (40 - 10.3) (15,5 - 43.8) (2092 - 3568) 


1000 


800 


600 


400 


Call duration (ms) 


200 


Site 


Fig. 6. Box plots for durations of advertisement calls at recording sites of Crinfa glauern (hatched boxes) and C. sigmifera 
(open boxes). Values are corrected to 11° C where slopes are significant. The boa indicates the interquartile range and the 
included horiyvontal line is the median. The vertical lines outside the boxes (whiskers) connect to the last data points within 
+ 1.5 x the interquartile range (the fences). The asterisks indicate outliers (values lying between + 1.5 and 3-0 x the 
interquartile range) and the open circles indicate extreme outliers (values beyond 3 x the interquartile range). 


110 M. J. LITTLEJOHN & J, R. WRIGHT 


Number of pulses 


Site 


Fig. 7. Box plots for numbers of pulses in advertisement calls at recording sites of Crinia glauerti and C. signifera, See Fig 


6 for explanation. 


50 
40 


30 


20 


Pulse rate (p s“) 


10 


Site 


Fig. 8. Box plots for pulse rates of advertisement calls at recording sites of Crinia glauerti and C. signifera. Values are 
corrected to 11° C where slopes are significant. See Fig. 6 for explanation. 


CALLS OF CRINIA GLAUERTI AND C. SIGNIFERA It 


presented in Table 4, Values for combined samples of 
each species are also presented in Table 4. Box plots 
of these values at cach site are presented in Figs 6-9, 


Correlations of attributes of advertisement calls with 
body length 
Calling males of C. glauerti are smaller than those of C 


signifera (Table 5), with mean snout-urostyle lengths of 


16.31 (range = 14.0- 19.0; n= 29), and 21.31 (range = 18,0 
- 25.0: n = 16) mm respectively (t-test; p< 0.001). There 
is no significant correlation of carrier frequency (corrected 


5000 


4000 


3000 


Carrier frequency (Hz) 


2000 


to 110° C; Table 2) with snout-urostyle length when all 
collected specimens of C. glaerti are included (v= - 
0297; p = OV 18; n = 29): but there ts a significant 
negative correlation when only those specimens thal were 
measured following preservation are considered (r = 
-0.440; pp = 0.032: n= 24). For C. signifera, however, there 
is a highly significant negative correlation between carrier 
frequency and snout-urostyle length (r= -0.796; p > 
0.001; n= 16, all specimens were alive when nieasured), 
A scattergram of the correlation of snout-urostyle length 
and carrier frequeney is presented in. Fig. 10. 


Site 


Fig. 9. Box plots Cor carrier Frequencies of advertisement calls at recording sites of Crint glaweri and C. sienifera. Values 
are corrected to 11> C where slopes are significant. See Fig. 6 for explanation, 


Tanne 5. Vales fin aun) for sneuturesivle lenuths of males of Crinta eliuerti one Cl signitera cellected after ther 


advertivement cally had been recorded. 


Species Site Sample size Mean 
C. vlanerti | 8 17.2 
4 6 14.6 

i 7 lo.2 

| 3 (5.5 

5 5 17.6 

Total 249 16.31 

C. signifera 8 7 23.4 
os) 9 197 

Toul 16 31 


Range Standard Condition ol 
deviation specimen 
16.5 - 19.0 0.80 preserved 
I4.0- 15.5 O58 preserveil 
15.5- 17.0 O49 preserved 
IS.0- Lo.0 0.50 preserved 
17.0= 16.0 0.55 live 
I40- 1o0 1.25 
22,0 - 25.0 1.17 live 
18.0 - 21.0 141 live 
18.0 - 25.0 231 


a ——————— 


112 


Geographical variation in advertisement cally af C. 
vlaueru 

Analysis of variance, with Site as the grouping 
factor, indicated that there ure no significant 
differences Jor number of pulses (p = 0,074) and 
curmer frequency (p = 0.411). There are. however, 
significant differences for the means (adjusted to 11° 
C) of duration (p< 0,001) and pulse rate (p < 0.001), 
Across the distance of about 450 km covered by the 
five sites (Fig. 1), there is no consistent trend in 
duration (Fig, 6, Table 4), although a multiple 
comparison (Tukey test) indicated that the means for 
Sites 1-3 are significantly lower than those at Site 4, 
and Site | also differed from Site 5. There is a cline 
of increasing values for pulse rates (Fig. 8, Table 4), 
A Tukey test showed that the following means for 
pulse rate differ significantly: Site 1 trom Sites 3, 4, 
and 5; Site 2 fram Sites 4 and 5; and Site 3 from Site 
a) 


Geographical variation in advertisement calls of C. 
signifera 

Sites 7. % und 9 wre within the continuous 
distribution of C_signifera (see maps given by Brook 
1983: Tyler 1985) 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 & (big, 2). 


5000 


4500 


4000 


3500 


3000 


Carrier frequency (Hz) 


2500 


2000 


12 14 16 


18 


M.J. LITTLEJOHN & J-R. WRIGHT 


Analyses of variance, with Site as the grouping 
factor, indicated that there were significant 
differences for all four attributes of the calls (p < 
0.003), Multiple comparisons (Tukey lest) revealed 
the following significant differences in meuns 
between sites; call duration - Site 6 Irom Sites 7-9: 
pulse number - Site 6 from Sites 8 and 9: dominant 
frequency (adjusted to 11° C) - Site 7 from Sites 6 
and 9: pulse rate - Site 7 tram Sites 6, 8 and 9. 


Compearixon of advertixement calls of C. glauerti ane 
C. signifera 

Results of analyses of variance, with Species at the 
grouping factor, indicated that the advertisement 
calls of the two species differed significantly (a < 
0.001) in all four attributes. From a consideration of 
the combined samples for each species (Table 4), 
culls of C. glaverti are much longer. with means of 
durations diflering by 3,4 times, and with no overlap 
in ranges of yariation (Pig. 6), The mean value for 
number of pulses in advertisement calls of C. 
glanerti is twice that of C. signifera but there is an 
overlap of ranges between 7.5 and 10.3 (Fig, 7. Table 
4). For pulse rates, although the mean for C. glauerti 
is only 64% of that for C. sigaifera, the ranges 
overlap extensively (Pig. 8. Table 4). particularly for 
the closest sumples (Sites 5, 6). The mean for carrier 


26 


22 
Snout-urostyle length (mm) 


20 24 


Fie, 1. Seattersram showing the correlation between carrier frequency ancl snout-urostyle length for mules of Crinia 
glaverti and C. signifera that were collected after thei’ calls had been recorded. 


CALLS OF CRINIA GLAUBRTEAND € SIGNIL ERA Iie 


TALL 6, Physical clnvacteristies af seven yquelching calls uf Crinia clauentiy fron the sue set ofcally presented it igure >. 
PAR Sn 


Individual ‘Temperature Duration = Number ot Low pulse High pulse Overall pulse Currier 
Gind tape (ey (ins) pulses rate (ps) (ps') rile (ps!) frequeney (He) 
feferenee) 
a 
i (R4+4E-S) LIB 475 av) 43,2 205,4 13.2 3704 
bh (R441-1) 131 138 ce 31 794) 44,0 A7A6 
& (RAQ42) %.6 324 a7 467 125.0 OR? 4392 
dl. (RASTA) 13. S04 ‘I Abs Lift 52.7 AN40 
& (RANT-0) Ih. 478 2\ S74 02,3 dis 4220 
1 (R44 A} Os 604A a5 Hy O17 4.8 4263 
gw. (R4Q7-7) h,7 #32 24 29,7 Wa 27,0" +048 
Means Hat 359.0 40.0) ST XO 115.20 S455 4086 
Runges 6-141 435% - X32 21-50 297. 46,7 617-2059 27. - 103.2 3703 43402 


—— —— ———— 


‘ower than low pulse fate becuse of the four long breaks in the tains 


frequencies is higher in C. glawerti, by L452 Hz, bul 
with an overlap of fabges between 3229 and 3568 Hz 
(Pig, ¥, Tablet), 


Srructire of the squelching call of C, lauerti 

These culls are highly variable, as is mdieated by 
ihe selection of wavetorms presented in Fig. 5. The 
pulses are sometimes in groups within weall and the 
pulse rate can vary greatly through a call (Fig. 5). 
Values for four attabutes of the seven calls presented 
in Fig, 3 ave given in Table 6, The pulses are of 
similar structure to those of the advertisement call 
and (here are also lwo peaks in the frequeney spectra, 
us in the advertisement call. For the seven calls 
presented in Fig. 5, the following data apply (see 
section on pulses of advertisement calls for 
methodology): duration - mean = 3.77 ms (range = 
3.1 -5.1): carrier frequency - mean = 3955 He (range 
= 3481 - 4427); upper lrequency peak- mean = 3976 
(range = 3703 - 4414); envelope modulinng 
frequency - meun = 877 Hz (n = 5: range = 735 - 
1062); lower dominant frequency peak- mean = 3260 
Hy (n = 5) range = 3100 - 3464), The difference 
between the means of peuks oF upper and lower 
Hrequencies of 716 Hz is consistent with the 
explanation advanced Jor the pulses in the 
advertisement calls - of the lower sideband of aa 
envelope modulating frequency, The pulses may be 
grouped within a call (Fig. 5) and the pulse rates can 
differ vonsiderably between groups mone call (Table 
0). 


Geographical variation in frequency of vecurrence 
of the squelvhing call of ©, glauerti 

Reeorded sequences of calling by 25 of the SI 
individuals included squelching calls. Beeause of 
confusion fron: the calling by two or more close 
individuals, only those of 21 could: be reliably 
assessed. classified und counted (Table 7). A 
geographical trend is evident, with the frequency of 
production of squelching calls by individuals being 
ivher in the southern and south-castern sumples 


(Sites 4, 5, Table 8). No attempt was made to 
determine the extent of variation in the squelching 
culls of individuals. nor overtime for & population. 


Structure of the encatmiter calls of C. signifera 
Littlejohn ef ak (1985). by playbiek af 
advertisement calls of CL pariosiguifera and ©, 
vigniferd iat peak sound pressure levels (0 dB re 20 
Pa) above €. YS und 104 dB respeetively. evoked 
encounter calls (= territorial culls sens Hawels 
Littlejohn 1977) from eight males of CL srenifera at 
Willowmavyin wi south central Vietoria (37°16" 8, 
I44°54" E), The encounter cull (Hig. 11) ts alse 
pulse train, with w regular pulse rite (Hawel, 


TAWA 7. Numbers af cedvertivemens calls. interiresiate 
(transitional) calls and squelching calls, and proparitanys if 
the fatter (ofall cally of (het iuadividiueal), produced by 21 
moles of Cringe ghivert, 

Note that beeaise of overlap wilh neighbours. ealls of + 
Individuals Could not be unilysed, 


Site Individual Nauiber Number MuseDs Proportiitt 
ct) i “ 1) 

advertisement iterpwidiite squelching — sqielehing 
wills calls culls calls 
| | MI 0 ] (001 
2 hit I2 2 7 (33 
2 2 9 | I ()52 
3 3 Ih 4 4 (15 
3 3 27 j 4 (h?] 
3 V alt | 4 (10 
3 7 2| + I 4) 
4 4 }3 ) 7 (48 
+ s in 2 | 0.20) 
4 2 10 | 4 (1,22 
a) lu Is : \2 (38 
3 af in 5 4 (h2I 
3 aT {5 6 4 ih 
5 di 24 (| { 04 
5 Su O} 0 if) vn 
3 ( ‘I iQ) 13 WOU) 
2 \h ' 5 0,27 
2b i] 4 3 (105 
5 s) 15 I } Q43 
5 th It 0 | (34 
5 Sh tu 5 \3 (Af 


Ld M.S, LITTLEIOUN & dR, WRIGHT 


Tabne §, Miaibens ind proportions of miley of Crinit glauert) prodietie yqueleline calls at each vite, 


————kn— — 


Sie Number of nrales 
assessed 


Number of males 
producing squedchine calls 


Proportiun of males 
produemyge squelehing calls 


I i0 
2 i 
3 2 
4 r 
5 \ 


td 0.200 
A (273 
5 OAL? 
1 0.500) 
it 1.000 


ee COO OCC 


Litlejohn 1977), A wavelorn of an encounter call 
from Site 3 is presented in big, TL Values for means 
und ranges front the original date for advertisement 
cally and evoked encounter calls of the same five 
individuals discussed by Littlejohn er al (85) are 
presented in Table 4, 


u 700 200 300 400 500 
milliseconds 
Mis Th. Waveform of an encounter call of OC) sieaifera 


(References: R4AO8-9) Sire 6. etfectiye lemperatures. wel 
bulb air 1 1.3° CL water 12.5" ©), 


TABLE 9. Compariven ef attributes of udvertvenenr cally 
cn eneounte? culls of Crnia signers from Willoynenin, 
Vater (frome date af Litdéjalay etal. 198s), 

Meuns and ranges (in parentheses) are given (n= 5) Datu 
are not cofrected Tar possible effects of temperuiure. 
Reflective lomperatures ranged frond 10.7-13.27 © (ean = 
12.04), 


a 


Atirihute Advertisement call Encounter call 
Duration (ins) qa 192 
(56-108) (| 79-2 ]4) 
Pulses/note 43 17s 
(4-5) (16-19) 
Pulse rate 40.6 RK_| 
ips!) (30),K-53.7) (742-1901) 
Upper dominant 4390 3217 


frequeney (He) (3125-3642) (2425-4020) 


Production ef encaunter calls by males af C. 
stoner 

I wu subjective evaluation during playbuck of the 
lape recordings of ©. yfignifera, involving 53 
individuids and some 3527 advertisement calls. three 
Interactions, presumed to jpvolye production of 
encounter Calls. were noted. Otherwise. the recorded 
sequences of 47 imdividuals consisted only of 
advertisement calls, 


Comparisan af squelching call af C. wlauerti and 
rerriorial call of C. signifera 

The variable squelehing call of C, glauerti (Fig. 5, 
Table 6) ts longer, contains more pulses, is of bhher 
average pulse rte, ine of higher dominant frequeney 
(han the tereitorial call oC. signifera (Fig 11, Table 
9), Even so, they are both pulse trains whieh are ol 
similar cartier freqaeney bul of longer duration, 
contin more pulses, and are of bigher pulse rate than 
(heir respective advertisement calls (Pig. 3, Table 4), 


Discussion 


Siruchure of the advertisement culls uf hath species 

The advertisement calls of both species are or 
similar structure, cach consisting Of a quasi-periodic 
pulse Lain, The pulses are short damped oscillations, 
each with a sharp attack and an exponential decay. 
There are more pulses inthe calls of ©. eleueerti. und 
These are produced ut a lower repetition rate. The 
frequency Spectra are of sittilar shape. with most 
individuals having two peaks, the upper being 
aliribuited to the fundamental (= carrer frequency} 
while the lower 1s presumed to be produced as the 
side band of the envelope modulating frequency. The 
carrier frequency is higher in C, glawerti, and this 
may be correlated with the smaller size of males ol 
(his species ~ a usdal characteristic Of the calls of 
anurans (e.g. Robertson L986) There ts a (rend of 
Increasing pulse rate from north to south-east in the 
samples ol C. elawert and the durations are lower in 
the samples from Sites 4 and 5. Values for samples of 
the other two auributes display no obvious pattern, 

Advertisement calls from Site 6 in the 
geographical isulate of C. yégeifera on lower Eyre 
Peninsula are Jonger and contain more pulses than 
those to the cust in the main distribution of ©, 
Nignifera. Thus the westernmost sample of © 
Nignifera (from Site 6) is more similar to those of C 
giver than ace the others, The populations of © 
siumifera on Eyre Peninsula have probably been 
sepiraled from the main distribution singe the sea 
rose Lo its present level at the close of the last elacial 
period of the Pleistocene Epoch rome) 12.000 toe 
6.000 years ago: sce Littlejohn er al 1993 for a 
summuary and references) and this isolation may have 
contributed to the divergence. 


CALLS OF CRINIA CE ALLERTE ANDO, SIGNIPLRA Ws 


Quendaal es at (LY86) recorded a sample of 
advenisenient als of Co vignifera from the same 
locution on Yellowmun Creek (ther Recording Site 
No. 5: our Sile No, 7) and aver a comparible range 
ol efiective (wierd temperatures (1K) 122° Cy, 
Yo - 41.2 © - this study). Por all four attributes, 
however, thei values ure Higher than those obtained 
in the present study. The use of other protocols anil 
abalytieal techniques may account for some of the 
differences. bur at This stage. no explanation can Pre 
offered, 


Frequency of varemaier cally iC, slavery) 

Moretecent observations by Litleyoha (unpub.) 
indicated that C. glaverd oecurs in syntopy willl © 
subinstgnifera (Littiejoln) near the eastern limit of 
Ws distributing (eg. Site S. Pig. ty) Crea 
swhneni/ercd bus an advertisement call that soureds 
ike u “eng low-pitebed squelch” (Lildeyohn 1957. 
959) Bor two temporal utierbutes of the 
advertiseient calls of 37 individuals of C. 
whinsiguiferd conected to-an effective lenperalure 
wl 1 the mean curation rs 540 ms (range = 420 
- O60) und the mean pulse rare ys 174 (range = 129 - 
10) (Littlejohn M961). As these canges overlap those 
of the squelehing call of €. glawert? (Table 61. 
explunations alber (han repradiedve charieter 
Uisplicement aust How be sought! for the higher 
frequencies of becnmrenve of squelefing cally ip the 
soattraind south-east but none can be provided) at 
present. Clearly, there is a need for further 
investigwuons, including playback expeciments with 
ddvertisement culls and squelehing calls as stimuli 
tinder coatealled conditions. Such studies should be 
preceded hy jhe (locumentanou of frequency ol 
veeurrenee oof squelehing calls in natural 
ussembluges. und the context in which they are 
produced, The meusurement of the sound pressure 
levels of calls of conspecific neizhbours ts also 
required so that the appropriate stimuli can be 
applied (Linlejohn eral LYSS). By varying the levels 
OF sfinglation, thresholds could then be determine 
wd geographical patterns may he revealed. 


Relationships of C, glaverti 

Prom the presented data, appears that the nearest 
populations of CL vlawerti and C. yignifer CSites 5. 
6) have the mose similarly structured! wdyertisement 
calls, Phe main difference helween the culls of the 
two Species is i cunier frequeney which may be 
devounted lor by the difference in sizes of the twe: 
tase (Table 3. Fig. (0). The similarities in call 
Structure ave Consistent with the postulatcd close 
relationship of the (Wo taxa suggested) by Viarin 
(W957). and subsequently supported by the 
multivariate numerical analyses of morphology ine 
fearures of Whe history carried oul by Bhike (19734 


and Thompson (1981). Althoush consistent in 
showing a close ussechition between them. 
molecular studies do nol help ay resolving the 
relationships of Co efamerte and C. sigaifern, The 
albumin iimunolomed! wialysis of Daugherty and 
Maxson (1982) places © sfgnitera clusest aC 
riparia (1S WD units), followed by Co efadiert and ©. 
patinsigniferd Cooth 2+ 1D units) anu C. gearg iene 
(29 1D units). The clidisne analysis of allazvmes 
curried oft by Burendse (1984) offers several 
interpretations and appears incondlasive ahout the 
relauonships: in one scenumy Co glawert’ and C. 
signifera ure grouped with CL georgian. Burendse 
(1984) did not inelide CL niperia in his study 
Roberts & Watson (1994) have reviewed the recent 
litvrature on relationships within some eraups of 
Austrian frogs. 

Three of the specres of Crini described since the 
work of Main (1957) - ¢. biliveue Martin. Tyler & 
Davies (Murctin ef al WSO). Co renia ‘Tyler & 
Parker (Tyler & Parker PO74) amd €. rear 
Lidejathn & Martin (Littiejohn & Martin 1965) - 
have cleurly pulswile advertisement calls. Tyler & 
Parker (1974) noted the sinvilurity of the 
advertisement culls of © reniota and ©. efaierrn, buy 
they did not provide information whout the recording 
temperatures. As the recording of the call ob © 
remafa Was obtyined ut Morehead, Papua New 
Guinea in January. the ambient temperatures 
presumably were much higher than those apply iig te 
recordings analysed in the present study. Hence, 
divect comparison cannot be made woh the cally of 
C. glanert) obtained ab (emperiiores ob 13.4 C and 
lower. Blake (1974) noted thal Crit ripurte bucks y 
tympanum and columella and placed the taxon into a 
different species group ulong with Co fasmenitensis 
(Gunther), Because of the laek of a distinee 
tympanum in ©. remote (Tyler & Parker L974), 1 as 
suggested frat there may not be a close relationship 
between this species und ©. elanern 

As the name indicates, the advertisement calbal ce 
ilingue is stronaly biphasfe (Martner al W880), In 
this species, males commence a calling sequence 
with short calls (<545 ms) of high pulse rate t>76 ps!) 
and then gradually change over (a long calls (23580 
ms) oF ow pulse rate < 54 ps!) Again, the high 
recording (amperatures (wet-bulb atir = 243.4 - 26.6° 
C) mean that it i not possible to make a proper 
comparison with the advertisement call of C 
clunerti. Evel so, the aidiospectrogrum of the Shor 
cull appears (ode simikic to some of the variations im 
the syuelching calls of Co elawerte. Martin er al 
(J980) considered the Functional significunee of the 
two disuoct culls of ©. bifimgite and rised the 
possibility thatthe short culls of higher pulse rade are 
milting calls, und that the long calls of tower pidse 
rile: are territorial walls, the converse af the situation 


Has Moa DIP IRMOTIN & TR. WRIGHHE 


for pulse rate in O) siuaiferea, The long call of €. 
Hilingue Comiuits about Iwiee as aiminy pulses. (227) 


as ub the advertisement call of iC) glanert tele TF 


allowance is made for the ditlerence my lemperalure 
(by Using QQ) Of 2), the pulse rare of C) ile ul 
HC woukl he about 20 pos band eould Overlap the 
pulse rates mm the advertisement calls of ©. e/awertt. 
Crinty bilineda possesses a Cyinpanuin bul possible 
close alfinities with ©. wanerti were nol considered 
by Martin etal (1980). 


Acknowledgements 


The field) stadies a7 Western Australia were carried 
gut under Licence Numbers 1274 (1986) and SF Lk6 
(1989), issued by the Departinent of Conservation 
and Land Mingement, The field studies were 
wirried oul during fiekl research leave front the 
University of Melbourne. Equipment was provided 
by grunts from the Australian Researeh Council 
(ARC) and the Upiversity of Melbourne, Recordings 


of CL slenifera Von South Austrailia were abtained 
diving the tenure of a research grant from the ARC, 
Number A T8831316 (1989). Some of the ueouste 
and stitistical analyses. and preparation af parts af 
the manuscript, were carried out ducing the tenure al 
other grants tam the ARC, Small grants: Seheme 
(Gul Numbers SG 0935514, 8 OS94A7578) cod the 
procedures that were developed have provicee 
protocols Tor acoustic and statistieul analytics! 
techniques of Wider application, Mrs PG, Litllejohn 
assisted with recording al all af the sites iin Westem 
Australia and at wea of the South Australian 
localities. Dr T. CG. Litthejohn assisted with record ine 
at the other two South Austtalian localities, Mp bh. A, 
Smith, Western Austin Museuin arranged for ihe 
cunition ol The preserved specimens and kindly rite 
them available for subsequent exanminahion ane 
newsurement during a subsequent visit, Dy M, J, 
Keough provided jalvice Of stuistical aualyses and 
Dr Hh CL Benner-Chak provided an interpretation of 
the speetral amd temporal siete ot pulses i the 
ivertisement calls, 


References 


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(Anurin Myobitrachidae) i souhern Australias A 
Phylwenciic ghulysiy ol qlovying dita supporting 
vhdemic speekiiii i Southwestern Australian Avelution 
38, (248 1250 


Biakh ALT DL 11979) Taxonomy and relativnships af 


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Brows, Woot & Witsna., EO, (1956) Character 
ddsiphiceniont Swit 4and S$, 49-t. 

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Leiba. MOP Ch9S7) A new species of frog of the penis 
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(1958) \ new species of frag of the wenus (rine 

Tsehud) from soutlecaster) Austtatia Pree, Linn See, 

NSW 83, 227.226. 

(1959) Call differentiation ima complex ob seven 
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yh S14 S36 fe Blair WO Chea Vertebrate Speesation” 

(Liversiby at Fesus Press, Aust), 


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Uifferentintion Wi Cefniy siaaifard Ebalytion UB, 262 
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(1970) Chea rasanertitersiy (Ate: 
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anuriis: An iiteerated and evolutionary approdelt pp. 
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Reproductive Binlogy af Amphibians’ (Plenuny Press, 
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Tharrison, POA, & Mac wan RO © ¢7a85) 
Inferspecihe acouste interactions jn synpatrie 
populitions of Raidedke signifercd and Ay partistenifere 
(Ane Leptodaeciylidaes pp. 287 206 In Cirle Ciy 
Shine, R. & Bhan bh (eds) “Biglowy oF Austedasian 
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Chipping Norton, 

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 ROKERES, JD. Warsos, G Boa Davies, M, (1993) 
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Main. A. BR, (1957) Studies in Australian Amur I. The 
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three genera of Austratiun frou, Byoluiion 12, 224-233, 

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Homhwestecn Austin, Capere TOBO, Yay 


CALLS OF CRINIA GLAUERTI AND C, SIGNIFERA 17 


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of the frog Ranidella riparia. Anim. Behav. 34, 1836- 
1843. 

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2A Amphibia and Reptilia’ (Australian Government 
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and the role of vocalizations in the Australian frog 
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STRAUGHAN, LR. & MAIN, A. R. (1966) Speciation and 
polymorphism in the genus Crinia Tschudi (Anura, 
Leptodactylidae) in Queensland. Proc. R. Soc. Qld 78, 
11-28, 


THOMPSON, M. B. (1981) The systematic status of the genus 
Australocrinia Heyer & Liem (Anura: Leptodactylidae). 
Aust. J. Zool. 29, 93-102. 

Ty er, M. J. (1962) On the preservation of anuran tadpoles. 
Aust. J. Sei. 25, 222. 

— (1985) Biogeography pp. 225-229 /n Twidale, C. R., 

TyLer, M. J. & Davirs, M. (Eds) ‘Natural History of Eyre 
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& PARKER, F (1974) New species of hylid and 

leptodactylid frogs from southern New Guinea. Trans. R. 

Soc. S, Aust. 98, 71-77. 

. SMITH, L. A. & JOHNSTONE, R. E. (1994) ‘Frogs of 
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amphibians. Anim. Behay. 25, 666-693. 


THE BREEDING BIOLOGY AND 
ADVERTISEMENT CALL OF LITORIA SPLENDIDA 
TYLER, DAVIES & MARTIN 


By GRAEME F. WATSON* & H. CARL GERHARDTT 


Summary 


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. 


Trascenods of the Royal Saucier at §, Awe (P9972, PIL VIM TI 


THE BREEDING BIOLOGY AND ADVERTISEMENT CALL OF LITORIA SPLENDIDA 
TYLER, DAVIES & MARTIN 


by GRABMP F. Watson & H. CARI GERIARDT: 


Summary 


Watsons, GP & Geniskor HC, (1997) The breeding biology and advertisement call ol Litnria splenditls Vy ler, 
Diyies & Martin. Trey. R, Sec. 8, ast T2104). 119-124, 28 November. 1997. 

Breeding biology and deseription of the advertisement call of (iroris splendida uve presented, Analyois of a 
calbol i. caerilen is also provided anid slows that these sitar, syalopre species have structurally siniikir calls 
wad thus presurmiibly show sinifegal yeoustc jolerachions in mixed choruses 


Key Worns; Lituria split, Laurie coerilec. logs. calls. breeding biology. 


Introduction 


Litaria splendida Tyler, Davies & Martin, L977, is 
one Of the largest fadult body length 82 - Loe ain 
(Hyler M82), und most beauaglul Australian frogs. 
The species is thought to be sparsely dystribuied 
throughout the widespread escarpment country at the 
Kimberley Division ot northern Western Australia 
(Tyler 1992). Literia splendida is notable for: 
hypertrophied parotid and rostral glands (Tyler & 
Davies 1993). a characteristic it shares with 
schescent members OF the closely related Litevte 
crerdteag an ability to ulilige ghindular seeretions as 
a Waterprool covermg (Tyler & Dayies 1993), and 
(he produciton of phirmacologically aetive caerins in 
(he skin secretions (Tyler & Davies 1993). 

What dite is known of the reproductive biology of 
L. splendida is based on reproduction by  cuptiye 
mdividuals (Pyler 1994), fn wo aquarium, a female 
laid 2000 eggs, depositing them in discrete clumps ot 
up lo 200 eggs. which Tyler (1992) has suggested 
may reflect.an adaptation oF females m the wilil to 
lay batehes of ezes in several temporary ponds to 
maximize the likelihood Of at leas} some of (he 
offspring surveying the larval stage. Maximum 
length of the tadpoles was 54 mm. (Tyler 1992), 

Literia splendid is Known to call. and presumably 
breed, alter heavy rams in the early wet season (C. 
Done, Department of Conseryation and Land 
Management Kununurra unpub. obs.). During the 
numerous field trips to the Kimberley Division 
undertaken by one of us (GPW) and colleagues from 
the Liniversity of Adelaide over the past 20. years. 
however, calls of the species have never been heard 
nor has breedimy been observed. Jn fact, the speeres 


Departnivntiot Zoplogy, University of Melbourne Parkyille View 
Wd, 

Division of Biologie Science diners of Visser (abumabye 
Missohin USA GS5201 


is rarely envountered except when associated with 
iuTificial habitats. for example ralet blocks. where 
CoOL, moist sites. iire readily available (Tylor 1992), 

Dining January. (997. we yistled the Kununurra 
urcu und en¢uuntered a breeding chorus of L. 
sendida, Here we deseribe the call of the species 
und provide brief notes Gn associated behawvieur, and 
alse a description of (he cull of syntopie L. ceerulec, 
au phylogeneticully closely related and ecologically 
similar species. 


Methods 


Recordings ot culls of Lo splendida were: ynade 
using a Sony TCD-5PRO cassette recorder (tape 
speed 4.76 ems!) and Beyer M-S& cardioid dymainic 
microphone. For comparative purposes, one call ota 
syilopic L. ceeruled was ohtamed trond a video 
sequence Of the breeding chorus (Canon Digital 
camcorder. Hi & mm tape). Air wet-bulb 
temperatures (the effective temperature of frogs 
sulling on land) were measured at the calling site of 
euch individual using an electronic thermistor 
thermometer (Takara Digimulti Model D6I1). 
Recordings were analysed on a DSP 5500. digital 
Sona-Graph (Kay Elemetries Corp.) using the in- 
built set-up #00 [Sampling rate (samples See!) 
10240. requeney runge! 4+ ky] with playback on a 
Nakamichi Dragon cassette reeprder, Overall 
variations in tepe speed (i.e, from recording to 
playback) are estimated ap less than 0.5% and 
frequeney responses of all audio-eleetronic 
components ure close ta linear within the relevant 
lrequency range (bused on manufacturers’ 
specications) 

Por each call, theee prigmary attributes: were 
determined: (1) duration, ws the interval from the 
beginning of the first pulse to the end of the lust pulse 
(ms): (it) number of pulses per note (direct count). 
and (ii) dominant frequency (Hz). as the maximum 


10 ti. hb. WATSON & HC. GERHARD 


vilue of the specteatn of power across the whale call, 
In addition, a derived characteristre, pulse repeliion 
tate (pulses 8). was caleulited as FOO) (irl 
pulses /duration ins, Levels oF resolution were less 
than | jis for tepporal eharacteristies and less than 
40 Ve for Trequeney, 

Because there is no possibility of misidentifying 
(hese large vod distinclive Frogs und in the iiterests 
of their conservation, voucher specimens of recorded 
jnales were pot taken, Video ahd 35-mmn- 
photographic records OF (he culling niles were 
obtuined, 


Results and Discussion 


Breediiw vite and breeding behuavienn 

Breeding and calling were observed, following 
hreayy aftemoon fain, On the pight of P81. 1997 in an 
uren Of sandstone escarpment adjacent to a darge, 
temporary pond near the main ear park within the 
Hidden Valley National Park, Kununurra WA. The 
pond was formed within an ibdefined watercourse 
(hat ran along the base of the cliffs and Wis fed by 
ton-off from a number of temporary waterfalls that 
How down the clit face after heavy rainfall. 
Occasional cally of males al other nearby sites were 
heard but chorus behaviour was confined 10 this one 
sie, Four species of frogs. Litorta splenedida, 1. 
caertlea. Le rebella and Limnadynastes arnciis, 
were culling aruund the pond. Amplectunt pairs of 2. 
slendida and 1. caerulea were observed in and 
around the pond. No pairs of the other two species 
were seen, ullhough no exhaustive search wats curried 
out, The Foumy egy masses of /. arnatiy were 
scattered over the pond as were farge. floating, 
single-luvered sheets of hylid eges, presumubly 
(hose oF Le splendida and L. ceeruled, although we 
did netobserve any pairs of either species depositing 


cogs. Nevertheless. contrary to the suggestion of 


Tyler (1992), on this occasion all L. splendida would, 
Of necessity, Wave deposited their entire eps 


complement in the One pond because Ho other 
aquatic Habitits were available in the vicinity of the 
vhorus, 

Males of 4. vplenedida called [rom exposed 
positions either on (he pean vertion) eh face or on 
natin! ledges upon the rocky surtice. The tye 
recorded males were calling approximately: 1.5 und 2m, 
respectively, above the pond. Several other 
individuals and amplectant pairs ol L. splendid 
were Observed in simiku positions, Males of / 
cuertiea called from similar sites oi the foek face as 
well as from elevated positions i surrounding trees 
und on the ground near the pond, Liroria vahelle 
culled from ground-level sites near the pond and 
Limnodynustes ornaluy called whilst Moating in the 
water, 

Although we did not observe pair-lormation ut 
ego-laying. pairs Of L. splendid coun remain iw the 
(rating embrace tor prolonged periods, at least Wp to 
94h, We observed several umplectant pairs silling in 
the open, or in cliff-faee ereviees, throughout 
daylight hours before the night of chorus wetivity 
deseribed above, Presumably these frogs had entered 
amplesxus during the previous night, 


Deseription af call 

A wave-lorm display and spectrogrun of the cull 
of L. splendida ave shown in Pig, 1, Table | lists the 
values of measured call allribules, The call is a fons, 
pulsed and apparently well-tunwd call that is 
rewulurly repeated (maximum call rate abserved wats 
56 calls min’), The call is broad-band but has a tonal 
quitlily because its relatively high pulse cate (which 
exceeds the Lemporal resolution of the human 
auditary system and henee our ability to detect 
pulses at this rate) is perceived us a comples tone, 
The call is characterized by a very slow rise in 
amplitude witha rapid cut-oll after nuatnui 
intensity is reached, The calls of both individuals 
Jisphiyed a number of frequeney peaks in the power 
spectrum (Fig. 2, Table 1) with-an iaterpeak interval 


PARED LoSnmeary of cll attributes af Linnie splendida recorded i Hidden Valle National Fark, Kannativrea, Western 


Australia 


Values ure baead yp analysis of five calls of two individuals, Frogs were calling on a sapdstone chil tree between 1S and 
2 ynabove a pond. Tempercitiires al the calling site were A,= 25.5" Cand Ay=25.1'C. Values for the first three attributes 


show the meant iid range (in parentheses), 


Call No. of 
Tridividuial Duration Pulses 
(mst 
Coe we bob, B28 
(25-703) (79-89) 
Cin #2 THAG $22 
(047-831) 74-94) 


Pulse Domiminil Other Notuble 
Repetition Frequeney Frequencies 
Rateips ) (Ha) (Ha) 

1238 {280 400, S20, H4t) FRO 

(1204-1 25,2) 920, 1040, 1160, 
L440, 1560, L680 
Has 1-H) 520, 640, TOU, BSU, 


(LOOK TTY. by HOO, 1260, 1520. 


1O40, 1760, THRU 


CALL OF LITORIA SPLENDIDA 121 


kHz 


100 300 500 700 
msec 


Vig. 1, Wave-form (upper) und audiospectrogram (lower) of the call of Litoria splendida recorded in Hidden Valley, 
Kununurra Western Australia, Wet-bulb air temperature at the calling site, 25.1" 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 ms on the audiospecirogram is an artifact of the printing process, 


Panne 2. Auribires of a veprescrnaive call ef Litoria caerulea recorded in Hidden Valley National Park, Kumunrra 
Western Australia 
Temperatures al the calling site were A= 25.5" C and Ay= 25.1° C. 


Call No. ol Pulse Dominant Other Notable 
Duration Pulses Repetition Frequency Frequencies 

(ms) Rute (ps!) (Hz) (Hz) 

210) 32 147.6 1440 440, 580, 720. 


1140. 1300. [S80 


[22 ©. T. WATSON & H.C. GERHARDT 


4 0 2 4 6 8 


kHz 


Fig. 2. Power spectrit of the call of Liveria splendida aeross (wo ranges of Frequency; 0 - 4 KHZ, Wo stow details of energy 


peaks and 0) 


of upproximiately 120 Hz, Many of these frequency 
components result from amplitude modulation of at 
harmonic series generated by the vocal cords 
becuuse the interval belween components is nearly 
identical to the pulse repetition rate. Because of 
resonating and filtering characteristics of the sound- 
producing structures of the emitter, some of these 
frequency bands are emphasized, particularly those 
uround 520 (? the Fundamental frequency of the call), 
640, YOO, 1280, 1420 and 1540 Hz (slight variations 
uround modal values occur because frequency values 
are measured in 20 Hz steps on the digital 
sonagraph), These spectral modifications make it 
difficult to determine confidently which components 
ure purt of the harmonic series and which are side 
bands arising from amplitude modulation. The 
frequeney band with most energy (dominant 
lrequency) dillered between the two recorded males 
(1280 for male #l and 1400 for male #2). with no 
within-individeal variation (within the resolution of 
this analysis) found in the five calls ofeach male that 
were analysed. Although the call includes speetutl 
energy peaks across a large frequency range (from 
around S00 ta 4000 Hy), little energy is present 
above 4 Ki (Mig. 2), 


Choris struenire 

Although only three other species were calling: in 
vhorus with J, splemdida, the chorus structure is of 
particular interest because the morphologically, 
bebaviourally and ecologically similur species. / 


4 kH4, to show that there is relatively litle energy in the call above 4 kHz. 


cderiled, Was a conspicuous component (Fig. 3), 
Auributes Of a representative eall of syntopic 4. 


caerifea are listed in Table 2 and a wave-form 
ot “wo 
ot z = 
= s 3 P: 
2 OG s Ee 
o a a] o 
g 2 5 4 
@ 2 g x 
3 3 a 2 
= 
/E 
“ ~ 
a | 
i} 


kilz 
~ 


secands 


Fig. 4, Audiospeetrogram of part of the ehorus ob toy 
species teeorded in Hidden Willey. Kununuira Western 
Australi. In (his reeardine Lira splendidi lis the 
loudest call, with backwound culls of f 
hibelhy and the very short cull of Linnie vinastes arn tay 
Eflective lemperatives for calling tales were: wet-bulb 
aie Lemiperatiine (2, splerutidia, th. ribettas 
25.1'°C gnd water temperature (2. aca) 26.47 


coeruled, 


coerulea Tt 


CALL OP LITORIA SPLENDIDA 134 


kHz 


100 


300 
msec 


60 
40 
dB 
20 
1@) T T t 1 
O 1 2 3 4 
kiIz 


Pig. 4. Waveform, audiospectrogram and power speetrum of the call of Literia ederilea recorded in Hidden Valley 
Kununurra Western Australia. Wet-bulb air temperature at the calling site. 25. L" C. Note: (i) the different temporal seales 
on the wave-lorm ahd spectrographic displays; (i) that the ordinate of the wave-form display is not labelled because it 


depicts & relative linear scale in volts, 


display, spectrogram and power spectrum are shown 
in Pig. 4. The call has w similar dominant frequency 
and broad spread of peaks of energy as that of L. 
splendida, but it is considerably shorter, has fewer 
pulses. a much faster rise time and less abrupt cut- 
olf, as well as a higher call repetition rate (130 calls 
min'). To the human ear the call of 4. caerulea 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 for significant 
avousue interference between them, 

Our observations of calling and breeding in 1 


splendide 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 Kununurra area in the 
preceding four weeks (pers. obs.). During our visit, 
heavy afternoon rains fell on most days and this 
simulus appeared to trigger calling and breeding in 
L, splendida, Successtul reproduction also requires a 
continuous aquatic habitat for larvae to complete 
their development and it is likely that L, spleadide 
will suceessfully recruit new individuals to the 


124 G. F. WATSON & H. C. GERHARDT 


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 
development would have been impossible. From this 


experience of the unpredictable rainfall patterns of 
this area, even in the “wet” season, it is possible that 
successful reproduction in L, splendida is a relatively 
uncommon event, 


Acknowledgments 


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 (S19711493). The work was 
undertaken under Department of Conservation and 
Land Management Licence No. SFO02003. 


References 


Tyzer, M. J. (1992) “Encyclopedia of Australian Animals. 
Frogs” (Collins Angus & Robertson Publishers Pty Ltd. 
Sydney). 

(1994) “Australian Frogs. A Natural History” 
(Reed Books Australia, Sydney). 
& Daviks, M. (1993) Family Hylidae pp. 58-63 


In 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. 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL, 121, PART 4 


SEA-LEVEL INDICATORS FROM A HOLOCENE, 
TIDE-DOMINATED COASTAL SUCCESSION, PORT PIRIE, 
SOUTH AUSTRALIA 


By E. J. BARNETT*, N. HARVEY*, A. P. BELPERIOT & R. P. BOURMANE 


Summary 


Barnett, E. J., Harvey, N., Belperio, A. P. & Bourman, R. P. (1997) Sea-Leyel 
indicators from a Holocene, tide-dominated coastal succession, Port Pirie, South 
Australia. Trans. R. Soc. $8, 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. peronii. 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. 


Transeo trons of ilte Revel Society ae S. Mast, (1997), F214), 125-155 


SEA-LEVEL INDICATORS FROM A HOLOCENE, TIDE-DOMINATED COASTAL 
SUCCESSION, PORT PIRIE, SOUTH AUSTRALIA 


hy E.L BARNETT. N, HARVEY, A. Po Briprrig’ & RP. BouRMAN: 


Suramary 


BARN LEO. Haewiy, No Bhbrekio, AvP & Bouman, R.P(1997) SeieLeyel indicators froun i Holocene, 
tide daminwed coastal suceession, Port Pirie, Sour Australia Trans, A. Soo. S. Aust, W204), 125-135, 26 


Novernber, 1997, 


Peritidal Holocene sediments at Port Pitie in the horthern Spencer Gull of South Australia comin several 
indieators of set-level change over (he Fast 7,000 years BP. “The clevahons of present subtidal, [arterial and 
supratidal envirvaments and eorresponding sediment facies were surveyed in order lo establish critical 
boundaries rehitive tothe tidal peer. The subtidal Posidonie ticles vecurs ator below meio low qatler spring 


(MEWS) Gide: intertidal sandflat, min 


grove dnd sumphire facies oecur aver specie miervals berweea MEWS 


Hide and mean high water spring (MHWS) ide. Back facies is clearly identifiable in the subsurface. with 
intertidal Sandflat facies particularly characterised by fa sifecarticulited bivalves Anepetla eyeleedan atid 
Katelysia scalar or K. perouit, A combination of several palucosea-level indicators from different lida Gigies 
best defines Weal sea-level change over the millennial timescale, 


Kay Wottos: Holocene sea level indicators. Uidal zonuticn. progridimg coastal sequence, fiers hoinilanes. 


Introduction 


Tide-diminued coustlines commonly generale 
prograding coastal sequences with excellent 
preservation of intertidal and shallow subtidal 
sedimentary facies (Belpeno ef af [988; de Boer er 
al PORK: Fleteher ef af. 1993), Such sequences can 
reveal Nigh-resolution records Of past sedimentation 
Often Containing a variety Ot palacosea-level 
iidicatoys — (Ferewsidt L988). A thorough 
understanding of the relationships of present 
Wdicators ak sea level, or invadition level, is 
required if correct iMlerpretations of pust relative sea 
levels ave to be achieved. With eritical appraisal of 
ihe present-day distribution of Titertudal facies, flora 
and fauna, pulawosea level history Tram subsurface 
steiraphy Gun be inove confidently interpreted, 

The northern Spencer Gull, South Australia (Pig. 
1) provides an excellent example of a wide, 
prograding coastal sequence in uw mesolidul 
environment with an identifiable eonation of 
Higloceue coastal Uepesitional cnyionments, “A 
number af coastal stadies has been conducted 
previously in this area (Firman 1965: Bure 1982: 
Burne & Colwell 1982; Belperio et af l984a,b, 
988: Gostin es al 1984. 198K: Norrish et ad. LOX6), 


Mowsoh Ciliates Cone fat Danieninenial Sduhes, GC iersily 
al Adlefnicle: And, SOUS, 

© Ponnerly Mines sid Gneegy Resouncus South Atserabi, POV Bos 
1S) hasewotl So Ana 5064, Civendy Minpiir cole fa 
Ghitetone St Pallarton S. Mast 8004, 
Seftow) of Liv itorntinenhilaintl Reeruadiony Minjageent. baealiy: ol 
Hoeeeriy and Devitonmvnt Pieveryity af Seo Atetealie 
Winrerti ub Mike LevelaS. eMiiah 804s 


In particular, Burne (1982) identified several 
important palavosea-level indicators fray beach 
ridges, the lop ol the subtidal Pasidonia seagrass 
facies and base of the intertidal sandflar facies. and 
Belperio ef al, (M9Sdb) demonstrated the presence ol 
well-defined boundary between Pasideariia seagrass 
and intertidal sandtlat facies. Related stratigtaphic 
studies in nearby Gull St Vincent inclutle those by 
Cunn & Gostin (1985), Belperto ef al. (1986. LO88), 
and Belperio (1993, 1995), At Port Adelaide in Gull 
St Vincent. Belperio (1994) confitmed that the 
boundary between fhe intertidal sanme{fal ane 
nuingrove facies was a relhible palacosen-level 
mdicator, Prom all these studies, it is apparent thet 
there tre local and regional differences ins the 
reliability and distribution of various sea-level 
indicators. This paper provides a critical appraisal ol 
the dilferent palueosea-level maulivatars ina mesotidal 
environment, 

The wide progruding sedimentary sequence of the 
northern Speneer Gulf region. which forms the apes 
ula large relatively shallow inverse (or negative | 
estuary, is a direct response to the modern coastal 
environment, Warm temperatures ind law rainfall in 
ihe region promote high mites ob evaporation utd 
salinities whieh are often higher than iwerige for 
seiwialer, in cacess Of 40% and as nybel ats 48 
(Bye T98T, Nunes & Lennon 1986). Seawater 
temperntires for the northern gull vary lypically 
berween (2° and 24°C (Nunes & Lennon 19k6), The 
tides are mostly seini-ditinal, with spring and peap 
hidal runges at Port Pirie of 3.8 im and 4m 
respectively. Die to the length of the gulf ana 
relatively show dea sealevel oscillations, wind 
stress can further jyercase the astronomical te 


126 E. J. BARNETT, N. HARVEY, A. P. BELPERIO & R. P. BOURMAN 


b Port Augusta B * 
32°39 Telowie Beach * 


AUSTRALIA 
ADELAIDE 


Weroona 
Island 


a | Le 4 
tt _ _ : 
Km | ____.138°00' 7 : 


Fig. |. 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). 


SIME. INDICATORS AT PORT PIRTE 17 


sieniticandy. Th Speneer Gulf and much af the 
southern coastal Australia (Nunes & Lennon 1980), 
there is a hinge spring-Heap tidal movulition die te 
Khe dearhy equal lunar ane solar semidiarnal 
cunsditvents causing 4 dodge lide onee every 
lortght when With: tidal variation gees. Sehlater 
er atl, (1995) have postulated that this simile 
wmnplitude ool the major semiediuenal  (idal 
constituents gives rise to particuhkie shallow qatter 
Hidal intereetions ti the upper gull, which promote 
sinphire and munyroave colonisation. 

Given the progradin Hdal sequence and (ie close 
prosimly of a tide piuge with reltible lony-term 
revords ut Port Pirie, this study was undertaken te 
deerme the vlevatiins OF goastal sedimentury 
vones velutive lo diode sea level and to jdeutit’y 
uppropriie jodern sediments asx anilogues ol 
Subsurface Holneene sedimentary lagies. This 
approich prayides sun opportunity to denuly the 
most reliable palawoseaslevel indicators iy a 
mesoudal environment aid ta develop a 
methodology for subsequent studies of relative 
land/sea movements at sites where bistorie clad lea 
valet. ‘Phe stidy is unigae tia that highhielits a 
Humber Of sedimentary facies, surveyed relative to 
modern sen level, ahd tlentifies critical equivalent 
iNnciwtOTS In the weologieul revord. Th does nev 
depend on one midieatar ti isolation, bul tises a suite 
of subtidal to supratidal indicaters 16 identify sea- 
level change. 


Methods 


Th Was vieeesstiry fo Survey the modern coastal 
covironments un detail to establish elevation 
Wilferences of the tidal zones relative (o local fidal 
datum. This wos done csi both basen and atomic 
spit levelling aisteumicnis. The first had ai ert ot 
Jess Tih | ennover 200 0), and neasivenients were 
kepl to within 400 in the, + 0,02 m). sine spint 
levelling. (he distance between each reading was less 
than 1OQ am, whieh weneraily kept vertical 
nehsurement errors to less than + O.O1 pi Taine 
based sites wathin the sandtlat, tmrave ane 
sabiphire Zajies were surveyed foo third-order 
Austin Plemht Datum CAHD) benchmirks. The 
supyeying wits conducted rnostly in the Pork Pirie 
eowstal Vicinity de Well ae Tuethee (athe noectheast 
within the Telowie Beaeh eoustal region. in order to 
decess all ol the modern-day tidal sethings (ig. 1), 
The present-day levels of the seagrass une sand thie 
gones al Port Pirie were veastred (ron water levels 
Mehabye to the Port Pie (de sauge., At Port Pine, 
ders Lidia dhithen (TI) is correlated! with (he lowest 
usiponumical Gide am related fo AID using i 
vorreelion of 1923 mi isurveyed | 7-03.10%4) Seaith 
Australie Ports Surberiny). Sea-level tdicators 


including seawrass. shell aid mengrove remains 
within und at the top of cach tidal zone were 
idenificd and recorded Tar later comparison with 
subsurhiee equivalents, 

In order to establish and simple the subsurface 
straliraphy, a totil of dhirty-live sites was selected 
within the broad coastal flats of Port Pirie Wig FA 
Vabrocorer was uscd to obkun cores 7A mul an 
diameter undp to 4m indepth At of the vibrocores 
were vorreeted Tur sediment eompuction by 
recording: penetranion depth versus core recovery 
length ane apply og correenon factor ta the 
thickness of the sedanents, Coring peripheral to anid 
Whi mangrove woodlands Was Carried cub Hsin a 
peut duger A baek-hoe wis used hy excavate 
scuimernts jit the sapratdal reson, Using this 
method, ny correchon for sanipling compuehon wits 
necessary. Surfieve and subsurface elevations in land- 
based vores anid excavations were surveyed to AMD, 
Murine-based vores were surveyed to TD uind taken 
Willi a few Kilometies of the tide giuiee to reduce 
the effects ol tidal lag and meteoralogical conditions, 
Samples were tiken back (othe laboratory, where the 
sediments were logeed with) particular attention 
being diven tothe elevations of facies boundaries and 
the presenee of sea-level tidreatlors. 


Modern depositional tidal environments in 
the Port Pirie aren 


The costal environment adjacent lo Port Pinte 
(Fig, 2) is a tidally-dominated lowland. Subtidal. 
Intertidhil and supeandal zones were distinguished hy 
the extent of marine influence or exposure and hy 
(heir vesetution assemblies and sedimentury ficies, 
Broad. shallow subtidal seagrass meidows pass 
laterally oshorewarnd frit ttertidal sands, 
miungrove woodlands, sarmphire-ulyal murshes una 
suplatidal evaparite Tat ehvitonnients, Chis 
association ab perhidal environments and their 
yegetition zones has. tow hinge extent, controlled the 
sliveessiae development al the coastal plain areund 
Por Pirie, A sehematie suminiy of the teal Zanes, 
assOciated vewetation and their pelaglopiships. to 
elevation OF (UNdaLON levels is given O) biguee 3, 


Tle Sublidtal zone 

Ii othe browd.  shallow-nniae envicaninent 
northwest of Port Pirie township, seugriss meadiws, 
composed lairwely ol Pavidertia auivieedis grow (rent 
around amcan low water spring (MLWS) tide level 
vere TD) 1a LO below TD (igs 3, 4), Aayteatio 
daorradéy eat only survive liiited periodas ol 
emersence so that, al is upper growih limik a is 
generally patehy am restricted i below 0.25 m TD. 
At depths below 4 om PDL Pesidemia sins 
dominates the seavrass assemblage, Mawiderlea heal 


126 bE, J. BARNETT, N. HARVEY. A, P. BELPERIO & R. P BOURMAN 


sheaths and rhizomes are resistant to decomposition, 
and seagrass fibres become incorporated into and 
bind 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 aceumulation of 


sediments in this environment, 


The intertidal zone 

The region extending (rom MLWS tide ta mean 
high water spring (MHWS) tide is defined as the 
intertidal zone (Fig. 3). This zone 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 


seayTusses cun grow ubove MLWS, Posidenia 
australis struggles to survive and is replaced by 
Zostera mnuelleri. Farther shoreward, bare sandflats 
are dominant, These sandflats are host to numerous 
epibenthic organisms including the intertidal 
molluses Barillarig sp., Veneridae sp.. Tellina sp.. 
Clanculus sp. Anapella cycladae and Katelysia 
scclarina or K. péronit and foraminitera that live on 
and beneath the sundtlat surtuce. These organisms 
occasionally accumulate in shallow tidal channels. 
Intertidal sandflats are replaced by mangrove 
woodlands above 1.32 m TD. Only one species of 
mangrove, Avicenna marina var. resinifera, has been 
recorded growing in South Australia (Butler e7 al, 
1977; Gostin et al. 1984; Cann & Gostin [985) (Fig 
5). Around Port Pirie, mangroves haye formed dense 


Fig. 2. Aerial photograph of the Port Pirie coastal zone. The subtidal and intertidal sandflats have been colonised by 
seagrasses (Seagrass meadows), Further inshore. mangroves form dense woodlands along the coastal margin and grow 
along dendritic tidal channels. Samphire communities occur more landward in the intertidal to supratidal zone. In the 
supratidal zone, the yexetation 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 ot 
Natural Resources, South Australia, Mapland, telephone (08) 8226 4946. 


SEA-LEVEL INDICATORS AT PORT PIRIE 129 


communities within clearly defined tidal limits. 
While the lower limit of mangroves is close to mean 
sea level (1.75 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 flushing of precipitated salts 
(Chapman 1975), A mangrove-algal association 
occurs at seaward levels of mangrove growth and 
along exposed tidal channels. Cyanobacterial mats 
also extend on to wide sandflats and into samphire 
areas in intertidal and supratidal zones. Numerous 
other organisms are associated with mangrove 


NW 


Subtidal 
Zone 


Intertidal Zone 


TD Elevation (m) 


woodlands, including the small mud crab, Helice 
haswellianus, which burrows into the substrate and 
promotes oxidation of the upper sediments. 
Gastropods, bivalves, polychaetes, 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). Sarcocernia — quinqueflora, — Sclerostegia 
arbuscula, Halosarcia halocnemoides and Suaeda 
australis are the main samphire communities present 
in the Port Pirie environment, followed by minor 
occurrences of Maireana  oppositifolia and 


SE 


Supratidal 
Zone 


(WW) GHV 


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: MHWS - mean high water spring tide 


Fig.4. Shallowly submerged seagrass meadow of Posidonia 
australis in the subtidal zone photographed during low 
tide, Width of field approximately 3 m, 


Fig. 5, Landward intertidal mangrove margin with intertidal 
samphire communities. Only one species of mangrove, 
Avicennia marina var. resinifera, grows in this southern 
temperate latitude. Dieback of mature trees along the 
landward margin can be observed, which generally 
indicates marine regression, The dead mangrove in left 
centre of the photograph is approximately 1.2 m tall. 


io Io, BARNETT, NOHARVEY A OP BELPERIO & Ro BOURMAN 


Hetlosareta indice toward the supratidal margin. 


The supraddal sane 

Above MEWS ude elevation (3.2 m TD). the 
supratidal zone (Pig, 3) 1s flooded on only the few 
vecusions when enher high or king tides combine 
with storm surge activity, predominantly from the 
soulliwest, or during and shortly alter extended 
periods of rainfall, Consequently. this zone is 
dominated hy evaporative processes und their 
associated sediments, In some ponded reas, algal 
mas we Well established und form cyanobacteria 
Hlats. Although this region consists miuainty of bare, 
poorly draining saline and gypsiferous flats. same 
samplires andsaltbushes survive (Fis, 7). OF these, 
Halosarcia heatloenemeides, Atriplex peliidese: 


Halosiucia welica wid Atriplex vesicciria are most 
thundint, Within this zone, variations in: elevation 
ive crewed by aeolian deflation and: formation vl 
eypsilerous dunes between remnunt tidal channels. 


hig, 6) totetidal samplire zone inchwing Sarencarnia 
quingqueflora, Sclerosteera crbuserla Heloseareica 
hadlocieiiaides ard Sudeda uatstrelis, The mangrove in 
the ripht foreground is gpprasinrately ooo lagh and the 
samphire bushes dre up to d0-enr in height, 


Mig 7, Supratthil samphire gone Tneluding oocascopal 


Hirlowipeda Tretlacnemotlen, \iriples Paluadase, 
Hiloyearier theica and Anipfes Yestera und hire 
exphinses of Salli, The simphire species are upto son 


iW herght 


Luneties hive also formed on the leeward margins of 
sabkha flats or salt lakes. 

The distribution of coustal environments around 
Port Pirie is shown in Figure &, Six distinetive tidal 
zones IWansecting the cost have heen identitied 
From seaward to landward, these are; 1) subudal 
acuprass micadows (not shown in figure), i) low 
intertidal bare or Zesrerg-covered sandflats. jd) 
intertidal mangrove woodlands, iv) high intertidal 
samphirealgal marshes, v) supratidal evaporative 
flats, and vi) supratidal ind extratidal clay and 
vypscous dunes and lunettes. Aerial photographic 
interpretation of (he mangrove woodland reveals that 
only minor change i its distribution ts apparent for 
the last 40 years or $0 (1957-1993), Mangroves have 
prograded seaward ito intertidal seaerags/sund (lal 
areas on the northwest peninsula of the Port Pirie 
River, between First and Sceond Creck and along the 
margins of he Port Pirie River itsell) This isin 
contrast to vapid seaward Mangrove colomsalton Ubi 
has occurred al Port Gawler (Cann & Gostin 1985) 
and landward colonisation in the Port Adelaide 
revsion (Burton 1982: Belperie 1993). 


Kvidence of depositional tidal sediments 
in the subsurface 


Much of the sedimentary stratigraphy at Part Pie 
represeats aggradation and progradution ol 
sedimems in peritidal environments since the hear 
stabilisation and slight fall in sea level trou) 7.000 
yeurs BP lo present (Belperio 1995), Holovene 
sediments and Pleistocene alluvial sediments of the 
Pooraka bormation underlie most of the area, 
forming an undulating boundary with the overly tig 
tidal sequence, In some places, the upper sections of 
the Pooraka Formation show evidenee of being 
altered or gleyed by marine parewuters. The coastal 
sediments record an upward change jn sediment 
Facies that vorresponds with the lateral olamee i ike 
tidal zones 

The subtidal Pasfedenia facies is Wie Wiest extensive 
Holocene Udal facies in the region, Th consists of 
mostly grey, poorly sarted  terrigenous. and 
caleareous sandy mud. with numerous fibres of 
Posidania australis and Vragmentary molluses (ee 
Spisule osp., Phasianella sp. Cantharidus sp, 
Dosinta sp. and Batillaria sp.) and foraminauera. Its 
thickness varies from greater Wan 4 man the present 
subudal zone but thins inland underlying intertidil 
and supratidal sediments to belween O und 2 m, 
depending on undulations in the surface of the 
underlying Pooraka Founavion. The landward extent 
of this Tietes indicates that much of the present 
coustil environment was a shallow marine 
environment during the early Wo mid Holocene 

The intertidal sundial fieies is warrey Co listht yrey, 


SEA LEVEL INDICAPORS AL PORT PIRIE WI 


poorly seried, terrigenous and calcareous shelly 
muddy sun fe occurs extensively inland beneath 
much ol the study area Having developed im response 
to upward showing of the subtidal sedimentary 
environment. Tn mest ool the Port Pirie region, the 
intertidal sandflal faeies is overlain by sanphire 
lucies. This is im contrast io The presentaiey: cidlal 
A0malion Where a transition fronn sandflit to 
Mangrove woodland generally oecurs, 

The intertidal mangrove facies consists of brown or 
hluish grey. mostly fonedleuredus sedinwnrs with 
fragments of roots. sheaths und fibres. tis largely 
resincled lo (he present diy distabution of mangrove 
wondlands, Le progriadational deyelapment and 
preservation of strata tive heen limited and 
Aviewiia marina sain resindere woodhinds upped 
to fave developed i telatively recent times, Where it 
is undeveloped. modern mangrove cools penetrate 
inte the underlying Facies, 

The sediment facies of the sample far Loris ut 
(hin veneer Over extensiveareas of sundtlac facies at 
fhe coastal plain, Ik consists of pale browat to yght 


33°05" 137055! 
a Ht 
l 1 i i [ J 
henry 
Spencer Gulf 


SSS eee... = dt 
ST KILDA FORMATION. (UndiHerentialed 
Holocene marine and coastal marine sediments) 


Supratidal anc extratidal clay and 
gypsum dunes ond luneties 


WH Supratidal flats. Gyps@ous dlays 


JI Stranded beach ridges and coaslal dunes 
Shellgrit, shally sand and line sand 


Samphire-algal marsh 
Garhonale anc lerrigencus muds 


arey, Often Mottled caleaeous and tertyeenotis clay 
nich muds with occusional small gastropods, biyalyes 
wid foraminifera. Sow) plant fibres and thin tubular 
robles are upparent in some regions hut absent in 
others, dependiag of whether plot rater wats 
originally present ami/or preserved, Gypsuai content 
is variable. due largely to elevadion and evaporation 
History, With eypsurende dune sediments preserved 
atthe highest elevations of the supraddal cone. There 
is liltle clistinction between rarertidal and Sapa 
sumphire sediment facies, und the two are considered 
to form a single unit. While particule samphire 
species gan be identified grawing in wither the 
infertidl or supratidal gone, in the subsurface, 
sumphire roetlels und remains gunnel be dentitied te 
speeies level, 

In addition to the sediment facies ubove, several 
(HCPOCHVTOU MONS Of sUBLaeies Beele it the region 
(hal tive Conterponiian arilogues. In particular. 
pockets of cyanebacteral fieies are evident 
Vhroughout the intertidal ti supruodel ones. 
Wherever cyanobacterial Nuits ape present in the 


| 
438°00' 3 


Germein 
Bay 


Mangrove woodland. 

Organic, terrigenous mud 

Bare or Zostera-calanised law Intertidal sand 
or mud Hat, Mixed shell and quartz muddy sand 


= Organic/shelly sand/mud of mangrove 


woodland and sampnhire-algal marsh 
ABC Range Quartzite 


Pie} Cristal cofaey mpi ite Pon Preis eoriplivg) ron the South Avacaliie Geotoey Gutubuxe Mime anil [ney ay 


fesuurces South Aust ealit, 


[32 Ei. J. BARNETT, N. HARVEY, A. P BELPERIO & R. P ROURMAN 


intertidal or supratidal zone there are active sites of 
sediment aggradation, In the intertidal zone, storm 
ridge facies, or Cheniers when developed over muddy 
sediments, haye been formed during periods of 
combined high or king tides and storm events, 
Ridges are generally aligned parallel to the shoreline. 
Only one storm ridge is preserved in the western Port 
Pirie area, although several others occur to the cust. 
The northerly orientation of the coastline yenerally 
protects the area from dommiant southeasterly slovm- 
ridge forming events. 


Palaeosea-level indicators 


The specu significance of the northern Spencer 
Gulf is that the peritidal coastal suecession eantains 
a Well-preserved tevord of pulacoseu-level change, 
The sediments include various palueosea-level 
indicators that hive heen used, with uppropriate 
elevation dati, 16 reconstruct palacosea-levels, 
Although present-day tidal environments and 
equivalent sediment ficies may range over 
simificaot vertical elevations, the contuet between 
each sediment facies is generally more restricted, 
Subsurhice facies contacts can provide relatively 
precise estimales of pulaeosea-levels. given geourate 


Tidal 
Datum(m) 


‘Supratidal 
flat facies 


Samphire-algal 
facies 


Mangrove 
facies 


Intertidal 
sandflat 
facies 


Subtidal 
Pasidonia 
facies 


Vio. Palaeoscu-level uriteria for the Port Pirte Coastal renin, 


D-E contact 


C-D contact 


B-D contact 


B-C contact 


A-B contact 


surveying: of the vertical extent of present 
sedimentary facies and thei’ contacts, Once the 
elevation range of a particular sedimentary contact is 
known, a height correetion for that contact can be 
made relative lo present sea level. This establishes 
the elevation at the time of deposition and indicates 
whether sea level has subsequently risen or fallen. 

We have established that, in the Port Pirie area, the 
boundary between Posidonia facies and overlying 
shelly intertidal sandfat facies provides a palaeosea- 
level datum corresponding to an upper limit of 0.25 
+ 0.25 m TD (Fig. 9). Consequently, the subsurface 
occurrence of distinctive, massed. fibrous Poyidonta 
facies in Jand-based sediments at elevations higher 
than 0.25 + 0.25 m TD implies that relative sea level 
was previously higher than at present, 

In a similar fashion, the intertidal sandflat facies 
generally occurs between 0.25 and 2.2 m ‘TI relative 
lo present-day sea level, However. a more precise 
palucosca-level estimate is provided by the sharp 
contact belween intertidal sandflat) facies and 
overlying miunyrove facies that cyuates to |.32-+-0.2 m 
TD (Fig. 9). At Port Pirie, the mangrove facies 
mostly occurs directly beneath the present mangrove 
woodland, und confidence in using Us contaee with 
the top of the sundflat factes is realest where 


Contemporary 
Elevation 
(Tidal Datum m) 


General Dascription 
of sediment contacts 


Change frorn calcareous clay to gypseous, 
300.4 clay-pellet, structureless sediment 
Change trom organic, rooted, peaty clay 
26+0.4 to cream calcareous clay, laminated to 
weakly rooted 
2.2+0.5 Change from coarse shell sand or 
coquina ta cream, nalcareous clay 
132+02 Sharp change from coarse shell sand or 
a coquina to organic, rooted, peaty clay 
Sharp change trom poorly sorted, fibrous, 
0,25+0,.25 shelly sand and mud to cleaner, better 


sorted shelly sand 


SLA-LEVEL INDICATORS AT PORT PIRIE ae 


niussed wrteuliwd wadves of acelin credudie and 
Ketelivia sedlariic ork. perdi ure present, 
INUICALE frevete Post-norteny, preservation Where 
(he marwrove lictes 4s bseol, the contiel bemvcen 
the sundflat und samahine facies ts also stip, 
althoigh the present-day boundary herween the 
Heol sali gund Sgimphire Zones ds not well 
delined in the immediate wieiiity oF Pork Pirie, The 
upper lait ol the sindthial facies with samphire 
facies geeurs around 2.240.5 nm TO (Pig. 9), 

While the present Port Pirie tidal zones extit ia 
triosilion from mangrove woodlands to sample 
nirshes at 2-6 © On TD, this as nol commun 
Observed ih the subsurfuve sediments due tr the lack 
oF proseqdypondl development of this: strutiaraphic 
horizon, Consequently, the leyel at whieh the 
sunphire facies oecurs in the subsurface proves 
only an upproxmmite estamite of palacasea-level 
relayive Wits present elewauenr range aF 22 be A on 
TH, Although different samphite species are closely 
related Lo small elowition chanees, these are nyt 
bbserved at the mucro-level jn ihe subsurhice. 


Discussian 


Several factors must te adilressed when 
inturpretiny the evidence for palieusea-level change 
fron progetcding. peritidal sequences. In purteular, 
the relationship of ewel) indigalor ju sea level at the 
fime ol ils formation must be established. Al Port 
Virie. the subsurface presenee of iv sie fibros 
renwins OF the Scagrass Puyideie austradig indicates 
thar sea level was above this site ut the time of 
deposition, “Phe (ausitipn between Posie bacies 
und overlying intertidal sundllat facies is a more 
powerlulindicutor ol paulaeosea-level. vurrespondiig 
i) O95 +4 025 wy presentalay TD. Siilurly, 
mingroves grow will a Rurky broad: intertichal 
range, but their contagh well the imlertidal same 
facies provides durum of 132 = 0.2 m. Mangroves 
have previdusly heen cited as one al the more 
reliable figed, vie site palacosed-level indicutors 
(Hopley & Thom 183, Thom & Roy (983). For 
northern Spencer Gull, Burne (1982) reported: a 
range opm the elevihioo of seaward thmgrove 
colomsation from ES to 2.9 mM TR (0.4 ta 10m 
AHD) und us previously mentioned, we record a 
lower level ob mangrove volomisutionat 132+ 0.20 
TD. Clearly, the level of seaward colomsaton of 
mangroves depends prmnwily on local coastal 
dynamics or coustal orientation, and will ovcur a a 
vuriely al clevulions relative to the tidal spectrum 
(Allen 1995). Therefore, i follows that the height of 
the contac between sundial und mangrove liwies 
will also vary. IL 1s apparent front the differences in 
elevation that the ise of mangroves as palilensed- 
fevel midicators can only be applied locally. where 


present-day clevalions uf mangrove seaward erowth 
we owell defined Even in’ this case. the 
palacoenviranment muy huve differed from the 
Moder environment, producing different teal 
ninges and mangrove distrbations. 

The boundary between samdflal and either 
Hngtave or samphine facies has the potenchil te 
define palacosea level. purtivularly singe its contiel 
in the subsurface is sharp. Hewever, the conmunelrtiin 
a Port Pirie is thik while the present-day zonahon 
from sandilar to mangreve waedland is exrernsive 
his (ransiian ts not easily observed in the 
subsurtace, Furthernore, while the present-day 
Lainsilinn belween sundial aml samphire zones is nol 
well tepresented iit Port Pirie, tis eowraee ii 
subsurface Sediments ts Widespread, Near Port Pirie, 
the present cleyalion ob the supdlflat/samphire 
houndary is 2.2 + 0.5 pm TD. To the northeast il 
Telowie Beuel, Unis boundury oewurs ue 2S 23m 
TD, a slightly higher elevation thaw for Port Pine 
possibly due to local geomorphic factors ane 
sedimentary processes in the lee of Weeroona tstand 
(Five. 1). The elevation of the top of the sand 
vanes depending on whether it is suceceded by 
mhwwroves of samphiee, Thus, athouely the conmuet 
can be used as a determinant of seci-level change, 
there ts wide range in ils elevation. This problem 
niuy be mimimised by curelul field surveying of the 
local region, 

Beach ridges and the lop ool Payidonuios seag rigs 
deposits are relatively good indicators oF pulaeossi- 
levels. However as wilh imungraves. beach ridge 
clovition dig) cunnot he used on a reg tonal busts since 
the elevations to whieh such ridges aire constructed ure 
highly dependent on local wave resimes, In regard to 
seagrass us a-sedevel indicator, Posidenta anstrals 
presently grows to 0.25 in TD (1.68 m AND) at Port 
Pirie, but elsewhere in northern Spencer Cull, on 
clevalion of nt mW TD (2-2 m AHD) fas beer 
observed (Burne (982), These differences muy he best 
explained by varying Comskul oriengiion, waye regime 
and coustal circulation patterns. Although a shore- 
parallel zonation of sediments and vegetition is 
common throughout the harthern sult, cach sediment 
and floral or fauna) community, whether Mosidenia- 
dominated seagrasses. Anapelia or Katelysig sp. 
bivalves, mangroves or hulophyles and silthushes, bas 
a broad regional range in elevation, Henees it ts 
impenttive that Toca! elevation eontrals und docu 
coniitions be used in assessing sei-level dala rather 
than applying regional values, 

Given that the cleyation range of bidi) facles atid 
sea-level indicators cur vary, vrealer aceurtey: on 
reconstruction of pulacesed-level is aehieved if 
several different indicators aire used. Each muicator. 
either relational or fixed, will provide evilence thut 
either Sapports or challenges andicators fram othe 


134 E. J. BARNETT, N. HARVEY. A. P. BELPERIO & R. P. BOURMAN 


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 Holocene, but rather, the region has 


been uplitted 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 littke 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. 


Conclusions 


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 subtidal Posidonia and intertidal 
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 Holocene 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, Sarcocernia 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. Jn sin 
articulated shells such as Anapella cycladae and 
Katelysia scalarina or K. 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 + 
0.4m TD. 

Good precision in palacosea-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 palacosea- 
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 the 
region. 


Acknowledgments 


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. Fotheringham 
assisted with surveying and vegetation identification. 
Access into Pasminco Metals/BHAS was granted by 
A. Gilbert. The aerial photograph was supplied by 
the Department of 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. 


SEA-LEVEL INDICATORS AT PORT FIRIE 135 


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THE RESPONSE OF GALL-INDUCING SCALE INSECTS 
(HEMIPTERA: ERIOCOCCIDAE: APIOMORPHA 
RUBSAAMEN) TO THE FIRE HISTORY OF MALLEE 
EUCALYPTS IN 
DANGGALI CONSERVATION PARK, SOUTH AUSTRALIA 


By P, J. GULLAN* 7, P. S. CRANSTONT* & L. G. COOK* 


Summary 


Gullan, P. J., Cranston, P. S. & Cook, L. G. (1997) The response of gall-inducing 
scale insects (Hemiptera: Eriococcidae: Apiomorpha Riibsaamen) to the fire history 
of mallee eucalypts in Danggali Conservation Park, South Australia. Trans. R. Soc, 8. 
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 Riibsaamen, 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 1n 1985, although the two 
commonest Apiomorpha species differed in their responses to fire history. 

Key Words: Fire history, mallee, galls, Eucalyptus, Coccoidea, Apiomorpha. 


Tran neny afte Bowel Socrery ef. Avan 1997), ERTL LTT Ul, 


MALLEE EUCALYPTS IN DANGGAL] CONSE 


THE RESPONSE OF GALL-INDUCING SCALE INSECTS (HEMIPTERA: 
ERIOCOCCIDAE: APTOMORPHA RUBSAAMEN) TO TILE FIRE HISTORY OF 
RVATLON PARK, SOUTH AUSTRALIA 


by PL GuLLAN *. PLS. Cranston’ & LG. Cook" 


Suiinary 


Guitan. BI. Cragston. PS. & Cook. LG (007) The resprnke of gallanductiy scale ciseets (Mennytera 
Lrovoccnlie: Aprominphe Riibsaamen) to the lire history of mallee cuealypis in Dungwali Conservatron Park 
South Australian Tray. WN, S Ads. TBA), 137 P46. Ts Novernher 199? 


Mullee communities. espechilly the pliant components. dre often considered! to be fire-adapted bur there tne 
iiformating on how effectively any phytophigods pisces re-establish Then populations alter a wilttire We 
adilressad this issue hy studyie the sedle insect genus Aprmoernle: Baibsuamnen, in whieh species midltice 
CAfspiclious. SeSOdHY CIMA phe Balls Of species specitic morphology on Avcrvpnis species. Tt early (990 we 
surveyed Thespecios richness anid abundance of Apiearacenyaltes aaulls Wi relaticn ta fire history and species al hast 
encalypt in mathe veretilion at Pangea) Conservation Park, South Australia, bbullol the Fourteen stiles surseved 
laut heen burnt by waldfire in tate [9SS, whereas rhe orher seven sites Hate por heen burnt for al least 45 years, 
Only the iyo caummonest af mine Aviiimeepii spemes showeel little or no Host-plint specsienty. Leue-unburit 
sites id nat difler in speies rightiess ner in tani abundinee Of Apremanpa palls frond sites huent in Tess. 
clLodeh (he Avo Canmenest ieaiapla species differed in their responses to fire History. Gully of fb. 
metecucoly Gutlan were equally ubundgat ul long-unburdt and burnt stles. whereas valk of ovicelandtes 
Hepper) were on sverage four dimes more abundant al lome-unburnt (har al burnt sites; this differences nity relate 
ty Uiflerential dispersal steatewies of che (st cistur nymphs. We conclide that mallee willfires sb intervals al 
Hoare thi 10 yours wauld be unlikely to impact detrinentalby ba the long-lero survival ofa uliwerse A ptirneipilis 
assemblage provided That some longer Onburn’ areas (rerigi) remain to serve as souree’s far the calonising 
ny tiplis 


hy Worps: Dare history, nmaitles gulls, Eaenhipria, Cyceailew, Aprenoryltia: 


Introduction 


Tire is i significant factor i Australi ecolory 
and inmuch of the continent the flora is fire-adapted 
(Barlow JRL) Many ophints, tiecluding inst 
Bicalyprs Lo He, species (Myttaecic), possess fire- 
protected structures (epicormic buds or Nenotubers) 
from whieh new growih sprouts, or have seeds thal 
serminite aller fires (CH MY8 tab: Hodgkinson & 
Griffin 1982: Noble 1982) Sueh regeneration 
ubihties ure purteularhy characteristic ab pints i 
mallee vevetulion, thal is. woodland eormmunities 
Uonhimited by imultiestenmead cuealypts, which are 
themselves also called mallee (Noble 1982) Kor 
inuny cugulypls, espeerlly millees. fire cua fer itate 
seed germination (Wellington 1989) or elimiale 
phytophigous tisects (Noble 1482) une parasitic 
Inisileloes (Call (Yb 1a) Gucalypts dominate most vl 
Australis forested ecosystems ind have cormples 
rehwonships with mative ands. ineluding many 
iisects (Greensinde & New 1991). Habind 
MaaeeMent using five whether Comastent wath 
History (ee. lo reerese purported past Aboriginil 
lind mitnuwement practice) or far havard reduetion, 
has effects (hat are rehitiyvely well understood fo 


Iyision of Motu aid Avcloy. The Suxtrahiin Mater 
Ciel Cadberna ACT 0200, 


' Suinath Rewuntoes amb Biewliversiey Programe Bevisiony at 
Dtoiintagy CSEROEGIPOD Hes DITO irbotie wel Tne 


vepetulion bub the implicutions foe many other 
portant Organisms, especially invertehrites. wre 
largely unknown (Bradstock ey af, LYYS: Friend & 
Williams 1896). The few Australian Studies of the 
effects of fire on invertebrates have coneentrated on 
soil and littsmdwelling organisms (ee. Campbell & 
‘Tanton TY§ 1; Neumanis & Tolhurst 991; York 1994) 
or have sampled a wider invertebrate ussemblige 
usm only pitfall traps: (ag. Prend & Williuns 
1006), Arboredl insects probubly survive fites less 
well tha epigcie and Hypogeie speeies (Whelar 
1995) und, provideck sampling biases can be 
addressed. should be good candidates For study ine 
the cflects of fire on dnveriehrate populivons. tn the 
presehe sty we examine the effects of fire history 
on cndemic Agsttahan scale aiseets (denmiplera 
Coceoiden) that five only fa the caopy oF cuealy pts 

Seale insects of the genus Apiemearpha Rilbsaumen 
(hriococeidie) Tive within galls That they induce on 
iheir eucalypt hosts (Gullan 1984acb) | tn 
Apioniuphe. couch adult female resides ia large. 
oflen symmetrical und woody gall usually on the 
stem but sometimes on leaves, buds or traits of the 
host cuealypt (Guatht 2984), These are easily 
recognised as Cocco walls because there ts a small 
apical orifice through whieh the female can chiminidte 
her waste honeydew and also mate with the rate 
Her offspring or firstinstur nymphs. eulled crawlers. 
inake thei exit from) the muternal gall dhrough: this 
sine apeniny and then disperse fo inithite new eulls 


{3K PUCOLLLAN. I SOURANSTON & LG, COOK 


on sullble foliage, The sadls of tales ave much 
smiuler than (hose of the females, rarely more than 
une centimetre Tong. and are tobolae with an apioul 
orifice unl sometimes un Gulwardly-dirceted Tange 
ab the apex, The shape of the gall of the female ts 
pecullir to lhe Apiemoreia species thal induces 
revardless ofthe identity of the host eucalypt, Most 
Aplomorpha specics show sane degree ol speemieny 
Wea restriged range Of Encalyptis species (Galhin 
ital Apiamorphe hus 39 deserthed species 
(Cjullan 1984a; Gullan & Jones 1989), each ol whieh 
Haims Chavaeterisnic., sexudly-dimorphie galls, 
Generally. Apdomarphe can be identitied Lo species 
Jevel in the field, even by Hotespeehilists with 
minal Grass, 

Seale inseels are otlen claimed lo have low yagility 
(lor reviewosee Hinks & Denno M99) heeause the 
wingless adult females usually spend) thei entire 
lives ona side host phint and oviposit thece, adult 
males are short-lived and weuk-fMving. and all 
dispersal is duce to the movement of the vrawlers, 
Which dsually remain on their natal host (conditions 
une Havourible (Gireathead (990: Nested eral, 1995), 
Ih Apiomer phe. our held observations ih sclerophyt 
wowthinds have sugeested (bal there is lower 
diversity and abuidanee of galls io aes sabypeetcd te 
Hequent or severe burning, This reduetion seers 
Linrehited to the suitability OF post-fire enealypts for 


gall development, since the epicormic Push oF 


foliage that follows a fire resembles the preferred 
plant material utilised by Apromenphe in leony, 
inburnt areas. His more Tikely that afer a fire kills 
the galls and their occupants, i takes time for 
reenlenisation by erawlers fe oecur and if fires are 
irequent, OF potential sources of colonists ate distant, 
locul populitions Wiiv Wet resestiblish. This 
hypothesis. renwuis speculitive at the ubscnee of 
quuntilication of ny differences in species diversily 
wid ubundidmee between bunt wad) lange cnibernt 
ureus, We fovestinated re-colonsation alter lire hy 
surveying the species rivhness and uhundunee ol 
Apromaepha it rclatien to both the fire histary ane 
the species OF host eucalypt in mallee vewetution in 
South Australia, 

Thos study was undertaken tn mmilee for several 
reasons. Firstly. mutlee is a) typicully Australian 
vegetation that has beer ti seclous deeline for the 
ust 150 yeurs threugh kd elewranee und other 
forms of degradation (Land Conservation Council 
1987: Cheul JO8G Harris 1990) Secondly. it his 
heen hypothesised that Mallee plant aud) animal 
commniies are muintiined by episodic fire (Noble 
1982. 1989) hut no research has been done te 
investigate how efecuvely any phytophapous msecey 
peestablish their populations in matice aller fire. 
Thitdhy, itis easier to count the galls on mallee 


ehealypts Wan on the Giller forest ang woodkind 
species. 

Seale disects of Apiomearpla are good canchdites 
for studying the effeets of wihtfire because, unlike 
more mobile wsects (Whelan ered 1980). they are 
fob ubleold move From the thair host plants to aver 
the Tames and. although the thigk Wall Ob the galls 
Hits been suggested loa be an adaphition for fine 
protection (Koteju L986). the bit intensity of mullee 
wildlires. usially hills the overstorey  lobiage 
(Bradsivek 1990), Ty Gontrist ta the wellek iow) 
banes of pilhalling (rapping and other methods ol 
sampling soil-or fitter Hain (see Whelan 1995). our 
subvey method lor prone potentially assesses 
Ihe tanh nuinher oF galls on each tree Further 
Advan tiges of the Use ab Aponte galls bs that 
their ubundinee vires Tile wallh season of survey 
singe (he Temules of most speaies probably live tor 
much longer Thun a year (L.A. Cook unpub.) une 
walls can remain on the trees for several years: (ue 
the death of the occupant This providing a recon ol 
the presence of (he species ul a site. Lastly. in order 
to elucidale | eausul relationships berween 
iwerlvhrate abundanee patterns and: fine, scleetoel 
invertebrate groups need to be examined at a tines 
level OF taxonomic resolution (han the order oy 
family level that ts used inmost studies (Pricid: & 
Williaits 1996). Foom this perspective. apioneeplre 
is a ideal study genus because gulls can he 
identified readily to speeies level, ever ny We Geld, 


Methods 


The work Wis curled out ih Danggal) 
Conservation Park whieh wits estiblished tn 1976, 
heciine Australia's Mist Biosphere Reserve in (077 
andis now parcul ihe Bookmark Biosphere Reserve. 
This quarter of 4 million heewire reserve js about 90 
kin north oF Renmark in South Australia and tes in 
the northern halPor the Murry biasin, adjacent (or the 
New South Wales border (Pig, UT), tb inelides hol 
arid ind mallee lam systems and allows aevess (4 
some speclicnlar old-growih wallee. OF purtientar 
sivnifieance ty our study is the doeumented (re 
History of Danggali Conservation Park (rom 
National Parks and Wildlife Service, Seutlh 
Aust. File Tis Hol been tiscd as a puapennent 
tool th the inaintenwnee of tis mallee for 
egoservation: recorded wildlires either have heen 
mulurally-ovcucring, Tollowing lightning strikes. or 
the result oF burma acuents, Many areas have niu 
evidence of burning for u considerable perid 
perhaps for over a century, Or ul least noLsinge either 
JOP? or [951 when extensive wildtifes burn muel ol 
the region, (loweyer, of special releyange loo the 
prescnl stinty isu inajor lighining-induced tire thin 
burnta huge eential area of the reserve i December 


GALL-INDUCING INSECTS AND MALLEE FIRE HISTORY \39 


no 


SECU HO ce eee eer) 


PU te “ we ~ os 
"Morgan Vale" 
Ruwis 


eT 
Mt Morgan 


AUSTRALIA 
WALES 


UU PUREVINUULEEUEADO LAAN Hain ne 


SOUTH 


SOUTH 


NEW 


HUTTE 


“Cangpus" 
Homesicad 


FT Re 


7h 
fn, “Aypurna 
th, 4 
“eg Homestead 


mn Park boundary 


— Road or track 


@ Landmark Bayi c0d 10 OHH UTU OHNO EELAAU PRAHA LANA nan 


@ Study site 
Extent of 1974 fire 


i Extent of 1979 fire . “mn 
0 5 1a 20 
SEE | 


fea Extent of 1985 Fire 


Fig. 1. Map of Danggali Conservation Park, South Australia, showing the extent of recent wildfires and the locations of the 
14 study sites where galls of Apiomorpha were surveyed, Inset shows location of Danggali Conservation Park. (Figure 
based on map and aerial photographs from the National Parks and Wildlife Service. South Australian. 


eH) 


1985 (Fig. 1). Since then, (he burnt eucalypts live 
regeneruled from (heir mates Hanotubers, although 
the dead stags of past sdbstuntial limbs still protrude 
from the now-lMourishing mallee teerowth, 

We made conparisons belween the Apienerplee 
galls associited with eucalyprs in the old-prowth 
mallee (“long-unburnit’ sites) and in the mallee that 
had regenerated after the L985 fire Churn” sites). 
All Fieldwork Wits conducted in April 1996. Fourteen 
sites were chosen (ig, Ty seven iy cach treatment 
type (burnt versus long-unburnt), with the site co- 
ordinales as given in Table 1. All enealypts at the 
burnt sies had been burnt completely inthe 185 fire 
tind sites 6-9 were completely surrounded by post- 
fire reveneraled mallee. We only used burnt sites 1 
the eentral aren uf the reserve hecause Wwe were 
vertuin dhit vegetation in this area had been 
vompletely destroyed in 1985, whereas the 
Veeeuilion around “Tipperary Dany and “Morgan 

fe Ruts and in the owe smaller areas also burnt 
i F985 on the periphery of the reserve (Fis, 1) 
appeared 16 hive heen wore patehily burnt Five 
hiirntand five longeunburnt sites had an understorey 
Wilh porcupine gruss, Trindia irrirany Re Be. 
(Ciramineae), sometimes ay the dominant ground- 
laver plant. Che ater four sites either had reasonably 
hire Shon Ora mixtire of shrub species, 

The jiallee cuculypl species al the study sites were 
Hucelyplas incraysata Sieber ex DC. (syn. 1. canine 
Pm Muell, & Bel ex fo Muell.), & didmayea Cun, ex 
Oxley, 2 gracilin T Moell,, Bo aleosa P Muell, ex 
Mig, dnd E. seciadéiy b Muell, ex Mig, Ideniilication 
Wis Indde using Costermans (1994), except tat the 
name E. (erassetr (syn. 2. cayiefe) is retained (MLL. 
Brooker pers. comm), The prapertions: of each 
species varied aecording to locality but usually three 
ol the five mallee enealypis were present at ull sites 
(Table 1). We chose sites i dong-unburnit or post fire 
regenerating patches to provide 10 multi-stemmed 


12, 


PT, GULLAN. BS. CRANSTON & LC. COOK 


inallee eucalypts. of between 3 and 6 min height, per 
site These 10 trees were chosen us representative al 
the proportions of caul speevies inthe local ares. "he 
height was inposed by the need to inspect (he total 
foliage hoth front the ground and from a 3 in ladder 
pluced in the centre of the tree, 

For cach tree we vounted the number of walls 
conlaining live and dead Apiomerpha females, Tie 
surveyed galls varied in size fron a few iim Cyouns 
or aborted) to more (han 4 em lone. und in shape 
from eylindrical and bud-like to urn-shaped or ovend, 
Galls were revorded us vootaining dew females i 
they were old and brown or showed signs of atuck 
by predators Or parasitoids, Usually, the presence of 
u living coecoid Was Confirmed by (he presence of 
White powdery wax atthe gull orifice, Only voucher 
material and galls of uncertain identity were 
vollected. so the survey Wos ?elatively non 
destroetive, Voucher specimens. of galls: and slide 
mMotinted sects ol Apiomarplia have been deposited 
ithe Austratiain National Insect Cullection (ANIC), 
CSIRO. Canberra. 

All ditt analyses pertain to galls contdinine five 
plus dead female jaseets, unless otherwise stuled 
The gills of dead inseels were jneluded in eounts 
becuuse any sucvesstul initiadon ol uw gall was 
evidenve thar josect had reached the site and that the 
Wee Wis at Suitable hast, Host speciieiy within 
Apioniorpha was examined by calenliting the 
percentige of (he surveyed teees Of euch eucalypt 
Species ial supported galls al femules of quel 
Apromorplia species. The response of Apromerpli 
Species 10 fire Was evaluated using one way analysis 
of varianee (ANOVA) Lo compare burnt and Jong 
Unburnt sites in terms of the total numbers of palls of 
females (of all Species summed), total numbers of 
gulls af the seven least common species (Le. walls of 
AL mallecacote Gullan and Ay ovientaides (Vepper 
exchided), and lolal Apiomerpha species found aut 


PANE 1 Site localities (omm GPS reclining) with qire histany flan unit daa) perses Dit in LORS CBE) aed Hee umes 


af tives af each species sampled ar euel site. Ko dy — E. dumusu. E. go> bk. gracilis, Boi = BE, inerussata, Ban 9K, 
Oleost, Bos, = D. soetalis, Nawihery en the fant of each spectes ealunin “hr tor the fatal Fniien Of ees warwyveu fie 
(het specter, 
Site History Latinde Longitide oad, Eee Mai, E. 0. EL 
| tu 33°17 13" moras’ a " a it] 4 
2 lai ABT" 14oras'uy" 4 [ {) 4 2. 
x 6 a ee Issa" {) 2 () Oo 2 
+ ob 17" 140) a5 08" a 3 oi 4 it} 
5S leu Aa Luray [4() 38°27" 6 it] 0 () + 
o b 33°16" lath ya ye 3 | I | 4 
7 4b AM Gd" 140 43'08" 6 u (1 () + 
yooh FAA Ay" Ltd's” 4 | 0 {) 3 
ae 33 lesa" 1g s7” | 7 {) {) + 
Wh dew 38°34" 30" ln arsy” i) 5 \) ‘) 5 
Woodew 34°05'02" l4or42'40" 3 + ‘) i) 4 
12 lr 33 17! 0" 4040's" a a 4 if] 2 
Mi lu 43074" LAPS! 4 J i 2 ) 
I4 ob 33° 19/20" Ly 35" ay" h 2 0 0 2 
q 5 7 a w 


GALI-INDUCING INSECTS AND MALLEE FIRE HISTORY M41 


each site (14 units total), Analysis was carried out at 
the site level, not the tree level. because there were 
Iwo or more different (ree species per site and many 
trees had no oor few galls. Two species. A. 
malleedcola aod AL eviceloides. were common 
enough fo examine their mdividual responses to fire 
al the site level using ANOVA but the data were 
transformed [ly (x+1)] fo correct for skewness, The 
responses of AL malleeacela and A, ovicoloides to 
fire also were evaluated using individual trees as the 
Units Of analysis. In this cuse. binary presences and 
absences were analysed using Chi Square tests 
because gully were not abundant or widespread 
enough to satisfy underlying statistieal assumptions 
wt this level, ATL amilyses were carried oul using 
IMP™ (SAS Institute Ine., © 1989-91), 


Results 


Apiomorphia species recorded 

A total of nine species of Apiemerila was 
recorded fron) the sites surveyed, ‘These were: A. 
calyoing (Tepper) A. densispinese Gullan, A. 
karsehi Ritbsaumnen. A. mMalleeavolu, Ay miinita 
matleensis Gullun, A. avieolaides. Al regilaris 
(Tepper). A. strombylose (Tepper) and AL arnedis 
(Tepper). All of these species have been collected 
previously from mallee vegetation in southern 
Australia (Tepper 1893: Gullan 198). 


Host-plant specificiy af Apiomorphit 

One species, A. densixpinesa, was recorded fron 
just three sites and solely on E. dsmose but only six 
gulls were found and none of these contained a live 
coccoid. Six other of the nine species ol Apiomarplia 
showed some degree of host-plant specificity (Fig. 
2), Galls of A. cahycina were found on four of the 
seven surveyed trees of BL prerassete as well as ou 
two other cucalypt species, Galls oF AL Kerselii alse 
occurred on three eucalypt species. whereas A. 
munita malleensis, A. regularis, A. strombylosa and 
Ao urnalis each were reeorded from only two 
cucalypt species, However, 13 of the 14 trees that 
supported galls of A, drnalix belonged to E. gracilis 
and nine of the 10 trees with galls of A, strembylove 
were EL socidlis, Only one species. AL ovicoloides, 
was recorded on all five species of eucalypl it was 
the commonest species, occurring On 52 of the 140 
surveyed irees, The next mast common species wits 
A. malleeacola which was found on four species ol 
eucalypt and on 37 of the 140 surveyed trees. 


Species richness and abundance of Apiomorpha in 
relation ta fire history 

All nine Apiomorphe species were found at both 
burnt and Jong-unburnt sites, Burnt and long-unburnt 
sites did not differ significantly in the number of 
Apiomorpha species recorded on survey trees 
(F, 9=3.57. p=0.08) (Table 2), ANOVA of the total 


Vania 2. Meany per site # ESD, orwith range in parentheses, and significance of differences due to fire hisrary tiediversity 
of Apomorpha species, number af galls ef-all Apiamorpha species, monber of galls of A. malleeacols, mune of galls 
ofA, ovicoloides aif number of gally of all species excluding \. malleeacola ave A, ovicoloides, (Foy A. mallecacola 
ful Av avicoloides, means and rnges ure from the caw deta bur F and yp values cre from transformed cette, as 


indicated by!) 


Long-upburnt 


Burnt 


Fis Value p Value 


i (sites) 7 


Mean number of 
Apiomerphe 5.34.1 
Species per site 


Mean number of 
Apiamrearpeltt 
gully per Site 


374495 


Mean miunber of 
A mullecacula 
galls per site 


9,4 (2-12) 


Mean number of 
Av nvicolotdes 
falls per site 


13.3 (6-30) 


Mean number of 
Apiamoarpha 

gills per site 

with A, wielecacald 
and AL eniealatides 
excluded 


16,6 £544 12, 


7.6 (0-21) 


BAH) 


40414 AST 0.08 


24.34 148 3.96 07 


40! 0.54! 


av 


OU! 


Id. ().34 (3a 


“o Of surveyed trees with galls present 


°s of surveyed trees with galls present 


“e of Surveyed trees wilh galls present 


% of surveyed trees with galls present 


60 


50 


P. J. GULLAN, P. S. CRANSTON & L. G. COOK 


A, calycina 


Ea€&9 Fi Eo, &s, 


Eucalyptus species 


A. malleeacola 


Ee, Eg El BG ES 


Eucalyptus species 


A, ovicoloides 


E.d Eg Ej} Eo Es. 
Eucalyptus species 


A, strombylosa 


Ed Eg Ei Eo Es. 
Eucalyptus species 


So of surveyed Irees wilh galls present “s al surveyed trees with galls present % of surveyed lrees with galls present 


% of surveyed trees with galls present 


A. karschi 


Ed Eg El Eo, Es, 
Eucalyptus species 


A, munita mallensis 


Ed &g9 Ei Eo Es. 
Eucalyptus species 


A. reguilaris 


Ed Eg EF Eo. Es, 
Eucalyptus species 


A, umaills 


Ed &.g Ei E.0. Es. 
Eucalyptus species 


OALLSINDUCING INSECTS AND MALLE FIRE HISTORY | b5 


nuibers of galls of females (live anel Jeu) recorded 
from) cach site todiedied that the fire history of the 
afles did fot affect ull abundinees af Apiomerplng 
specigs Ub) =3,.96, pe0.07) (lable 2), Burnt and 
fons unburolsiles did not dilferin the percentage of 
live fo dead Apemnarplin 230) oF galls at lone- 
unhurt sites and 24% OF gulls i burnt sites were 
estimated to contain live coveaids. EXcept fur A, 
mofleewcala and A, oavienlondes lhere were 
insulficien| daly lyr siarstieal analyses bused on 
individuail Species. although for the therd most 
ubuman) species. AL nrauliy there were compuruble 
populations at both burnt and lone -unbuy mt sites, 

bor Al maleeucol, ANQVA of the trinsformed 
ubundunes [I (xb bd} ah cach site indivated that 
numbers ab galls did not differ benween burnk and 
lutig-ubburnt sues (FP) =0.40, p=). 34) (Table 2), Chi 
Squive atilyscs OF presenee-ubsence dale for A, 
malleracela on individial trees alse indicated that 
Tire History did notafieer the likelihood of finding A, 
mallecaola YaMls QA) \,=0.04, p=0.84), with walls 
being present ai 27% ob all (ees sanpled in dong: 
Gabum) sites awd 260% OF all trees sumpled al burnt 
Stes. TP trees oF EL irerassara. on whiel A, 
maflecacale was never Tound, were exeluded, the 
resuills renmained very similar Cg?) =U,214, prst,04: 
gulls presenton 30% ob lone unburnt trees and 26% 
4) burol (rees), Burnt ind long-unburnt sites differed 
onty slightly in the percenniwe of dive to dead A, 
matlecacolen Ad OF galls ut king-unburnt sites nid 
S90. co) 4ills ah burnt Sites were estimated to contain 
live coccoids, Along the burnt sites surveyed there 
wits no evidence that galls of A, medleedonle were 
mone abundiunt we sites close to (range 1- Lb wah 
mean OfG-7 gulls per site), compared with distut 
from (range M21 with mean of 4.43 gulls per siter, 
long Unburnt aresis, 

Ii contrast, for A, orteulerdes, ANOVA of the 
transformed abundance [h, (k+1)) al eich site 
indicuwed (hit numbers ef galls were significantly 
lower at burge sites (PF) = 114A 7, ps0,01) (Tuble 2), 
Burnt and long-unburnt sites differed only slightly in 
the pereentive of live to dead. avicalordess (O06 OF 
gulls ut Jong-ontuent sites and 1% oF gallsat burnt 
Sites Were estitmied Lo contin live coecoids. Chi 
Square amilyses ol presence-ubsence dati for 21. 
ovicoleides on individual trees also tdjcateal that fire 
Instory ullected the likelihood of finding A 
ericolofiles, gully Qe) .=9.04, p=0.003), with gatks 
being present on 44% of all trees sampled in long- 
Unhurnt sites, hur on only 24%) of hurnt trees. Among 
(he burnt sites surveyed, walls of A. auiediaidey were 


Slightly nore suuree at sites FOU-200 ni trom unburn 
Veachition (range 0-3 with mean ol LO gulls per sites 
than atosites several kin distant trom lone-unburnt 
areas (rime 1-8 with mew of 4.5 galls per site). 

IWohoth A. mallemeeda and Al ovicaliides were 
excloded tron the anulysis GF total numbers ol galls, 
ANOVA of the gall abundance at cach site indicated 
that numbers uf galls of the other species combined 
did net differ hetween burtand long-unburnt sites 
(FP) =0.39) pets) (Tuhle 2). 


Martealiny fuctors 

The orttaal occupants of imny of the Apioanaples 
gulls (hal we recorded during our survey erther had 
been Killed by parusituids. probubly wasps. vl 
removed by preditars, prohibly cockatoos <add 
parrots. Some walls lac a single, large eit hole the 
wall, providing evidence of the enicrgence af w dart 
tailed Wasp. Crrterantel/ta Dalla Torre (Proromitlidie | 
(Tillvard 1926; Buudek 1988s Nauman [90d aud 
other walls had taby tiny ememence holes, Twelve 
salls tad one side removed whieh is the typleat 
appearinee of a gall Gpeded by a bird. Mortatiny of 
he datter Kind was twice as vommion in the long 
LDU sites asin the Buri sites, but There were tou 
few walls datnavedl ta this way lo etermine whether 
(he difference had statistiow! signifiownee. Murry 
other galls were browi auid obviously daud= but 
wenerally we could not determine the cadse at death, 
A few other galls were deformed by inquiines. thal 
is, Ober iisects had wecupied the pall Liste or the 
cuvily hut had nol direetly killed: the Apienrarplie 
Jemale. ‘The identity of the inquilines was not 
dvlermined becatise those still oceupylag the gill 
were citer diptenin or Hhymenopleran larvae and me 
rearing Lo adulls was arlempeed. 


Discussiun 


Th the even! of fire, season of burn is believed to 
huve the greaiest influenve on the phint composition 
of mallee vonimunities. with frequent aucuinn fires 
Causing substantial iortality of apajlee cucalypls 
(Noble 1982, 1989), Continuous canapy growth at 
miullee cuealypts can occur after a summer wildtire 
and may be die to the absence of phytophigaus 
Tseets (Noble 1982), hire frequeney (the interval 
between fires) is believed (0 Haye the inost importunt 
Jong-term effeet on amullee fauna beeause most 
animals aduplrot to Hire itself but ty the floristi¢ and 
suuctural leitures of the plant communities that 
resull from different fire rogues (Land Conservation 


Fig. 2. Hosi-plint speciticity of Aplonmepha plored us the perenne ol the -stirveyed trees of cach etedlypt species thit 
supported galls af females oP each Apiemorpaha species; Hvalyptiny specs: bok = Edun, Eo = BO ergedliv, Bon 


=e Meraysetie, Boa = Oleava, Gos = 0, sevralis, 


I-44 PS. GULLAN, ES. CRANSTON & LG COOK 


Counc 1987), bor jivertebrates, whieh generally 
AHIbU ScusOdul aCHViLy, Tapproprluike Li and 
puTivulachy Prequeney oF fire cun have daimswine 
consequenees for populauons (Land Conservation 
Coun) (987: Prichd & Williams 1996), Bor at least 
some Jrfodio nudlee communes, jh hws been 
sugpested thal the nattral fire frequeney is unlikely 
tu vaceed snore tha one fire every 15-200 yeurs 
heenuse of ihe time required for fuel loads te 
acemulate (Noble 1989). At Dangguli Conseryatiin 
Park. many of the qiillee sites that we stueied bac a 
Tindia undersiorey und i tad been 10.5 years ste 
wiltfire had destroyed our burnr sites: Thus. 
popu ktiotis OF Apfomorpha species ih hurit sites 
cont be shown to fave recovered to pre-byry levels, 
the probable natural fire frequency oF 15-20) years or 
mire Would heounilikely to have any lume-termn 
detrimental effeets on populations of lpieniu pee. 

He main Hinedings of oop survey were that the long- 
Unburnt sites dishnot daffer id species rretiness cor in 
litahubundunve of Apivenrpia galls frum sites burnt 
JOS yours age. but Chat the mediunm-terie celleer af 
fire ly vary for diflerent Apronarpler species. bat 
the wer species (hal were common enough fe allow 
dnilysis oF their abundance ie relation ta fire bistury, 
He A melecacolod) was equally abtumdint in burnt 
and long-unburnt sites. whereas the ather (A 
pyreutides) Was siznifieantly less: abundant a the 
Rurni sites, Indeed, for AL eyteafeides, re 
estublishinent al burnt sites was low even where a 
sOlirde a pelenril colonists wis just acrass a 1 nm 
wide proud, Par this species with slow or finited post- 
fire re-establishment, the effcet of another fire iy the 
moat live to J0-years might be yirtail extinction, 
eapeciilly TP no old-gvowth matlees, (hat my serve 
as fine refugia lor such insects. survive the fire. The 
consecvalion oF fong-Uoburntareas Of mallee should 
be management priority. ‘Stmibaly, Feiend & 
Wilhuins (196) have emphasised the inportance ol 
Hie ORHIge Ten) lo protect fire-senaiive species ind 
habitars fron tio-trequent fires in nmitee-heath 
shrublands of south-western Ausoatia, 

fy contrast. Ay madiveaeote and the third most 
conuin species, A, Henalis, bad re-cstublished 
Populations eqpvatent to those recorded Th long- 
uobuent sites ab siles burnt TOS years previously 
(although the number of records fora. wraaliy were 
insulivient for statistical analysis), (addition, there 
Were no obvious Fibe effects on re-establishment of 
Ihe opher six Apvemarple species: bul nuiibers: of 
aalls observed. at bath burnt and long-unburnt siies 
Jar eich species, generally were low compared with 
Ao nalleeucola Reeeolonisution fad ocwurred: at 
burn sites tt were LO Kim or inere distant from the 
nearest stands of long-unburnt matlee, For vagile 
species. the postfire flush of growth tiv he 
henelieuil to gall establishment. This suggestion is 


Supported by the vhservaiton that an athe 
environments, galls al Apiomarphe appear to he 
more ubunedunt eh youn ane resvowth ehealypts 
than oon the foliage ol older trees (heBrolon &e 
Vaurwerk 1994; Pod. Gulkun pers. obs. Although 
(his Tipression thay he creatod hy sampling hits (its 
jis casier lo search low fofage of saplings and small 
trees Than the canopy of more mature treesd. uy’ 
ghisshouse rearing as showin that the first-taster 
nyinphs will itive galls only un the pew Folie of 
uetively rowing Shoots. In another stady (Yeu 
1989), the abunidaree of phytophagous tHseets. 
especially sup-sucking species, hits been shown 0 be 
Nigher ou coppiee thar mature nitlee. perhaps 
hecause young leaves end shoots ure Mere HUTFILLOts 
(hin old Jolie dod Coppice Trees hive more yours 
erowlh (han mature matlees, 

Some o\pranurae speeies exhibit lost-plant 
prelorences lar certan cuealypt spewes, Both 4 
rovitinls and AL strambviase Oecurred only on iyi 
cuculypt species, , socvedis and b, aleavg whielt are 
closely-related) species - beth ure in the series 
Subulate of Lnvce/yprs (Chippendale: 98s), Cults 
Of AL munita melleensiv were lound oily on Be 
digneoe and bo wecilis, whieh are in different series 
(Chippendale (988), Two very oclesely-vefaiel 
Apiamerple species Ay culyeoim sind aA, aerate 
(Gullan’ PO84a, LC Cook nnpub shiwed very 
different hast preferenees. Thirteen of the 4 
surveyed (rees vith galls OF A, (dfis were 
erapilia. wherens gills ol Ac cafbetae were ties 
common on 20 7erdsseme aml pever neeurrecdl on fy, 
eretix Simee We twa inost commun Apion 
apreeies, A, marlleede Ale and Aa evealoedles, alse had 
the browdest Host-plant mutes. differences jo the 
pecurrence vind abundance of tes fw species 
relavian to the fire history of sites cannot he 
Hithibuied ta uiny varnation i the composiiian of 
euculypl species amnone sites, fnstead, population 
differences amone Apiomeryia species in rehatien fo 
the fire history oF sites may be best explained by 
differences in their propensity to disperse, Che 
eriwiers of some Apiary species may disperse 
inure readily to new frees Than those of other species 
There is dimple evidence that Hest-imstir seule sects 
of other wrotips are dispersed passively by the Wind 
and, even though morality is very high, may be 
curried for distances of a lew to several kr, aid 
more rarely a lew hundred key from the natal trees 
(reviewed by Pedgley |982. Greathead 1990; Hunks 
& Denno 1993), Some seale jnseets crawlers hive 
been reported to orientate dowrwind amd stand on 
thei hind dees with ootennue and fore lays 
outstretched = (Washburm  & Washburn 1984. 
Washburn & Primkie }985, Greatheal 1990), Sueh 
hehayviour probably cosures (heir dislodwenent qd 
dispersal by wind, The eriwlers ob a lew species ot 


GALL-ASDUCTNG INSECTS AND MALLEE HISTORY 145 


Aplottorpia taye been observed displaying similar 
behaviow under ghisshause condinuns Ch. G. Cook 
uTpub.. Apiomeanpha species may differ in their 
Propensity Cither Lo disperse actively Cour the host 
Plank or vera op the ated trees In contrast to (he 
NeYOriLy OF Apiomerphi species, Tt is extremely 
Giffin Wy estublish infestations Of A, matlecdeolea, 
A valved andl. wrnedix by releasing crawlers on to 
polled wuealvypis in ud glasshouse (lL, Gy Couk 
Unpub); this suigeests (hat at Teast some crawlers of 
Ile laller Species muy exhibil obligatory dispersal 
beliviour dnd. Hh the glasshouse. may suicide by 
uctively departing from the only suitible host plant, 
Under natural conditiolis, however, mall lees 
probably would he surrounded by mther suitable 
bosts, especially i mative vevetiion, 

Dispersal ability may tehilte Lo morpholowiedl 
uduplations as well as behavioural ones, ‘The 
Hatlened bodies iad (wo or more long. filamentous 
ciidal selae of scale inseet crawlers ure heliewed to 
enhance ther dispersal potential (Wainhouse 9x0: 
Pediley 1982) Thus differcnces up body size and 
shuipe may partly account for differential dispersal 
winone species, The crinvlers of Apiomierpha have 
Hlatlened, Oval to Subeirculue bodies fringed with a 
conrinuons cow Gl margin setue (Gillan (984), 1) 
addition, he surkice area of cael) irginal seta is 
extended by a thin sheet of waxy segretion 
(NwOOUS Tn general UpPpearaniee lo The wabe froin 
ihe shaft ofa feather), ‘The Frst-instir nyiphs ot A. 
iMicaloidesy are about the same length as. but 
narrower ((95-225 pina widest part) than, those of 
A, mallecucola (265-280 pm wide). although. the 
marginal setae are approximately equal in lent 
(34-44 pint on bath species (LG. Cook unpub, i 


dispersal wbity in lpiwmorpled is conrelated with the 
surface arent of the body oF the crawlers, the 
differences in abundances ol AL weatleeccala ands, 
eeoloides in burnt-and long-unbuunt sites may be 
attributed at leastin parka diflerences in he size and 
shape of their crawlers, 

Selection for both active dispersal behayiour anu 
body morphology thal fiveurs passiwe urilt tay 
gecur in scale mMseut species that Gecupy 
Unpredictuble or lemporary habitats, vis las bee 
sugeested far armoured seule insects thal feed on 
short-lived versus longedived host plants (Greathead 
1990), TP this hypothesis ts valid) fur Apreniaplea, 
{hen seme species. such ak. wailleewoote. ean be 
postulated ( be better adapted to the vagaries of fipe 
in the mallee environment, 


Acknowledgments 


We wish to thik Mo Collott for assisted inthe 
field und M- Osborne, the Rimger al Dianpeali 
Conservation Park. for help and information 
provided during our fieldwork, KR, Pullen kindly 
arrunged, (ind the National Parks aod) Wildlife 
Service Of South Australia provided. the scientific 
permit for this study. The Culperuin/Bookmark 
Survey of Invertebrates Prajeet furided by of the 
Australi Natute Conservation Agency (new 
Lovonment Australia) provided the travel funding 
tor Lis work, PD, Conper give advice onan BOG L. 
Mekie curried Gal the statistienl ainihysis, A, 1, 
Austin, ROK, Blanehe, GM, Clarke. M. I, Collofl’ 
and Ro B, Floyd made useful comments on cartier 
versions OF the manuiseript 


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AN INTENSIVE MONITORING STUDY OF TWO WETLANDS 
OF THE RIVER MURRAY IN SOUTH AUSTRALIA; 
PHYSICO-CHEMICAL PARAMETERS AND 
CYANOBACTERIA CONCENTRATIONS 


By A. M. OLSEN* 


Summary 


Olsen, A. M. (1997) An intensive monitoring study of two wetlands of the River 
Murray in South Australia; physico-chemical parameters and cyanobacteria 
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. 


Transactions of the Royal Sector ofS. Aust, (1997), 12144, 147-155, 


AN INTENSIVE MONITORING STUDY OF TWO WETLANDS OF THE RIVER 
MURRAY IN SOUTH AUSTRALIA; PHYSICO-CHEMICAL PARAMETERS AND 
CYANOBACTERIA CONCENTRATIONS. 


by A. M, OLSEN 


Summary 


Orsps, AJM. (1997) An intensive monitoring study of nwo wetlands of the River Murray in South Australias 
physico-chemical pagumelers und cyanobacteria Concentrations, Trans. R, Sec. 4. Agst T2104), 147-155. 28 
November, 1997, 

Ovuanitive data were collected on physico-chemical characteristics of suirtuce waiter temperaruires, ph. 
turbidity, conductivity, dissolyed ummonia, dissolved reactive nitrate and total phosphite of Bannock and Loch 
Luna wetlands from 46 sumplings jo cach wetland over a 20-month intensive monitoring study, Coneenmritions 
of the various physico-chemical parameters were within the ranges found in similiar freshwater River Murray 
wetlands. 

Weather factors, such as sttomg winds. heavy rain rdnott and lightning. produecd pertubations it turbidity. 
conductivity, dissalved reactive nitrate ane total phosphate levels in the bwo wetlands. 

Nutrient concentrations in excess of 0.36 mel! total phosphate and 4.0 mel’ dissolved reactive nitrate with 
Vistig waler temperatures were related to rapid cell multiplication af the cyanobacteria Atahuena spp. Vhree 
Vrabeene spp. were predominant in the two wethinds aml reached their greatest nombers (23,700 cells mb!) in 
Loch Lami trom fate December 1994 to mid - Junuary 1995S, 


Rey Wokos: Wethinds. River Murray. monitoring physicochemical parumeters. cyanobucteria. South Australia, 


Introduction 140° 22' E) opposite Bunrock Station and about one 


kin upstream of Lock 3 weir (ANCA 1996). Banrock 


In 1990 the Murray Darling Basin Commission 
through its Natural Resources Management Strategy 
(NRMS) funded a preliminary study of the water 
chemistry une aquatic invertebrates and land 
vertebrates of 10 wethinds of the River Murray 
Noodplains in South Australia (Goonan et al, 1992), 
The survey was conducted during May ~ June 1990. 
Between May 1990 and February 1992, a second 
more Uetailed physico-chemical and biological 
survey Of cight alkaline freshwater wetlands was 
curried out on the above - mentioned floodplains. 
Five of the eight wetlands were located between 
Clover Luke (Culperum area) und the Berri 
Evaporation Basin und the remaining three were 
between Rameo Lagoon (Waikerie) and Lake Carlet 
(6 kin upstream of Mannum) (Fis. 1). These results 
were reported by Suter e/ al. (1993), 

Banrock Station [loodplain (Section 662, 681 und 
682 - Hundred of Moorook) and associated wetlund 
fig i about the middle of the [OQ km (approx.) 
stretch of the river between the tive upper and three 
lower Wetlands studied by Suter er al. (1993), 

Hlow regulation of the River Murray at Lock 3 in 
1925 helped create two permanent lreshwater 
wetlands in the region, ane located in the floodplain 
of Banrock Station (34° O8' S. 140° 20! EB) and the 
other inthe Loch Luna Wetland Complex (34° 12" 8, 


Th Orchard Groves Newnan S, Aust S074 


Station Wetland has a 90 to 130 ha area depending on 
water depth (20 em - 1.1 m) and was created in the 
1050s by damming the upper section of Banrock 
Creek. The wetland behind the dam wall is gravity 


Pe fi. 
yf | “S Jy 


Swan Ren | 


ih 
j | a) 
¢ 1 ~élosan 
3® Blaneherewa 
| 


i : 
South Australia 


Mariuit — 
—_ 7 
Pawiray 

Uri ee 


t 


140 F , 


Shale (kin) 


Hig. 1, Location map of wetlands studied River Murray, SA, 


148 A. M. OLSEN 


fed 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 lo 
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 1991-2 there had 
been visible blue green algal blooms in both 
wetlands, 


Materials and Methods 


Sampling 
This monitoring study of the two wetlands of the 
River Murray began in November 1994 and ended in 


Banrock 
Sampling 
Stations 
1-5 


Banrock — 


(34° 08’ S, 140° 20’ E) 


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 flow 
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 


34° 10'S — 


Loch Luna 
Sampling Stations 


Loch Luna 


(84° 12! S, 140° 22’ E) 


140° 20 E 
Kingston on Murray 


Fig. 2. Banrock and Loch Luna wetlands showing sampling sites, Note direction of flow. 


PHYSICO-CHEMICAL MONITORING OF TWO WETLANDS 149 


Task lt. Numbers of cells of Anabaena spp. ane TABLE 2. Numbers of vells of Anabaena spp. and 


concentrations of dissolved reactive nitrate and total concentrations of disselved reactive nitrate and total 
phosphate - Lach Lana 1994-96, phosphate - Banrock 1994-1996, 
Date Cells DRNitrate Total phosphate Date Cells DRNitrate Total phosphate 
(mg!) (mgt!) (mgt) (mgl!) (meh!) (mgl') 
Noy 3, [994 + 0 0.16 Noy 3, 1994 14 {} 0.20 
Nov & 0 0) 0.13 Nov 8 0 OA O17 
Nov 15 Q 0 OA7 Nov 15 + (44 0.27 
Novy 22 56 0.44 0.20 Novy 22 Q ) 0.30 
Nov 29 41 Tr. 0.25 Nay 29 1.460 Tr. 0.27 
Dec 6 434 0,44 O.13 Dec 6 2.480 3,5 0.41 
Dee 13 1.230 0 0.27 Dee 13 1,560 Tr (0.60) 
Dec 20 5.590) 0.44 0.47 Dee 20 303 Tr. 0.70 
Dee 27 23,700 2,20) 0.16 Dee 27 125 0 ().23 
Jan 3, 1995) 10,600 Tr 0.46 Jan 3, 1995 96 Tr. O15 
Jun 17 14,500 6.1 O19 Jan 17 245 4a) (133 
Jan 31 1.050 0.44 0.45 Jan 31 146 2.2 0,21 
Feb 14 3,560 Tr. 0.07 Feb 14 373 0 12 
Feb 28 97 3.42 O10 Feb 28 311 7.9 0.06 
Mar |4 437 O88 0 Mau 14 R83 0.88 Tr. 
Mar 28 172 22) ().23 Mar 28 5 0) 0.33 
Apr 1& 425 2,20 0.36 Apr 18 123 0 0.27 
Muy 16 909 0 0.25 May 16 189 Tr. O33 
June 13 100 1.32 0.17 June 13 wetland drained 
July 18 11 0 0.17 July 18 50) () 0.20 
Aug 16 8 3,52 0.19 Aug 16 0 0 Q),24 
Sept 5 a] 0 0.30 Sept 5 + 0 0.40 
Sept 19 2 Tr. 0.07 Sept 19 + Tr. 0,25 
Oct 3 a) Tr. 0.53 Oct 3 Q 0 0.38 
Oct LO 8 if 0.47 Oct 10 0 0 O39 
Oct 17 6 0) 0.43 Oct 17 I4 0) 0.35 
Oct 24 12 0 0.95 Oct 24 0 0 0,23 
Oct 31 I 0.44 0,04 Oct 31 a 44 0.27 
Nov 7 24 0) 0.20 Noy 7 24 Te 0.26 
Nov |4 6 44 0.21 Nov I4 27 0 0.340 
Noy 21 450 0 O17 Noy 21 225 () O16 
Nov 28 1183 0 0.23 Now 28 626 3,52 0.30 
Dec 5 1,220 7.AS 0.30 Deu 5 461 Tr 0.30 
Dee 12 L.ASO Tr. O40 Dec 12 455 ) 0.36 
Dee 19 15 Tr. (0.67 Der 19 135 Tr. 0.43 
Dec 26 1,270 22 0,20 Dev 26 304 Tr. O18 
Jan 9, 1996 997 6.6 0.20 Jan 9, 1996 2.300 44 21 
Jan 23 3,190 44 O17 Jan 23 1,040 1.32 OAS 
Feb 6 2,530 Q 08 Feb 6 2,170 () 0.02 
Feb 20 368 Tr. 0.25 Feb 20 240) 0) O41 
Mar 5 63 contaminated = contaminated Mar 5 71 0) 0.60 
Mar 19 444 0 0.23 Mar 19 328 0 (40) 
Apr 14 57 0 0,12 Apr 14 28 () 0.12 
May 14 0 () 0.41 May 4 0 0 0.33 


Jun 9 0 0) 0.06 Jun 9 (0) 0 Q.24 


150) A.M. OLSEN 


April to September on the dhiles given i) Tables band 
y 


suinple was ikem and bulked ia five fire plastic 
bolle with swnples taken fron the other four sites, 
Aliquots of the bulked sample were lrinstepred: to 
diriree S00 ml polyourhonate plastic serewlop 
bottles for Subsequent physico-chemieal analyses at 
(he Scienve Section, Glossop High School The one- 
litre willer samples lor Count cyanobacteria Were 
wansterred to 1-25 | plastic serewlop bollles leaving 
u 250 ml headspace. The suples were keprolitled 
uncdil delivery to the Australian Cenme tor Water 


Quality Reseweh, Bolivar SA for enumeration of 


eyarmbacteriy cells (HIMSO 1990), The special 1.25 1 
plastic bottles for the water simples were suppliced 
from the Water Quality: Laboratory, 

A tloadhe “Der Grune Punke’ No, 7428, blue 
Wleoho! colin thermomerer was used to record the 
slike Wate Temperatures at each site, A mein 
surtiwe Water tenmperutire wis then ealenlated: for 
cauh Wetland A plashe bodied minimum - mixin 
(hermometer, (30) 450°C) with pressure adjustment 
for indivalors on the mercury Column Was used) to 
record MINTO ad MAXIM Willer (emperatures 
henveen tiites Ol consecutive sanipling visits, Tt was 
suspended |S cn beluw the surface. 

There was ur $3 June 1995 collection in the 
Banmek wetlam because this wellind was drained 
for taintenudee work On the irigarion pimps uxee 
for Iehlind vineyard irrigation, The Loch Lam 5 
March 1996 sample wus discurded heeuuse ol 
Suspected CONLUNTAGOD, 


Chemical anilyyes 

Witter saiiples were heldait 4 C andil required ane 
i Most iWstahees were upalysed within 48 bh. 
Measurements ot turbidity. dissolved anninionta, 
Uissulved reactive nitrite und total phosphate: were 
imide using 4 HACH DREL/S instrument ond 
premixed reapents (Hach 1084), Bor disselved 
aminonia cach sample was fbered through a Double 
Rijies 20) filter paper to remove suspended solids. A 
TS mbabquol was meashred (loa Clow) Llass sample 
cell and tml ol Nessler reagent added (Lhiech 1992), 
The misture was eft far colour ly develop, In thts 
case the blink was distilled water (25 mip with bom 
of Nessler readent fneluded, Measurements were 
(ide of 425 ny andl recorded us mgh! aller applying 
a conversion fretor of mualipheation by 1.29 (Much 
1942), 

Vor dissolved renelive nitrate (ORS) Gach scape 
was filtered through ip Double Rings 207 paper to 
remove clay particles: A 25m) whiqnol was measured 


Niel Cre Crlends OA 


AL cueh site (Fig, 2) a one-fitre surface water 


ito aelea glass sample cell and the contents oFone 
foil suchel of premixed Nitraver S reagent added, 
This inixture was rapidly agitated for one minute 
then tele for five minttes for valour to develop. 
Another filtered 25 mil sample was used us u hlank. 
Measurements were made it 500 nine pecayded us 
mel! after applying a conversion factor of 
multiplication hy 4.4 (Mash 1992). 

Poe totil phosphate (PP) a digestive process was 
dised (Oo convert all forms OF phosphate to the solute 
orthophosphate form (bhich 1992), Fifty nib ot the 
sample were measured into a@ clean [25 ql 
Kvlenmeyer Mask aluog with 4 rl S.25 8 HSO) and 
(wo foil sachets of KGS O, (Hlieh 1992), "The wixtune 
wits fealed jn a boing water bath fer 30 mites, 
alloweel to cook bo room temperature and then ml SS 
NWOH were added, The sample wis split inte two 25 
Wit portions tH clown glass sample cells, Que foil 
sachet of premixed Phosver 3 reayent was added to 
one container and the colour allowed to develop, The 
other was used as a blank for spectrometric amilysis. 
Measurements were taken at 700 no and recorded vs 
mel! after applying a conversion factor (division by 
4), These DRN and TP resis complement dais 
collected lor eight SA wetlands by Suter ef al 
(JOU4), 

pH was measured using a Hanna HE &424C pH 
ineter, The probe was rinsed in distilled water 
bebween cach test and left in the saniple doth the 
highest stable reading was reached, 

Por turbidity measurements. each saniple was 
ugihiled vigorously and a 25 mi aliquot was added to 
4 clan gluss sumple cell and placed in the DREL/S 
spectrophotometer, Measurements were made at 450 
pm and recorded ws Nephelonmetric Turbidity (NP) 
unity. A blank of distilled water was dsed to vero the 
TISEPUITICNE, 

Condactivily meusurements were made Using a 
EDT PE 200 conductivity meter calibrated to 14 ts 
Eleetrical Conductivity (2C) units between euch usy, 
The calibration solution was prepared by diluting 
74.55 2 of oven - dried RCT in LOOO int distilled 
water, The probe was rinsed with distilled water prior 
fo-each vse and the Maxinuin reading taken euch 
Line th Was used Restlis were in EC units: 


Results 


Mean surfeve water tenperdinres 

Throughout this study the mca surhice water 
Lemperatures. in’ the wo wetlinds showed alimest 
identical trends, Ia generul, water temperatiires were 
higher Wy the More exposed shallow Banrock wethend 
han in (he slighily more protected Loeh Lun 
wellanu. The highest! surtaee water lemperatures ty 
Banroek, 28.57, 28.6" and 264° © were recorded tn 
(he summer ol 199465, corresponding tamperarnires 


PHYSICO-CHEMICAL MONITORING OF TWO WETLANDS 151 


in Loch Luna were 27.4", 28.5° and 25.3" C (Fig. 3). 
The lowest mean surface water temperatures were 
10.3° Cin Banrock and 10.2" C in Loch Luna on 18 
July 1995, 

In Banrock the greatest range in minimum and 
maximum waler temperatures between consecutive 
visils was 20° C (15° - 35° C) between 3 and 17 
January 1995 and the greatest range in Loch Luna 
was 145° C (14° - 28.5° C) between 22 und 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 
difference between minimum and maximum water 
femperalures between consecutive visits in) Loch 
Luna was 0.5" Con 18 July 1995, 

Lock 3 Pool surface waler temperature SOQ rn 
upstream of the Banrock Station Intake in 1994 was 
slightly higher than the mean temperatures in 
Banrock and Loch Luna (Fig. 3). 


pH 

The pH values ranged up to 9.56 at Banrock (27 
December 1994) and 9.04 at Loch Lana (3 January 
1995) (Pig, 4). Lock 3 Pool water registered a 
maximum pit §.62 on 14 November 1995 and a 
minimum pH 7.51 on 26 December 1995, 


Turbidity 

The shallower Banrock wetlind mostly recorded 
higher turbidity yalues 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 
[4 March and 28 November 1995 while m Loch 
Luna the highest turbidity was 170 NT units on 5 
September 1995 and the lowest LO NT units on 14 
March 1995. Turbidity in Lock 3 Pool ranged 70 NT 
units on 14 May 1996 to 20 NTP units on 26 
December 1995, The mixed River Murray - Darling 
waiter had a (urbidity yalue of 48 NT units on 9 June 
1996, 


Conductivity 

The initial high conductivity of 1898 EC units in 
Banrock on 3 November 1994 decreased to 871 EC 
units in 6 weeks und more slowly thereafter to 500 
ERC units by 9 June (Fig. 6). In Loch Luna the initial 
conductivity of T095 EC units on 3 November 199+ 
deereased Wy 687 EC units on 17 January 1995, rose 
slightly before falling toa minimum value of 327 EC 
units On 19 September 1995, Within a fortnight there 
Were two sharp increases In concentrations (2720 EC 
units on 3 October 1995 and 1759 EC units on 31 
October) before conductivity valuies decreased to 
511 EC units on 9 June 1996, 

The conductivily values of the River Murray water 
decreased slowly from 686 EC units on 31 October 
1995 to 51) EC units on 14 May 1996, The mixed 


Temparauea 45) 


en | T Hey! 
‘ oe bat Mar Ape May din 


Fig. 3. Comparison of mean surtace water temperatures at 
Bunrock and Loch Luna, November 1994-June 1996 und 
River Murray, October 1995-June 1996, 


Lip vin 
‘ fpuee Ranity 
' iy 
ry ' 
4 j ! ' 
heey a 
tha 4 iar \ 
h ' 
iy 7h | Pe . 
ink are) m ) ou \e 
¥ , . » th ‘ 
’ w » 
| 
’ —— | i} 
¥ Pek Win Ape May dun Jub 2iyy Seg et tee TT ‘" 


Fig. 4. pH. Banrock and Loch Luna, November 1994-June 
1996 and River Murray. October 199S-June 199%. 


big. 5. Turbidity. Banrock and Loch Lunia. November 
1994-June 1996 and River Murry, October 199S-June 
1996. 


152 


52 A.M. OLSEN 


River Murray - Darling water was 438 EC units on 9 
June 1996, 


Rainfall 

Weuther factors, such as heavy rain und 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 to he representative 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, 29 nim on | May, 53 mm on 25 
October 1995, 38.5 mm on 2 January 1996. 25 mm 
on 27-28 February and 20.1 mm on 3-4 June 1996. 


Dissolved aniunania 

The patterns of dissolyed ammonia (Fig. 7) were 
similar in both wetlands. The range in Banrock was 
from 0.17 mgt! on 28 November 1995 to 3.10 mgt! 
on 7 October 1995 and in Loch Luna from 0.30 mel! 
on 3 January, 28 February and 31 October 1995 to 
3.48 mgl! on 21 November 1995, 


Dissolved Reactive Nitrate (DRN) 

In Banrock there were occasional high DRN 
concentrations, exceeding 4 mel!, in the period 
November to March 1994-5 and again on 31 October 
1995 and Y 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 1995-6 there were four occasions when the 
concentrations exceeded 4.0 mel! (Fig. 8). 


Total Phosphate (TP) 

The highest TP concentrations recorded for Loch 
Luna were 0.95 mel) on 24 October 1995 and 0.67 
my! on 19 December [995 (Fig. 9). At Banrock TP 
peaked at 0.7 mel! 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 mgb!. 


Cyanobacteria 

When water sampling started in Banrock on 3 
November 1994 the total cyanobacterial count was 
I4 cell ml! (Pig. 10). By 6 December 1994 
cyanobacterial cells peaked in Banrock at 2480 cells 
ml! and then declined steadily to 5 cells mb! by 28 
Mareh. The second but smaller cyanobactenal 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 of 


2300 cells ml! before declining to 71 cells mb! on 4 
March 1996 (Table 2). Three Anabaena species were 
predominant in the Banrock wetland. 

In Loch Luna between 1994 and 1995 the same 


Fig. 6. Conductivity. Banrock and Loch Luna, November 
[994-hiine 1996 and River Murray. October 1995-lune 
1996, 


ao —— — 
af = Lon Luna 
/ BAe i 
{ 
4 
~ ' 
2 | 
4 | ; 7 
fac \ ' 
J \ 
2 ron f y 
gae fi oy VY } a 
> aie 
a Aid ! : > 
wah | J ¢ 
i nly i 
Thy ear 3 ' 
iy! i , | 
= } i fA * 
oy a / ' 
a oo 
By toi ood \ Tr yyy 
Nou Ome ior Fab Mer Ag Muy dun Ju) Ang Sep Onl Now Gb von Fel Mar Anh May. die 
of ' a5 ' ab 


Man) Yew 


Fig. 7, Dissolved ammonia. Banrock and Loch Luna, 
November 1994-June 1946, 


= kook Lunn 


© teantinil 


ah 


CAAA leuid coded pte nitrate (tral! 
+ 


ao 


pire © oe Coe an 17 ay he 
fv Hen dare Fete dan Fee mar App Muy: dun 
a oa 


edie 


Monin Year 


Rig &. Dissolved reactive nitrate, Banrock and Loch Luni, 
November 1994-June 1996, 


PHYSICO-CHEMICAL MONITORING OF TWO WETLANDS 153 


* Loch Luna 


Acanroch | 


ail i | ae Gow." 
Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jur 
95 ith 


Nov bec Jan Feb 
or) | i 


Month - Year 


Fig. 9. Total phosphate. Banrock and Loch Luna, 
November 1994-June 1996. 
\ 
| 
«| ||, 
: ih 
* || 
a ara 
| a 
w | 
} oy 
+ | 
| 
| Lf fs 
| iN 
a pene ytel  RR eh eo ating” Tasso ae | 


is 7 pened T 
Nov Dec Jan Feb Mar Apr May Jun JU! Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun 
a | a5 | 96 


Month ~ Year 


Fig. 10. Cyanobacteria (blue green algae). Banrock and 


och Luna, November 1994-June 1996. 


g. 
L 


three Anabaena species were predominant through 
the 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 cells mF! on 27 December 1994 
before declining to 97 cells ml! two months later 
(Fig. 10, Table 1). Anabaena coiled species was 
predominant until 3 January 1995 after which 
Anabaena circinalis displaced it until mid - 
February. The coiled species again became 
predominant until June 1995 when all three 
Anabaena species were present in low numbers until 
late November 1995 when multiplication of 
Anabaena coiled species began again. This species 
peaked at 2530 cells mb! in early February and a 
month later cell numbers fell to 63 cells ml! (Table 
1). Cell numbers rose slightly to 444 cells ml! on 19 
March 1996 but no cells were detected on 9 June 
1996, 


Other cyanobacterial species identified in the water 
samples, although occurring only in low numbers, 
were Anabaenopsis elenkini (6 December 1994 - 20 
February 1995), Aphanizomenon sp. (22 November 
1994 - 13 June 1995, 23 January - 19 March 1996), 
Oscillatoria sp. (3 November 1994 - 18 July 1995, 9 
January - 19 March 1996). Cylindrospermopsis 
raciborski, Planktothrix spp., Arthrospira_ spp.. 
Microcystis aeruginosa and Pseudoanabaena spp. 
were identified from time to time. 


Discussion 


Surface water temperatures, pH, turbidity and 
conductivity levels followed similar trends in both 
Banrock 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 ef a/. (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 
Banrock on 3 November 1994 was caused by a 
blocked inlet pipe into Banrock; with the clearing of 
the blockage conductivity values in one week 
dropped to 1507 EC units. Turbidity in Banrock 
increased from 104 to 155 NT units between 5 and & 
November 1994 due to turbulence from the rush of 
water following clearing of the blockage. Seven 
wecks 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 Bonney 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 POISI7 in Nockburra 
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 Banrock was 
0 - 7.9 mg}! and in Loch Luna 0 - 7.48 mgt!. The 
high registrations occurred mainly after heavy rains 
but there are also nitrate contributions from time to 


Ja4 ‘AM OLSEN 


tite Tony water tron the River Murry. and fron 
avriculliral dritos, town effluents amt sewage 
Jischurzes us well as Jovaleed autolvie breakdown 
of nitragen - fixing ble green alee (Anahecine 
spp. There isin tnknewr DRN puteent lapae fron 
huge Hoeks oF pelicans (> LOOQO birds) swans. 
vormorants and ducks and lesser nunibers of ather 
water birds resident in Baoraek and Laeh Lane 
Wetlands und on the banks of the River Murty, 
Nitvates are also prodiwed by lightaine (Sith 
1996), 

The highest TP level (0.95 nib ) recorded in Lach 
Luna veeurred on 24 Outober 1995 the day aller 53 
ni at yar felhat Barrera and Sk mm at Banrovk 
Stabe (Co Lb, Rohliehl pers. comm. 1995), Such 
heavy rain aml consequent runol cause Potton 
Jisturhinces in shallow wethinds whieh redistribute 
Jisselyed unaite flosphorus compounds und 
iWortatie phospheris bound to suspended ar 
Jistivbed bolo) organic putheutates: 1) the wiler 
coh, Brees ef a, (1985) have alse drawn 
dllenion to the complexity of chemival relauiashipes 
withon werkinds aml the elects of weather fietors. 
SUCH os Wands and feinpemdure, On Water chemistry 

No blooltis of eyanobacteria were observed in 
Banrock wetland, “The multiplication of 
eyaiobueterial wells je Baaroek during December 
IWS wis hulled by the inercased witer inflow 
ollowiig removal of the blockage Hi the titer pipe. 
Whe eyunobucleriie were (Mlushed oul preventing any 
further development ofa eyarmobacterial bloone in 
Birroch wetland that scar 

IW Loch Lune evinobacterial cells reached a 
utr of 23,700 cells tal! oon 27) December 
helore declining wozere by 1+ Mareh 1905, After the 
oollapse of the cyanobacterial populariain ia Lach 
Lunn December (YO. a small bub visible blue 
areen abil bloam developed dowtstreaaiy i 
Februaey vind Mareh 1995 along the eastern hank ol 
Ihe River Murray wujacent te TLoek 3 weir Nis thely 
(hil this bloor had gis genesis in the November - 
December 1004 cell mulfiplionion te loch) Land 
Upstreain und pn the same side af the River Murry. 
Small blue ereen olval hloors Had ocetrred mH this 
sales THcadon Lr previous yes (hock 3 staft pers, 
cop, LOO8 |), 

Bowling (1994) repurted the occurrenee ane 
possille eases OF a severe Anefaen etre inulin 
blow tit Lake Corellign NSW oir fate 190-0 | 
wher cell numbers exceeded (004100 vellsiml! The 
physicu-chenneal levels i the lake fi 1990) had 
nines close to the 1994-4 values i Loeb Lun far 
Wailer lemperiures, pil, turbidity and conduetiviry 
hil lower Wasi in concentrations af TP and DRS 
fan were Toupd dor Loch Lane. Bowling (1o94) 
capressedt The view suboul the Like Cirgelleo bloent 
hat “wiilnueh severe! underlying eanses ol this 


bloom are probable, the elevuled  nuirient 
concentrations, espechilly of tatal phosphorus. were 
major Taeturs that contributed ta an He drew 
aiention to the Taek that mase physicoeehemical 
stidics af cyanobacteria blooms were slanted ufler 
the blooms had oceurred. 

To this study of the physive-chemeal properties of 
The two Wwetlinds a seach far blot - Torning tosie 
cyanobacteria species Was started betere any cell 
MuUplication Hac commenced jimh a seasonal 
pullern is) described. Cynnobacterid: celhy may 
romain dormant Ti eolk waters cand grow: best ut 
wuller lerpperitures excecding 13°C. with Gptinnl 
erawth rales a) 25° Cor higher (kobarts & Zolury 
JOSS), The efleek of wiler lempernitures on 
eyunobuclerial cell nurmbers in Loch Lune is shove 
wilh dita front Table} which shaw that Blears only 
occurred between November une Pebruary, Nurmhers 
Ot Aikimeecha spp. tose suddenly fran cero on $5 
November 1994 (20.1 ©) toy 23,700 nmil-l on 27 
Decensher 1904 424.2" OC) and thea began subsidioy, 
ever Uneil the wiatttr teniperauores were high, 
perhaps because of OxHUstian OF WUUPENES. 

The rise (und fall fa cell numbers iy aso be 
rehiled to the concentrations of the two mitrieitts, 
DRN aml TP recorded dutiog the vrowth and decline 
ol the }994-5 vind 1995-6 blue ween ulgal outbreaks 
Mable 1 However, sivee the concennyiot af 
dissolycd reactive phosphorus is nol known, (hese 
relationships must be teeatod with caurion, 

There were Jour ceeusians (20 December 199d, 4 
and 31 Jani 1995 and F& April 1995) when TP 
concentrations in Loch Luniowere at or above 0.46 
mel! oun DRN was present even tn law 
cnneenreations (Pr - 2.2 mel. Aten vaeh ol these 
events there was a vise in the nunbers ar 
cyanobacterial cells. hy October 1995 (mean 16,9 °C) 
(here were four PP coneen(nitions between (44 and 
1.95 ggl! hur DRN was low or absent and ng vel 
mulrephiearion developed, 

A patrern rehued ta totrivat availability anu 
lomperniture Gin be seen in curly 1996, Alter an 
Wnerewise ty 3190 cells on 23 tunuary 1996 the cell 
Hmbers began to deelhe possibly die te very lov 
eonecntrations Of DRS aed Mretuiting values of TP 
Wothe walter colin, Phe eyanobieterie populiion 
reughed vero on 14 May (Y9G. "The mean wattere 
lemperiture onthad chite was 14 oC whieh is dear the 
MIT MAT Temperate range Tor eeowel al many free 
living Gyanubucterin species, 

Prom (he data obrincds i this study ibis stimested 
Nhat outhreaks af cyanobacterial bloenis id jet 
secur in the supers of [9945 gad [995-6 beeuuse 
{here were muadequitle concentrtions of LP and 
DRN ni (he wiler columim during the period of 
lavourable growth for eyanobacteria Dane fram this 
study indivare that LP coneentradin abowe O.S6 gt! 


PHYSICO-CHEMICAL MONITORING OF TWO WETLANDS 155 


and DRN concentration at or above 4.0 mgt! im the 
welland may provide for continuous growth in the 
Anabaena species. 


Acknowledgments 


This study was funded by the Murray - Darling 
Commission through the Natural Resources 
Management Strategy, Project 53128. 

1 am extremely grateful for the assistance [rom the 
following contractors: Mr M, C. Schultz, Glossop 
High School for the physico-chemical analyses, Mr 
P. Christy and staff Australian Centre for Waiter 
Quality Research, PMB Salsbury SA for 
identification and enumeration of cyanobacterial 
species and Messrs R, [. Gropler and J, A, Pillar for 


their regular and supportive help with Held 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 for preparing 
the manuscript and Mr G, Wright, SARDI West 
Beach for the preparation of the figures. 

Mr B. T, and Mrs E. L. Engel former owners of 
Banrock Station and BRL Hardy Ltd, the present 
owner ire thanked for allowing access to the 
wetland. Mr C. L. Rohrlach, Manager Banrock 
Station and his staff gave valuable assisiance 
throughout the study. 

The author thanks the editor, Mrs J. Bird and the 
two unnamed referees for their constructive 
comments On an earlier dratt, 


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Management Strategy Wetlands Management Monitoring 
Program. Project No. S002 Final Report. Australian 
Centre for Water Quality Research Report No, 7/93. 


ASPHONDYLIA ANTHOCERCIDIS, A NEW SPECIES 
OF CECIDOMYTIDAE (DIPTERA) INDUCING FRUIT GALLS 
ON ANTHOCERCIS LITTOREA (SOLANACEAE) 
IN WESTERN AUSTRALIA 


By PETER KOLESIK*, REBECCA WHITTEMORE & HELEN M. STACET 


Summary 


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. 


Line tans ap the Ravel Seerene ap. Aust, (F907, Pad, TS7-1Od, 


ASPHONDYLIA ANTHOCERCIDIS, A NEW SPECIES OF CECIDOMYIIDAE 
(DIPTERA) INDUCING FRUIT GALLS ON ANTHOCERCIS LITTOREA 
(SOLANACEAE) IN WESTERN AUSTRALIA 


by Prerer KoLesin®, REBECCA WiirrreMoRb? & Helen M. Sracet 


Summary 


Rolbsnk. Py 


Wireromort. Ro & Sixvck Te Me (1Q97) Ayyhondvlia qaimthevercilis, a new species of 


Cecidomyndae (Diprera) ihducing (rait galls on AnMiocerciy fitferca (Solunaceue) ty Wester Australi, Zens, 


RK. Soe, So Aust. D2, US T-161, 28 November, 1907. 


The (unt walls onthe Western Australian yellow tatlower, Admocercis inored Labill. (Sokindeeae). reditee 
the reproductive porential rf this plant, bub their etusative agent his uatiT now, been inkrown. Ourresedrelt his 
shown that ai new gill mide species. Asplondyiitmrhocerchdis. muuces these walls The farva, pupa, Male aid 


lemuleol the new species are deseribed and dlestrated. 


Key Won: Diptera, Cecidomyridae. Asyplomivia aamoceraidis, Anthacercis litte. Western Australi 
I b / : 


Iniroduction 


The gill midge fauna of Western Auswahl is 
poorly known, with only (We species having been 
desoribed previously (Gagné 1989), One of them, 
Tponnit barnemissca’ Colless, is a species which 
presumably feeds on fingi growing inthe seiland jn 
jeal Witter (Colless 1965), The biology of the second 
species,  Eveinelicornia austrdfasian Felt is 
unknown (Pelt 1915). although (his species is tikely 
to be a phint leeder considering that its congener £ 
Malweskii Kolesik, causes walls on Eirealyprs 
faycieulosaan South Australia (Rolesik }99Sa), 

The yellow tailllower Aathecercis fittorea Labill 
(Sohkinaecae). a shrub which grows lo 3 im. 1s 
endemic 16 the south-west coust of Western 
Austialia, primarily on calcareous sands in disturbed 
habits such as recently burnt areas, roadsides. Fire 
breaks ind cleared Jots (Purdie ef al. 1982 
Whittemore!) The froi valls on A. /itere have been 
known foe some time (Purdie ey af 1982 ) bul their 
causative agent has remained unknown, I uly 1996, 
one of us OR. W.) collected froit galls from A, dittorce 
containing larvae and pupae, from which adults were 
reared, The salleinducer proved to be a new speeies 
of adh midge whieh is described below. The 
development of the galls and their impact on the 
reproduction of A, fittarese are deseribed by 
Whitlemore!. 

The genus Ayplandyiie in the context of this paper 


DepAP enn ob Phen clita. Vili Tite ane Erertalopy, Miwulty oe 
Avrioulluni ond Satin Resodree Scieheds. The Lineversity ob 
Adelie PMB Glen Ounomd S Suet Sherk 

Departiaent ot Beraiy. Greversity al Western Austitlie Neotunds 
W Aust 6407 

Warr sokn RC hQ86) Aspects of Ihe dnscetendiowd fri) galls 
and repraddetve: biahhgy al anoeoreds fiterec: (Sulimacaae 
BSe (Hone) thesis. bfiversity of Western Aastratia (anus) 


is defined by Kolesik (1997) "The new species ty lo 
he attributed to PAK. 


Material and Methods 


Stems ol Aniiocerciy littarea bearing fruit galls 
were collected al Hillarys, about 20 km nortiecast ot 
Perth, on 23.971,1996, A small number of galls was 
dissected and the larvae und pupae preserved in 70% 
clhanol, Larvae and pupae retained within galls were 
reared lo adults on stems avhieh were kept in plastic 
viitls. Larvae pupated within (he galls, Emerged gall 
midges were preserved tovether with pupal skins in 
70% ethanol, Canada balsam mounts of the type 
series tur microscopic exalnination were prepared 
according to the technique outlined by Kolesik 
(19S). All measurements refer Lo the type series, 
The type specimens and other material retained in 
70% -elniel are deposited in the South Australian 
Museum. Adeéluide (SAMA) and Australian Nutional 
Inseet Collection, Canberra (ANIC). 


Asphondylia anthocercidis sp. nay, 
(PIGS 1-15) 


Haloryper 2. Hillarys, Western Australia [41 48" S, 
115°45' Ej, emerged 28.vir L996. Ro Whitlemore. 
reared from larva rom fruit gall on Anidiecerers 
littorea Labi. gull colleated 29 vii L996, 121435 
|SAMA}. 

Paraivypes: 2 8S. 3 99 5 pupal skins (SAMA, 3 
29.3 F F.4 pupal shins [ANIC], all same date but 
emerged 28-30.v11, 1996; 2 larvae [SAMA]. 2 larvae 
{ANIC], all collected with holotype, 

Other mererial [all SAMA|: 2.6 8.5 2 9,2 pupal 
shins, 4 pupac, wll same chit as paratypes: 2 larvae, 
collected wilh holotype 


158 P, KOLESIK. R, WHITTEMORE & H, M. STACE 


Figs 1-8. Asphondylia anthocercidis sp. nov. | - 4 male, 5— 8 female. |. 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. 6. End of 
ovipositor in lateral view. 7. Last five flagellomeres. 8. End of abdomen in lateral view. Scale bars = 100 jim 1-3. 5-7: 
50 um 4; 500 fim &. 


A NEW SPECIES OF ASPHONDYLIA 159 


Figs 9-15. Asphondylia dnthocercidis sp. nov. 9 - 11, 13 pupa; 12, 14 larva; 15 infestation..9, Anterior part in ventral view. 
10. Anterior part in lateral view, 11. Prothoracie spiracle, 12, Head and first thoracic segment in ventral view. 13. Last 
abdominal segment in ventral view, 14. Last two abdominal segments in dorsal view, 15, Fruit gall on Anthocercis literea 
Labill. [redrawn from Rippey & Rowland (1995)|. Scale bars = 500 wm 9,10; 50 pm 11; 100 pm 12-14: 10mm 15, 


1Ov) PO ROLESIK. 2. WHITTEMORE & WM, STAC 


Description 


Mule (Vigs 1-4) 

Colour; sclerotized parts of body reddish-brown, 
noselerotized parts of abdomen grey, 

Head. Antennin scape broadest distally, 17-20 x 
breadth at distal end, 2.62.7 x length af pedicel: 
pedicel width 12-14 x length; first Magellomere Ls 
Joy Tengith of scape. fagellomeres evenly 
evlindrical: citoumfita dense, equally distributed 
along flagellomeres. Eye favets hexagonoids eye 
bridge K-48 facets Wide, Prons with WO-17 sete per 
side. Lubella prominent, laterally with 7-10) setae. 
sclulose. Manithury palpus 3-seamented, seenjents 
successively and progressively longer: 

Thorax, Wing length 2.4 tim (range 3.13.7). 
widih 13 mm (hd-l4) Se cell pigmented 
proximally, Claws of all legs subequal ih size and 
similar in shape. ws long as empodii, 

Abdomen. Genitalia: gonostylus with two fare, 
apical (eth of same lengths aedeazus elongate and 
narrow, reachine tniddle of gonosty tits. 


Femiule (Pigs 5-8) 

Frons with 9-20 sere per side. labella with 7-4) 
selue laterally, Circunfila comprising twat 
longitidinal and two short transverse bands. Wing 
Janeth 3.6 mim (2.3-3.9). width bd mm (i235), 
Seventh abdominal sternite 18 ().0-2)2) 8 leneth ol 
sith. Genital ovipositor 1.9.8 C8 -2.0) length of 
seventh sternite; basal Jobes an ovipositor broud i 
dorsal view, divided in posterior thitd) pecially, 
fused vere: phibrous, 


Pupa (Pigs Sell. 14) 

Colour brawn. Total length 4.00 nn (3.64.5). 
Anennal horns not serrated, 242 pri (237-247) long. 
Obe Upper and three lower frontal horns. Profhoracie 
horn slightly curved, basal part about 2 x width of 
lenmminal third, terminal third setose. Abdoeniml 
dorsal spines sitiple, straight, with 2-7 pitirs Gn list 
sexment curved laterally, 


Maiure larva (Figs 12. 14) 

Colour: yellowishayhile, Total length 3.4 min (2.6- 
41). Mead cupsule strongly pigmented. postero- 
huteral extensions not developed, Spatula with four 
witerion teeth, miner pair smaller Thun outer shit 
narrow, broadened both at mid-lengih and) base, 
suroudded anteriorly and daterally by extensive 
pigmented area, Hach side of spatula with triplecane 
pair OF fiteral papillae, all setose. Six terminal 
papillic present, one pair corpiform, 2 pairs with 
short setae, other papillae as for Ayplaned ylrer (Molin 
1955) 


Gall and biology 

This gall nidge induces delornyution oF Fruits of 
Anthavercix lillaren, The upiearpellite: Ovaries are 
transtormed into glabrous, spherical to ovare, 
nippled galls, 7-18 mim long and 7-13 mm wide (Pig. 
45) and bright green to purple in colour, Inside the 
gall a chamber. about 3 mm Jong and 2 mm wide, ts 
oveupied by one larva, The chamber is always lined 
with fungal mycelia. Although the fungus was 
abundant in the many gals examined, no sexual 
stages were observed und the fungtis remains 
unidentified, Visible seeds are rarely produced in 
alls although pollination is esscatatd) to vetain the 
oull on the plant. The numbers of galls ine (ilerea 
populations are ofien very high. with the galls 
outnumbering the Hormuallysdeveloped fruits by tp tt 
38 limes (Whittemore), 

Pupution takes place within the gall, AC the end ot 
iis development the pupa cutsan opening. in the gall 
and Lifts mast af its body outside the gall. Phe pupal 
skin then splits open ahd the adall enierges, At 
Hillarys in 1996, the adults emerged throughout the 
entire bost plant lowering period, Le, fom April to 
September. 


Distribution 


Ayphandylia anthocercilis sp. nay. ws syonpaten 
with A. filtered aeross (he entire geographic 
distribution of the host plant, whieh tanges tron 
Kalbarres [27"50' &. LIASO7 KE] it the north te 
Israelite Bay [3427° S. 119923" Ey) tn the soul 
(Whittemore), 


Envimology 
The fame is derived from the generic name of the 
host plant, 


Remarks 


Asphondvlia isa worldwide genus wilh six spectes 
previously described [rom Austratia. The file history 
of three ol thent is known: AL dedonecae Kolesik 
induces walls on leaves of Dodonaea viveose Jacq, 
subsp, apadiudade (Sin West. and AL m/tever 
Kolesik and A, ericiformiy Kolestk Induce galls on 
branch segments of Malosarcia peruranulate (Black } 
Wilson subsp. pevgeaidane amd A. indica subsp, 
leiosmchyer (Benth.) Wilson. tespectively (Xolestk 
1995b, 1997). Life histories of (wo other species, A 
loewe Skuse and A, evbienndd Skuse. are unknawn 
(Skuse PSS8, 1890), The remaining species. A. fill 
Ldwards, has been reported to indice galls on the 
stent of an unidentified pling (Bdwards 1916) 
Aspondvlie Ai, As loewi and AL rebierrda ape: (ot 
considered in the present paper. The deseriptians of 
these three species were superlietal and therefore itis 


A NEW SPECIES OF ASPHONDYLIA lol 


Hol possible Lo compare them with each other or with 
A, dedonaeae, A. inflata, A. eriviformis or A, 


wuhocercidis, A review of the Austrahan species of 


this venus is planned by PL K. 

The new species differs from Asphondylte 
dedonacae in the longer adult scape. the wider teeth 
on the gonostylus. the unserrated antennal horns and 
the presence of both upper and Jower frontal horns in 
the pupa. the shafted spatula and the presence of a 
piginented area around the larval spatula. The new 
species can be distinguished from both A. inflata and 
A. ericiformis by several characters, In A. 
cithocercidis. the wedeagus reaches the middle of 
sonostylus, pupae have three lower frontal horns, the 
prothoracic horn is setose at the distal third and is 
about twice as Wide al the base as is the distal (hird. 
At least two of the dorsal spines on the last pupal 
segment are curved literally and the spatula has four 
anterior teeth. In both AL faflare and A. erictfarmis, 
the uwedeagus extends beyond the middle of the 
gonostylus. The pupa ol A, inflata has one lower 


frontal horn and that of A. erfciformiy has none. In 
both species. the prothoracic horns are asetose and 
about four times wider at the base than at the 
(erminal third. In the pupa of A. inflata, only the 
prominent pair of abdominal dorsal spines on the last 
segment is curved laterally: in A. erieiformis all 
spines are straight. In both species the spatula bas 
two unterior teeth. 


Acknowledgments 


The field work Was supported by the University ol 
Western Australia while R.W. was completing bet 
Honours degree, We are grateful to J. D. Gray. 
Department of Horticulture. Viticulture and 
Oenology University of Adelaide and R, J. Gagné. 
Systematic Entomology — Laboratory USDA 
Washington DC for their comments on an carly dratt 
of the manuscript. 


References 


Coctess, DL bh (1965) An extraordinary Australian 
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Eowakos, FW. (19760 Two new Austaihan Diptera Ann. 
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herr, EL P.(1915) New genera and species of gall nidyes. 
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GaAGNE. RoI. ( 1989) Family Cecidomyiidae pp. 152-104 In 
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Press and Rod, Brill, Honolulu). 

KoLbstk. P.(1995a) A new species of Fovineticoria Fell 

(Diptera: Cecidomyiidae) on Eucalyptus fuseieulase in 
South Australi, A Ads ent Soe. 34, 147-152, 
— (1995b) Asphonidvita ledenieae, a new species of 
Cecidomyiiae (Diptera) damaging leaves and 
branches of hop-bush, Dodonaea viveosa (Sapindacene) 
in Australia. Tran. Ro See. 8. Ads EIQ 17 L176. 


(1997) Two new species of  Asphamnidylia 
(Diptera: Cecidomyitdue) fram  Hilosareie spp. 
(Chenopodiaceae) m South Australia, (hie. 121. 59-66. 

Monn, E. (1955) Beitrige zur Systematik der Larven der 
Honididae = (=Cevidomytidae, Diptera) 1. Teil: 
Porricondylinue und Itonidinae Mitleleuropas. Zedlovien 
105, 1-247. 


Purpiz. RoW... Symon, D, B, & Hara Lb. (1982) Family 
Solanaceae pp, |-208 fn George, AS, (hd Flora of 
Australia” Vol. 29 (Australian Government Publishing 
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Rirrey. E, & Row asp, B. (1995) “Phints of (he Perth 
Coast and Islands” (University ol Western Australia 
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Skuse. FA. A. (1888) Diptera af Australia, Part 1, Proe, 
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(1890) Diptera of Australia, 

Supplement |. /bid. 5, 374-412, 


Nemitocera - 


FIRST RECORD OF THE ORIENTAL LATRINE FLY, 
CHRYSOMYA MEGACEPHALA (FABRICIUS) (DIPTERA: 
CALLIPHORIDAE), FROM SOUTH AUSTRALIA 


BRIEF COMMUNICATION 


Summary 


Blowflies are well known for their ecological, veterinary and forensic importance’ but 
they are also significant medically as mechanical vectors of dangerous pathogens’. 
The Oriental Latrine Fly, Chrysomya megacephala Fabricius, 1794’, 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* and have been implicated in the 
transmission of viruses”, protozoans’, enteric bacteria’ and helminths*. The larvae are 
also known as facultative parasites in traumatic lesions in humans and other animals’. 


Tis tans af tlre Royal Socteny aS) Avast (1997), Wy, 164 The, 


BRIEF COMMUNICATION 


FIRST RECORD OF THE ORIENTAL LATRINE FLY, CHRYSOMYA MEGACKPHALA 
(FABRICIUS) (DIPTERA; CALLIPHORIDAE), FROM SOUTH AUSTRALIA 


ny 


Blow/lies are well Koowir Cor their eeologiesl vetering 
dod Jorensie importance! hut (hey are alsa significant 
medically us tneclanical vectors af dangerous pathoweny: 
The Oriental Lutrine Ply, Cheywonwd ievecepheata 
Pubrighis. [P94 is fotorigus mn this regard, Adulls of rhe 
aynanthirapic Lorn ol this species (see helow) unc attracted 
fo Toodstulls, Hunan and ainiinel Gieees and carrion! and 
have been Tipheaed in (he (ansmission of viruses’. 
protozoa! entero haeler iy and belts. The larvae see 
WO KAQWI TS fHcuTLative parasites i triumatie lesions i 
HUM shel other guiiagels:, 

TPrepart lure the first record oo eueeey ede in South 
Australia, It was eaueli ai lire numbers ina liver-baited 
Lap on The Nort Terrie campus or tie University ol 
Adib in April. 997. 1 had Wot previously encounluredt i 
uP this sie. despite periodic upping of blowHies over the 
preceding seven years. Neither hac | encountered it 
elsewhere TH South Austadin There are pe specimens uf © 
Hievecephale Ton this sake in the eitoiiologival collection 
abthe Soult Australian Miscuimauil the closest record vil iy 
fo Soul Avsttuha oo that ool three fentiles at 
Murrcinbatenian, New South Wales (s4 58'S, taued2" be), 
approvimalehy 950 kaye the east al Adulane UK, R. Norris 
Divisional Entomol CSIRO pers, comm, [Y7). 

Ulsewhere in Aust G. megerephaliy occurs weross Ue 
fie north ol the eentinent. down the Gast codsts of 
Oueenshid und New South Wales and in south-wese 
Westen Alisttalia, W lias not beee recorded frei Vietoria or 
Rist! 

Clinyyomye iegueephuli is probably a 
HEPOUCTON to South Austria, as il is actively ex pamting 
ies Fe iT GHer parts AP Wark Sittce the TOTS HL hue 
invaded New Aeahana!!, several parts of Afhied und arens 
HH South. Cantal! and North America! Ut is alse fined 
i) lapin and ds widespread throughout the Oriental reign 
sod (he Ausiro-Mahiyan and Polyresin subrewions al the 
AUSTiiasian reson, 

(Cheyanne negaveplidto Oceuis 1 Lwe harms which are 
Horpliglowiehy duel ecolusicdlly different. "They hive been 
defined by Kurahishil cs the “norm aid “derive lt’ 
Faris The Horni) (arn ts resteicred to tropredh faresis on 
Sout Pacific hands from the Bismurek Atehipelaust to 
Suro, Has believed tebe the plestonarphie Lorne eb the 
species, The derived fori is synantiivapic and dispersive 
HHO is Thouwwht Wo have orieinated jit Papua New Guiness ain 


Ween 


farzinelioglu, AZ, (06) Glowlles  iRichiinwad 
Publishiie Co Lik Shout, 

‘Greenberg, Bo (1971 "Flies aid Disehae. vol 2. Bialowy 
und Discuse Transmission” (Priicetaa University Press, 
Pringen 

Wabrictus, fh ©. (1794) “Enromuleia systematics 
crendiliy ch aueGi Secunda elisses, ardiney. wenera, 
species unheeds sy munis. loci bservationibus, 


descriptiomibus’. Tome +00. G, Pratl. Maroiaes, 


the westerh bodndary of the species! ynewseal 
Uieuiburion! “The individuals eolleeied from Adebnide are 
ofthis (orm, 


‘The derved form can be distinguished tram the normal 
form hy the greutly-enkirged onimatiche in the upper (wee 
(hinds wf the eve Th males. “Phe noenal form hus aaly 
shighlly enhurged onamuidia in this vesion % Sine the two 
fornis differ so markedly in there eeologicul preferences and 
WH SOME AApeets Of their merphology. they may be 
sufligivutly divergent genenediy lo warnint subspeeitie vr 
“yen specitic status. This miehl berevweuled by buwhenicul 
analysis. bs creed aut recently for ather closely-rclaled 
blow(lios af the genus Culliphera'. although a detailed 
Morpholosicul comparison would abso be required betore a 
conelusion about tverr status could he teached. 


Boil forms af) megacephete can be distineiasted fron 
ullathee known Austin species of Chryyannya hy the 
following Motphotogicul characters: Hairs on prevlar kaob 
black. loner thio height ef Roob: anterior thoracic 
spineless bhickish brawny legs bhick) subyibrmsal sealae 
bhicks eyes Hr males willl oniauitidiae ig (he upper Uwo-thirds 
enlaived and sharply demuncuted from the small ones in the 
lower thirds [rons in females wider tn the middlehie tl, 
However, the species is similar physiologically (ae rithet 
species al Chrysomved in that it is Mermophilie, ant 
(herefore, in South Australian will presumably be active 
OOTY ducing the waemer months of the year that is benween 
Oetober and April. 

Because al its habits, Co itegdcepheald deserves serious 
ubention front it publie health perspective and its 
distribution ih Atistralig: should (herefore be monirered, tts 
larvae could also be cneountered iy forensic cases in the 
Adeltide revion, but because their moarphalagy bears 
Superficial similarity to that ol seme other commun euriion- 
hreeding species al blowlly. they eGuld be iistaken bor 
them. Sinee the pile of development al Co mega lela ita 
gWen temperature differs significantly from these orbper 
species? misidentificdiion Of this important fly could 
Ted to serious errors TH extinates of (he tiie singe death ot 
DUT Corpses 


Uihank Drs Do A. Duekhotise, RoR. Nortis ane ALB Bird 
for theiy user commenr on the munserph, Tho ahi 
also thamks Dr Norris for his helphil advice and casisiince 
iouttaigry the loa of specimens. 


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World” (Butlerworths. London), 

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‘Harris, A. & Down. Ho A. (1440) Ani Prop, Med, 
26. 7AY-ROG, 

‘Lima, M.1. POS. & Baie, KOO) Achy Biol, Manan, 20) 
Oless 


164 


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°’Spradbery, J. P. (1991) “A Manual for the Diagnosis of 
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Herman, T. J. B. (1990) Weta 13, 12-13. 

“Braack, L. E. O. (1991) Onderstepoort J. of Vet. Res. 58, 
311-312. 

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. 
Kurahashi, H. (1984) Dispersal of Filth Flies through 
Natural and Human Agencies: Origin and Immigration of a 
Synanthropic Form of Chrysomya megacephala pp. 37-63 
In Laird, M. (Ed.) “Commerce and the Spread of Pests and 
Disease Vectors” (Praeger Publishers, New York). 
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'7Wallman, J. F. & Adams, M. (1997) Aust. J. Zool. 45, 337-356, 
'SDear, J. P. (1986) Fauna N. Z. 8, 1-86. 

“Wells, J. D. & Kurahashi, H. (1996) Med. Entomol. 
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~»Wijesundara, D. P. (1957) Ceylon J. Sci. 25, 169-185. 
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J. RF WALLMAN. Department of Zoology, The University of Adelaide Aust. 5005. 


THE STATUS OF CYCLOSTRONGYLUS 
MEDIOANNULATUS JOHNSTON & MAWSON, 1940 


BRIEF COMMUNICATION 


Summary 


Cyclostrongylus medioannulatus Johnston & Mawson, 1940 was originally described 
by Johnston & Mawson’ 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’ revised the genus and considered its relationships 
with related genera. She found the type species of Cyclostrongylus which had been 
erected in 1939°, to be identical with that of the type species of Oecsophagonastes 
(Johnston & Mawson, 1942) and placed Oesophagonastes in synonymy with 
Cyclostrongylus. 


Treason of the Royal Seeter: aS. Aust, (1997), T2165, 


BRIEF COMMUNICATION 


THE STATUS OF CYCLOSTRONGYLUS 
MEDIOANNULATUS JOUNSTON & MAWSON, 1940 


Cy lostonevtis oedivcmilunis Johnsen & Mawsen, 
IM40 was onwingtly deserbed by Johnston & Miewsion! 
from three females found int the stemmed af Waeropas rents 
(ae) now Clivlvvale thediy collected trom the Burnett River 
distviel in Queenshund. The authors commented that the 
words differed trom other speeies of Cyelosmene vis 
fohinston & Mawson. 1939 only in having a iareosy 
Supporting ring dround the bucest! cuyily, Mawson? revised 
the genus and considered tts relationships with relitedt 
gener, She found the type species of Cyelasmenowes 
which lid been erected in 1439 to be mdenteal with that ot 
the type specivs of Gesephavornaytes Sobroston d& Mawson 
1942) and placed Cesoplemanasies in syaonyinw with 
Cy lusenevios, Maawsoie listed the valid species ol 
Cyclosiomeyliy us €) wellahiae Johnson & Mawson, [949 
(ype species) eecurring in Wallahia biculor. Co gallerélr 
Johnston ok Mawsan. 1939 Geeurrine Tn Mew reps 
rufourtsens, ©, kertune (Miowson, }YS5) occurring in Ay, 
euxont ull ME rifeerisnin CC heprtons iMawsen, 65) 
oeourrm im MW. dieseliy and Co parina Joliston & 
Mawson, 1939 uccurring in Ad, pervs. 

Ol the other species previously 
Cyelosironeviis, ©. déysanilis, lohrston & Miawsen. 1958 
veeurring in We Bivelor owas referred tw 
Manmpostoneylnides (Yaron, T6b and Co cledeeds 
Johnston & Muwson. 1939 oecurmmg in A defer now A, 
yivdHels Waa new gents later deseribed as Alecesromd 
Mawson, P974° Subscquedthy ©. valle: wos eedesenbed 
is Wellabmema gallard!, 

othe case oF OC) medina Mawson! Pound (har the 
specimen libelled as the Pype was i femule Rageplicaryins 
qtralis (Monning. 126), abyiously placed there in error. 
Sinee the original material could Hat be found she declared 
the species a speeles frnynivencde, 

Mawson? also revised the genus Macrapesrang lie 
Yorke & Maplostone, 1926 and ereeted Hiree new fener 
Including — Pepevestrongylis, — Pupovesmronwuytiis yas 
chuetensec by an oesaphawus With eV lindrical Corpus. a 
qurrow isthiusindh ending ina bolh; anda buces! capsule 
whieh is thickest ar midength and has an anterior border 


aussie tn 


‘Johnston, TAH & Mawson. FV (1940 broc, Linn. See. 
NSW 65, 468-476, 

‘Mawson, PM. (1977) Trans. BR. Soe. S. Aust, LOD, 19) 20. 
Vohnston, TH. & Mawson, PML (193) Prac. Linn, Soe 
NSW 64, 514-536, 

‘Beveridge, b. (984) Aust. Zool. Suppl. Ser YL. 1-8. 


Without projections, The species icluded the type species 
P wellabiee Jolston & Mawsen, 1930), accurting iM, 
puifegrisens and Wo hivalen Po peaevani ovcurring in 
Petrowale turevalis., M. vageml and Me puilieriseas and 2B 
iomecoccurring in ML tema, Bevenndye® then uiended the 
coneriv diagnosis, extunied the host niige ut FE pearvens to 
inchile M. fidivinesis wind described two new species. 7! 
mocrupodiy ocvurriny 1 ML pigeniteis, ME cufies und ME. 
robustus, und FL diviegele oveurring ww Thivlogale 
vinmatica, UT hrunii and Petrogale peryephom Beveridge’ 
commented thal Modivlevele Oecurred commonly in 7 
vigimetica ti northern Queevishin, anid i Perweale 
penephone tram central Queensland hit had mut heen 
found in 7 artgeninien or 2 thers in southem Oueenshind, 
Spratt ev af! disted ©) tedioanaildtus as a species ol 
Popovastronglis. 

Reeendy a slide (HC 2660) labelled Cyclosmennlies 
mrecdiodneataiies to PL ME Mawson’s handwriting, was found 
in the South Australian Museum. Adelaide. The slide 
COUsISDs Ol pitees Of bWwo Tenmale worms Henibed uy resin 
und has allkached to jl another band-wrillen note by Mawson 
Indicating that the specimens are two ol the anly three 
females ever vollected. The charueters of the oesophiyiis. 
buccal vapsule, female tail and ovejector thal can be 
Jistinguished in these two femules are consistent with the 
characters al Hapovestinteyiins, Purther. the bea! capsule 
has an unnulirihiekening around the middle as 15 lound in 
Potlivlagete. The eesophagiis (O07) Mn is sharin in O. 
medivaniilaius than in Polivlawale (09-094) mim. Phe 
measurements OF the posterior end of CL nreetiocmiaiiy 
Ure OL Cong rue WHE those of A iivtowle. Vulva te fall up 
0.35 compared to 0.7240, 80. nd Wil 0.24 compared to 0 45- 
0.59 mim respectively, The viawini vera oof ©. 
mediocmoulatin. however. ts Ure sie tenth as that ot Je 
Nvlegale. Cvelostronuyluy medioannidanie. Weretore. is 
leary wspectes of Fapovestueyils anel is mest similar to 
Po thyfowale. Addition materiah including: male 
specimens. iA needed betore un exact determunition can be 
thade. No other specimens of Popovasinertio, Towever 
have been Found in 7 iferty bo date 


Mawson, PM, (1977) Trays, R, Soe S Aust TOL 51-62. 


"Beveridge, 1. (1956) Bull, Mus, Nati Plist. Nal. Parts 4 
Sen 8 (4, 2), 257-265, 


‘Spratt, D.M., Beveridge, £., & Walter, ILL. (1091) Rec, 
So Aust. Mus. Motion Ser 1} 105. 


LAR SMALES. Departinent of Biology, Facully of Applied Scienve Central Queenstind University Rockhampton Old 


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