Skip to main content

Full text of "Transactions of the Royal Society of South Australia, Incorporated"

See other formats 


VOL. 126, PART | 
31 MAY, 2002 


Transactions of the 


Royal Society of South 
Australia 


Incorporated 


Contents. 


Wasson, K. A review of the invertebrate Phylum Kamptozoa 
(Entoprocta) and synopsis of kamptozoan diversity in 
Australia and New Zealand - - - - - 
Murray-Wallace, C. V. & Bourman, R. P. Amino acid racemisation dating of 
a raised gravel beach deposit, Sellicks Beach, South Australia 
Bourman, R. P., Alley, N. F. & James, K. F. European-induced environmental 
change in the Adelaide area, South Australia: Evidence from 
Dry Creek at Mawson Lakes- - - - - - =- - 
Pledge, N. S., Prescott, J. R. & Hutton, J. T. A late Pleistocene occurrence of 
Diprotodon at Hallet Cove, South Australia- - - — - 
De Silva, P. & Riley, I. T. Aspects of the survival and reproduction of Anguina 
microlaenae (Nematoda: Anguinidae)- - - - - - 


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


39 


45 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 126, PART | 


TRANSACTIONS OF THE 


ROYAL SOCIETY OF SOUTH AUSTRALIA INC. 


CONTENTS, VOL. 126, 2002 


PART 1, 31 MAY, 2002 


Wasson, K. A review of the invertebrate Phylum Kamptozoa (Entoprocta) and 
synopsis of kamptozoan diversity in Australia and New Zealand = — 
Murray-Wallace, C. V. & Bourman, R. P. Amino acid racemisation dating of a raised 
gravel beach deposit, Sellicks Beach, South Australia 
Bourman, R. P., Alley, N. F. & James, K. F. European-induced environmental change in 
the Adelaide area, South Australia: Evidence from Dry Creek at 
Mawson Lakes -— = = = = 
Pledge, N. S., Prescott, J. R. & Hutton, J, T. A late Pleistocene occurrence of Dipretedor 
at Hallet Cove, South Australia - 
De Silva, P. & Riley, I. T. Aspects of the survival and reproduction of Anguina micro- 
laenae (Nematoda: Anguinidae) 


PART 2, 29 NOVEMBER, 2002 


Spratt, D. M. & Nicholas, W. L. Morphological evidence for the systematic position of 
the Order Muspiceida (Nematoda) 
Shattuck, S. O. & McArthur, A. J. A taxonomic revision of the Camponotus wiederkehri 
and perjurus species-groups (Ifymenoptera: Formicidae) - -— — 
Dutkiewicz, A. & von der Borch, C, C. Stratigraphy of the Lake Malata Playa Basin, 
South Australia - - = 
Dutkiewicz, A., von der Borch, C. C. & Prescott, J. R. Geomorphology of the Lake Malata- 
Lake Greenly complex, South Australia, and its implications for late 
Quaternary palacoclimate 
Styan, C. A. & Strzelecki, J. Small scale spatial distribution patterns and monitoring 
strategies for the introduced marine worm, Sabella spallanzanii 
(Polychaeta: Sabelidac) 


Brief Communications 
Taylor, D. J. First records of two families of freshwater Amphipoda (Corophiidae, 
Perthiidae) from South Australia 


Insert to Transactions of the Royal Society of South Australia, Vol. (26 Part 2, 29 November, 2002 


A REVIEW OF THE INVERTEBRATE PHYLUM KAMPTOZOA 
(ENTOPROCTA) AND SYNOPSIS OF KAMPTOZOAN 
DIVERSITY IN AUSTRALIA AND NEW ZEALAND 


By KERSTIN WASSON 


Summary 


Wasson, K. (2002) A review of the invertebrate phylum Kamptozoa (Entoprocta) and 
synopsis of kamptozoan diversity in Australia and New Zealand. Trans. R. Soc. S. 
Aust. 126(1), 1-20, 31 May, 2002. 

Kamptozoans are tiny suspension-feeders superficially resembling bryozoans or 
hydroids, but phylogenetically affiliated with spiralians such as polychaetes. All 150 
of the described species undergo budding, either to form clonal aggregations or 
interconnected colonies. This review provides a synthesis of current knowledge about 
Kamptozoa, updating the last general English-language description of the phylum 
provided by Hyman in 1951. Kamptozoan morphology, reproduction, and 
phylogenetic relationships are characterized. Finally, each of the three major 
kamptozoan families is described with examples drawn from Australia and New 
Zealand. Currently 37 species are known from this region, but many more remain to 
be discovered. The Australian fauna is unusually rich and varied and includes the 
world’s largest kamptozoan species. 


Tramyacnons af the Revel Sacre ofS, Anse (2002), ooh 1 20 


A REVIEW OF THE INVERTEBRATE PHYLUM KAMPTOZOA (ENTOPROCTA) AND 
SYNOPSIS OF KAMPTOZOAN DIVERSITY EN AUSTRALIA AND NEW ZEALAND 


by KERSTIN. WaASSON 


Summary 


Wasson, be (2002) A review of the invertebrate phyllim Kaniptazed (enloproeta) and sy copsis oF kaniplozead 
diversity im Australian New Zealand. Pres. R, Sec. 8. die. 12606), 1-20, 31 May 2002, 

Kamplovouns dre tiny’ suspensivn-feeders superficially resembling bryocoais ve hydreids. hub 
plylowenetically alitated with sprraligas such vs pulvehactes, AI TS0 of the deserhed species: undenmu 
hadding, evher to form clonal ageregauions or inicreemmected colonies, This review provides a synthesis of 
cirrent knowledge dbuut Kamptozea, dipdating wie last general benulisti-lagguase deseription af the phylum 
provided by Plyman iq 19S). Kaimptozain morpholory, reprodiehoo, and phylogenctic relationships are 
characterized. Finally, cael ol the three major kamptovoun fantilies is described wah esamples drawer fan 
Austalin and New Zealand, Currently 37 speetes ire know (rom this region. but many more roman ta he 
discovered. ‘Phe Australian fauna is unusually meh ond yard und ineludes the world's hireest kumptovean 


Spevies. 


Introduction 


Kumplogoans are tiny, fentaculate suspensian 
fecders that live in all oeeans of the world. Clonal 
averegations Of independent zoids (Mig, lay are 
foun on iyertebrate hosts, while colomes ol 
interconnected voids (big. Tb. ¢) grow on various 
substrata, Each zooid has the shape ofa wine glass: 
a bowlLshaped calyx. is supported by a slender, 
Hexible stalk that attaches basally le lhe substratuin. 
The calya ms ringed hy a horseshoe of ciliated feeding 
fenlacles and containg aw U-shaped gut a smull 
fangliin, a pair al protanephridia und one ar two 
pairs OF gonads. The space ettclosed by the teritac{es 
forms-an atrium, the deepest park of which serves as 
u brood chamber for develope embryos, 

Kaniplozoan soaids actively bend and twist. Their 
characteristic monon is reflected ins the phytin’s 
scientific name (Creeks Acwptested — (er bend) and tts 
common nume, “nodding beads”, Another name for 
the phylum, Entoprocta.is less appropriate because it 
suggests an aliihation with the Eetoproeta (Bryozid) 
andi imiplics Crraneously thatthe anus is complets ly 
enclosed by the (entacular cilation. Ramptozouns 
hour only vd superfierl resemblance to bryozoatis. 
wilh owhieh they were  Gnee grouped, 
Developmentally, Kamplogouns are spiralians bul 
their phylogenetic relationships to other metazoans 
remain enigmanie. 

About 150 species have been described worldwide 
but kamptovoun diversity probably exceeds S500 
speeds (Nielsen 1989) While they are widespread 
ane are quile-abandant i some microhabilats, most 
of the world’s kamplozoans are poorly characterized 


Hk hom Stowe tutional Pstimineg Research Resenye, Royal Okay 
CAYS TOO 1 SAL R-inails rescurchimelkhanislouh are 


oy not known at all, because most species are tiny 
und casily overlooked, Kamplovoans occur in all 
oceans. from the intertidal cone to several hundred 
metres depth. A few colonial species live in brackish 
water, and one ih freshwater, Representatives ofall 
three mayor lamiltes (Loxosomatdae, Pedicellinidae. 
Barentsiidae) haye been found in every marine 
reyton that has been thoroughly Surveyed. Phe fourth 
faanily (Loxokalypodidae) has been found only one. 
ii he northeastem Pacific. 

The main purpose af this review is to synthesize 
current knowledge about the Kuimptozou. The last 
general Eniglish-linguage deseription of this phylum 
was provided by Lynn (1951), and there have been 
many udvanves in nur undersianding sinee that time, 
In summarizing what is known about kamptozouns, | 
draw heavily on work by two recent pioneers in 
kamplozoolowy. P. Erschermann lee. bnsehermann 
1972, 982) and C. Nielsen (ue, Nielsen 1977. 1996: 
Nielsen and Jespersen 1997), A second objective of 
this review qs to highheht the rch und unosual 
kuimptozoan fauna of Australia and New Zealane. 


History of study 


Kamplozoans were frst iustraled by Ellis (17546), 
Pallas (17740, b) described the first species as 
Brachionuy cemmus. placing it ina genus of rotilers 
The same sapeties was pluged in the new gens 
Pedieclline by Sars (0835), who considered it a 
naked bryovean, Vai Beneden (1845) contributed 
the first Worough monogriph of kamplozoan 
morpholoyy and reproduction, Phe genus (rneiedte 
wus described by Leidy (1831) and Levan by 
Referstei (1862). Allman (1856) pointed oul the 
uniqueness of kamptozoan calyx and tentucle 
structure. Nitsche (1470) conceived of Pedicellau, 


x) K. WASSON 


sterile fertile 
segment segment 
stolon 


| 
sterile fertile 


segment 


stolon 


Fig. 1. Structure of kamplozoan zootds. (a). Lecosomedlfar sp. 


Urnatella and Loxosama as a natural grouping. the 
Entoprocta, and separated them from all other 
bryozoans, the Ectoprocta. Hatschek (1S88) first 
raised the entoprocts to the level of phylum. Clark 
(1921) proposed the name Calyssozoa to distinguish 
this phylum further from the bryozoans; Cort (1929) 
agreed with this intent, bul changed the name to 
Kamptozoa, since the name Calyssozoa had already 
been applied to another taxon (the enidarian 
Stauromedusac). Late in the 19th century, a number 
of prominent scientists investigated kamptozoans, 
emphasizing embryological and phylogenetic 
questions (e.g. Barrois 1877; Harmer 1885; Seeliger 


a 


segment 
WY 


“SA St ° stolon tip 


\ 


a 


3 on sponge. (b). Pedicelline whitelesvii. (c). Barentsia sp. 1, 


1890). Since then, only a few researchers at any one 
time have focused on kamptozoans. 


Morphology and physiology 


External characteristics 

Kamptozoan zovids ure generally constructed of a 
stalk, basal attachment and calyx (Piz. 1). The height 
of individual zooids ranges among species from 0.3- 
30 mm. The stalk develops as an outgrowth of the 
calyx to form a exible, roughly cylindrical support. 
Clonal forms (Family Loxosomatidac) have a 
specialized basal organ (either a muscular suction 


A REVIEW OF THE PHYIUM KAMPTOZOA (EN TOPROCTA) 3 


disc or a differentiated “foot” with an associated 


; gland (Fig. Ja)) with which they attach to 
ak. invertebrate hosts. Beneath the stalks of most 

XY , Ss as ealoni: ; : Pamilies Pedicellinidac 
WY AO aD a ) colonial forms (Families Pedicellinidace and 


Barentsiidae), stolons (Fig. Tb. ¢) adhere to various 
fig, 2. Locomotion hl Lotuscma cite. Modilied {rors living and non-living substrata with cuticular 
Nielsen (1960). adhesions. The cup-like calyces range in height from 
0.2-1.2 mm and are ringed by a horseshoe of 
tentacles. The mouth and anus ure at opposite sides 
POSTERIOR of the calyx, regarded as anterior and posterior 
respectively (Migs 3, 4b). The calyx is bilaterally 
symmetrical: a vertical plane through mouth and 
anus divides the calyx into right and left: mirror 
images (Fig. 3). The region above the stomach is 
sie z ventral (this region was below the stomach in the 
( larva): the bottom of the calyx and stalk are dorsal 

LErT ; RIGHT 1, ; 

SIDE = =e SIDE (Pig. 4b). 


teptenc ilar 
ern biter: 


emo 
wane 


Body wall, musculature.and support 
} ‘ The body wall is a single-layered epithelium, 
a covered by a glycoprotein cuticle containing a trace 
(0.06-0.45%) of chitin (Jeuniaux 1982) but no 
collagen (Emschermann 1982). The cuticle is 


{rales vepshtindriin 


il ANTERIOR generally thickest.on the stalk, which may be darkly 
pigmented. moderately thin and transparent on the 
Vig, 3. Diagrammutic lop view ola pedicellinid calyx, calyx, where the internal anatomy can be readily 
a b VENTRAL 
tentacular contracted 
membrane A tentacles Sxpaheee 
; entacles 
testis 2 mouth 
oesophagus 
ventral | / 
stomach ! f- stomach 
“roof” 
: anus 
{ mouth tentacular 
Ls fe membrane 
stalk 7 | rectum 
? 4 ovary = 
rf oesophagus 
] embryo 
A ; ANTERIOR POSTERIOR 
=500um veniral intestine 
stomach stomach 
A , “roof” 
star-cell organ 
fertile sterile 
segment. segment =100”"m 
at DORSAL 
stolon 


Vig. 4. Structure ofa kamptozoan calyx, (a), Anterior view of contracted male Pedicel/ina whiteleges? void. (hb). Side view 
of expanded temale 2 whifefegesi calyx. 


| kK WASSON 


observed through the body wall and thiinest on the 
inner (frontal) side ol the tentacles (Nielsen & 
Jespersen 997), 

Strong longitudinal musele fibres beneath the stalk 
epithelium praduee the characteristic hending 
inouons Ol Kamploeodn vooids. Circular muscles are 
Jimited to the tetlacular membrane yind sphincters 
hefween purls oF the gul The structure of musele 
libres has been deseribed by Unischermann (196%b, 
1982), Reger (1969) und Nielsen and Jespersen 
(997). Katiplozoans Jack a coclom. The cavity 
surrounding the culyeal organs and extending inta 
the tentacles and Stalk is filled by a loose Muial mutris 
of nesenehyme cells which acts us a hydrostatic 
skeleton and, together with the cutiele. lends the 
stalls prendity (Brien 1959). 


Locumonon ded nN 

Al karnptozeans have larvae thal swan or creep by 
cilary action, While larvae represent the main 
dispersal mode tor most vatorial and perhaps many 
solilury Species, some specs ure mobile at other 
sulaes m the lile-cyele. In some loxosomatid species, 
Hewly released, asexually produced buds cur swir 
with (heir stalk forward. propelled by their tentacular 
villas fad few loxosomatids. adults may alsa be 
capable of such switining (Atkins 1932; Ryland & 
Aust F960) Wielser (966), [fh loXxosoinatid spevies 
Whose adulis cunaltach repeatedty to the subsiraium, 
passive driffiny of dewehed goods may alse serve 
fut dbspersal Mose colonial fornis are sessile as 
adulis. hut in the freshwater species Upynitel/e 
wraciliy Lewy 185). shork propagation stolons of bwo 
or three goaids often break fram a lamer colony. 
leading lo rapid colonization ota favourable area by 
fiagmenis of the saine original colany which have 
spread by drifting (Emsebermann 1987), 

In some species in the genus Loyexaina. zovids 
employ their basal suction dises to somersaulr across 
Uwe substratuin CAssheton (9) 2: Nielsen 1964), 
Siovings ie a manner fascinating and unique by a 
senies uf vymnustie ellorts, whieh combine the 
aanlity of the luingaros and the deliberition of y 
neometer caterpillar’ (Assheton, 1972). The oui 
bends down until the calyx atliches by four lony, oral 
lenlucles io the substratum: the suetion vise then 
detaches fron the substratum and fips over the calyx 
fo reathiwh some distance from its orienral site: the 
void then returns Loan Upright orientation (Pig. 2). 

While adult locomotion jecurs jy only sone 


species. the men-locomotory bendims motions of 
attached Zooids are churacterisue OF all members of 


the phylum. Although the rapid ind vigoraus 
nodding of kumplozoans inimediately catehes the 
observer’ seye. the mechanisms and stimuli invely ed 
liave out been thoroughly examined. Bending of the 
Suu results from shortening of lonyttielina museles 


on one side (Aricn 1959) AO Stronger bendine 
response ys Obtiined by stimulation of calyces than 
ol stalks (Cort 1936). The nodding and weithine may 
help zouids escape predatars, may djminsh 
overgrowth by fouling organisms. op nay peevent (he 
ealyces tram repeatedly flterina the same avaler, 

Finally. individual calyees have a ebaracteristic 
response Lo disturbunee. When jeritured, the tentacles 
curl irewards and are ghelosed by a delicate lnyer of 
Ussue, the lenlaculir membrane (Figs 3, 4). whieh 
lithlens like a draw-string purse by means of efeular 
musculuture, This intolling of the tentacles 
reseinbles. the contraction of a sea anenone wore 
than the felaetion w bryeoun lophophore. 


Peveling ane cigesiee seater 

Kamplovoans are suspensmon feeders on 
phytoplinkton and other particulste food, Lact) 
lentacle hak five longitudinal raws of ciliated eetls 
(Atkins 1932: Mariscul 1¥65; Nielsen & Rosiguard. 
1976). On the sides of each tentacle (Pig 3). large 
lutera! eels bear compound cia that heal urweards 
the tentacle’s frontal midline (Nielsen & Rostaaurd 
1976): these cilin “enerate the feeding eurrents. 
Water is drawn between the Wentucles from below the 
lanticular erawn. then sent upward aveny tron, the 
calyx (Atkins 1942) The tater) ili also caplire 
partieubile food trom Ure water currents they ereates 
kaniptozoans employ a downstream cellecting 
mechanisin (Nielsew & Rostaadrd 197). Inside the 
rows Of lateral cells, rays of narcow bterafroital 
cells hear short cilig thal presumably transfer food 
from lateral tor frontal etl (Mariseal 1965). Ihe 
fronuil midline of cavh tentacle has a single raw al 
larze frontal cells bearing short cilia and small 
mules vesicles: these cilia beat with the elftetne 
Stroke towards the base ol the tentacle. and transpert 
captured partieles ina band of mucus te the base ol 
the tentacles (Nielsen & Rostyaard |976), Pood 
particles then travel in ciliated gutters, the right aye 
leli arial grooves (Pig. 3) te the mouth CAthies 
193), 

Some kiumplozouns apparently: trap crhales and 
other organisms by rapidly contracting the rentaeular 
crawp (Atkins 1992). One Sotaretic kanplozoun hus 
special multicellular estrusive organs (lime-twin 
glands”) that discharge hollow, sticky threads, 
presiimably fo capture larger prey alems that 
supplement its diet of suspended — particles 
(bmpchermann 199236), 

Kamplozouns have a U-shaped gut wilh both the 
moulh and anus opening yeotrally (igs 4. 4b). Ve 


digestive triets of larvae and adults are Siniple tubes 


of ciliated epitheliuns divided ina four regions, anil 
lave heen eharacterized by Beck (1938) and Nicher 
and Jespersen (1997), The crescentshaped muuth 
(hig. 3) leads toa funneltike bueeal cavity, then to a 


A REVIEW OD TIE PLY LEM KAMP TOMOA CN DOPROD TAY £ 


Harrow oesophapus that opens into a voluriious 
stomach filling much of the calyx (Pig. 4b). Ingested 
particles are embedded in strands of mucus thal are 
kept in constant rotation by cilia in the stomuch: the 
ul (teks musculature except al sphineters between 
rewions and food is Wansported entirely by ciltury 
geo fBecker 1938) The strands pradually 
consolidate into clumps as they pass towards the 
infesting. Digestive eheymes are Seereled by 
whindular cells in the ventral “roo? of the stomachs 
absorption aveurs boll in this region of the stomach 
und i the intestine (Beeker 1958). The stomach 
leads to a short intestine. and then to the rectum, 
Which progeets above the Moor of the atriuny (rigs 3, 
4h), such thal lavees released into the lenticular 
Walter current ure swept away from the calyx. When 
the tentacles are contracted, the rectam tolds [u-like 
over Lhe alo. 


Cirenlatary aud respiratory: swatenes 

Since kamptozoan calyees are ony. dilusion is a 
sufficient transport mechanism: na special organs 
(agililale circulation within the calyx, Loose 
mesenchyme surrounding the organs allows [or the 
free cireulauon of dissolved vases and nubrents. 
Conmmitry to earher indications (ee. Uyman 1951), 
there dre mo free -ameeboeytes. enhancing nutrient 
Ininsporl within the  mesenehyme matrix 
(Emsehermann (9699). In loxosomatios, Murds also 
juins treely between the calyx and the stalk. helped 
on their way hy museular movements. I rin 
colommal kaniptozoans, diffusion may aor suttice tor 
circulation throughout the zooid because Ure stalk is 
often much longer than in loxesemiutids and ( parity 
separated from the edlys by a cutienhir septu. 
Podicellinids aid barentsiids have a efretilatory 
structure, the slareell organ (Fnischermann 96%). 
AS stack of flanened. stellote cells spans the narrow 
sone belyeen the stitk and calyx (big. 4h). The 
topmest cell contracts and expands. like a pipette 
hulby rhyiimie pulsations of the stacked cells purnp 
fluids beiween ealyx and stalk (Emsehermano 
)OOUn) 


Ayerendo 

A pan of flame-bulb protonephridia, located just 
posterior of the oesophagus (Pig. 3}, apparently 
functions mainly i fon pegidation and 
osmoreenlation (limschermann 1982), bach 
protonephridivm is composed of tour mulliciinted 
vells. Two of the cells form a terminal organ. witha 
Hiltrauion area where they interdiyitates the third and 
aun cells encircle the nephridial lumen, and the 
lourth cell forms the nephridiopore (Franke 1993), In 
loxopomalid calyees. the two protonephridia open 
separately into (he atrium. while tm slolorites: Ley 
open through a conynon nephridiopore (lV ramke 


1993). The freshwater Kamptovoun Credrelle 
wucihs has a more highly developed excretory 
system, wilh 30-40 prolonephirdia in the ealya, and 
inany others in the sialk (Finschermann 1965). 

Excretion of metabolites likes plice ia the venral 
stomach “poot? (Pig, 4), a region that is often eye 
calching because itis conspicuously coloured by the 
pigments of consumed phytoplanklon, The Jarse 
vacuoles of cells in thts regton contain precipitated 
uric acid and guanine as well as algal pigments 
(Beeker 1938: tanscheriiaan = 1965). These 
jntvacelular inclusions are eventually expelled ini 
the slomuch and voided 


Nervans svstenr an sense ovat 

A large, dunibbeli-shaped ganglion fies ventral lo 
the stomach. just posterior to the protopephiricia (hig 
3). Nerves radiate Trim this subenterie ounglion ta 
the Lenticles. to other parts of the calyx, and to the 
sialk, Many kamptozouns have unicellular tactile 
receptors on the tentacles and on the surface of the 
calyx (Nivken & Rostesard 1976). In adiition, 
JoXosornatids often have & pair ol lateral sense 
organs consisting of ciliated papillae on the riglit and 
lefl sides of the calyx, There are no nervous 
connections between zooids in a colony: earlier 
suggestions (Hilton 1923) ofan miterzooidal nervous 
aystein have been rejected (Fmschermann 198?) 

The larvae of many loxosamauds have a pad of 
eyes, each consisting of a cup-shaped pigment cell, a 
Tens cell ind a Sensory cell. The structure of the eye 
isunusual in that Gieht enters perpendiear to. rather 
than parallel lo the long axes of the sensory cilia 
(Woollacott & Eakin 1973). No adult kamptozouns 
are known lo have eyes but zoids of some species 
contract in response ty Sudden exposure ta bright 
light (Eiischer ai L982), 


Reproduction and development 


serial reproduction 

AIL kamplovoaos derow by budding. bn 
loxosomiulids, which hve on otter pnverlebrittes, 
buds form in two anterior or anterolateral regions of 
the calyx, oflen roughly level wilh the lop of ihe 
stomach (Migs la, Sac b, Sa. 6). Buds pray be 
produced. alternately or Sintuftaneously at the twa 
budding sites. The basal part of the bud’s: stalk 
develops ai altichment organ, Che bud may remain 
wltached io ils “patent” for some time, feeding and 


even becoming sexually mature, but it eventually 


breaks away often attaching to a nearby speton the 
invertebrate host. 

Colonial kamptozoans also bud ut the anterior lace 
of zooids, but budding occurs earlier in the Hile of 
aooids that i loxosarnatids (Brien 154) ‘The 
cous producing buds are offer themselves stilf tiny 


6 Kk. WASSON 


=200um 


Fig, 5. Asexual reproduction. (a). Calycal budding in Loxvoseme/la sp. $5. (b), Calyeal budding in Loxosoma sp. |. (e). 
Budding at the stolon up in Burentsia metsushimana, (d). Budding at the stolon tip in Pedicellina pyriformis, 


buds; cach stolon tip is a bud primordium forming 
anterior to the next youngest bud (Figs |b, 5c, d). As 
the buds grow and differentiate into fully formed 


zooids, they are separated by intercalating growth of 


the stolon. Eventually this growth ceases and a 


septum with a central opening forms on each side of 


the zooid, partitioning the stolon into fertile (zooid 


bearing) and sterile (without zooids) segments (igs 
Ib, c, 4a). Because of this pattern of formation, the 
anterior side of every zooid along a stolon faces the 
growing stolon tip. Colony form can be more 
complex in some barentstids, which bud trom 
specialized stalk regions. In some species, resting 
buds (hibernacula) are formed at stolon tips, These 


A REVIEW OF THE PHYLUM KAMPTOZOA (ENTOPROCTA) 7 


undifferentiated buds are enclosed in single or 
multiple chambers and are covered by a thick cuticle, 
They germinate only after the stolonic connection to 
the rest of the colony is severed, and following 


exposure to low temperatures (Toriumi 1951; 
Emschermann 1961, 1982). 
Pedicellinids and  barentsiids, unlike most 


loxosomatids, can regenerate calyces, Old calyces 
degenerate and are shed and are replaced by a 
budding process at the apical stalk tip comparable to 
that at stolon tips. Injured barentstid zooids can 
regencrale new calyces and stalks even from basal 
stalk and stolon remnants (Hyman 1951; Brien 1959; 
Mukai & Makioka 1978). 

Patterns of bud formation at the histological level 
are very similar in all kamptozoans (Secliger 1889, 
1890; Brien 1959), An epidermal proliferation of the 
anterior body wall of a zooid results in an 
evagination that forms the bud primordium. Budding 
is essentially an ectodermal process; while some 
mesenchyme cells migrate from the “parent” into the 
bud, no endoderm is contributed, At the apex of the 
bud primordium, an invagination forms, then 
constricts into an upper and lower yesicle, which 
become the atrium and the digestive tract, 
respectively, A narrow passage connecting the 
vesicles becomes the mouth, while the anus breaks 
through ata later stage. A constriction soon separates 
calyx and stalk and the latter elongates. Eventually 
the atrial cavity breaks through, freeing the tentacles, 
and the bud begins to feed, 


Sexual reproduction 

Most loxosomatid calyces are protandric, with a 
discrete male phase followed by a female phase 
(Nielsen 1971; Emschermann 1993a); calyx 
gonochorism has also been reported (Ilarmer 1915; 
Prenant & Bobin 1956). Barentsiid calyces are 
typically gonochoric (Wasson 1997). Some 
barentstid colonies are gonochoric, too, containing 
calyces of only one sex; other barentsiid colonies are 
simultaneously hermaphroditic, with both male and 
female calyces formed along the same stolon (Mukat 
& Makioka J980; Emschermann 1985; Wasson 
1997). A very few barentsiid species have 
simultaneously hermaphroditic calyces (Johnston & 
Angel 1940; Wasson 1997). Some pedicellinids have 
gonochoric calyces in gonochoric colonies (Marcus 
1939); others have gonochoric calyces in 
simultaneously hermaphroditic colonies (Dublin 
1905); sull) others have — simultaneously 
hermaphroditic calyces (Brien 1959; Emschermann 
1985). 

The reproductive system is rather simple in both 
sexes. Gonad rudiments derived from mesenchynial 
cells first appear above the stomach as a pair of tiny 
oval translucent vesicles (Mukai & Makioka 1980). 


These grow into large ovoid sacs, consisting of a 
one-layered epithelium which ts the germinal layer 
from which the gametes arise (Brien 1959), In 
simultaneously hermaphroditic calyces, a pair of 
testes lies posterior to the pair of ovaries. Each gonad 
feeds into a gonoduct, and ihe right and left 
gonoducts merge at the ventral midline to open 
through a common gonopore posterior to the 
ganglion (Brien 1959), 

The testes grow rapidly and may fill much of the 
calyx (Figs 3, 4a). The spermatozoa have elongate 
heads (Emschermann 1982; Franzen 19836). 
Spawning has rarely been observed; apparently a 
cloud of sperm is released following a sudden 
contraction of the calyx (Dublin 1905). 

All kamptozoans brood their embryos and release 
fully formed larvae. The ovaries remain much smaller 
(han the testes (Fig. 4b), with only a few germinal 
cells at any one time differentiating into oocytes. The 
small (40-80 tm) but yolky eggs (Franzen 1983a) are 
fertilized in the ovary, then discharged into the 
deepest part of the atrium, the brood chamber (Cori 
1936; Marcus 1939; Mukai & Makioka 1980). A 
glandular region of the oviduct secretes a pliant 
envelope, which encloses the embryo and extends 
into a cord which tethers it to the floor of the brood 
chamber (Marcus 1939; Brien 1959), The ovaries 
release one or a few eggs per day in alternation, the 
youngest embryos pushing the older ones farther 
from the gonopore (Brien 1959), The tethered 
embyros, like a varied bouquet of balloons, can 
occupy a substantial portion of their mother’s calyx 
(Fig. 4b). The brood chamber contains many embryos 
in a regular succession of stages from cleaving eggs 
to contractile larvae. When larvae hatch out of their 
envelopes, they remain attached to the atrial wall by 
the cord, with their mouth and ciliary band upward, 
allowing them to feed on particles in their mother’s 
current (Brien 1959; Mariscal 1965). Swimming 
larvae are released about a week after fertilization 
(Mukai & Makioka 1980), 


Embryology and development 

Kamptozoans show typical spiralian, determinate 
development (Barrois 1877; Hatschek 1877; Harmer 
1885; Lebendinsky 1905; Marcus 1939; Malakhov 
1990), Cleavage is spiral and the 4d cell is a 
mesentoblast cell that proliferates — loose 
mesenchyme in the interior of the embryo, 
eventually giving rise to the muscles (Marcus 1939), 
The arrangement of cells at the animal pole 
resembles an annelidan rather than a molluscan cross 
(Marcus 1939). The larval mouth forms very near to 
the anterior margin of the blastopore, which 
eventually closes; the anus forms secondarily as 
well, There is never any hint of coelom formation 
(Marcus 1939), 


ss 


kK WASSON 


Pig 6, Pholophores. Gai, Grabevos gnd htevue oF Ceveseviiig sp. §,(b), Larvae ol Pedicelling wiittelewedi, (2). Larva of B 


wracilis yar, siypley ALL Taures 20 sume seule 


apical organ— 


frontal 
organ 
cillary_— & 
ring 
# 
mouth* 
atrium | anus 
foot 
a 


stomach 


anus 


tentacles 


stomach 


stalk 


d e 


Vig. 7. Schematic representation of metumorphoses in Pecdice/ling ceria, (a), Swimming larva. (b), Newly settled larva, 
(oc) Period of vigorous anteriar growil, td), Zooid wilh separation between still and enlyx; tentacles forming. (eo) 


Pocding vooid. Moditied Irom Cori (1929), 


Kainiplozoun lirvae are vencrally hat-shaped (Pips 
6. 7a), Salvini-Phaowen (1980) suggested the name 
thalophora (Cireek: Mielas ~ dome; tholia — straw 
hat) for them. There are a number of detailed 
deseriptions of larvae (e.g, Burros 1877; Con 1920: 
Marcus 1939; Mariscal 1965; Nielsen 1971) from 
various regions of the world. The hyposphere of the 
larva is deeply Jndented into the prominent, hat-like 


cpisphere when the larva is swimming. The curve of 


the L-shaped gut is a the upper part of the hit; 
mouth and anus open on the ventral surface (Vig. 7a). 
‘There is an apical organ atthe top of the hat, a frontal 
orean ut the front of the hat. and a@ ring of Tony 
compound cilia around the brim, just above the 
mouth (Figs 6. 7a). Below (ventral to) the mouth, 
there is a second band of shorter compound eilia in 
the shape of a horseshoe, with the opening of the 
horseshoe at (he wnus: the band ts also broken behind 


the mouth, These two ciliary bunds beat in 
Opposition and cuplure particles that are then 
Iransported to the mouth by short cilia i (he atrial 
urooves, Which ran between the two hands of longer 
cilia from anus to mouth on both sides, as in the 
adults (Mig. 3), Often there ts a ciliated creeping loot 
in the ventral area between mouth and anus (Pig. 7a). 
Some tholophores show Unusual features (stalked 
vesicles, a spiderweb pattern of ornamentation, an 
adhering layer of detritus, etc.) that are not yet 
understood (Nielsen 1971). 

Tholophores resemble the trochophores of some 
spiratians (Balfour 1885: Cori }936: Nielsen 1971, 
1995, Emschermann 1982). The downstream- 
collecting ciliary bands of tholophores are similar to 
those of trochophores in cell-lineage, structure, and 
funchon (Nielsen 1995). The apical organs of 
Iholophores also resemble those of trochophores, Bul 


ARV IRAW OD TER PEAY TOM KAMPTOZOA LEIS TOPROL TAY 9 


unlike waehophares, most Lholophares havea frontal 
orwvin and a ciliated foot, and their hyposphere is 
deeply indented jnte the episphece when the larvit is 
swimming, A few loxosomaticl farvae lack the frontal 
organ and feet and have a mere pronaunged 
hyposphere, thus more strangely resenrbling 
trochophores, but these forms are considered 
denyed, not ancestad within the phylum (Nielsen 
1971) The strongest resemblance of thalophores is 
lo adult kamptozoan calyees; larva und adult shore 
the sume Shape, Struvture of the digestive system, 
ALU WIT abt grooves. and avery siinilar ciliary 
fvediug mechanism: 

Larvae from only a few Austialian species ere 
known, One Lerayemie/ta larva (Vig. 6a) 1s clonvate 
in the anterior-posterjor axis, Witb adhere particles 
and a well-developed fel. The Peedfeelfien 
iwitedleved Johaston & Walker 1917 larva (Fig. ob) 
is Lub in the ventral-dorgal axis, covered with) i 
remarkably dense laver of detritus, dnd lacks a foot 
{Wasson 1995). the Barevtyararaeniy larva (iw 
fe) is relatively big, vccupying a large portion of the 
parental wilys. [bis about as high as wide and ts free 
of adherent particles. 

Most tholaphores appear capable of both 
swimming und creeping; il is nol known ta what 
vatent the larval period of most species ts pelagic or 
henthie, Most thalophores are feeding larvae with a 
functional wut. However, the larval period ol tiary 
Kam plozouns appears lo be extremely short - hours bo 
days (Mielsen LY712 Emsebermatin 1982. Wassuny 
1998) -s0 the larva’s Teeding while still in the brood 
chamber may be more important than feeding after 
felease, On the other hand, some Lovesonee kirvae are 
Olen caught inthe plankton andar presumed to have 
along pelagic phase (Rigersten 1964: Nielsen 1966). 

Metamorphosis has been warefully deseribed i a 
few kaniplozoan species (Bartois 1877: Marther 
Das?) Cort 1936; Marcus 1939; Nielsen 1971: 
Emsehermann (982), Phe larva etceps ou the 
substratum, testing ik awith the frontal organ, be fare 
iiaching by the reeion around the frontal organ. 
settling: on the anterior side (loxosometids) or by 
Wiiching by the foot resto, selmi anh the 
eiroumlercuce of the retmeted wentral eviary girdle 
(pecicellinids aod barenistids), The atrium becomes 
enclosed by a vonstiction af the episphere dorsal to 
the ciliry girdle (rig. 7h). The atrium and dizestive 
tract ure rotated upwards as a result of rapid growth 
wl ihe anterior region of the episphere (Piz. 7c). 
Nesta separation forms between calyx and stalk and 
the Javier elongates (Pig. 7d). Cillated tentactes form 
as eclodermal protuberances at the periphery of jae 
atrium (Fig. 7d). roughly in the location uf the 
degenerating larval ciliary bands. [nally ihe utrium 
breaks Open, releasing the tentacles, and feeding 
bouis (Fig. Te). 


While in all colonial and many elonul species the 
lurva does metamorphose direetly ima the ada, 
some lovasomatids have precocious buddmpe in 
which the larva does nol metumurphose, bul instead 
dies as the buds it bears grow and are released 
(Harmer 1885; Jawersten 1964: Nielsen 17h). In 
effect, the larval bud. father than the larva jtself, is 
the roule to adulthood in these species, Ia (he most 
extreme eases, the larva is completcly consumed by 
an internal bud that lorms while the larva is still 
within its parent, and the larval tit isabsent (Nielsen 
JY71). Seamhe remurkuble species disphyy further 
heleroehrony: (he hids themselves already haye buds 
in tum or even ure sexually mature while stall 
camained in the larva (Jigersten (964) 


Phylogeny 


Poasvil record 

Kamptozoans fossilized by bioimmuration occur 
in upper Jurassic rocks mm Greal Britain (Pode) & 
Taylor 7993) and northern brance (J. ‘Todd, pers. 
camm. 199s) The  strocture of vance 
unambiguously identities them as members af the 
extent genus Burentsia, These Mesozoic Fossils sera 
minimum time for the divergence of what is 
probably the most derived tumily, suwvestiny that 
ancestral members al’ the phylum may date hack 
much turther. 


Relat with amher hveviehrare teva 

Listerivally, there hive been several proponents of 
close relatronship bepyect Kamplocodns and 
bryaroans (ee. Harmer 1883; Marcus 1939; Prenant 
& Bobi 1956; Nielsen 1971, 1995). Zo0ids of bath 
taxa have a Loshuped gut and are ringed by ciliated 
tentacles, Budding and hiberngeula occur tn both 
taxa and neither has an endodermal contribution 
from “parent” to bud. in boll) groups. kuval evespats 
have sensory cilia oriented at right angles te the 
invaming light (Woollacote & Lakin |973), 
However, jiny other workers egjeet we close 
evolutionary alliation at Kamptozoans with 
bryogouns (eg, Allman USS: Latschek Pass: Cort 
1936; Hyman T9S1; Brien 1959: Jiversten 1972; 
Emsehermann |982). They attribute the similar body 
plans of adults to common suspension (bediny hubits 
and tiny budy srs. Budding and hibernacula are 
found in many sessile taxu and lack of endodermal 
contribution to buds is found in pterobranchs and 
some asvidiais as Well as kaniptosouns anal 
bryovouns. The simihinty of the furval eyes is 
striking but, since the eyes are constructed somew hal 
diflerently (Weollacott & Gakin 1973) they are hot 
necessarily hontolagous. 

Beyond useribmy similarities to convereeiee, 
epponents of a close rckationship between 


te) K. WASSON 


kamptozoans and bryozoans emphasize — the 
differences between the two taxa. Kamptozoans have 
no coelom; bryozoans do, although it is. rather 
unusual. Kamptozoans have protonephridia and 
vonads; bryozoans do not (Emschermann 1982), 
Kamptozoans retract their tentacles by curling them 
inwards and pulling the tentacular membrane around 
them; bryozoans retract the whole polypide and the 
lophophore shuts like an inverted umbrella (Brien 
1960). Kamptozoans have downstream-collecting 
ciliary bands, while bryozoans have upstream- 
collecting ciliary bands (Nielsen & Rostgaard 1976; 
Nielsen 1995). A key component of the bryozoan 
body plan is the box-like cystid, absent in 
kamptozoans. There is little 
communication or nutrient flow — between 
kamptozoan zooids, or of polymorphism among 
zooids; these features are characteristic of bryozoans 
(Brien 1960). Kamptozoan nervous systems are 
limited to single zooids, while bryozoans have 
colonial nervous systems linking = zooids 
(Emschermann 1982). Kamptozoan metamorphosis 
usually involves retention of the larval gut and other 
larval structures; bryozoan metamorphosis is a 


“catastrophic” reorganization without retention of 


larval features (Brien 1959), A recent molecular 
analysis of complete 18S rRNA sequences (Mackey 
etal, 1996) provides further evidence against a close 
relationship between kamptozoans and bryozoans, 

If kamptozoans are not closely related to 
bryozoans, with what group of animals are they 
allied? Based on embryology (Brien 1959; Nielsen 
1971, 1995: Emschermann 1982) and molecular 
sequence data (Mackey ef a/. 1996), affinities must 
be sought among other spiralians. Some authors have 
been impressed by similarities between kamptozoans 
(especially loxosomatid larvae) and rotifers (Barrois 
1877; Harmer 1885; Davenport 1893; Hyman 1951), 
or turbellarian flatworms (Salvini-Plawen 1980). 
Haszprunar (1996) proposes a_ sister group 
relationship between kamptozoans and molluscs, 
emphasizing similarities such as a chitinous cuticle, 
a circulatory system with sinuses, and a ventral, 
ciliary gliding sole (at some stage In the life-cycle) 
and a pedal gland. Alternatively, kamptozoans may 
be more closely allied with annelids (Emschermann 
1982), Until further evidence resolves the question, 
the precise phylogenetic position of kamptozoans 
remains an enigma. 

The similarity between adult kamptozoan calyces 
and tholophores has led to the proposition that the 
phylum originated by paedomorphosis. This 
hypothesis is developed in depth by Jiagersten 
(1972), who envisages the original kamptozoan life- 
cycle as consisting of a planktotrophic trochophore 
larva and a benthic creeping adult with a ciliated 
foot. In this paedomorphic scenario, the original 


evidence of 


motile adult was eliminated but its ciliated foot was 
retained by the larva, which became sexually mature. 
This larva then gave rise to a secondary benthic 
adult, which retained the same ciliary feeding 
mechanism as the larva, although the ciliary bands 
eventually were drawn out on to tentacles, The new 
adult developed a stalk, an attachment organ, and the 
ability to bud. Haszprunar ef a/. (1995) recently 
presented a similar scenario of a paedomorphic 
origin for the phylum, but beginning with a 
lecithotrophie larva. 


Key to the orders and families 


| (a) clonal: new zooids budded at calyx and then 
released; musculature continuous between stalk 
and calyx: star-cell organ absent; larva usually 
with paired frontal Organ oo. eceeeeteseeeeeseeeetees 
BetsiZonerpeceensraeh O. SOLITARIA, F. Loxosomatidae 
(b) colonial: new zooids budded at base of older 
zooids or from stalks and remain connected to 
CACH OTHE... cece O. COLONIALES; 2 
(a) zooids connected by non-septate basal plate; 
musculature continuous between stalk and calyx; 
star-cell organ absent; larva with paired lrontal 
OTEAN ve ceeeeeeeseeeeeeeeeees Sub.O. ASTOLONATA, F. 
Loxokalypodidae [known only from North- 
eastern Pacific] 
(b) zooids connected by septate stolon or rarely 
(Urnatella) septate basal plate; musculature not 
continuous between stalk and calyx; star-cell 
organ present; larva with unpaired frontal organ 
Wensvi nguteensecadesrgadaenh dhietigars Sub.O. STOLONATA; 3 
3 (a) stalk of zooids with continuous longitudinal 
musculature, fairly wide throughout whole 
length, stalk and calyx often with cuticular spines 
Eh CSdarccite Sepa del os yied goigegy coi eentrety F. Pedicellinidae 
(b) stalk of zooids alternating between wide 
muscular nodes and narrow rigid rods; rods often 
with cuticular pores; stalk and calyx generally 
without cuticular spines ....... asansteag FP, Barentsiidae 


to 


Systematics and Australian diversity 


Order Solitaria Emschermann, 1972 
Family Loxosomatidae (Hincks, 1880) 

The order Solitaria contains only a single family, 
the Loxosomatidae. Nevertheless, it is the largest 
natural grouping of kamptozoans, with about 100 of 
the 150 described species. Three loxosomatid genera 
are currently recognized (Nielsen 1996); 
Loxesomella, Loxomespilon, and Loxosoma, and are 
distinguished primarily by their basal attachment 
structures. About 20 species of loxosomatids have 
been reported from Australia and New Zealand but 
only seven of them are described (Appendix). Many 
more species certainly remain to be discovered; until 


A REVIEW OF THE PHYLUM KAMPTOZOA (ENTOPROCTA) V1 
a . b 


Fig. 8. Loxosomatid diversity. (a). Loxosomella sp. 3 showing foot, (b). Loxosomella velatum. (c). Loxosamella sp. 1 with 
larvae at top of calyx. (d). Loxosoma sp. 2 showing basal muscular disc. 


1? kK. WASSOMN 


more thorough surveys are undertaken, it is 
impassible: to ussess the rue diversity Australia's 
loxasoniatids, 

Loxosoimatids, whieh form clonal muyeregutions by 
cilycal budding. are considered the most 
plesiomerphie group of kamptozoans (imsehernunn 
1972), The highly contractile zovids are often very 
sitmall (lens than bimim high). Calys and stik are nol 
Sluirply separated und longitudinal musculature is 
eontindous between them. The ealyx und tentacles 
ime generylly driented obliquely to the stk (Figs ba, 
h). Lhe calyces are Olen Compressed in the anterior 
pesteriorusis, sometimes so strongly that the zoeids 
tesemble puddles. 

Ii Lavesamedlfe, the basal purtiol the stalk al buds 
is differentiated (nto a structure resembling a human 
foul (Fre la. fap The heel ot the loot is anterior and 
conus a conspieuous gland. A groove lined hy 
uceessury ahund cells runs frond the |ieel to the 
posterior Loe, Where TLopens, When a bud is ceoleased 
(rom ils “parent”, it allaches to the substratum by ils 
Tov, In some species the good retains the whindulor 
foot lor its chtire existence and is able to defuel and 
reytlach repeatedly over its Hiletme. tn other species, 
the foot ef the bret deeenerates ler atiachment anil 
the adull besos permanently cemented to the 
substratum (bigs &b, e). Zouids of the monseypie 
pons Lovamespilon have a very reduced stalk and 
foot byt otherwise resemble Lorosenela cootds 
(Bobin & Prenant 153; Nielsen 1996). Seven 
deserihed aml eight undeseribed species of 
Loxasomella are known Tron Australia and New 
Zewund, and most of the species in the Appendis 
Whose basal atuchinent structures could not he 
ussessed (and So ure listed merely as “Loxosomatic 
spo) probably belong to Loxasemedls as well. 

In Lavoseme, each Zoold 1s attached by a muscular 
suetion dise at the base of the sulk (Pig, Bd 
additional suction dises may oecur pusteriorly and/or 
al the hase of the lentacles (Nielsen 1996), Zoos 
retain fhe abilipy to detach and reallach, sometimes 
roovrog: wetivelw aeross the substratum (Pie 2). All 
known Latesene larvae have stalked vesitles on tit 
episphere and undergo buddihg rather than a nornial 
metamorphose  (Nieken 1996). Only three 
(Undeseribed) Loxeasonr species are huowo tron 
Australivand New Zeaband, 

Most loxosomitids dwell.on other jovertebrites. 
Australia and New Zcaland they have been reported 
Irom ovurious spolges, a Sipunculon, varius 
palyehuetes, two berudineans, a squat lobster, two 
priwns, und Wiriots bryevadns (Appendix) As tore 
patented hasts my this region are examined for the 
presenee of loxosemalid symbionts. thes lst will 
veriainty grow. Bach loyosoniatid species appears To 
Ihe enher a sinele Hust species on at limited set ol 
Poll host species. Lurvite, aml pussibly alse buds 


and motile adults, ci colomae new hosts: it is net 
known whether propagile preferenee on diferente 
frortihity on different host species is responsible tor 
the lafer disiribulion ofadults. Association with other 
invectebrates has clear benefits for the loxssonniid. 
The vooids are-ofien located jn the pathway. of the 
host's feeding or respiratory walter currents, which 
they may use for their own ciliary eeding (Nielsen 
1964). Vhe host probably offers the frawile saaids 
profechon from predation or o(her damage. Whether 
the presence of loxosomutids negatively affects theyr 
hosts ts net known: Williams (2000) has shawe that 
host epidermis may be modified by loxosorutid 
symbionts. 

Worldwide, many loxosomatid species (about 
50%) live on polychaeles; they ure Sound on or 
between the parapodia, on the wills, on the sete. ur 
under the elytra oF menibers of ten polychaete 
families (Ni¢hsen L989), Laxesoneller eiapeatricule 
Willams 2000 and seyen undescribed species of 
Joxosamatids are known fronr polyehuers in 
Australia and New Zealand (Wigs 3a, bh, Auk 
Appendix). 

While loxgsomtatid species diversity is highest ov 
polychuetes, loxosamatid density is probably tizhest 
OW sponges. Loxosomatids inay form sirikiiely 
dense avyregations on Sponges — sometimes 
LO0.000) zooids on a fist-sized sponge (huteler 
1968). Some oF these sponve-dwellime forms are 
unusiilly darkly pigmented. and an aggecation 
aginst the background ofa brightly coloured spatige 
can be eye-catching. Pwo undeserihed species of 
Toxosomele ave khown [ron spotiies ith Austialia 
und New “valand (Figs la. 8a). 

Sis lososomatid species in Australia CLecosene(ler 
heeve f. eireware, 1. civrifertun Le pusiltnin, b 
bevorin (all Tarmer IYIA) 22 spo} grew on 
bryozoans (Appendix). Most of these specns ane 
ornamented by odd cirrilorm- organs or papillae (Pie 
Sh, eh und share other sinilarities that suggest they 
comprise a clade; horh the ceolowy and the laxanonmy 
ol bryozoan-dwelling species ment further attention. 
Some bryovoun-dwelling loxosomarids, orivinally 
desenbed by Llarmer (1915) from Sihaed expedition 
material, Hive Hy very close association with tlnei 
hosts. One miniscule loxosomatid species even byes 
in the compensation sac of its hast: almast every 
compensation sae in an dtested bryoacuun woluny 
vontuins a loxosomaud zuoid (Harner 1915). 


Order Colotiates Lyysehermatin, (472 
Sub-Crder Stolonata Fpsehermann, 1972 

‘The sub-order Stolonaly is ihe other large natural 
grouping oF kamptozoans and exhibits the sevond 
hasie body plan within the phylum. The calyees ol 
siolonaies are wenerally larver than these oat 
loxdsomuatids, with stranver eiiary clirrents that 


ARE VIEW OF THE PIV LUM RAMPTOZOACENTOPROC TA) 13 


apparently free the zooids from dependence on 
hosts’ ciliary currents (Umscherimann 1972), 
Stolonate calyces are generally laterally compressed 
(Vis ducw 4h; Mies Wav. L0b) and musculature is 
reduced. offen to just a few longitudinal strands. the 
atrial relractor muscles, which extend from the base 
of the calyx fo the atrium aod serve to depress it 
(Lanschermunn 1972), Calyx and stale are separated 
by wcuticolar diaphragen and the calys-stalk junction 
is spammed by the circulatory star-eell organ 
(Pmavherann }969%a): the longitudinal musculature 
ofthe stalk is tot continueus with that of the calyx 
Nhe stalk ollen bers culieulur pores or spines which 
vary in sive andl derisity with environment 
vondilions. Stolonaie zoids, as their name implies, 
row on cylindrical stolons iat are usually divided 
into fertile (vould-beuring) and stenfe (iro soos) 
segments by trapsyerse septa (Figs |b, c. 4a). The 
sepia may function to spice the zooids, thus ayoiding 
intecterenee in feeding, or may prevent damage by 
sealine olf (ntact sections from harmed ones. 

Stolonate kamptozoans are members of the sessile 
bemlye community and often grow together wilh 
hydroids and bryovouns. They are preyed upon by 
nudibranvh motluses. sovne of which appear to 
spective on barentsid species (MacDonald & 
Nvbakken  J978); predation by turbellarkin 
Hhalworms has atsa been observed (Canning & 
Carllon 2000), Although seldonp conspicunus. 
stolonate kamptazoans are often fairly abundant. 1 
iave found sfolonates intertidally at every site 
surveyed in Australia dnd New Zealand by colleeting 
yanrous Substrata (mostly sponges, use dyin, 
bryovouns. worm tubes and bivalve shells) in the 
Held and examining then in the laboratory. I some 
lowalities, an astounding SO%-75"% OF all substrata 
searched were infested with slolonale kamplozouns, 
although the level was usually about 5-10 a other 
Sus, 


Fanily Pediceltinidwe (Johnston, 1847) 

The fannly Pedicellinidae is considered prere 
plesiomomphie than the Barcoisitee (ascherinan 
1977): pedigchinid zoids retain a Krly simple 
aooidal structure, with undiflcrentivtcd shulks that 
have continuous mittsculatuic five genera ute 
recognized hab four of these — (Chaaspite, 
Loxesomatoides, Myrosonn, Sanmeavellay) contain 
only one or two spectes, and have pot been reported 
trom Australia or New “Zealand. The larger genus 
Puheellina comprises ubout iwelve species 
worldwide. sis Gf whieh are Kaown Tron Australia 
dnd New Zealand (Appendia), 

To colder waters of this region, 22 a fit degeni 
Johnston & Walker J9re (Pius lh, 4a, b, Ge) 1s 
Ubiquitous aid ean be collected reudily frond costal 
habitats (Wasson 1995). This speeres is recounized 


hy its spination, by the distinctive, ghistenme, double 
rows of jurge cells on the tentiteles. and by its tall. 
purlicle-covered larya (Vig. 6b). lo warmer waters, /? 
whiteleee/i is replaced by another abundant species, 
Povanpoera Harmer 19 ls (Fig, 9a), whieh Js. 
charagicrized by short, squal zooids ornamented willl 
filitorm spines (Wasson (995), 

A rarer pedicellinid fram Otayo, New Zustand, and 
Tasmania is. Pediotline ppritoreis Ryland 19s 
(Pig. Yb) The stules gow ap tea 6 min high, and 
calyces can be almost | min hight this species is a 
gtint amone the avorld’s pedicellinids. Zoids are 
also invre densely clustered in this species than is 
other pedicellinids. Phe wide stolons tack septa: the 
absence Of intervening sterile segments allows 
2oaids to grow very close toxetbher along the stulon- 


Family Barentsiidae fingsehermann, 1972 

This family is characlerived hy the division of the 
stalk inte wide, Mexible. muscular nodes and array. 
rizid. AON-mMusculdr rods that are ollen perlorated. by 
pores (Pigs fe, We, ble), An incomplete cuticular 
seplum separales each node from the rod above 1, 
There is @ minimum of one basal node and one rod 
upies! ta ti, but many speeies have multiple 
alternating nodes. and rods, lending a segmented 
appearaice to the stalk, 

Five genera of barentstuls are recognized, 
Coriell. Pseadopedicellina. Pedicullmepsis and 
Urnereiia (the sole freshwater form) each contain a 
sindle Species: mast af the roughly thirty known 
harentstid species belong to the genus Bearentsie, 
Seven barentsytl species ure Known from Australia 
wid New 4eolwid (Appendiad, six in the genus 
Barentsia and one my the genus Perfieellinapsis. Vhe 
comman species of colder wikers, Barenisia spy | 
(Pigs le. Wa, bj, is characterized by small. celteale 
zudids only about] mm high. ustulhy with 1-3 series 
ol'stalk nodes und rods. In warner waters, sp. | is 
supplemented by ZF yenjenlaeer Hayiner 19tS (hig. 
We) which has many (average 4-5) serigs of stalk 
nodes and rods, In its seamented stalk structure. & 
germenive tesembles ihe cosmupolitun specacs &. 
bereden! (Moettinger TANT) (found i Australian 
harbours), Grom which ib can be distinguished by its 
Wider, Shority nodes and by the loss promunced 
Anterior orientation of the catys. 

Pedicvllinepsix frutivoye Vines 18s4 (Pre. [bbs 
a remarkable barentsiid apparently endentic tp 
souchern Austalian warers (Appendist. Zooids 
spiral around a hard centru! stem (Pig. tai fren 
which cach voonl is Separited by a septum. Lach 
sium resembles a tree-fern, with the newest zooids at 
the apical growing tips older reeions of the stem 
where Zooids have degetichited have spiral patteris 
of cow sears as de lower regions of tree-fery 
Irons, The thick, regi stems branch, forming bushy 


\4 K. WASSON 


a Cc 


¥ =250um =110ym 


Fig. 9 Pedicellinid diversity. (a), Pedicelling compacta, (b). 2 pyriformis. (c). Po whitelegsii. 


A REVIEW OF TTIE PHYLUM KAMPTOZOA (ENTOPROC TAY | 


=100um 


Uh 


Cc 


Fig. 10. Barentsiid diversity. (a) and (b), Baresia sp. | in side and anterior view, respectively, (eo), B. geniculate. 


colonies that may reach 30 cnr across, larand away 
the record fora kamptovoun. They are anchored to 
the substratum by a lush basal growth of tree 
stolons, which extend downwards to serve as 
rhizoids and secondarily back up the stem, 
becoming intertwined with it, tndiyidual zooids, 
Uthourh uUnscgmented, grow toa length of 6 mim, 
Phe nodes ire large and annulate (Mig. tle). The 
rods are a deep golden brown due to a very thick 
cuticle and make a striking contrast to the pale 


calyces und nodes. The rods are decorated with 
alternating rows of bubble-like pores and pairs of 
lateral cuticular ridges (Fig. 1b, ©), a pattern of 
stalk ornamentation not known from any other 
barentsiid, A large cuticular spine extends up past 
the stalk-calys junetion on the aboral side of the 
yoo (Fiz. 11h). With its long fist of unique 
features, Pedicellinapsis fruticusa may be the most 
highly derived member of the phylum Kamptozoa, 
It has yet to be observed alive, 


lo Kk. WASSON 


_— 
_ 


_ 


=200um 


Fig. 14. The barentstid Pedicellinopsis fiitticosa. (a). Colony, showing zooids spiraling off of thick main stem. (b). Calyx 
and posterior spine. (c) Stalk, showing large annulate node and regularly ornamented rod, 


Perspectives on the Australian fauna 

Reports of kamptozoans from Australian waters 
are scarce, and currently only about 37 species of 
kamptozoans are known from Australia and New 


Zealand (Appendix). However, the Australian 
kamptozoan fauna is unusually — varied, 
encompassing extremes of the body plan. The 
world’s largest kamptozoan,  Pedree//inapsix 


AREVIEW OF THE PUY E UM RAMPTOZOA (EN TORROC TAY 7 


frunivosa, dwells in these waters, as de sone of the 
world’s smallest kamptozoans, liny Lavesanela 
speeres Gn bryazoan hosts. Austrafian species may 
also bold the record lor the erealest density of zooids 
in colontes: Pedive//ing pyetfarmis packs inone giant 
foo alter another alone its peeuliar non-septate 
stolen, while tn Pedlce/napsis fititicosa, zooids 
spiral around a cigid central stem resulting fi a 
density of zoords anda growth pattern unknown in 
ather Kamplozouns.. 

Kuniplozeais in Australi are neither rare nor hard 
to find, The fauna of Australia is so poorly 
chaaclerized that new and unreported species (as 
well as those listed in the Appendix) probably can be 
vollucted. in only a few hours anywhere along the 
goust, Beyond taxonomic identity, we know vitor ly 
nothing about (he biology of Australian species. The 
litle we do know Jeads us to suspect that further 
invesnyatens hold much promise lor new msighes 
Ilo kamiplovoun eenjogy, symbiotic rekitionships, 
larval bielouy. biogeogruphy and phyloveny, 
Certainly, given the geoyraphieal dimensions aud 
ceologival diversity of dhs country. minmy new 
morpholoueal adaptations and life history. variations 
ave likely to be revealed when the Australi 


kamptoczoan fauna is more thoroughly examined. 
Acknowledgments 


Tam deeply grateful to Pe Arnold. b. &, M_ Barker 
P. Berents. D2 Gordon, Ko Lb. Gowlett-Holimes, S- 
O'Shea, S, Shepherd. and W. Zeidler for their 
hosoikilily and lor thei ussisianee in uceessing 
museum collections und carrying oul fieldwork in 
Austialia and New Zealund Environment Australia 
generously fupded jy Heldwork and, through the 
Australian Biological Resourees Study (ABRS). 
commissioned this review asa part af the Panne of 
dustvatia, Which series was closed prior to 
publication Of this chapter Eat indebted ta ALT. 
Newberry. ©, Nielsen, J, Pearse, amd 1. Potts lor 
their thoughttol comments that greatly improved the 
content, und to A. T.) Newberry m particular fir 
invaluable guiddnee with the Write. Pibhy, Hany 
(hanks to a team oF delighitlidly supportive editors: 
from ABRS. C. Clashy for initiating this projeet. and 
G, Ross and A, Wells lor lending to ils development: 
and, froin the Royal Society af South Austealia. J, 
Bird for blinvine ib lo trbeo, The leures ure 
published with the kind permission of ABRS. 


References 


ALLMAN Go 1 (ES80) “Monormph of the breshwaner 
Polvand” (day Secely. | ondin) 

Assil tos Re (i912) Lovosoma doxatine amd Laaveasaune 
veins - lavay New species: Guarn o/ icra. Sei Sa 
17-143 

Niaees. 11949) The ciligew feeding meelmnisay of the 
entoproch Palyvoa. ahd a comparison with that of the 
cetaproet Poly ani Mieh 75, 493-424 

Bay bovis, 1M S835) “Comparative Pmbryology?, Yel 7 
(Macmithin, | onulteny 

BArRO: J (1877) Memoire sur Menbryaligie des 
Bryovourres. Theves presacen ala Poonlte des Semweneus 
de dae AYO. 1-29. 

BeOK OR Oh (b9S8) Untersuchungen fiber den Darm tind aie 
Verda von Kanptozoen, Bryazoen dnd Phoraniden. 
/ Morph Okal Titre 3X, 72-127. 

Minis, Go & Pesan Mo (1953) Deus Tosxdsones 
Aouveaiy de Rescate aie, Lay, tren, Mt. 73- 
As 

Hetty. POL9S9) Chasse des Cidoprocies Ou ham plazeitires 
pp. Y2T-Tn da Grasse, Pd) WP de Zoalow ie 
Vol 5 (Masso, Pats). 

(1460) Le bourgeonnement eb la phy lwenese 
des Pndoproctes el des betoprocies: Reflexians-sur les 
promesaiis de Wb volition jiimale. Aah ch Ser deel 
Rey Bele (SU Ser.) 46. 748-76, 

Bist. G, (PRR) Report on the Palyzou collected ay 
HLM.S. Challenger during the vears [S73-1N7G, Part 
Hh. the Cyelostomata, Cleniostoidale. and Pedicel nes. 
Rep ved Res Vas MMS Challenger (fauks UP 1. 
ay 

Canning, My Ul & Cartnas. 1 (2000) Predatinn on 
amplozouns (Ee ptoproetay dover Bink WO ARG ART 

CLAARI DT. BO OS67) Miscelindes vodliengues Vo Sur le 
Hiveseum kefersiigne in sp. Rryovoaire mou do volte de 
Naples. tive Soe Nat Sth See. (4001) & 28-30, 


Chlagh A EL (1927) Anew chissieutian Olunimnis, flgte 
His, Oecanogn (Monin 4a, L-24, 

Cyn ©.) (1920) Karmptovcut pp. 1-64 fie Kiker. Wood 
Krumbach, | (kes) “Handbuch der Znologie™ Vol 2 OW. 
de Oruyier, Berlin. 

~ (1836) Rampteazoa pp. Th fi Pronk bhi, 
(Pd) tk bissen und Orinunen des Tierreiehs” Vol 4, 
M2. Book 4 (Akademische Vertuzsmeselfychult, 
Leipeig). 

Davinronel cB S99) On Donatella eeacilis all Mus 
Comp. acl, (Mutvatd Lin) 24, 1-44 

Dut. be b (180s) the fistory of the germ cells in 
Pedicetnramerteunea, dia NOY leet Sei Wy 1-64, 

Pibis.h C750) “Pssay towards a tatinil Histiry ab the 
conilines” (Millar Londen), 

EMSCIERMANWA. F Clh961) Ober Brutkarper ber dem 
kaimploroon Berciitvas wraciliy Sars. 4naf  dalirl 
(Physiol } 69. 395-438, 

(1Y6S) Dos Pidonephidiensystent var 
Grnatella sraciiy Leidy (Kawiplosonp fedb More, 
Okol Ticks 55, 559-9] 4 
all (19690) Lin Kreishiulorgan bei Kamptoyaen 
Zit, éelllursch YT, 576-607 
= (69h) Spralie qnecordhere Uijterembeiten 
moden A-Vikimenten der Kamptocoenoruskulitue. 
Lypericidia 28 975-76. 
— (1Y72) Derokedipiy saedadis Wen cb ap. mos. 

(Kumptoszog. Doxokulypoditdae liom nad, enn 

meuer  Rimmploxoentyp ues dem ndrdiichen 

Pavitisehen Oveun. Gin Versehtig 2ur Netlist 

der Kamptozoen systenmmatik, Afan Brad 12. 237- 

254. 

— (1982) Les hamptovogires, Ett setuel de nos 
coumiissanees aur leur dntitanme. dour développement 
Jeu bialowie et lour pasition plylopenetique. bul Aen 
Aol, by WT 317-444. 


18 kK. WASSON 


(J985) Factors inducing sexual miaturadon and 
Infuicncine the sex determination of Bareniie discrens 
Busk (Latoproct, Burenistidue) pp, 101-108 Jy Nielsen, 
Cound Larva dP (ids) *Bryozoa: Ordovician to 
Recent” (Olsen & Olsen, Fredenshoru, Denmark). 

(1987) Creeping propagation stolons ~ sn 
efieciive propagation system of (he freshwater entoproct 
Urnatella gracitis Leidy (Barcatsddae) eed, Hrurabiol. 
108, 434-448 

11993a) On Aniaene Mioprietas nematoeyst- 

Vike cirguns in ou loxosomulid. adaptive developmental 
strategies. hast specificity, dnd bipalur ocudrrence of 
species, Bfol Aull V4, (84-185. 

— (1993) Liime-twig Ghinds: a unique invention 
Aba Anturevie entsproet. Hid 185, 97-108, 

Pivnih, M. (1993) Lilbrasteuetire af the protonephetlise mn 
Lovosomella faivel, Bavenisia manushimane and 
Pedicellina cernua. Implivations [or the protonephridet 
nthe eeound pattern oF the Entoprocks (Ramptozei, 
Aienu, Mur ok, 7-38, 

Fianzes. A. (O83a) Bryozoy botopmmin pp. 61-509 fy 

Adivyod) KG. & Adivyodi RG. (hus) “Reproductive 
Biol oF hivertebrites” Volt (Wiley AC Sens, New 
Yorks, 
(YRS) ryozoa Entoproeie pp. 505-517 In 
Adiyad: Kar & Adiyed? Ri (hds) “Reproductive 
Biahiwy of Tavenehraes” Vol 2 (Wiley &e Satis, New 
York 

Cobrokh, PLT (1909) Conte bution to oul kaawledae af 
Australian Plirudines, Play rac. Linn Sac, NON We 34, 
12\-732. 

Goeknen P.O (1872) Biological relaiiinships ino an 
inweradial bryozoan pepulation il Var Mise 6. 502-314, 

— ® HAL VBIPPSL, WA CY) Cape Rodney to 

Okukart Point Muping Reserve: review of knowledge anid 
bibliguraphy to 1476. Tame (4 tnektun! Cniy tele 
Chih) ZZAsuppl.), 1 140, 

Hania. Soh OMRS) On the siietom une development of 
Lavenaiiece Quant 2 Mieroxe, Sed. 25. 261-337, 

(PRET) OW the lifeeiistory of Hodieediie, tid 
2}, 49-263, 

(1915) The Palyvon ol the Siboga Lixpeditiorn, 
Pi oT. Bitoproeta, Clenostomaie aid Cyelostonmta, 
Sthent, ayn. Rep, 2A, |TKO, 

Hasninos. AL (1932) The Polyvon, Ge Bern Reef Exp, 
PUIN LY OU Bet Rup, WV-ASS, 

Haswork, WoOA. IBY TL) Observatians-on (he Chlorgeniidie, 
with apecnil reference to eertaln Austcalian forms, Prec, 
Lian Sie NOS I 22, AL9356, 

Hasvmeinaky, Ch (19%6) The Mollusca coelonnite 
Hirbullaians ie inesenchympane unmtelids pp. [24° a 
Vaya D.C Origin and eye lirtonaey Racine yt 
the Maollusae” (Oxted Liniversity Press. bameny 
—_ SALWINEPAWEN, Dov Rab, RM. C1995) 
Larval plinktutrapby er primitive (rail i the bilateria’ 
lets Zocl, 76, \4- 154, 

Havrscues, H. €1877) donbryanilentwiek lunge und 
Krospune der Modk effin vellinitn 4ele Miss, Aval, 29, 
$(12-549_ 

UISSS) “Tehrbueh der Agnlowe (G2 Pischer 
Jena), 

HBLEY GC. (9b) AW cedlogienl sketeh at the Sydney 
Bepehes (Presidentialiddress)./ Brae Ava Seo NOS 
49, 15-77, 

Hinorspokn bo W, CbS88) On te Gecurrence of Peiowline 
i New Zealand, Cran, Deen Me dane ON, S137, 21 

Pivot, WooAL C828) A study of dhe movements of 
entaprochin beyoeouns, Cees. lates Mires, Sie, #2, 
145-141 

Hise. DOT NS 0) "A History Ol the Bribes manne Peshy ean” 

(Vin Voorst Landon). 


(1884) Contributians towards a generat! history 
of the marine Polyzoa. clin Mens Not Hixt (ser ay 13. 
456-364, 

Hyvian. db. EL (19S) ) "The Twertebrates” Yor 3 (Merri: 
Hill New York). 

JAGrRaT Es, OG. (1964) On the morphology and reprocie tan 
Of entproet larvae, Zool. Bie Upp, 36, 295-3 14, 

(1972) “Evolition ofthe Metazeai tite Cyele 
(Acitdeime Press, New York), 

Jebsnvwin. ©. (1982) Composition Chine compres des 
foymations squelertiques ches les lophapliariens er tes 
undoproctes. Mull See. Zaal fe 17, 229-249, 

Jonsvan, G. (187) 7A history af British zoophiles” 2nd 
vd, (Vin VYoorst, London). 

Jousstom, bbb & Wabki ke Mah Chol? A new species ol 
Peyhcellina Tron Sydiey Harbour Mine AL Sie. Ol 29. 
Of-O3. 


RAT, LM 948) Landoproetin Ba Not 
Anlare Res, Byp, LOMA UST Rep dS Be Zool a Bat} 
4, 2)5-231, 

Ki irasiii. Wo (1862) Uber Larosa sinuidare wen, eb 
sp... den Sehmaratver einer Anneli. Unterstiehunpen 
Uber niedere Seetiere. Zo7 Wisy. Zeal, 12, 131-142. 

RIRKEATRIOR, RO (TSU0a) Reports an the doolovicul 
collections made in ‘Torres Siriits by Professor AL 
Hliddon, PSS8-1884, (lydroida ind Polyzou, Sen Proc 
Rr Dahl See. €y 403-626, 

(890K) Polyzoa from Pom Phillip ae Aten 
Nit, Hist 12-21, 

Tin issiy, 1 (19S) Lie Taibryonalentwieklung der 
Pedicelling echinala Sars, Biol fent T5, 540-548 

Kitby dy (E831) On some American Polyaei, Prac deuedt 
Newt Sek Phi &, 3-922 

Mi Domathoa. Ro & Nydanien, 1 Wo CLO 78) Additional 
Notes on he food OF some California nudibranebs with a 
summminy OF lego Hood hibits OF Caliban iit sprectists 
boliwer 2A, TEUET TR, 

Mav Gintiviny, BoE USS 7) A catalogue of the miceiie 
Polyvoa al Vielory, Pree Aa Sine Moe 2a. 187-224 
Mackay, LY. WINN einecns, Boo Die Wariiin. Be 
BACKHLIAU, TL BMSCHERMANAL PA: GAREY JR (POY6y 
IRS RNA Sugeests thal Lotoprocks dre protostonies, 

vorelated to Ketoproctiuad Mele, Byel, 42, 562-559, 

MALARHONY. Y V(1990) Desenipbin of the development pl 
Jseapadamia divereta (Colonials, Burentisiidaey una 
didcussion af the Kamprodod stutus in the animal 
Kinwdon. [Pa Rilaaiiis Pnthiah Suni) Zoed Ali a 
20-30), 

Matisoan. BoM, (1965) The aetule and larval morpho} 
vid File History of he entoproct Aemernsie deeentis (M 
Sars. (835)... Morphol. 116, 411-338, 

(1975) Pitoproeta ppe 14d de Giese, AC, & 
Pearse. OLS. (ida) “Reproduerian oof Marine 
fivertebrites” Vol. 2 CAgaderivie Press. New York). 

Matettnn | (1939) Briozoarios mariihos brasileives. THe 
Bek Fac Pik Cie het (Uti. SO Poulop fond 3. ttt 
205, 

Mora Th Ae Manionkas To CIV 78) Stdies on ite 
rovenerulion OF an entoproel, Buywielon inert A fay, 
Anal. WS Ay \- 27h, 

—— (IYRO) Some observahwiny oa dle ses 
Jiflereniiion ofan eioprach. Rercutsie dente (PiaK) 
/hnd 2S, AS-59. 

NINLSEN, CE TMO4) Stodies on Danish fihaptocge, (yede 

1 LT, 
— (1966) On the lileeevele oof seine 
Loxosonuilidie (Lintoprowtad foie 4, 221. e 
(IY71) Enloproct Wieeveles und fhe 
enleprrel/ecloproct rlalionstip, fave, %, 209-341. 
(1989) “Prtoprieis’ Synopses of the Arist 
Foun, New Sertes, Val. doth al) Bet been. 


A REVIEW OF THE PHYLUM KAMPTOZOA (ENTOPROCTA) 19 


(1995) “Animal Evolution” (Oxtord University 
Press, Oxford), 

(1996) Three new species of Loxosoina 
(Fntoprocta) from Phuket, Thailand with a review of the 
genus. Zool. Ser, 25, 61-75. 

& JESPERSEN, A. 1997, Entoprocta pp, 13-43 Ji 
Harrison, E.W, (Ed.) “Microscopic Anatomy of 
Invertebrates” Vol. 13 (Wiley-Liss, Inc., New York). 

& ROSTGAARD, J. (1976) Structure and function 
of an entoproct tentacle with a discussion ol ciliary 
(ceding types. Ophelia 15, 115-40. 

Nitsciip, H. (1870) Beitrige zur Kenntniss der Bryozoen. 
Zeit. Wiss. Zool. 20, 1-36. 

PALLAS, P.S. (1774a) “Spicilegia Zoologica, quibus novae 
et imprimis obscurae animalium species iconibus, 
deseriptionibus atque commentariis ilustrantur™ Vol, | 
(G.A. Lange, Berlin). 

— _— (1774b) “Naturgeschichte 
hiere. Zoophyta” (G. A. Lange, Berlin). 

PrENANT, M. & Bobin, G. (1956) Bryozoaires. Pt I. 
Lintoproctes, Phylactolemes, Cténostomes. Faune Fr 60, 
1-398, 

Rear, J. F. (1969) Studies on the fine structure of muscle 
fibers and contained crystalloids in basal socket muscle 
of the Entoproct, Barentsia gracilis. J. Cell Sei. 4, 305- 
325, 

Reirziar, K. (1968) Loxosomella from Tedania ignis, the 
Caribbean fire sponge. Proc, U.S. Nat. Mus, 124, 1-11. 

RYLAND, J. S. (1965) Some New Zealand Pedicellinidae 
(Entoprocta), and a species new to Europe. Trans, Roy. 
Soc. N. Z. 6, 189-205. 

& Austin, A. P. (1960) Three species of 
Kamptozoa new to Britain, Proc, Zool, Soc, London 133, 
423-433. 

SALVINI-PLAWEN, L. Vv. (1980) Was ist eine Trochophora? 
Line Analyse der larventypen mariner Protostomicr. 
Zool. Jalirb. (Anat.) 103, 389-423. 

Sars, M. (1835) “Beskrivelser og lagttagelser over nogle 
maerkelige eller nye i Havet ved den Bergenske Kyst 
levende Dyr al Polypernes, Acalephernes, Radiaternes, 
Annelidernes og Molluskernes Classer” (T. Hallager, 
Bergen). 


merkwiirdiger 


SEELIGER, O. (1889) Die ungeschlechtliche Vermehrung 
der endoprokten Bryozoen. Zeit. Wiss. Zool. 49, 168- 
208. 

_ (1890) Bemerkungen zur Knospenentwicklung 


der Bryozoen, /hid. 50, 560-599, 


Topp, J. A. & Tayeor, P. D. (1992) The first fossil 
entoproet. Nafurwiss. 79, 311-314. 

Toriumi, M. (1951) Some entoprocts found Matsushima 
Bay. Sci, Rep. Tohoku Univ., 4th Ser. (Biol.) 19, 17-22. 
Van Beneprn, P. J. (1845) Recherches sur Panatomie, la 
physiologie et le développement des Bryozoaires qui 
habitent la Céte d’Ostende. Histoire naturelle du genre 
Pedicellina. Nouv, Mem. Acad. Roy. Sci. Bel. Let. Bruxy, 

19, 1-31. 

Wasson, K. (1995) The kamptozoan Pedicellina 
whiteleggii Johnston and Walker, 1917 and other 
pedicellinids in Australia and New Zealand. Rec. S. dust. 
Mus, 28, 131-141, 

bal, —— (1997) Sexual modes in the colonial 

kamptozoan genus Barentsia, Biol. Bull, 193, 163-170. 

_ (1998) Sexual reproduction in the colonial 
kamptozoan Barentsia hildegardae. Invert. Biol. 117, 
123-128. 

& SiePpHerp, S. A. (1997) Nodding heads 
(Phylum Kamptozoa or Entoprocta) pp. 975-992) In 
Shepherd, S. A. & Davies, M. (Eds) “Marine 
Invertebrates of Southern Australia” Pt 3. (South 
Australia Fauna and Flora Handbooks Committee, 
Adelaide). 

Waters, A. W. (1904) Bryozoa pp. 1-113 /a “Résultats du 
Voyage du S.Y, Belgica” (J. E. Buschmann, Anvers). 
WHITELEGGE, T. (1889) List of the marine and fresh-water 
invertebrate fauna of Port Jackson and the 
neighbourhood. J. Proc, Rov, Soc. N.S. W. 23, 163- 

323. 

WILLIAMS, J. B. (2000) Descriptions of two loxosomatids 
(Entoprocta), with emphasis on a relationship between 
symbiont attachment structures and host cuticle. Cancel, 
J, Zool. 78, 110-120 

Wooitacort, R. M. & EAKIN, R. M. (1973) Ultrastructure 
ol a potential photoreceptoral organ in the larva of an 
entoproct. £. U/trastruct. Res, 43, 412-425, 


Appendix 


Known kamptozoan diversity in waters around Australia and New Zealand. 


This appendix lists the 19 described and 18 undescribed species of kamptozoans known from Australia and New Zealand. 
The first column gives the species name. Undescribed species have been assigned a number. Those loxosomatids whose 
basal attachment (generic character) could not be determined are listed simply as “loxosomatid”, The second column gives 
the author of the original species description for described species, or a brief descriptive phrase (for loxosomatids, host is 
given) for undescribed species. The third column gives the citation for occurrence of this species in Australia or New 
Zealand. For new records (Wasson, this paper), the name of the collector is given in parentheses. The fourth column lists 
(abbreviated) the Australian State or the Island of New Zealand where the species was found. 


FAMILY LOXOSOMATIDAE (7 described + 17 undescribed species) 


(Harmer, 1915) 
(Harmer, 1915) 
(Harmer, 1915) 


Loxosomella breve 
Loxosomella cireulare 
Loxosomella cirriferum 


Williams, 2000 
(Claparede, 1867) 
(Harmer, 1915) 
(Harmer, 1915) 


Loxosomella diopatricola 
Loxosomella kefersteinii 
Loxosomella pusillum 
Loxosomella velatumn 


Loxosomella sp. | 
Loxesomella sp. 2 


on bryozoan 
dark zooids on sponge 


Loxosomella sp, 3 light zooids on sponge 


Hastings 1932 QLD 
Hastings 1932 QLD 
Hastings 1932; Wasson, this paper QLD 
(R. A. Birtles & P. Arnold) 

Williams 2000 VIC 
Wasson & Shepherd 1997 SA 
Hastings 1932 QLD 
Wasson, this paper 

(R.A. Birtles & P. Arnold) QLD 
Wasson & Shepherd 1997 SA 
Wasson & Shepherd 1997 SA 
Wasson, this paper SNZ 


(M, Barker & K. Wasson) 


30 


Loavxosamella sp. 
Loxosxomella sp. 


Ln ps 


Loxosomella sp. 6 
Loxesomella sp. 7 
Laxosomella sp. 8 


Loxesoma sp. | 


Loxosomea sp. 2 

Loxasoina sp. 3 

Loxosomulid sp- 
Loxosomatid sp. 
Loxosamatid sp. 
Loxosomatid sp, 
Loxosomatid sp. 
Loxosomatid sp, 


Ain lho— 


aD 


K. WASSON 


on polychaete Sthenelars 
on polynoid polychaete 


On Prawns 
on polychaete 


on polychaete Eunice 


on polychaete Copperingeria 


on polychaete Pectinaria 
on polychaete Axiothella 


on sipunculan Phascelosonte 
on hirudinean Brarnchellion 


on hirudinvan Pontohdella 
on bryozoan Amathia 

on squat lobster Thenuy 
on aquarium walls 


FAMILY PEDICELLINIDAE (6 described species) 


Pedicellina cernua 


Pedicelling compacta 
Pedicellina granilis 
Pedicellina pernae 
Pedicelline pyriformis 
Pedicellina whitelegan 


(Pallas, 1774) 


Harmer, 1915 
Ryland, 1965 
Ryland, 1965 
Ryland, 1965 
Johnston & Walker, 1917 


Hastings 1932 

Wasson, this paper 

(M_ Barker & K, Wasson) 
Wasson, this paper (R. Lester) 


Wasson, this paper (D. Gordon) 
Williams 2000 


Haswell 1891; Hastings 1932; 
Wasson, this paper 

(R.A, Birtles & P, Arnold) 
Wasson, this paper (J. Collins) 
Wasson, this paper (D, Gordon) 
Whitelegge 1889 

Goddard 1909 

Goddard 1909 

Harmer 1915 

Wasson, this paper (RR, Lester) 
Gordon & Ballantine 1977 


Kirkpatrick [S90b; 
Chitleborough!; Wasson 1995 
Hastings 1932; Wasson 1995 
Ryland 1965 

Ryland 1965 

Ryland 1965, Wasson L995 


VIC. SA 


OLD 
SNZ 
SNZ 
SNZ. TAS 


Wasson [1995 (and others cited therem) NSW, VIC, 


FAMILY BARENTSITIDAE (6 deseribed + | undescribed species) 


Barentsia benedeni 


Barentsia discreta 


Barentsia genioulata 


Barenitsia taxa 
Barentsia matsushimana 


Barentsia sp. | 


Pedicellinaps(s friicasa 


(Foettinger, 1887) 


[misidentified as B. gracitix| 


(Busk, 1886} 


Harmer, 1915 


Kirkpatrick, }890a 
Toriumi. 1951 


minute, delicale 7ooIds 


[misidentified as RB. gracilis] 


Hincks, L884 


Wasson & Shepherd 1997; 
Chittleborough! 

Wasson & Shepherd 1997 
Wasson, this paper 

(D. Gordon & S. O'Shea) 
Wasson, this paper 

(R.A, Birtles & P. Arnold) 
Kirkpatrick [890a 

Wasson, this paper 

(M. Barker & K. Wasson) 
Wasson & Shepherd 1997: 
Kirkpatrick |890b; Waters 1904; 
Hastings 1932: 

Hilgendorf 1898; Gordon 1972 
Hincks 1884; Busk 1886; 


SA, NNZ. SNZ 


QLD 


NT 
SNZ 


NSW. SNZ. SA 
VIC, QLD 


SNZ. NNZ 


MaeGillivray 1887; Whitelegge 1889; VIC, NSW, 
Hedley 1915; Johnston & Angel 1940; ‘TAS 


Wasson & Shepherd 1997 


| CHM LEBOROLGH, R. G. (1952) Marine Fouling at Port Adelaide. MSe Thesis, The University of Adelaide (unpub.). 


AMINO ACID RACEMISATION DATING OF A RAISED GRAVEL 
BEACH DEPOSIT, SELLICKS BEACH, SOUTH AUSTRALIA 


By C. V. MURRAY-WALLACE* & R. P. BOURMANT 


Summary 


Murray-Wallace, C. V. & Bourman, R. P. (2002). Amino acid racemisation dating of 
araised gravel beach deposit, Sellicks Beach, South Australia. Trans. R. Soc. S. Aust. 
126(1), 21-28, 31 May, 2002. 

The extent of racemisation (total acid hydrolysate) of the amino acids aspartic acid, 
glutamic acid, leucine, phenylalanine and valine indicates a minimum age of last 
interglacial for fossil molluscs occuring within a raised gravel beach deposit at 
Sellicks Beach, South Australia. The base of the raised gravel beach occurs up to 5.5 
m above Australian Height Datum (AHD) and possibly indicates 3 m of local uplift 
since the last interglacial maximum (c. 125 ka; Oxygen Isotope Substage 5e). 
Emergence of the gravel beach is attributed to ongoing neotectonic uplift of Fleurieu 
Peninsula. 

Key Words: amino acid racemisation, last interglacial, neotectonics, sea-levels, South 
Australia. 


Transactions of the Royal Society of S. Aust. (2002), 126(1), 21-28. 


AMINO ACID RACEMISATION DATING OF A RAISED GRAVEL BEACH DEPOSIT, 
SELLICKS BEACH, SOUTH AUSTRALIA 


by C.V. MURRAY-WALLACE® & R.P. BOURMANT 


Summary 


MurrAy-WALLACE, C.V. & BOURMAN, R.P. (2002). Amino acid racemisation dating of a raised gravel beach 
deposit, Sellicks Beach, South Australia. Trans. R. Soc. S. Aust. 126 (1), 21-28, 31 May, 2002. 

The extent of racemisation (total acid hydrolysate) of the amino acids aspartic acid, glutamic acid, leucine, 
phenylalanine and valine indicates a minimum age of last interglacial for fossil molluscs occurring within a 
raised gravel beach deposit at Sellicks Beach, South Australia, The base of the raised gravel beach occurs up to 
5,5 m above Australian Height Datum (AHD) and possibly indicates 3 m of local uplift since the last interglacial 
maximum (c. 125 ka; Oxygen Isotope Substage 5e). Emergence of the gravel beach is attributed to ongoing 


neotectonic uplift of Fleurieu Peninsula. 


Kry Worbs: antino acid racemisation, Jast interglacial, neotectonics, sea-levels, South Australia. 


Introduction 


A resurgence of interest in recent years in 
Quaternary emergent shoreline successions has 
arisen from the increasing ability to determine the 
age of these features due to technological advances 
in geochronology (Rutter & Catto 1995; Noller e/ al. 
2000). Similarly, an increasing awareness that 
coastal successions, particularly those deposited 
during the last interglaciation (c.125 ka), are 
sufficiently old to quantify even modest rates of 
neotectonism, has bolstered this research endeavour. 
Accordingly, the elevation of last interglacial coastal 
deposits has been widely used as a benchmark to 
delineate recent tectonic behaviour at continental 
scales (Murray-Wallace & Belperio 1991; Ota 1994; 
Bourman ef al. 1999; Zazo et al. 1999), In this work, 
the age of a raised beach deposit at southern Sellicks 
Beach, South Australia, is determined based on the 
extent of racemisation of several amino acids within 
molluscs from the fossil assemblage. In addition, the 
neolectonic significance of this deposit and_ its 
relation to other emergent shoreline deposits on 
Fleuricu Peninsula is examined. 


Materials and Methods 


Field investigations 

The elevation and lateral extent of the gravel beach 
deposit was surveyed to Australian Height Datum 
(AHD) usimg an automatic level. In addition to a 
general field description of the deposit, shell samples 


' School of Geosciences, University of Wollongong, NSW 2522. 
" School of Environment and Recreation Management, University 
of South Australia, Mawson Lakes, SA 5095. 


were collected for amino acid racemisation dating and 
to document the fossil mollusc assemblage. Species 
identification followed that set out in Ludbrook (1984). 


Amino acid racemisation analyses 

Samples of fossil molluscs for amino acid 
racemisation analyses (total acid hydrolysate) were 
collected from the gravel beach deposit. Shells were 
removed from the matrix of the deposit and their depth 
of burial recorded. Analyses were undertaken on 
specimens of Patella (Scutellastra) laticostata 
Blainville, Thais orbita (Gmelin), Svdaphera undulata 
(Sowerby), Nerita (Melanerita) atramentosa Reeve 
and Ostrea sp. Linnaeus. 

Sediment adhering to the surfaces of shell samples 
and diagenetically moditied aragonite, particularly 
chalky surfaces, were removed with a dental drill, 
followed by successive washes in distilled water using 
an ultrasonic bath. A dilute acid etch (2 mol HCl) was 
subsequently undertaken to remove the outer surfaces 
(c. 10-15% by mass) of the shells that had been in 
contact with the host sediment. Samples were 
subsequently hydrolysed for 22 hours at 110° C in 8 
mol HCI. Following cation exchange isolation of the 
amino acid residues, samples were freeze dried and 
derivatized. Chromatography of the N-penta- 
fluropropionyl D, L-amino acid 2-propyl esters was 
performed using a Hewlett-Packard 5890A Series II gas 
chromatograph with a flame ionisation detector and a 
25 m coiled, fused silica capillary column coated with 
the stationary phase Chirasil-L-Val. Full details of the 
analytical techniques followed in this work are reported 
elsewhere (Murray-Wallace 1993). Enantiomeric ratios 
were determined for the amino acids aspartic acid 
(ASP), glutamic acid (GLU), leucine (LEU), 
phenylalanine (PHE) and valine (VAL). 


22 C.Y, MURRAY-WALLACE & R. P. BOURMAN 


% io" RINE! 
Wee 


* River ill 
<i. Dairies 
> mf | st 

r 2 / 

- o / i . 7 | 
iy : eet Mt, | 
\ C . 

‘ v 54g Pivasant / 
‘\ para! River’ ; Mt re | 
) « 542 Gawler ef / 
7 Po ¥ 
f Op , _ / 
a / 
| rors wi A R io 2 f 
SOUTH qorrer : na @ 
al : ( 
AUSTRALIA Sf _ ADELAIDE ’ e 
| _ _ Yo a it, j r 
bem .) B y 
\ Af, _ a 5 = ¥ a 
ty ce | comk vt Noarlunga rae H / < 
Sy | Cove ; / Embayment on™ ; ny J 
F aM ing” ¥ ny 
ADELAIDE a? yore ! 4 / 
/ font iP /? / a 
hes / | sn Z \ f= 
2 ae mye / F ng x } e 
S rod i= ; 7 g 
s | Witunga /o ‘an = 
{| Embayment / 4 aR, 
2 fF im & 
18 4 7 
Mt, oy & .sa 
Magnificent / ee ; : > 
J, ap LF Y =>. 
" Finniss Bi “ i \ y 
Wer Y 


Beach jp han ae 
Mvp At +, f _—— 


/ K& 
“, ey, 
[earickainga R. ; % Currency f = S \ Lake Alexandrina 


nga p 
Fan ce a i 
ita 7 
Weymouth 
wi a cl 
ox Fleurieu is tj 
’ Cape : } 
- Jervis 


<. a Waitpinga 
“-Tunkalilia Beach 


Beach 


Scale in kilometres 


Pig. 1 Location of the raised gravel beach deposit, Sellicks Beach, South Australia. 


AMINO ACID DATING OF A BRAISED GRAVEL BEACH DEPOSTT 


rok, 


hiv 2. View lookin: south wong southern Selleks Beach 
tewards the southern Adchide Hills und coastal cliffs 
developed on Pleistocene allivial fin sieeessions, ‘The 
Tocution ol the niscd heael dupasit, whieh cers i the 
scarp foot zone rs indicated hy win arrow. 


Fig, 40,4 shore-nonmnal siew of the puised gravel beach 
deposit at Sellieks Reuch, Phe steeply aippmp Ochre 
Cave Vormatiuners visible nour the survey stall. The stall 
wineh is fully extended. is 5 mm long, 


Geomorphological Setting And 
Site Description 


The raised gravel beach deposit is srluated near (he 
Willunga Faull ul the southern-must part of Sellicks 
Beach (35° 215 09.8" S; 138° 261 07,5" EF), landward 
ofa moder, gently seaward sloping interlidal shore 
phitlorm (Pigs 1,2). The modern intertidal platform 
is approximately 20 mm wide in a shore-nornal 
Irtnsect, 1S partially covered with boulders and 
cobbles and representy a modern anglogue for the 
reliecL platform (Mig. 3). An accumulation of boulders 
and cobbles occurs at the foot of the modern clilFand 
represents a further modern analogue of the raised 
boueh deposit. The emersent eravel beach lavies 
rests On Uo Strongly eroded remnant of a shore 
plattodm that is developed on the steeply dipping 
Olwo-eMioeene Port Willunga Beds (Daily ef af 


tw 
“ 


Pre 3. View looking seuthwest towards the rised gravel 
heach depesi at Sellicks Beaeh, The gravels pest 
naconformably on the Olize-Miocene Port. Willutys 
Beds ane the Middle Pleistocene Qehre Cove bornnitian, 


The gravel deposit dips gently seawards. The 
umconformily surkice represents a rele interridul shore 
platform. The modern titertidal patton meeurs i thy 
foreground. dips gently seaward and is partially eovered 
by boulders and cobbles, The maxiniin difference i 
clevation between the (wo phitlorars 5 tras deteriniiicd 
in the most landward exposare, nat yisihle tn thos 
photograph, Small, isolated. sey stieks representing 
erosional renimints of the Jormerly mire extensive 
Pleistacene shore plidorm occur within Unis area (eg. 
“aT in the Middle distinee), 


1976), and in part, & steeply dipping portion of the 
Middle Pleistocene Ochre Cove fetmation (Ward 
1960; Pillans & Bourman 1996; Fries 3. 4, 5). The 
gravel deposit occurs within a former searp foot zone 
excavated ata time of higher sea level, and abuts 
fanglomenites of the Oclire Cove Formation (May & 
Bourman |O84). 

The bedrock surface on whieh) (he gravel beach 
facies resis, grades ina seaward direetion from 3.55 
into 4,95 in above Australian Height Datum (AHD), 
The platform extends oul seaward fron) the deposit 
sume I-15 in forming a well-defined bench (Fig, 6), 
The vravel beach lieies erops out over a shore- 
parallel distance of approximately 50 m, and ranges 
in thickness between | and 1.5m (Fig. 6). 

The grovel deposit is poorly sorted and conprises 
Subsroulided to subangubi clasts of silistone, 
quartzile and bryozoal limestone that range from 
boulder lo pebble size, aldiough the modal chist size 
is boulder-cobble (700-70 mm), The lithoehists are 
tightly packed. Numerous entire and fragmental 
fossil molluses oceur within the granular matrix of 
the gravel deposit. 

A pale grey, elean, free-Mowing sand ts: thinly 
draped over the gravel deposit and the underlying 
funwlomerates and extends up to 2.5 np above the 
upper bounding surface oF the deposit. The sand alse 


4 COV MURRAY WALELACI & RB. P BOL RMAN 


Fig. S. Detail trom figure 4 showing the hehtly packed 
urrangement vat lithoelasts. 


Fis. & View Tovking eustnortheast showing par of the 
shore-parallel lateral estent of the raised beach deposit. 
The letler “Wo denotes the general level of the pravel 
deposit which is approximately 4 abave phe wrivel 
covered tbotsinpe ofthe small chit i the foreground. A 
planared surlice representing rennants of an intertidal 
shore plallorm as visible on the seaward side of the 
deposit. Pletsiocene fanelomerates are evident in the 
upper riuht-hand prtion of the phetograph “bh, The 
naised beach deposi is averlin by adhin veneer al sand 
Which also partially covers the fanglomerule, bul is 
(iMcult to discerns fa this photograph. 


oceurs WiTN the Upperinost part of the matrrs of the 
gravel bed neur the contact between the gravel and 
the overlying sand. A thermoluminescence ave of 
34,0229 ka (W2317) Was previously reported for 
this sandy uni} (Bourtnai ef od. 1999), hr additon, a 
radiocarbon ave (minimum age) of -30 ka (Gak- 
6095) has previously been reporied for molluses 
trom the gravel beach deposit (May & Bourman 
1YR4). 


Results and Discussion 


Mallise axyenthlege 
The gravel until contains w relatively diverse 


assemblage ool fossil molluscs. principally 
pustropods, within the sediment matrix, Molluscs 
include Patella (Scutellasira) laticostate Blainv lle. 
Mactra rufescens (Lamarek), Ostredt neers 
Sowerby, Monadonta (duyirecuchlea) constrieni 
Lamarck, Nerina (Melanerite) atrainentasa Reeve, 
Cunariela fesuewm? Iredale, Conny sp. Linnieus, 
Dilume (Chlavedilama) adelaidea (Philippi. 
Bemluchim welimestme (Gmelin). Svdephera 
undulata (Sowerby) and opercula of Yiwho sp, 
Linnacus. Many of the shells also oecuh as hinge 
frawnents, filshly abraded and ol unrecognizable 
allinily. Collectively, the fossil assemblage indicates 
deposition in an environment comparable to the 
modern Goust ab Souther Sellieks Beach, with 
molluses. found in sand or attached ty poeks, in i 
felatively sheltered setting oF the lower Hiloral zone 
(Ludhrooak TOX4). 


Dati 

A owenerally high degree of  rucemtsation 
(expressed us a D/L ratio) is evident for the five 
dilferent cnantiameric amino acids measured i 
cach af the fossil molluses from the gravel heach) 
deposit (lable 1), Phe relative extent of racemisalion 
lor the dillerent amino acids. within the single 
mmotluse sumples, generally follows the relation 
VAL@LEUSGLUSPHILSASP. Similar trends are 
reported for fossil molluses from United States 
Pacific coastal plain sites (Lajoie ef a, 1980), 

Three specimens of Patella  (Seutellestra) 
faieastate (samples UWGA-695. 696 and 763) 
reveal Goud concordance in measured enantiomeric 
ratios (Le. betweensshell D/L. ratio variation) with 
coeticrents of variation less than $6" forall amino 
avids for the combined data (actual values include 
VAL. 2.2. LU 5.6, ASP 0.3%, PITE 10% and 
GLU St), The consistently lower degree of 
racemisalion forall amino acids in the specimen of 
Patella sp. (sample UWGA-697), compared with the 
outer three Priel stmples is possibly due to the 
diffusive Joss of the more highly racemised. lower 
molecular weight peptide Traction trom the shell 
curbonate matrix. Accordingly, the degree of 
mcemisation as determined in the total acid 
hydrolysate, Would be disproportionately werhled 
towards the less rucemised, hivher molecular weraht 
peplide residues that remain awithin’ the shell 
aragonite matrix. This explanation is consistent with 
the poorly preserved nature of some ol the molluscs 
withit the gravel deposit (c.g. chalky appearance). 

The high extent of racemisation measured. in all the 
fossil moltluses: from the raised beach deposit far 
exceeds values typically determined in’ Holocene 
fossils (Murrity-Wallace & Bourman 1990; Murray- 
Wallace & Goede 1995; Murray-Wallace 2000; 
‘Table }). The extent of racemisation inthe molluses 


AMINO ACID DATING OF A RAISED GRAVEL BEACH DEPOSIT 


iw 
Ww 


TABLE 1, Extent! ef amine ack racemisation (otal acid fryvdralysate) in fossil molluses fron a raised gravel beach 


deposit. Sellicks Beach yd other localities for comparisan 


Lab. Code 
or reference 


Species & Location 


VAL 


Sellicks Beach, 

raised heach deposit 
Thais arbita (columella) 
Svdephera ancdulaty 


UWGA-733 
UWGA-736 


O.284 00121 


0.30920,008 


Patella (Sentelasira) UWGA-697 
laliou ster 
Patella (Seutellasiru) UWGA-0U6 0412 


featiceys dialer 

Patella (Seatetlasiva) 
fetion stata 

Patella (Neuellastead 
faticustata 

Neriter (Melamenitiay 
ulramentosa 

Osirea sp. 


UWGA-605 O.40510,.003 


UWGA-763 (0.42 3-40,008 


UWGA-706 O.38640,008 


UWGA-T68 O.36540010 

Lute Pleistovene, 

Glanville Formation, 

Normanville, SA 

Mectra australis Bourman ef af O.28310,011 
(1990) 

Uindmarsh Island, SA 

Mercerra ausialis 

ort Wakefield, SA 

Anackive Wapesia 


0.26 40.0003 
Murray-Wallace — 0.320.06 
et eal 
Katelvsia Hivuiphore (1988) (1.3210,04 
Holocene 
Three Rivers Creek, 
King Island, TAS 


Pateller laticostiata Murray-Wallace 0.01 
(T90L6U yr IP, & Ciovde 

SUA-2927) (1995) 

Sir Richard Peninsula, SA 

Danay delinices Murray-Wallace (070.01 


(22604140 yr BP; 
SUA-2881) 


& Rourman 
(1990) 


Amino acid D/L ratio! 


LEU ASP PIE GLU 
Q.369 0,55640,015 S 3 
0.333 ),322+40,036 


OS7TOMOOLR 0,61310,033 0541-0006 0.540 O.057 


OSS200.021  O.800+-0.004 0, 79820,029 — O,606) 0.008 
O55) 0.007 079910007  O777TEOOLF — 0,55720,002 
0,52020,008 O804L0.001  0.77000,009 — (.6b1+0,009 


OSFTILON0G  O70240,029 O.59920,005 L671 0,006 


OANSHOMNS O83S#0.023  0.72720,008 0, 78940026 


QI975=S0.012  O.59000.010 - 0333 <0.006 


Q.3720.002 — 0,5640,001 O3600,002 


0,51+0.02 Q.5440,038 4820.0) 


0.31007 (4640.02 - OSS+0.04 


(054002 = 0.042 0,00) O.05=0,.00] 


- 0.2710,01 OTOHLOF O12 60,005 


"amino ueids: VAL - valime: I.BU - leucine; ASP - aspartic acid, PIE - phenylalanine und GLU) - glutamie avid. 


lrom the Sellicks Beach deposit also exceeds that 
apparent for representative examples from the Late 
Pleistocene Glanville Formation at Normanville and 
Hindmarsh Ishind. two localities with comparable 
current mean annual air temperatures and, as a 
eorollary, two deposits likely to have experienced 
similar diagenetic temperature histories to the 
Sellicks Beach deposit. given the caveat that the 
shells [rom cach deposit remained buried at depths 
>! m for much of their diagenetic histories (Murray- 
Wallace ef af. 1988; Bourman ef al. 1999: Table 1). 
The Glanville Formation, as originally defined in the 


Adelaide region (Ludbrook 1976; Cann 1978) has 
been correlated with the last interglacial maximum 
(125 ka: Oxygen Isotope Substage Se) based on 
thermoluminescence, amino acid racemisation and 
uranium-series dating of correlative deposits from 
other parts of the South Australian coastline 
(Belperio ef a/. 1984; Schwebel 1984; Huntley ef al, 
1993, 1994; Murray-Wallace 2000), 

Although the fossil molluses trom the raised beach 
deposit at Sellicks Beach were obtained from near- 
surface contexts (<50 em), the geomorphological 
and stratigraphical evidence sugyest that for part of 


ie (CV, MURRAY WALLACE & R.P BOURMAN 


hen dtiwenehe history. the fossils were more deeply 
buried (he. at lease |), However, these mollases 
Will have experienced a higher integrated diagenetie 
iemperature than for fossils that have remaived (i 
more deeply bured contests (Table |), Current mean 


annualair femperrtures (CMAT) at Sellieks Beach, 


Normunville and Mindmarsh fsland) are all 
approxinmitely 16° Cy and 17°C for Port Wakefield, 


The extent oF racemsation for the tajority of 


amino aids ts significantly higher in the molluscs 
from Sellicks Beaeh eompared with those fron 
Normanville and Vindmarsh Ishind (Table Up. The 
difference in extent of racemisation is less 
pronounced when compured with the molluses from 
Port Wakefield which have experienced a higher 
(ingenetic temperature (Viable 1). AS carrent men 
annual temperature al the Port Waketicld site fs 
approximately 1° C warner than at Sellicks Beach. 
ind wiven that rates of Maeeniisation are known tu 
iNerease by up to 20 per cent for such a temperature 
difference (MuCoy 1987), the implication is that the 
shallow burial depth of the shells al Scllicks Beach 


has contributed to the high degree of racemisation of 


aiting acids within these lossils, 
Amino acid D/T. ratios for the molluses from the 


Sellicks Beach deposit range from the enyclope of 


values representative of dast interglacial age to 
potontially The penollimate inteyghacial (e. 2200 leas 
Oxygen lsotope Stave 7), us revealed ia plot ofthe 
CXLONL OF PACCMMISAtiON ABAINSE CURFeNE Meal annual 
temperature (and as a corollary, latitwde) (Fig, 7), 
The lack of clustering and ehronologival consistency 
ol the data suggests a diagenetic basis ler the 
observed varuition in enantiomeric ratios rather than 
a Benge age variation between shells, The range in 
D/L ratios for (he shells from Sellicks Beach exeveds 
that typically found fora single matupie staye 
|Murray-Wallace 2000), 

Although racenisatiou rues are kigwit lo be 
genuds-specHic (Miller & BrivhameGretle 1989) (his 


jo UU Kely toaceount solely for the bicher degree of 


favemisauon fh the fossil Patella (Seutellastra) 
faticustuter Trom the Sellieks Beach deposit, 
compared with other genera trom the Glanville 
Forman, fees therefore coneluded that ihe hihes 
deerce of raeeimisation i the mollises front Sellicks 
Beach 16 due to faster rales of ragemisation, die ta 
then shallow barial depth davies bute ditenesis 
resulting: from de proeressive exhumation of the 
deposit, und a genus-elleel on racemisation, 

As the shells have been subjected lu variible burial 
depths during lute chagenesis, the integrated tate 
expression for ravemisation was rearranged with 
femperatire us the subjeet to oassess whether it is 
Possible to [duce the Pigh extent ah paceiisalion at 


umbient diagenclic lemperulures over the course ol 


the Holocene. As the amino acid analyses reported 


1a ® Katelysia 5p) 
© Anadara tapezia 
4 Patella (Scutellastr) lalicestata 
A Nenta atiatnentasa 

of () Thais rrbita 


@ Osea sp 
w Stage * a 
= 8 Sellicks Beach ee 
= data * — a 
= ° . 
= oa Ch wr 
oa 
i Suib-arnge Sn 
ea tee anvelape 
u2 BibOiady) OP 
(SUA-3102} 


ee 


W213 '4 416 We 17 18 19 FO Bt oF 23 


C.M.A.T.(C} 


Pik 7 The extent of wiline paeemisirion (romil quid 
hydrobysatey in feassil mrathises ai lasy pnterchigia) wee 
(Osyzen tsurupe Substage Jey (ream southerts Aust 
plored (Aust eHren eH HONueE aie emperaiece (oe) 
1 iistrate the Sellicks Beach dam within a brouder, 
rewiinal wonterr, Detuils of samples trom clewhere i 
southern Atstralia ane repericd hy Murriy-Wallice & 
Belperio (1991) ured Muteriy-Wallice ed af (128%) The 
mnie aeid data Car the last Tterghica! mollises gre i 
aeeord Wilh the exponential trend OF ierenaing extent ab 
MeO SOLON With Hivher diawenetic temperatures. und is 
a corolliry, higher enerent mean viug) lemperutares 
The fowsit inallises (rai the raised beach deposit al 
Nellicks Reach reveal a heed cinge on extent ol 
hiwemisation fram the cnyelapeal the haat itengkienidh 4a 
ilues consistent wil u penultimate teterghicn) ape 
(Msyen lbotope Suse 7 oe. 2b kay. Simin acid results 
lr Halogene and Stage 7 inalluses are preswimted us 4 
Hrumewark Tor comparsan, 


here were undertaken on diferent fossils from (hose 
used tor the radiocarbon assay (30 hea age; May & 
Boorman  J9K4). the integrated — diageneue 
lemiperdure owas. ealeulted to examine the 
possibility that the radiocarbon ave was the resullol 
vhinee sampling, of reworked Pleistoeene shells 
within a Holocene deposit. A tinimum age of 7000 
years Was selected for the ealeulation, ropresentidg 
(he Citi oF the culaination of the post-ulveral 
marine Wansgression in southern Australia (Belpene 
ef ol 2002). anc. therefore. the oldest ave likely for 
an uniisturbed FHloloeene coastal deposit. The 
raibonale for this is thal the early Holocene iy the 
only time in the Late Quaternary, apart from the last 
interglacial maximum, that sea level was sufficiently 
high potentially to fori the raised beach, Present sew 
level is not sufficiently high to form the deposit, 
Furthermore, interstadial sea levels of the Late 
Meistwueene (Chappell ere’. 96) were significantly 
below present sea level and would imply rates of 
tectonic Uplift of a magnitude inconsistent with the 


AMISO ACTD DATING OF A RAISED GRAVEL BEACTT DE POSTI aT 


well-established (celonic: Trimework lor the cegion 
(Bouman erat 1999: Belperio ef af. 2002). 

An average diagenetic temperature required to 
iNduee the degree of racemisation measured i the 
Jossils assuming wo age of 7 ka was determined thus: 


, = 5039 
(14+D/L) (1) D/L) 

IS 77-loe In - 1h 
(1-DAL Sf (1-D/L ye 


a 
Where [is the absolute temperature (“K), D/L, and 
L/L,, are the enantiomeric ratios of the fossils and 
ther modern equivalents respectively, 7 ts an 
assumed age Ge, 7000 yeurs)and 15,77 and 5939 are 
constints derived from the empirical rate comsuint 
expression (Wehmiller f982, 1993). Accordingly, an 
averuge diapenetic lemperature of 24° © would he 
Necessary to jnduee the extent of ravemisation 
measured in the three specimens of Pareles 
(Neufellanirad laticostiitla (UWOA-695, -696 und 
-763) from the raised beach deposit i they were only 
7T oka, A diagenetic temperature of this value is 
unlikely, hawever, given that the current mean 
uu! air leniperature at Sellieks Beach is 
approximately 167 C, A prolonged. higher mean 
winmual temperature by as much as 8? Cis unlikely 
over the course ob the Holoeene for this region 
(Chappell 99h) ‘Thus, the exten of racemisation 
incasarcd in the fossil fmolluses fram the raised 
heawh deposit could nol have been ullained during 
the Holocene. A penultinine interglacial ape ys also 
not favored. us the gravel beach deposit is utlikely 
lo have survived erosional processes of the last Wwo 
glavial cycles. Orr Uris basis a dust interglacial age is 
favoured for the raised gravel beach deposit ul 
Scllieks Beagh- 


Neolectonivs 

Vie raised pravel beach deposit at Sellicks Beach 
provides a farther oppertunily to examine the 
neolectonic behaviour of Flearicu Penimouli, 
Mrevious investizations have revealed that the region 
his exporioneed “culogivally recent uplifl as 
indicated hy the elevation ol last inferghiead coastal 
deposits (Bournaner af. 1999). 

Although many vravel bevel) deposits represent 
relational sea-level indicators (Le. ahways form 
above tidal datunry) and are therefore of only modest 
rehubility (Chappell 1987), several attributes of the 
deposit wt Sellicks Beach render it more celiable Jor 
quantifying rates of neoteetonism. The adjacent 


modern intertidal plitlorm has clearly formed within 
a narrow range OF tidal datum and represents un 
analogous feature to the Pleistocene eyiivalent. The 
tipper reavhes of the modern shore platforiy are 
covered by boulders and cobbles presumably 
accumulated during storm events. However the 
steep backilig slope of the cli? prevents houlders cr 
finer clasts from being deposited al uny sughifioantly 
hivher elevation above tidal datum. 

Pstimates of o plucio-eustatic sea level Por the hast 
iterglachl (Oxygen batope Substage Se) from Eyre 
Peninsula suggest a value of 2 m2 ATID, and 
represents a particularly reliable datum given the 
relative tectonic stubility of the Gawler Craton upon 
which much of the Eyre Peninsuly coastline has 
developed (Murray-Walhice & Belperio 199 )). Thus, 
uplift of the Sellivks Beal deposit by as much as 3 
mis indicated based on the elevation of the contac! 
heiween the gravel deposit and the underlying 
erosional surfiice of the relict shore plattorm, 

The amount of uplifl since the last intergtieil 
maximum, tiferred front the deposit al Sellicks 
Beach (© 3 mm) is less than that observed ut 
Normanville (c. 10m) to the south of the Willunga 
Paul (Bourman ef af. 1999), The uplilt is attributed 
lo the conbined elfeets oF ongoing eelonie uplift ot 
He Adelaide Wills und crostonal unloading ane 
assockited ertistal isostatic compensation, Further 
research is required to jnodel these processes 
geophysically, 


Conclusions 


The extent of racemisation for several anime acids 
in fossil molluses from a raised gravel beach deposit 
al Scllicks Beach, South Australia, indicates thar the 
deposit is oF Late Pleistocene age. and most likely 
formed during, the last interglacial maximum (e125 
kas Oxyeen Isotope Subsiave Se) The deposit 
indicates up ta 3 om of uplilt has qecurred in lis 
region since the lost interglacial and suggests that the 
region is stil Undergoing, Heotectonie Uplitt. 


Acknowledgments 


‘This research was supporled by the Research 
Centre for Landscape Change al We University ol 
Wollongong, (. Sloss und N. Riggs are thanked for 
luboratory assistance. The digures were prepared by 


R. Miller and © Crothers. This puper is a 
contribution to [GCP Prajeet 447, “Coastal 
Environmental Change Dorin Sea-Level 


Highstands: A Cilohal Synthesis with Tinplicutions 
for Managementot buture Coastel Change’ 


28 CLV, MURRAY WALLACE & Ro BOURMAN 


References 


Briain ALP. UaArvew, No & Bourwan, Re PL (2002) 
Spatial und tainporal variability in the Holoeene palace 
sed-level recerd around the South Australian coustline, 
Sec Geel, (in press), 

— Swbiy Be W. Poracik He A. Nrpraurias. 
CoA. Dr Masire, Do . Prescot, fR PIAn sd. BR. & 
Gustin, Vo AL (L984) Chronological studies of the 
Quaternary marine sediments of nocthern Spencer Gull! 
South Australi. Mens Quel, OL, 2605-296, 

Bovraian, RoE BE Lekio, A.B Mukiay-Wartacr, CoV. 
K CANS, HL (1999) A ast faterglicial embayorert fill 
al Normanville, South Australia, and its neatectonic 
Mplicutons, Trans A Soe Stasi, 123, 1-15, 

Cann, JU (TOFS) An exposed velerence seelion for the 
Clanville Formation, Quarterly Crevlogloul Nores 
Crealiuleat Surver of South Australia @S, 2-4. 

CHAPPELL. L (1987) Late Quaternary sea-level changes in 

the Australian region pp. 246-3451 dn lonley, Mod & 

Shenmun. 1 (Puls) “SegeLevel Chunpes™ (Oxford, Basil 

Blackwell). 

(199 )) Late Quaternary environmental changes 

() vaste wad eenteal Australia, and their elinatic 

Interpretation, Guat Sev Rew (0, 377-390. 

— , OMURAL AY BSat, bh, MeCh irom, M,, 
Pasooih hy, Ora, Yo& Pirnass, Be Cheya) 
Reconciliation of fate Quarernary sea levels denived fom 
coral terraces at Huon Peomsula with deep sen oxygen 
isatape records. Avid Manet Sei Lew, (4b. 227-236. 

Daity, B. Finwan, 0B. Poms, BO ac bINpsas. dM 
(1976) Gealany py 5-42 de Pwidale CL Ro, Vvler, MI 
& Webb, BP (Rus.) “Natural Plistory oF the Adelaide 
Region” (Royul Society of South Australia, Adelaide), 

Horney Dey Phe rros, hk Ty ad PReseari dR. (1993) 
he stranded beaeh-dune sequenee ol south-vast South 
Australia: atest ofthermoluminescence dating, 0-R00 ka, 
Gat Sei Rev 12, b20. 

’ & (1994) Further 
thermoluminescence dates Crop ihe dune sequenee inthe 
souihivast af South) Australia. Joie. 13, 201-207, 

Dior KR. Werivitiri 11 Se Keay, Gil. (Loma) 
Inter- and intra-wenerié trends [i apparent faceniization 
Kinetics oF ating acide in Qualermiry mollusks pp, 305- 
340 J Mare, PE. Uvering, fe ©) & King, KR. (ids) 
“Biogeochemistry oF aming veils” (Wiley New York). 

Liopikoos. NS. PL 976) The Glide Bremation af Part 
Adehide, Quaerly Gealagical Natey, Geological 
Sunvey af South Australia ST, 4-7 

(1984) Quaternary matluses of South Australia, 
Departinent of Mines and Energy, South Australia, 
Handbook No. 9 

May RM. d& Bobleaiis, RP (P884) Coastal lincdstuo pin 
tm Pletstowene sediments wl Sellicks Beach. South 
Australi. Trem, Ro Sac. 8. lian 18, 82-04, 

McCoy, W.1 (1987) The preeision af yamine acid prea 
chironoluwy ate paleathermomeny Gees, Sen Rev 6, 43- 
Sel, 

MILLER, GL TE & BarotaAaM-Giil red ClUsyy Amine ae 
LeOchronology: VesOlihOn Afi preesion. in Curbonale 
hoses. Cain Ltinrrre!. VV T 28. 

MURRAY-Wal i aci ©, VtT99S A review oP he application 
ol (he amino avid racemisttion reaetion te qrehavolowical 
citi, Pie oleefeet 1, 19-26 


(2000) Quaternary coustal aminostraligeaphy: 
Australian ditt jn oa global contest pp, 279-3200 tn 
Goodlriend, G, A, Collins, Mod, opel My L.. Macko, 
So AL & Wehmiller do (ids) “Perspectives i amine 
acide und protein geachemistty” (Oxford University 
Press, New Works. 

& Bremekie ALP CLOO}) The Jase imurghicial 
shoreline in Australia —A review, Quan. Sei Rev WM, 441 
461. 

& Toons, Bo (i990) Direet tadiveurbon 
ealihrarion Ton anime edd racenmiation dating. olan ob 
Heil Sei, 37. 365-307, 

Berrbelo, AB, Bou ratsn, BaP Comin he db & 
Mech, DL. oM, (1999) Pawies arelieerie ul a last 
atetehieial hare oa model lor Quaterniey barrier 
develapnent fron the Coorong ty Maunt Ciambier 
Coastal Main, southeasten Australian Alen (iret PSR, 
177-195, 

& Coun A. (1005) Aminosiratizeiphy and 
clectran spin resoniince duting af Quaternary coastal 
neotvetonism Th Tasimdia and (te Buss Streit pts 
tast A ket Seg 42, 51-67, 

_WIMBER, ROW, L., Bom tio. ALB & Gas tin, Vv. 
WS. (958) Arminostatignphy af te last ivteriiient tn 
southern Australia, Severed IY, 33-36, 

Noten LOS. Sowiksy i ME & beers. WR 2000) (Ras) 
‘Quaternary Geachranolagy: merhods id appligutions” 
CAmericun Cieopltysical Unien, Retorence Shell 4, 
Washington, DC.) 

ny, VC 94d) Last Interclacial Shorelines ie the Western 
Maciic Rind Gea 103, 809-827, 

Pirtass, Bek & Bateman, ROP (1996). Fhe 
Brunbes/Matuvinit polarity transition ats at chrone- 
stralivraphic marker in Australi regelih) stdies 
TSE Tas Geol & Cleaphys, 16. 289-294, 

BUTTER NM. Wo de Cora, MLR (L995) (ids) “Dating 

methods for Quaternary deposits’ (Geological 

Association of Candi, GO Text 2), 

Hwbibe, DOA, (L984) Quaternary stratievaphy und sea 

level Variations tn the southewstof South Australia: pp. 

WIA fe Thom, 1. (Ed) "Coustal geomorpholopy 

in Australia” (Agadenie Press, Svdiey). 

Wank. WoT. (1966) Geology, geomorphology and soils of 
the south-western part af County Adehude, South 
Australia. CSIRO) Soils Pubtication Na, 23, 

Whustinobky, de Be (982) AU review ool qinine eid 
racenivation studies tre Qaaternacy  motliskes: 
stritivraphic ancl Chronologe applications mp eoastal wee 
Interghivial sites, Pociie and Adkinitic Coasts, Linited 
States, United Kingdon, Gatfin dstund dnd tropical 
islunds. Qneit Sek Kei b 83-120 

(1993) Appliculivns of organic geochermstry 
for Ouutermary pesearehc pminostealigriphy and 
unmnochronolony pp. 735-783 dy Engel, M.& Maeko.s. 
(ils) “Organie euovhemistry” (Plenum, New York). 

PAd0, CL SUVA, POO, Gay, La Tb AriiaManene, C., 
CIAL EB, Ba LAR I BAROAI Lara Alt a, ALC I999y 
Coast! Uplilt iy vontinental collision plate boundaries: 
dant fiom the Last Interghicish murine feraees af the 
we Strit area (sQuth Spain). Teeraiaplty, Ad YS 

OW, 


wa 


EUROPEAN-INDUCED ENVIRONMENTAL CHANGE IN THE 
ADELAIDE AREA, SOUTH AUSTRALIA: EVIDENCE FROM 
DRY CREEK AT MAWSON LAKES 


By ROBERT P. BOURMAN*, NEVILLE F. ALLEY? & KRISTINE F. JAMES* 


Summary 


Bourman, R. P., Alley, N. F. & James, K. F. (2002) European-induced environmental 
change in the Adelaide area, South Australia: Evidence from Dry Creek at Mawson 
Lakes. Trans. R. Soc. S. Aust. 126(1), 29-38, 31 May, 2002. 

Post-European Settlement Aggradation (PESA) sediments flanking the course of Dry 
Creek at Mawson Lakes reflect land clearance and agricultural activities in the twenty 
years or so following the establishment of European settlement in 1836. 
Sedimentation in this lower section of Dry Creek occurred in response to accelerated 
erosion on upland slopes related to land clearance and burning activities. A tree trunk, 
dated at ~400 years BP occurs at the unconformable contact between the PESA and 
the underlying Pooraka Formation of last interglacial age. Although this might be 
attributed to Aboriginal firestick farming activities, the discovery of a European 
artefact from the 1850s favours the view that Aboriginal practices were not 
responsible for the accelerated erosion and sedimentation in the Dry Creek drainage 
system. 

Key Words: Accelerated erosion, sedimentation, channel incision, European 
settlement, urban drainage. 


Treamactiony }f the Rovel Soviene af S Atsp (2002), L2AC 1 24 Bet 


KUROPEAN-INDUCED ENVIRONMENTAL CHANGE IN THE ADELAIDE AREA, 


SOUTH AUSTRALIA: EVIDEN 


“E FROM DRY CREEK AT 


MAWSON LAKES 


by RoBERE P. BourRMaAN. Nevitte PALEY 


& KRISTIE F JAMES 


Summary 


Batieman, RP. ALY, N. E& Jaws. ke F (2002) Ruropedi-induced envivoninental change in the Adetitide 


areca. South Australia Kvidenwe from Dry Creek ut Mawwsen Lakes. Dan. Ay See 


May, 2002. 


~ Mase, V2001), 29.38, 31 


Post-hurapean Setllement Agaradation (MESA sediments Hanking the course ab Dry Creek at Mawson Lakes 


ro 


relleeh kind) clearance ant aerieulioral actrvities Ta the twenty years or su followings the estiblishiment of 
European setlement in E836. Sedimentation in this lower sevtion ob Dry Creek occurred a response to 
aveclerated erosion on upland slopes related co land clearince und burning aclvities. A tree dunk, dated al 400 
yeurs BP occurs alihe dneonlormuble contict beoween the PESA and the viderlying Poorika Pornuition of last 
interglacial age, Although this might be attributed to Aboriginal firestick farming itutivities, the discov ery of at 
hurrepean urtelinel from the FH50s Ravoure the view that Aborsinal prictiees were Hot responsible for the 
accelerated erosion and sedimentation ithe Dry Creck dninage system, 


kay Worns, 


Introduction 


Luropean-mduced accelerated evasion, immediately 
downstrenin af te Mant North Road crossing of Dry 
Creek had exposed 6 mt deep verlical sections in 


Quaternary alluvial deposits over @ distance of 


upproximately | kdometre. Given tht channel 
stabilisatian Of this section of the creek was to be 
underiken in assocniion with the development of 
the Mawson Lakes housing estate, we deciled to 
exgunibe and describe the cypascd sections prior to 
thei’ destruction, a process which is now camplete, 
The aim of the remediation was to reduce erosion 
und downstream sedimentation, and to remove deep 
vertical banks (hat might present a hazard lo people 
iin dirbarised aren. 


Drv Creck drainage bast 

The study area lies on Dry Creek (hig. 1). which 
drains an area of approximately 109 kin’ jo the 
Qortheny and north-eastern suburhs of Adelaide. 
South Australi. and is bordered by the catchments 


 Solomebol Las ironmennd aie Reereaiion Manggeneat University 
of Sauth Attain Mawout fakes Cinpis Mawson Botleyued 
Minit babes SA S095, 

HOfnee oF Minerals aid Breny, PIRSA, GPO Bos 167], 
SA AHL. 

PRK AVION MPS) ACP AGI HUCTEILOTN: ASSOC RYTON WHT 
Wii bing & PARSE UNSW PRY LOD Oo) iy Creek and 
Lith: Mien Calehments Inieusiteal Calehinent Water Man urenent 
Men Hackorennd ood Opportanition (iPS luvironuiment & 
Introsriioture, Adwhaidie) Valine 2. 

BO Posi ok Asset pares ClO80) "Hydrology, Die Creek 
Draniie Basin Prepared fur the Corperations af the (yes af 
Sulishiey. Tea Pree Gilly oti hiveld. (He Lontkin & Aswecues, 
Wilolailes 


v\cleliniale: 


Accelerated erosion, sedimeniion, channel incision, Wuropea seulement. urban drainege 


ob the River Torrens to the south and the Gillle Para 
River tothe north. The drainage divide between Dry 
Creek and the bathe Para River is extremely subdued 
and difficult to delineate with prevision. A series of 
non-integraled streams such us Cobbler Creek drains 
from the western side ol the Para Fsearpnient (Fig, 2) 
and disappears into drains or the alluviuin of the 
plains. Dry Creek rises at the northeastern exwenity 
of the basin, some 400 fast on the Bden Esearpimnent 
from whieh many first order streams How, initially in 
a westerly direction. These sireams unite and flow to 
the southwest along the fault angle depresstou 
between the Para and Eden fault blocks, eventoally 
culling a bedrock gorge at the westerm edge of the 
ara block to debouch on to the alluvial North 
Adelaide Mains. Originally the stream, like so many 
others of the Adelaide Plains. probably dissiputed 
into the allivial deposits and rarely reached the sea, 
Today artificial driins carry discharge front the creek 
through mangroves and samphire fats into the 
estuarine tidal eavicoument of Barker Hlet 

Where tt crosses the Main North Read, Dry Creek 
uppers to be relatively imsigniticant and it ts 
surprising, to vote that it drains some OM oF the 
Adelaide suburban area, Because itis so intensively 
urbanised, there have been many inipacts on the 
drainage busin that have required remedial works to 
inhibi erosion The eatehment oveurs — pre 
domimanthy witht the Local Government 
Authorities of the Cities of Salisbury (50 km?) and 
‘Toa Tree Gully (S51 ky, A small area also oceurs 
within the City of Port Adelaide/Enfielkl (4 kin’) 
(PPK E & 1& Willing & Partners 1997) BC Tonkin 
& Associates 19s?) The development of an 
integrated eatehment water maniugementl plan in 


30 BOURMAN. R-P., 


ALLEY, N.F. & JAMES, K.F. 


V y oo 
\ \ \ 
. : 
Re \ 
\ yo" \ | 
\ w N—'h \ 
\ & } \ 
‘ : ot | 
“ 
F River : oy ! 
wlier 1 . 
A Gay ge ; 
} / v Vy} Para .. 
J x S { E RS 
\ % e 
AUSTRALIA \ e / Fault PS ‘ : 
' \ id ; ‘ Block & 542 Pleasant / 
™ ae para Rive: poke Mi: a / 
AN we | bo Omvier | 
. ) a \ | ry | 
Study Site —>© on? f- ae I 
y 1 / 
‘ at / 
Ke we | & @ J 
| rte 7 / & 
| jerRan | ew ADELAIDE © / 
| AUSTRALIA % 
| / g 
Mt. | 
— / 726 Lofty 
Z - | 
i) | = 3s A Pd 
/, ° RR 
Hallett at Noarlunga ‘ “ i / a 
Cove / Embayment a? 7 ny / 
et = J s i] \ \ / 
Pm / 202"in9 r | / 
pone i/,/ 
/ } 7 \y > 
Sg te y x 
aie | J o 
Ll {x \ x 
3 Willunga / My. 7 s 
ia) Embayment / “P8, 
& CY | & 
S ‘$ %, ® 
~ ans Mt. } _ & Ss 
“= Magnificent / 2 
ro 7a 82 7 aa. 
~a ely, : i == / 
7) Sellicks \°-% inni Fre va 
Beach /*~ . hora Flomig River *f. LS 
e. i pod 
$ rae Myponga 444 v] . ae 
@ / Hill oa zy / Ts 7 
“ey Cc \ 
Carrickalinga R. “a, Currency CF, \\ ( Lake Alexandrina 
Bing ns ae . 
hg ay. & he Kay : 
Fp os = & \nman sa5 / Hindmarsh) SK AE ey 
f Weymouth | a \a ‘el . Ellio ioe 4 ? 
7 Hit oni Victor in uN : 
{6 Fleuriev ‘ _S Harbor : 
ape : y 
* Jervis Aik! é f 
ee Lis! —“W sitpinge ‘ Encounte, 
 Tunkalilla Beach. Bay 
a Beach 
0 10 20 30 40 
— _ | =) | 
Scale in kilometres 


Fig. |. General location map of the study area. 


EUROPEAN-INDUCED ENVIRONMENTAL CHANGE 31 


__ Salisbury 


‘py prayaend pod 


Grand 


— ae 


Dry Creek Catchment 


Hope Valley 
Reservoir g xine D 4 
a en | 


Pig, 2. Map of the drainage basin of Dry Creek. 


1997 involving community and _— technical 
stakeholders and local drainage authorities should 
benefit this urbanised creek and its ecology. 


Drainage network 
The main channel of Dry Creek is 28 km long, with 
major creeks (81 km) and major drains (54 km) 


comprising the remainder of the drainage network of 


163 km (BC Tonkin & Associates 19807). It is in the 
lower parts of the catchment that artificial drainage 
systems have been installed in’ response to 
urbanisation, but in the upper parts of the catchment, 
upstream of the Para escarpment, drainage occurs 
mostly in natural creeks that are generally protected 
by flanking reserves. Nevertheless, there has been 
some development on flood plains, and interruption 
of watercourses by roads, buildings and other 
constructions, with the risk of flooding increased by 
culvert crossings and creek enclosure (BC Tonkin & 
Associates 1980?). 


Climate 

The Dry Creek catchment occurs in a region of 
Mediterranean climate with pronounced warm, 
dry summers and cool, wet winters. The rainfall 
pattern is strongly seasonal and evaporation rates 
are high, The annual average rainfall derived from 
gauging stations located in or around the Little 
Para and Dry Creek catchments is 531.8 mm (PPK 
E& | & Willing & Partners 1997!'), with a 
tendency for higher rainfall in the eastern part of 
the catchment. 


Land use . 

Information derived from a digital cadastral 
database indicates the following land uses in the Dry 
Creek catchment: mining and quarrying (2%), 
industrial (3%), open code (3%), commercial (4%), 
recreation (4%), public utilities (5%), primary 
production (5%), public institutions (6%), vacant 
land (23%) and residential (45%), which comprises 
the largest land use of the catchment (PPK E & 1 & 
Willing & Partners 1997!, Some 20 years ago, BC 
Tonkin & Associates (19807) reported that more than 
*...90% of the catchment comprises either existing or 
proposed urban development”. 

In light of the large drainage area, its considerable 
modification especially by urbanisation, and the 
strongly seasonal character of rainfall, there is little 
surprise that accelerated channel changes have 
oceurred in the lower reaches of the Dry Creek 
drainage basin. 


Modification of urban channels 

As with many other watercourses 1n South Australia, 
Dry Creek has been significantly modified along its 
length. Some modifications have been directly 
imposed. Other changes relate to indirect impacts in 
response to human occupation. In particular, increased 
urbanisation has resulted in elevated discharges, 
reduced stream loads and accelerated erosion where 
there are no protective works, and this has been 
particularly exacerbated downstream of artificial 
knickpoints. This has resulted in accelerated 
sedimentation even further downstream, 


32 BOURMAN, R.P, ALLEY, N,B. & JAMES, K.P 


The battering of steep bluffs and their landscaping 
are common features of urban channels. A relatively 
recent example of this occurred on Dry Creek 
approximately 2 km upstream from the present study 
site and immediately downstream of the Bridgeway 
Hotel at Pooraka. At this locality, from a naturally 
eroding steep river bluff some 6 m high, Williams 
(1969) collected samples of detrital carbonised wood 
and carbonate for radiocarbon dating in order to 
establish the age of the Pooraka Formation, The 
Pooraka Formation is a very widespread alluvial 
unit, which underlies much of the Adelaide Plains 
(Sheard & Bowman 1996), including the present 
study site, Bourman er a/. (1997) were not able to 
sample from exactly the same site as Williams 
(1969) in order to date the alluvium, using the 
different technique of luminescence dating, as the 
steep river bluff by then had been battered, 
contoured, rock protected and landscaped. A drilling 
mg was required to collect samples from 
approximately the same horizon as the samples of 
Williams (1969). As well as impacting on research 
activities, the engineering works have also destroyed 
the usefulness of the locality as a teaching site, 


Materials and Methods 


In carrying out this investigation standard 
sedimentological and stratigraphic techniques Were 
employed. In examining vertical sections, sediment 
samples were collected every 10 cm. Detailed 
descriptions of the sections are provided in ‘Table |. 
Wood incorporated within the upper suite of 
sediments was dated by radiocarbon techniques at 
the Radiocarbon Dating Laboratory, University of 
Waikato, New Zealand. All exposed sediments were 
carefully examined and collections were made of 
foreign materials incorporated within the sediments. 


Results 


Site description and field observations 

The study site occupied a one kilometre section of 
Dry Creek, downstream of its crossing with the Main 
North Road (Pig. 3). At this locality channel incision 
and widening had exposed a 6 m deep section of Late 
Pleistocene and younger sediments, These recent 
channel changes have been related to human 
interference, with the construction of concrete drains 
under the roadway mentioned above and_ the 
construction of an erosion drop — structure 
immediately upstream of the actively eroding zone, 
causing accelerated erosion. Prior to remedial works 
being undertaken, both channel deepening and 
widening were continually exposing fresh faces. The 
extensive urbanisation of the Dry Creek catchment, 
reduced sediment loads and increased water yields 


Fig. 3. Aerial photograph of Dry Creek taken in 1935, 
illustrating that the present course of the creek, 
downstream from the Main North Road had been incised 
and established by then. The straight artificial channels 
downstream of the study site are clearly visible. ‘The 
length of the section of channel from the Main North 
Road to where it crosses the next fence line downstream 
is~ | km, (Source: Commonwealth Government). 


have also contributed to the accelerated erosion, 
Unfortunately, the development of the Mawson 
Lakes housing project, occurring in the lower part of 
the catchment west of the Main North Road. has 
resulted in these informative sections being 
destroyed or obscured. Consequently, this paper 
provides the only written account of these formerly 
exposed sediments. Exposures of the Pooraka 
Formation are critical to future research on the 
antiquity of humans on the Australian continent, 
investigations of the past magnetism of the earth and 
climatic change. Thus it is disappointing that the 
trend is to destroy natural exposures of rocks and 
sediments in urban areas vital for earth science 
research and teaching activities. 

The general elevation of the land surrounding the 
study site varies between 15 and [1 m, Natural 


EUKOPEAN-INDLCED ENVIRONMENTAL CHANGE hl 


Med, Section th riche bank of Pry Creek in the study aren, 
Depth af exposure is approwumately Gm. The upper part 
ol the seetion (= 2 my comprises MESA deposits, A 
proponnced leached A horivon, which peers (wo thinds 
vb the Way up the seuion marks the rop oF the Moora 
bormarnon, TH averlies a red/brawn clay Behorzon 
coniiiniug Meeks of catenin carbonate. Chis fenine a 
wavy bovndiry wboye wricher calcareous zone. A second 
calcareous zone eeeues in the base of the seefion, 


hi 5, 
sediments overly ain Pooraket Pormtion allay iden on (ie 
Jele bank OF Dry Creek, The contael isapproximitely at 
(he position of The feel oF the persor on the fiddler, 


Post Buropoin Settlement Avgeradution (PRSA) 


levees Hankin the stream externa 2 to 3m above the 
level ob the suirrounding alli fin deposits. Che 
Jower part of the exposed section, From the channel 
Noor up tod level about +m ubove the channel Moor 
ismarked by deposits of the Pooraka Formation (Fig. 
4) recently dite ats a hast interglacial age, which ts 
wpprosimately 125.000 years BP (Bourman eft af. 
1997). This alliivilim was deposited during. time 
when global sea level was approximately 20 higher 
than ail present (Marriy-Wallace & Belperia 1995) 
und the climate was warmer and welter thar now 
These climatic conditions would have favoured the 
avyradation of sediments washed front our of the 
Mount Lotiy Ranges. During this rime wnt. fossil 
Marsupials, approximately the size ofa rhinoeeros, 


PwBLE b Averend in te beth of Deal Creek 

-235 em = Younger grey brown alluvium 

- 400m Grey ailty chiy. Sour sub: Qualiy pes-caprie 
(1..) bulbs deeur dew to depths of 40 ent, 
Sedimentation fias aecttred aver the bulbs 
Sediment tides Vesieuhur Aaraeter 

Light wrey clay sell, sliahely calenied. with 
catciin carbotate diffusing along root 
clamnels- 

Silly clay, bul Coloured and mottled with 
“VO TLINT Garhatiate Crirrclincn. 

Sedimenk cones vesicles. Wilh ant bests 
afd footholes to depths af AO cu 
Crey/hrowwn Chey sil wilh pods idl pockets 
of chaweou), Sediment is a little more clay 
richamd more lithified than abowe, Calejam 
Carbonless more pervasive than above, 
Dominwndy erey/brown -chiy displaying 
some sub hormwatal stratification with 
minor crosssbeddims, Ontye roimer quantities 
ob calcium carbonate ure present, 
Cirev/brown course enivelly sum, whieh 
extends gion at diseanlormity with) the 
under yime Poorakil Formation. 
Dysconformnine 

235 -S¥6 cm Pooraka Formation 

235 -200.0nm 
760 - 280 em 


40-40 on 


KO 130 cn 


10-1 700m 


170 - 220em 


2 - A eM 


Light grey To WItISh arty. willy sera. 

Bill cohotied, greyish aiottled sandy silt 
will) some clay, producing blocky pods as 
the material dries oul, 

PAN 292 em Rod brownechey, sllehtly motiled, 

292 -S52lum Lighter volouree gilewreous chiys, with 
loi carboriite penetrating: ine fissures 
and mol lines, 

Grey Jo bull coloured clay. with the upper 
Aen containing nodules and rhiveliths of 
calcium carbonate Seine verneal bleaching 
of sediments wong raet channels, 

Grev to butt coloured clay. with the upper 
Athen contaning nodules and rhivolihs ot 
calciuny carbonate See verneal bleaching 
of sediments along root channels 


572 -59Gem 


Ihe Diprofoden spp, roamed (he swampy, duende 
Adelaide  Phuins. Numerous discoveries ol 
Dipretodon spp. remains have been imade in. the 
Pooraki Formation (Tate 1879; Twidale 1968: N, 
Pledge pers. comm, (996) of the Adelaide aren, 
Pedovenic or soil-forming features are preserved 
within the Pooraka Formation (Figs 4 & 7). For 
example, at the top of the Pooraka Formation is 4 
lewelicd. bleached silly sand A horzon, which is 
underkiin by a dark ced brown chiy B horizon. This, 
in turn, is underlain by a Bea horizon comprising 
nodules aad eylindroids of caleium carbormate. The 
above soils typical of red brow carths, A seeoud. 
lower Bea horizon Wasteates ia halt io scdimentcan 
of the Pooruka Formation during us depusition. 


44 BOURMAN RP, ALLEY, NAO JAMES, Ke 


Fis. 6, Post European Setdement Aggeradition (PISA, 
sediments Gecupying a shallow channel cat into the 
Underlying Pooraka Pormation. The contact iy near the 
rop of he Midder. Phe height of the section iy fom, 


Multiple buried soils withitt the Pooraka bormanon 
dre common, such as in Cobbler Creek to the porth 
af Dry Creek. 

The distinctively coloured, red-brown Pooraka 
Formation with its leaehed. bleiehed whitish oA 
horizon is overlain hy up to 3 m1 Ol younger, grey 10 
brown coloured alluvial materiils (Fig. 52 Table 1), 
deposited as levees alone (he present channel In 
places the Pooraka Formation has been eroded 
vulling thraugh the soi developed on the Pooraki 
Formation and the chapnels we infilled with younser 
sediments (Fig, 6). The youne alluyium on the lett 
bank has been largely. alihoueh not exclusively. 
deposited as Overbunk deposits. whereas those on the 
night bank hive been largely deposited. as channel 
deposits (Rig, 7h The young alheviuin combs 
masses of chareoal, land snails and. possitly, a 


East 


14 m (AHD) 


West 
PESA 
vce e ces 
Pooraka 
Formation 
VE ECUCE ERE S Sako fF 


Leached, bleached A horizons 


== === Red brown clay rch B horizons 
freed Calcareous horizons 
— 
EE Overbank deposits 
— Channel deposits 


Fie. 7. Diagrammatic sketeh of section deross Dry Creek at Minwson Lakes. The width et the section iss 600 m 


big. & 
hetween 


free faink. posxsibly Fo larviflarens. sandwiched 
the underlying last intereincil Pooraks 
Formation ind Post Muropesin Settlement Ageridation 
(PRSAY sediments. Che outer part af the (rink wis 
natiowirbon died id ~ 400 yeurs BP. 


Pie 8) Pholoeriph ob the base ol bottle with a shallow punt 
wdhwilh the easenption (CW Coe foculedhal the contact 
Hetween the Poortake Pornarion and the PESA sediments 
The bottle is thought to have been manulienured inthe United 
Kihzdom diving the [850s TS00s. Phe penis }4 en long. 


FUROPEAN-ATNDUCEL ENVIRONMENTAL CHANGE 4s 


imivealith artefucr. In places there are ripple 
strictures al the aneonformity and the ripple 
stritures are preserved both on the buse of the 
sediments and the top af the unconformity. 

A ipee drunk (Pie 8. lying horizontally, wees 
locwted wl the canter between the undecdying 
Poordku Formation and the overlymne younger 
lovin the right bank iminedlitely downs trean 
from (he drop structure across the channel A sample 
of the ouler part of the tee trunk. which appeared to 
be Laneelyptiw largifleventy UR Muell). was collected 
for radiovarbor diting. The onter part af uve trunk 
Was sumpled to dite the youngest part ol the trunk, 

The study sire was revisited after winter rains, 
Which had ficilitated fiorther underenting. Retreat ul 
the channel walls had exposed more of the 
uncanfonigble contaet, revealing the presence ol 
numerous European artefiets that included parts al’ 
bottles (both glass and ceramic). cattle bones, 
feneing wire und other metal abjeets. Some ol these 
objects were exactly atthe base Of the tinconformity, 
In particulin, (he bottom ol a black elass boule with 
the inseription “CW, & Co? was recovered from the 
base of the uneonformiy (Pig. 9. The bottle base 
has an indentauion known as a Spunt’, “hieknp™ or 
hick (Lachenmann 200] ') 

In dhe ease of Dry Creek there 1s hulle evidenee of 
wecleiuted erosion pripr tu the deposiuon al the 
PESA Sediments. A very sell developed soil prolile 
af the Pooraka Formation sugeests thar landsespe 
slibilily favoured the operation of pedovente 
provesses. Only ing few minor instanees was there 
evidenee oF the Soil protile developed un the Pooraka 
Formation beme eroded prior tothe deposition of the 
PESA sediments. which have a miuximum fhickness 
of <3 m. 


TDseussinn 


Coses of etrannel erosion 

The jnitiativn of sullies and channel erosion is 
related to many fietors. According ta Beyin & 
Schum (1984), gully erosion occurs Whenever the 
power Of Tows exceeds a threshold value equivalent 
iy the resistanee of the valley Cloor This may be 
(fected by basiy wide external fietars such as 
chimate and catchment wide luaduse, These tactors 
will imitate erosion on relatively steep and narrow 
seeuons of the valley-looras these sites are closest 
to The threshold condition and will respond first to 
altered conditions, Gully development can be. but 1s 
not necessarily, related to anthropayenic influences. 


Pachbewivnn, Mo rZuuhi Pio Prat femdare accessed 5 Oeb 20h | 
(TEL Wipe ev WiMe WP UL beih Gn abhi) jb 

HOOEM AB. IR. POLE) Londtori Studies reap Viet Llarlinti, Gy 
(Hons) Wests, Mie Liitversity of Adelaide (api), 


Gullying has occurred prior to human inlerterences 
amd may be atiribuled to the effects of climatic 
vhange influenving vegekition and runoll, tectonic 
uplift including tilting of the land or custatically- 
controlled sea level movements, Schumm (1979) hits 
also emphasised tat changes can occur as a result of 
factors inherent within the geomorphic system, For 
example, an agurading alluvial fin surface mas 
progressively steeper ly such an extent thta-eritea| 
threshold slope ty uchieved when the Sireann may 
hesin to ineise iS Owe deposits without extemal 
conditions changing, Site specitic factors such us 
ploughing. bridge and culvert coustruetion aod 
drainage schemes can also initiate erosion especially 
Where the eliinges ure must severe (Bourman & 
James 1995), The potential! rales ol non-hurnai 
factors ail human tifluences af streany 
sedimentation and erosion al the study site will he 
apsessed, 


Timi. of sedinwatation ahd chanel exasion at tlt 
vtuely sole 

The tree wunk ot the ungenfarnity between the 
Pooraka Porniation aud younyer overlying allayiiien 
returned a radiocarbon age of 420+ 50 vears BPOWk 
S825). This pachocarbon cite might Sugeest that there 
was accelerated erosion in the Mount Lofly Ranges 
about 400 years ago, possthly rehited to Aborninal 
occupation and burniig for frestick farming. This 
interprefation i$ supported by the observation of 
charcoal in the vounger alliwiunr and ils oecurrence 
close tothe deontormiry, A-siotikur situation oecurs 
in the Gawler River, approximately 30 knv yorth of 
the study site. Radisecarborn dating of Wood and 
charcoal incurporated ina alluviim, wis rimdertiken 
by C.R. Twiclale of the University of Adelaide, Vhe 
samples al carbon and wool collected from wiht 
the wlivial deposits near the present day channe| 
were diled at 374 + 70) Years BoP CANT? Sample No 
204) and 235 + 70 Years BLP, (ANU Simple No 205) 
respectively (Bournaa }969 ). These data. ton. are 
highly suggestive of uccvlersted sedimentation und 
erosion velited to vewetation disturbance by 
Aboriginal baramg activities. 

Bushlires prior to European settlement may have 
Hee alate deamatic as jilustvated below. Our 
altention was drawn fo the followiny by B, Taylor, a 
descendant of one of the early settlers, J. W. Adams, 
who artived on the “HMS Ballilo” in 1836 and whe 
penned wi aecounl of his erly days inthe settlement. 
‘This included a graphic account of a major summer 
bushtire Te the Mount Lotty Ranges. The “Bulfalo” 
met the “Signel™ at Port Lincola ou 24" Deventher, 
[36 and they saded together to PokiGust Bay where 
they dropped anchor on the 27" December, [836 
(Adams 1902) “When the anchor was cropped the 
usual bustle commeneed for tarding. Before we leh 


MG BOURMAN, R.R, ALLEY, NLP & JAMES, Kl 


the ship we witnessed a grand sight. All the hills and 
sullics as Tiras we could see were on fire, and the 
rellection wis.so strong Lhal we could see every rape 
and the men walking the deck of the “Signet. She 
Was about halla inilé in shore from us, and we were 
About five Wiles oul P have seen many fires sinee, 
but nothing iy compare wilh thal for grandeur? 
(Adkins L902), 

In combinadon, the ~ 400 year radivenrbon date on 
the tree funk imcorparared within the cecenr alluvial 
sedirnents oF the plain, plus the first hand account a 
imensive havaijg on the adjoining Mount Lothy 
Ranges CAckiIms 1902), could suggest that the 
aceclerated crosiin and sedimentation may bave 
vecurred prigr la the arrival of Buropeimis iind hid 
resulted from fires shuted by Aboriginal peoples 
‘Vhere is no direct evilenee that the Sires of have 
Devember, (846 were started by \boriginal people 
and may have had natural catises. blowever the 
imense und widespread bushfires of more recent 
fines have oecurrect later in the fire season. usually 
in Tebrogey. This might support the view of 
Aboriinil influenees i stirling the fires of E836, 

Revardlass of the cause of the 1846 fires and the 
possible assocniion of Aboriginal burping aetivities 
with oeeelerated erosion and sedimentation, the 
discovery Of buropean artefdets in the younger 
mllovinrn, pariicularty the base of the glass bottle al 
the uneentommiiy, indicates thin the aevelerated 
landscape chanwe did not occur until some tine alter 
Europea setlement. The oveurrence Of a 400 year 
old tree (rink al the base of the youneer sediments 
Woes nol mean thar (he sediments were deposited 400 
years age, hut only that the tree died 400 years aye. 
Atlempts made to identify the bortle base with the 
uiseription “CW & Cor unequivocally have not been 
stcesstil, “Phere is no doubl (hat mh wis nol 
mnufachied locally os no bottle manufieluring 
firm) with this trademark hus exited an South 
Austrilia (Shiedrd & Tuckwell 1993). Furthermore 
Palen Shucird, ain wuthority on antique bottles, 
informed the writers (pers. comm, 25/10/01) that the 
bottle wes alfoost certainty a hal pint bottle 
monefaclured in the United Kingdom during the 
P8505-) 8605, and thatthe bortle predates the earliest 
bottle neanulucturing i South Austrade 

The Gecarmneve of buried saursoh bulbs (Qwelis 
pes vapree 1.1 ta depihs of 40 cm also provide dita 
ou the Himing of sedimentation, which post dated the 
Hitrnducton and dispersal ob Oyrea/és Trani Seuth 
Attics to South Austria, 

The ahove observations indicate thal the younger 
erey-brown alluvium is: aerually Post-Ruroapeon 
Settlement’ Averidatoo (PESA) and probably 
refloels sedimentition due ta accelerated erosian 
related to land clearance and buroing 1 the tipper 
caleliment zones, It alse sugeests that prion to 


Furopean Geeupation there was ner alhiyigiy 
overlying the Pooraka Formation al (his site sane (hat 
an extremely rapid rate of depesition tormed the 
levees Ut are up to J Thickness. The sediments 
Were not deposited at least until the P850s, given the 
postulated manufacturing date of The “CW & Cy” 
bottle, Furthermore, we Kaow that the presen) choy 
ineised ehanuel was established before 19300 as 
indicated by ao aerial photograph (Fig. 3) so that u 
HUNT fale oF Scali mentation building the levees is 
~ 23 em yr, 


Canyes af yedinentation and charted ineiviou at 
wed site 

AL (he study site there is ne evidence for mitirally 
oeeurring episodes of sedimentation und erosion since 
the Last Imerglacil (125 ka BPY when the Poorahi 
Fornatign was deposited. Only miner erosion of ihe 
Poomka Bormation fas oecarred. Preservation of a 
complete soil profile on the Pooraka Formation is 
common (Rigs ++ & 5). reflecting subactial exposure 
and landscape stability. Inthe wider Adelaide reyror 
the youngest alluvial uit Hat bas been recognised is 
naturally occurring os thal oF the Middle Holocene 
Wildeda Bormation (~ 5-6 ka BP) (Bourman ur al. 
19907), whieh has heen related to climatic change ane 
(slightly elevated sea level. Consequently, within the 
Jocil area. recent! accelerated sedimentitian said 
eresion is most likely die lo human laclors. especially 
ws scdiment Conlin’ Huropean artetaets (PESAD ts 
so Widespread within and Manking dhe Mount bolty 
anges. 

Hi has been susvested that valley side vegelilion 
vlearanve alone ts isulfieict to initiate channel 
drosion, which requires some fom ob chininel 
distirbanee such ds by stock gravis wid drainage 
works (Prosser & Slide 1994), Toweverucecleratcsl 
deposition of PESA derived: front Ure valley sretes 
Nitliwink elcuranve. mdnecthy leads ta erosiin by 


20 


Hig. 10. Mitre reeen) PRSA depeyity extending up in 2.2 4) 
ubove the gidly Hoon These sedimenis contained rece) 
Lurpean artefer svelte phinties, Tot) heuthh of 
acctiun = Am 


LUROPBAS-IN DUCE BAY POONMEN TAL CHANGE 5! 


burying Vewetution, killing i and steepening Slopes. 
‘This sets the stage foc channel erosion and incision 
Hhroush (he PESA sediments and into the underlying 
Unis ho fonger prometed by veretation. 

hollowing deposition of the Post-Kurapean 
Settlement Age radian (PES A) alluwium the strean 
col down throwh the PESA materials. into the 
Pooraka Tormation. developing a deep treneh-like 
channel. standing the PESA nitterials hich ap on the 
binky. Vhere is evmlenee af several phiises ah Post 
European Settlement Ageradation, based op included 
anfelogis dnd the level OF the PESA filling. At feast 
awn fords at younger PESA sedinments (Pig. 1) 
vecdr within the ehannel with the youngest 
containing plastic materials including bubble plastic. 
‘This rhalertal hid previgusty ti-filled parts of the 
channel rod depth of 1.5 m before renewed erosivi. 
Iris Uilfieule i determine whether these changes 
were cursed by setilement activities, or natural 
chanwes mn Hood periodicity and tlensity. 

Inilial incision of the channel Mllawed neeelerated 
sedimentition associated with fined clearance several 
duendes aller European settlement. Subsequently 
any other fietors haye infliteneed the sporadic 
erosion und sedimentation of the channel. Reduced 
bedloads and inereased water yields Lollowing 
Urbanisitin live impacted on the study Site chanel 
as have engineering works such as channel sation 
wil the constriction of arilicnl knickporits, 

Alihquely some of the vlurceul jo the PESA 
sediments muy have derived trom Aboriginal tres. 
such as i described by Adkins (L902), there is fer 
Cvidepce lo sugeest (hat the acegleribed erosion and 
averidation were related to Aborivinal activitivs. 
Punhermore. the unbtirnt. 400) year old: tree trunk 
may have lain around a the landscape fora very 
Jong time before bejoe jncolpergted inte the PESA 
sediments. This iplerprelatian supports (he vigays of 
Prosser (1990, (8b) awha moted pe Tnerease in 
Witespreud ugenmadon assockited with Aburivinal 
hiring a Wangrih Creek mm the Southert 
Fablehiids of NSW. Prosser (1990) aso noled ne 
evidence for invrensed frequency OF diiviiion athe 
Life ob tntensified kid use, 

A tiotulisingly similar study was produce hy 
Nelson (1965) Tron the Chemtog River Valley of 
New York ikl Penieyivania. He concluded that 
overhank deposition on the floodplain aveclerated i 
recent pedlogicul Lime, Mainly asa cansequenee of 
Numan iilerterenee, Cleric and culation 
increased paniolh, erosion sind Mood heights resulling 
in hicher stream sediment Toads jand mere nupid 
averbanh deposition vi tle lToodphin A piece af 
wood fevoyered LAK ty trom the sactaee and dated iat 
41+ ISO years GP jiwlignted a sedimentition rate ol 
O42 cm ye! With the appearinceat Buropean debris. 
there was a markedly (hereased cyte oF sedimentation 


to 1.7 em yr! Nelson (19635) kuwely attributed the 
IMpaats fo Europea settlement but emphasised the 
aericniiieal role oF the indigenous Midian iibaubilunts, 
Who probably inittvted the sequenve of changes 
Lenturics ayo. 

The shallow channels araded inte the Poorake 
Formation and the thick PESA sediinents sirinded 
high above the channel Moor favour the view that the 
inition oF ehwinel ieision any be related to PESA 
deposition. This would have baried former shallaw 
and vegetated channels. killed the stabilisuig 
vevelalion and sleepened gradients by deposition 
Once inated. variqus other Caictory woul! have 
vontribured ro channel erosion. This passible that the 
later Construction of gy drain cy the lower part of Dr 
Creek, downstream of the study site assisted Tuirther 
channel Hieision. An uenkd photograph taken on 
VAYTL/P935 (Pig, 3) shows the location of the 
(artificial channel andl reveals Chal the charmed inthe 
sticky site was incised prior lo extensive urbanisalren 
oF the gachrment. Consequently. inereased ranull 
related fo urbanisation can be dismissed as tun inital 
cause Of The channel inelsion. although it tas 
subsequently been important in causine ehaonel 
deepening and widening, as lave the placement ol 
the chiwmel in conerele conduits und the Construction 
of artificii! kntek pots 

Phe sequence oF events desccibed here (§ similar be 
those discussed by Sehurnnr (1977) who noted that 
there May bea sequenve or gascade of consequences 
Volloawing initial clearing of Gitehments for pastitre 
purposes, Such clearwmee results ta aecelermed. soil 
erosion On hillslopes, resulling to aggradation along 
drainage lines that can not aecommnudare the 
avalluble sediment load. Pvenruially, as the supply al 
erodible iateruils is exhaisted, jncreased ruprort 
fron) the valley side slopes conlinues and inevitably 
feads iy downstream eharnel incision. Onee the 
inital disturbanee of hillslope clearance has cecurred 
the SWHch from averidavon to mension could aceur 
without ficthe exterbal iM lienees, 


Conclusions 


Post-uropean Settlement Aggradation (PESAI 
Sediments Mankiiig Ue vourse ol Dry Creek i 
Mawson Lakes ure iilerpreted as the result ot 
European asveulural practices elie the pero 
TR36-1800, Accelerated erosion Qu upland sfopes 
Wis aispachied with sedimentition in the lower 
sechion of Dry Creek. Subsequently. in response to 
(he sedimemation killing stabilising channel 
vegeltion. channel jieiion was probably iitiered 
us scUiment yield reduecd and. rinatl mnereased, 

Aree trink oceirriny at he unconformity berweeyr 
the Poorake Formation aid the PES A deposits dated 
wl ~ 400 years BP. could be suggestive of Aboriginal 


3s BOURMAN, R.P., ALLEY. NB, & JAMES, K.P, 


firestick farming activities. However, the discovery 
of European artefacts of the [850s favours the view 
that Aboriginal practices were not responsible for the 
accelerated erosion and sedimentation in the Dry 
Creek drainage system. It is possible, however, that 
some of the charcoal from pre-European fires such as 
that described by Adams (1902) may have been 
incorporated into the PESA sediments. 

Finally, a disturbing feature of urbanisation has 
been the loss of many significant geological sites, 
which have been destroyed or covered in the 
interests Of aesthetics and/or public safety. Two such 


sites have been deseribed in this paper. There is 
clearly a need for local government authorities and 
developers to consult with geologists prior to 
underlaking major ‘restorative’ works, 


Acknowledgements 


The paper was improved by the contributions of 
the referees Professor Martin Williams and Dr David 
Dunkerly. Funding for the radiocarbon date was 
provided by the University of South Australia. Chris 
Crothers drafted the diagrams. 


References 


Apams, J. W. (1902) “My Early Days in the Colony” 
(Published by his daughter, Sarah Tilley). 

Brcin, Z, B. & Schumm, S$. AL (1984) Gradutional 
thresholds and landform singularity: Significance for 
Quatermary Suidies. Quer. Res. 21, 267-274. 

Bourman, R, P., MARTINAITIS, P,, PRescorr, J. R., & 
Benrerio, A. BP. (1997) The age of the Pooraka 
Formation and its implications. with some preliminary 
results from luminescence dating. Trams. R. See. S. Aust, 
121, 83-94. 

& James, Ko (1995) Gully evolution and 
management; a case study of the Sellicks Creek drainage 
basin. §. Aust. Geag. J. 94, 81-105. 

Murray-WaALLace, C. V, & Brbprrio, AL P. (1995) 
Aminostratigraphy of Quaternary coastal sequences in 
Southern Australia - an overview. Quai, Mirerhat, 26, 69- 
86. 

Nevson, J. G. (1965) Man and geomorphic processes in the 

Chemung River Valley. New York and Pennsylyania, 

Agsac. Am. Geograpliers Annals 56, 24-32. 

Prosser, 1. P. (1990) Fire. humans and denudation at 
Wangrah Creek, Southern Tablelands, NSW. Als. Geog. 
Suidies 28, 77-95 

(1991) A comparison of past and present 
episodes of gully erosion at Wangrah Creek, Southern 
Tablelands, NSW, [bid 29, 139-154. 


& Spape, C. J. (1994) Gully formation and the 
role of valleyfloor vegetation, southeastern Australi. 
Geology 22, 127-1130. 


ScHUMM, S.A. (1977) “The Fluyial System” ohn Wiley & 
Sons, New York), 


(1979) Geomorphic thresholds: the coneept and 
its applications, Jnsritute ef British Geographers 4, 485- 
315. 

SHEARD, M, J. & Bowman. G, M. (1996) Soils, stratigraphy 
and engineering geology of near surface materials of the 
Adelaide Plains. Mines and Energy South Australia, 
Report Book, 94/9, 


SHUEARD, H. & Tuck WELL, D, (1993) “Brewers and aerated 
Water manufacturers in South Australia 1836-1936" 
(Swilt Printing Services Pty Ltd... Stepney. South 
Austratia). 

Tare, R, (1879) The Anniversary Address of the President. 
Tras, R. See. S. Aust. 2. Xxxix-IXxv, 

Twipatp, C. R. (1968) “Geomorphology” (Thomas Nelson 
(Aust) Ltd, Melbourne). 

Wittiams. G. BE. (1969) Glacial age of the piedmont 
alluyial deposits im the Adelaide area, South Australia. 
Aust. J. Sci, 32, 257, 


A LATE PLEISTOCENE OCCURRENCE OF DIPROTODON AT 
HALLETT COVE, SOUTH AUSTRALIA 


By N. 8S. PLEDGE*, J. R. PRESCOTT? & J. T. HUTTON 


Summary 


Pledge, N. S., Prescott, J. R. & Hutton, J. T. (2002) A late Pleistocene occurrence of 
Diprotodon at Hallett Cove, South Australia. Trans. R. Soc. S. Aust. 126(1), 39-44, 31 
May, 2002. 

Despite Diprotodon fossils occurring widely across Australia, until recently, few finds 
have been adequately dated. This is due to several reasons, primarily the inadequacies 
of the radiocarbon methods. New dating methods, which coincidentally increase the 
datable age range, have been developed in recent years. One of these is 
thermoluminescence (TL) dating. Yet there are still few reliably dated Diprotodon 
specimens because they must be found and dated in situ. A chance discovery in 1992 
gave the authors an opportunity to test one of these new methods and at the same time 
solve a thirty year old mystery. An articulated portion of a Diprotodon skeleton found 
at Hallett Cove is associated with sediment TL-dated to about 55,000 years, and is 
also a possible source for a fossil tooth found on the nearby beach in 1971. 

Key Words: Diprotodon, Hallett Cove, thermoluminescence dating, late Pleistocene. 


Tramsnnetiions of the Reval Seaury ofS. Ange, (2002), 12601), 9b 


A LATE PLEISTOCENE OCCURRENCE OF DIPROTODON AT HALLETT COVE, 
SOUTH AUSTRALIA 


by N.S, Peenar . J, RB. PRreseort & J.T. TUTTO: 


Summary 


Probate N.S. PRiseos wh RO & Phvepos. LT. (2002) A late Plemtocene veeurrence ot Dipratodia al Mallett 
Cove, South Australia. Tras RoNoe 8 lust 1261), 39, 31 May, 2002, 

Despite Diprofodon fossils oecurring widely across Austria, until recently. lew Hinds fave been adequately 
dated. This is dite to several reasons, primacily the inadequacies of the ryhoearbon methods. New daira 
methods, whieh coieidentally inerease the datble awe ringe, have been developed in revent years. One ol these 
is thermaluminescence (ULY dating. Yet there are sul) few reliably dated Diprirodi specimens because they 
Musabbe found and dated desi. A chance discovery in 1992 etive the authors an opportunity to test one oF these 
new methods and al dhe same lime solve a thirty yeur old rrystery. An articulated portion oll a Dipratodan 
skeleton found af aller Cove is associated willy sediment TLedated to abou! 35 000 years, and ts also ce possible 
source fara losail tooth found onthe nearby beach un 1877, 


Ry Webs: Diproteden, Hallett Cove, thermoluminesence diving, late Pleistocene, 


fatroduction 


Many speemens of Dipromdun have been found 
Since its discovery by Major Mitchell an the 
Wellinyton Valley, NSW, Tir the carly 19 Century, 
and precise aves for Unis, the largest Known 
mirsupiil, have long been sought. Many, if not most. 
discoveries were made belore the development of the 
C-le+ jiethod of radiometric dating, Others were 
demonsurably beyond the ditable age ninge ahd 
hadioearbon dating oFolder material has been shown 
to be dnrelinble (Chappell era, L996: Roberts ev al. 
2001) Sal others could not be dated for want of 
sufficient preserved carbo. 

In }992 Mr Westlake, whilst walking his dog in 
uv newly designated council reserve at Hallet! Cove 


(hig. 1), 25 km south-southwest of the city of 


Adelaide, notiecd what appeared to be 4 large white 
bone (Fig. 2) eroding oul af an old) exposure of) & 
former private road. Closer examination supported 
this identificauon, and Mr Westlike subsequently 
informed the South Australian Museum, although he 
was sore thatthe relevant people would have knewn 
about iLalready, The o¢eurrence was not known and 
a visit was immednitely orsanised, 

On 26 June, 1992. Mr Westhike guided the senior 
authorand student Gavin Prideaus to the site. an ald 
roud-culling. through a spur of hillside overlooking 
the Pield River, not far from the beael ar Halleu 
Cove (about 35° 49" South, 38° 29° 8" Last). The 
bank was more (han 2m tieb, and the bore was 


Sourt Australie Museum, Marth ferme Adeluide Ss A S000, 

Deparment of Plivsigs tad Mhthentation) Physios, Whe Mencersity 
aH Adlelaithe SA 9005 

Dp Hel Hon dies! charrr (he arly: sties: eA? prepieneien Ob TA 
puprer 


exposed about |S no below the top and about 2 in 
above the surfitee of the nearby bridue. Across the 
road, the hillside fell steeply to the river about 5 m 
below. The bone was examined jv sie and appeared 
to be part atthe pelvis ofa large animal und, because 
it way Tossilised and se large, probably of a 
diprotodonlid. With some difficulty, the bone was 
excavated without greatly enlarging the culling and 
pluster-jacketted for Iransport. 


Materials and Methods 


The jucket contaming the specimen was opened in 
the laboratory and the sediment removed trois 
jround the bone by scraping witha small dental tool, 
often when the soil had been softened wilh) water. 
The bone was hardened piecemeal during this 
process, using, a dilute soludon of Bedacryhe in 
acelone, The stratigraphic section was measured 
aller the excavation, using a lipe-mensure, Other 
measurements were made by vernier caliper or ruler, 
as warranted. 

Sampling for thermeluminescence (TL) dating 
(Aitken 1985; Winthe 1997) wits carried out by 
Prescott and Hutton und several graduate students 
from the University of Adelaide Physics Department 
on 28 Auwust, 1992 (Fig. 4). 

Three horizontal auger holes were drilled into the 
bank (ig. 5) to brueker vertically the position of the 
bones, which had been removed earlier, FL samples 
FRIS/0.9 PRIS/IS and ERIS/2.1 were collected 
tor laboratory analysis, at depths below the top of the 
culling of O.9 mi. 1.3 theand 2.7 m, respectively, /a 
Vu gamina ray spectrometer meusurements were 
nade in the same holes trom whieh the Th samples 
were colleeted, at about 5 m depth into the exposed 
face oF the cutung. 


40 N.S. PLEDGE, J. R. PRESCOTT & J.T. HUTTON 


SOUTH)AUSTRALIA 


ADELAIDE } # 


Fig. |. Locality map; the fossil site is ina council reserve on the Field River. 


Gamima ray spectrometry gives a direct measure of 


the radiation dose rate due to gamma radiation under 
prevailing field conditions and subsequent data 
analysis gives the concentrations of K, U and Th. 
These are then used for calculating the total dose 
rates from radiation in the environment, and for 
comparison with independent measurements in an 
assessment of the likelihood of radioactive 
disequilibrium in the deposits. 


The age 1s calculated from the age equation: 
age (ka) = Equivalent dose (Gy) 
dose rate (Gy/ka) 
where doses are measured in grays (Gy) and ages in 
kiloyears (ka). 

Quartz grains in the 90-125 jum size range were 
extracted from the samples by standard procedures 
(Huntley ef al. 1993), 

The selective bleach method was used to find the 


LATE TIISTOCE NE IPROTODON THOM TALE EEE CONT lI 


Pie 2. The lossil bone as initially expesed in sandy lens 
benvern navel livers, blunnter ty S00) mim Jon. 


lip. 3. Vhe excavated fayparecene pelvis, fa da Tae 
skeletal Pragiment is upside dowel anterior uita Ute batik, 
bie card hy the Taner tandhe i 90ex 55 ten. Eb hell 
iselium: RL right tum Vovertebrae, 


hig. 4. J aprritivediua loxsil atte: wenera! view, casmvands, 
biel River ut mbt. Preparations being mude tor 
thepmuliniinescence during by Adeltide University 
Physics Department Still and students, (he Kile Dr donn 
Hilton second front mght, Prot) totin Prescott alariehl 


Vis 3. Lhallet) Cows Uhermotimineseenes sunmpliqg sthes 
Slosv ing jeeneral stritcraphy. cenhint hole mek the taser 
hormvon, Ja ie saan ray seigliidion coMnliie t 
Wuakirie place tn thie lowest tole 


equivalent doses (Prescoul & Mopirrabr (903). This 
method was deyeloped to reduce (he uncertainty in 
the level of solar bleaching, which resets the TT 
clock. The protocol uses optical filters Gy select the 
rapidly bleached component of the Th, The 
equivalen! dose is determined by comparing (he 
mitural “UL signal with one generated in the 
iboratory by a standard radioactive Source. The 


specific wicthod is known as “The Australian Slide’ 


(Prescoll ef wh 1993), 
Results 


The Jossily 

The main speeimen was found to comprise purhs ol 
both deh and tight pelves sill artieatated with the 
sacral vertebrae, plus an udjoininy luimbar vertebra 
id & frament of the first caudal (SAM: P33487) 
dnd js comsidered fo represent the rant miarsupral 
Diprotadan Owen (839 (Pigs 3), 

‘The pelves form a fairly Mac plate ata slg angle 
tu (he vertebral axis, The aectabulunt diameter is 
about LOU Wiad, the seni iteraectabular width (right 
side) fs about 200 dm, the saeral leneth about lau 
mim and the schio-iiae length is about 440 mim, The 
verlehrue dre nol well preserved except lor [heir 
weural arches: a lumbar vertebra has a eenteum with 
4 transverse diameter of about 80 mnt und length of 
OO nine These measurements ure within the range of 
specimens of Diprotocdoanspp. trom Luke Callabonma 
in the collections of the South Australian Museum 
und, aller comparison with We pelyis of a skeleton 
displayed in the South Australian Museum, the 
Hallett Cove specimen is considered to bea subadutt 
or female individual. Specific identity is not passible 
on the nuaternil preserved. 


Preservation is not good. The bones ure not 


42 MS. PLBDOIN WI KR. PRESCOTD Al bh HO ETON 


pettitied, being rather chalky, leached and 
unmineralised and held together largely by the 
supporting sandy loam. This cireumstance proved to 
be both a help and a hindrance during the preparation 
of the specimen, as the bone was Fairly easily cleaned 
bul had lo be consolidated und strengthened during 
the proeess because it would not support much 
Weight. The specimen is consequently fragile wath 
numerous fine cracks presumably associated with 
son movement. The lower surface of The specimen in 
the sediment (ihe animal's dorsal side) is Cairly 
complete wilh only seme crushing of high points 
such is the dorsal processes of the neuralurehes. The 
opposite Surface is less well-preserved with much 
Inissing bone and irregular ends, presumably where 
exposed, unburied parts had been croded by the 
elements or incoming sediment, Although soll and 
Susceptible to later damage, no cut marks, either 
from scavenger lecth oor hunter’s foals, were seen on 
uy hone but vo limb bones, which might have been 
Wore. wbractive targel, ure present. 

fy (he process of excavation, & few more bones 
were found in close association willl the pelvis, One 
of these bones. a fravment of immature lel mandible 
(SAM P35074), supported the dentification af the 
pelvis as Diproticdonw. This specimen, trom which 
erupting checketceth had fallen leaving only a barely 
worl ineisor, ts Only 42 mn deep al the first molar 
ulveoltis. ws compared to 100-710 mo or more in 
adult animals. Fragments of a title-work My ot 
Dipovedon were also found, Another bone was of a 
large kangaroo, Muerepus? sp, Several shells of 
ginal) saxivelous dand snails, Sioa asreales 
Verussac, 1821 (Suecinidae) and Pereloersiy arden 
Trodile, 19357 (Heleunonidae) (R. Hamilton-Bruce, 
pers. conn & daly 2001), apparently the first 
recorded fossil occurrence of these species in 
Australia). oceur in the fine sediment surrounding 
the hones, together with fragile moulds of fine stuns 
Such as are Seen in Cher-lmestones ins modern 
stream-pool deposits. 


Creiloauy 

The fossiliferous sequence appears to he ou 
marginal facies Of the Poorka Formation. cevently 
redated by Bourman et al. (1997) Much of the 
Dinratoden skeleton had been lost, either by 
disarticulition or vrosion before complete burial, or 


as oa result of road-buildiim exeavations. The 
remaining bones lay upside-down in a shallow 
depression Hlled with poorly-sorted coarse sand, on 
a bed al somewhat current-imbricated pebbles ol 
Precumbrian sandstone and shaly of apparently loow! 
origin (Vig, 3), The sandy horizon is lenticular and 
extends several metres on either side af the bones 
before pinchiny out. The pebbles, tanging up to some 
5 em in diameter, occur in beds 10 to 30 em thick 
above und below the sand and are subsneular lo 
subvounded, Similar beds, alternating with sandiec 
horivons, oceur throughout the sequence exposed in 
the cotting. Bedrock of steeply-dipping, slightly 
MelaMorphosed Proterozoic slates and quartzjws 
oecurs within FO om Juterally, and evidently forms 
puro! the original valley wall. 

lhe stratigraphic sequence wt the site of the hones 
is SumMMariscd below, 
Soil ul least O35 m al lop of culling, 
Mayvy, sheety, calerete-cemented coarse gravel, pale 
brown—().55 m. 
Marly silty sand, pale pinkish buff—.30 m. VL 
satiple FRE S/O9, 
Fine (up lo TO mm) bedded gravel lens. becoming 
coarser lo cast and west. bull—0.10 0. 
Marly silty sand, pale buli—0.200 om, pinehing out 
laterally, Bones ard saxicolous snails within this 
interval. TLisample PRUSSL.S, 
Coarse gravelly sand, angular clasts up to 50 om. 
roughly imbricated, Wghthrown 0.20 m, thickening 
cither side, 
Brown silly clay ono base seen. Pstimaled dept to 
bridge level lio 2m. Th sample URUS)2, | near top 
of this unit (see Fig.3), Height of bridge above 
standing water level about 3 11. 


Age 

Unfortunately, the guartz TL sometimes reaches 
dose saluratign at a relatively low dose level and 
here, the twa deepest samples. PRISALS and 
PRIS/2.) were approaching this saturation, A 
conpsequchue is the relatively large Uncertainty in the 
age oF PRISTL.S. A pilot measurement on PRIS.) 
showed thal ifwas unlikely ta-yield a date for the 
same reason and so dating: was not attempted, Mie 
pilot resulr is consistent with this sample being, the 
oldest of the three. 

Blemental analyses were obtined from feld 


Pasir LoCenpanenrs af tie aac calenkitian and the ages far the We deed capes. 


simple Lab. Code cyutvalent 


Duse (Gy) 


PRIS'O9 AUTLYSOU Sa=4 
PRISTS Ad TE da? hiv 
MRIS 21 


Doses 
(Gy hes) scimt 


LY4 =F 
7 7 TLE 
L753 ud 


Dase-rale ayerwed ue 
(Gry hat) XRS, (kui) 

NAA, alpha 

LASHUDG6 44-5 

VEF LO NY TT 
VL 


LATE PLEISTOCENE DLROTODON PROM THALLL TE COVE i 


“omimae ray seiotillometry for RK, U and Th: by X-ray 
spectrometry (XRS) lor Ks and by thick source alpha 
particle counting (TSAC) for U and Uh The thorium 
concentration was cheeked by feutron aelalon 
analysis (NAA) for PRESHLY and PRISMS Coad 
dercenientaniony (he methods. indicates that, wathin 
the unecerlumlies of measurenrent there ts no 
jadiaaeuye discquilbrium in the samples. 

Table | shows the components of the age 
cleqhition and the tes for the (wo dated samples, 


Discussion and Interpretation 


Comments on Table | 

The equivalent dose and its error ure output (orn a 
slitistical (filling programme. The errors aire 
reluuvely large becouse the inherent varrability ol 
quartz Thome the near dose-saturation of the Tl 
Hake precise curve-ditinge probleniatic. bor 
PRESO9. the Citing programme encountered ny 
difficulties and there was a satistictory dose plateau. 
Saniple PRES/).S was quite close to dose saturation 
ond has.a somewhat lareer uncertainty, 

There are two independent values for the dose rales 
(1) field gamma ray semtilometry and (2) XRS for 
Kk, NAA tor Th, thick seurce alpha counting lor UL 
Ihe agreement between thenr is gratityirg, A 
conlabution from cosmic nrys i included (Prescoll 
& Tulton 1994). Although no equivalent dose (and 
ho aye) was ineasured for FRIST. the dase-rate 
data are mehided for completeness. 

Fhe cero i age is detentyined almost exchisively 
by the uncerliinty in the equivalent! dose, [hie 
equivalent dose and age of PRIS/O.9 are well 
desenhed by the quoted Hgures. For PRIS/1LS the 
dose curve is approaching saturation. Although the 
cron quoled for the equivalent dose is objectively 
found by the lilting procedures, the ermor limits in the 
age areasyimmeiric. This asymmetry Hes within the 
linuits shown in the table which are one standard 
cfrok, AL 9S") eomfideiee, with allowanee tir this 
asymmetry, the ave tes within the interval 42-70 ka. 

Systemic errors jnelude variahility of saree 
content, becuse the dose tate depends on this. tn 
keepiig with Adelaide laboratory praccve. the aye is 
quated for the abserved wiler Content (le) al dry 
wentht for the whole profile), Vow much i may 
Nave waited fy (he past Ps a matter of profession 
judwenent, Porall levels a this sie, a 1!) inerease ip 
Willer resulls ina 1% decrease in dose rate. Vhus, i 
the avenge water coment in the past bad been (. 
Nigher then the dose rate and) the gneasured 
cyuivalent dose would have been lower and the 
present-day ave estimate would be yn low. Cosntie 


HP ASOS HAE US Diets dient domi ob Werth fe delavae) 1 
Vii 


ray variability provides another possible source of 
systemile error bal at this site, ik is ol ne 
consequence. 


Cedlagioul histery 

Soe LOO O00 years age, when world sewleyels 
were much lower and the Gull St Vineent was a 
broad plain draimed by the ancient River Vincent that 
foawved to join the River Murray to the east of the 
future Kangarod Island, the ancestral held River iad 
a steeper gradient. and had cui a ore hack inte the 
face ol the Mt Lofty Ranges. As the sea rose tron 
this lower level, the river's gradient decreased (and 
samlall may also have deereased) and the gorge 
besan Lo silt yp. 

It appears drat. possibly during a local Mash-flood 
some 35 thousand yeurs.uvo. a Dipratuden died andl 
Was. Sweph downstream with other bones that had 
been picked up wore the way. until (he stream 
velovtly dropped and/or the careass reached on 
ephemeral pool where it settled, Sand from the final 
Nush of oud water came to rest On and around the 
bods, which Was not completely buried. The pool 
silted up and exposed bone disintegrated under the 
effects oF the elements (Bebrensmeyer 1978) and 
possibly scavengers. Later: another Hood brought 
layer of gravel, in a process thal wats lo be repeated 
Jor centuries vs the valley eradually tilled with 
sediment 

The present worge/valley was probably ineised i 
the older sediments by 4 rejuvenated stream at the 
heivlt of the last wlacial maximum, when the 
gvadien! wus jguin increased, pr aim the early 
Holoeene. when rainfall inereased, It is possible that 
the very fumbled and beach-rolled isolated 
Diprofoden molar, found in 1977 by nine year-old 
Jonathon Dicker (Anon, 1970)! in beach eravels at 
(he mouth ef the Field River was washed out ar this 
Une. but it is tmore likely that it was uncovered 
during the coud-huilding operation earlier m the 20H 
Century und bull-dozed tite the creek, 10 be carricd 
by Hood-waters te the sen. 

The site of the Dirt bones has suice been 
marked with asmall adirnand plague by the Haller 
Cove Prowress Assouition. 

The uge ol this specimen. as presented here, is 
close to that of the putative arrival of (he first 
Aborteiies in Australia Chhorieees af (999: but see 
Bowler & Mauve 200i) Gillespie & Reberts 2000) 
A bumen factor has been suvvested in the Austraharn 
mepafsunal extinetion (Plamery 1494), either by 
dircet hunting or by environmental modification, and 
corti Srlas, ca, Cuiddie Springs, Horlhwestern New 
South Wales (Field & Dodson 1999), have heen 
claimed fa show evidenee of jrteriction bervcen 
hems und meealiuna, tis hits been challenwed Lor 
Cuddiec Springs (Roberts ef af, 2001). Phe tlitlett 


deb N.S. PLEDGE. J, R, PRESCOTT & J.T. HUTTON 


Cove specimen gives no indication of butchery, nor 
indeed of scavenging, with the remaining bones still 
articulated, [t cannot therefore be used as evidence 
either way. 


Conclusions 


Fossil bones found m Quaternary sediments in the 
bank of the Field River, Hallett Coye, represent the 
partial skeleton of, probably, an immature 
Diprotedon, which was buried in an oyerbank 
deposit of the ancestral stream. Thermoluminescence 
dating of the sediments has given an age of between 
42-70 thousand years before present at the 95'% 


confiderice level, This is close to the proposed date 
(Roberts ef al, 2001) of 46 400 years BP for the 
megafaunal extincuion event in Australia. However, 
there 1s no indication of a human factor involved in 
the death of this animal, 


Acknowledgments 


We thank the following for their help in the field 
and the laboratory; (3. J. Mctlenry. J. A. McNamara. 
G. J. Prideaux, P. Stamatelopoulos and 1) Westlake. 
The project was assisted by a grant to JP from the 
Australian Institute of Nuclear Science and 
Engineering. 


References 


Avreis, M. J, (1985) “Thermoluminescent Dating” 
(Academic Press. london). 

BEHRENSMBYER, A. K. (1978) Taphonomic und ecologic 
information trom bone weathering. Paleobiology 4, 
150-162. 

BouRMAN, RL P. Marrinarits, BP. PRescorp. J. Ro & 
Breperio, A.B (1997) The age of the Pooraka Formation 
dnd its implications, wih some prelinnnary tesults tron 
luminescence dating. Trans. R, Soc. 8. Aust 121, 83-94, 

Bow ek, 1. M. & MaAarr, 2. W. (2000) Redating Australia’s 
oldest human remains: a seeptie’s view... him, Eval. 38, 
719 726. 

Crapper, Heap, J. & Macre. J. (1996) Beyond the 
radiocarbon limit) in’ Australian urchaeology and 
Quaternary research. Antiquity 70, 543-352. 

Fiptp, 1 & Dopson, L (1999) Late Pleistocene megalauna 
and archaeology from Cuddie Springs, Southeastern 
Australia, Prec, Prehist, Sac, 63,275 301, 

Pransery, T. P. (1994) “The Future Eaters: an ecological 
history of the Austealasian lands and people” (Reed, 
Sydney). 


GILL spi, RL & Ropers, R-G. (2000) On the reliability of 


age estimates lor human remains at Lake Mungo, /. fiw. 
Eval, 38.727 732. 

Hurriy. D. J. Hurron, J.T) & PReseort. 1. R. (1993) 
The stranded heach-dune sequence af south-east South 
Australias a test of thermoluminescence dating, 0-800 
ka. Ouett. Sci. Revs 12, 120. 


Prescorr, J. R., Horii, Do & Hurron, J.T. (1993) 
Estimation of equivalent dose in thermoluminescence 
dating—the ‘Australian slide’ method. dreient Th UW. 
1-5. 


& Hurron, J.T. (1994) Cosmic ray 
contibtition to dose rates for luminescence and FSR 
dating, Rudiction Measurements 23, 497-500. 


& MoiAkrasl. B. (1993) Selective Bleach—An 
improved ‘partial bleach* method of finding equivalent 
doses lor thermoluminescence dating of quarte. dacient 


TL WW, 27-30, 


Roareis, R.G.. FLANNERY, T. P.. Aviabib, L. Ik. YOsHiba, 
K., OLteY, J. M., PribkAux, G. J. Lascerr, Ci, M.. 
Baynes, A., Smitil, M. A. Jonbs, Ry & Suir, BLL, 
(2001) New ages for the last Australian Mevalauna: 
conlinent-wide extinction about 46,000 years age, 
Science 292, [SS8-1892, 


Thorn, A., Grom, Ry Mortimer, G,, Spooner, N. A, 
Simpson, J.J. MeCurnoch, M,. Vaytor, L.& Curnor. 
Db. (1999) Australia’s oldest buman remains: uge of the 
Lake Mungo 3 skeleton. J. Ava. Evol 36, 591-612, 


WINTLE. AL G. (1997) Luminescence dating: laboratory 
procedures and protocols, Radiation Measurements 27. 
769 S17. 


ASPECTS OF THE SURVIVAL AND REPRODUCTION OF 
ANGUINA MICROLAENAE (NEMATODA: ANGUINIDAE) 


By PRIMALI DE SILVA* & IAN T. RILEYT 


Summary 


De Silva, P. & Riley, I. T. (2002) Aspects of the survival and reproduction of 
Anguina microlaenae (Nematoda: Anguinidae). Trans. R. Soc. S. Aust. 126(1), 45-49, 
31 May, 2002. 

Leaf galls formed by Anguina microlaenae in Microlaena stipoides were found, upon 
rehydration from natural dessication, to contain adults, eggs and juveniles that had 
survived anhydrobiotically. The sex ratio of adults in galls, excluding a proportion of 
galls that contained only females, was 1:1. Females in galls containing only females 
were apparently sterile as eggs were not present. Rehydrated eggs hatched over a 
temperature range 8-25° C with an optimum of about 20° C. Only limited egg 
production and deposition were observed in rehydrated females incubated after 
removal from their galls. 

Key Words: Nematoda, Anguina microlaenae, dormancy, survival, anhydrobiosis, 
reproduction, sex ratios. 


framsactions af te Raval Society of S. Aust. (2002), 126011, 45-49, 


ASPECTS OF THE SURVIVAL AND REPRODUCTION OF ANGLINA MICROLAENAE 
(NEMATODA: ANGUINIDAE) 


by PRIMALE bE SILVA’ & TANT. RILEY 


Summary 


Hh SILVA, Pode Raby LT (2002) Aspeets af the survival art reprdiicdon.ol), tiga idieralirende (senatody: 
Ansuinidacy. Tram. R Sue 8S dast PRACT), 45-49, 3) Mary, 2002, 

Leal gulls lormed by dagiimenierofaee in Mivrolaens sipotley were found, upan rehydranian from natural 
desiccnon: ty conkun gdulrs. cogs and juveniles that had suyvived aolycrobiatioally, Theses ratio al adults ia 
Halls CX cldiE & preportion al walls that cortained anty females, wis 1, Pamales in galls containing only 
femules wore upparently sterile as Gees were not present. Rebydrated eves latched over u temperanire tange 
B 28° Cowith an optimum of aboue 20° C. Only linited ese production sind deposition were observed in 
rehydried females neubaled aller comoval from their galls, 


Ky Words Nemutodin died diimoedac, dormancy, siirsiea lL anhydrabiosis, ceproduetion, Sex tlio. 


Introduction 


Ariong the Nematoda, members of the tamily 
Anuuinidae have remarkable abilities to survive 
anhydrobiotically (Antoniou 1989), Second slave 
juveniles (J25) of dnedine teinied (Steinbuch 1789) 
Filipjey, 1936 ure known to survive for nore than 30 
vears under dry conditions (Limber 1973), For most 
anguinid rentodes the survival stage is also the 
invasive sluwe and is a second, third or fourth sluge 
juvenile, depending on the species (Chizhow & 
Subbotin 1985). li two leaf gall species, dngnin 
australis Steiner 1940 front Ehrhertes longiflore Sim. 
(Riley ef af 2001) and Amine danthaniae 
(Maggenti ev uf) Breski JOR) (syn. Cynipenienine 
enmhonie Migventi, Hark & Pasinan 1974) fran 
Danthonia californica Bol, (Maggenti et al. 1973), 
the adults are the survival stage. Por these species il 
is nol known if the invasive juvenile stape ean also 
survive anliydrobiotieally. 

Angwined inieralacnen (aweell 1938) Steiner 
1940, a teal gall pematode of the Australian mative 
wruss, VWicroluend stipoides (Labill) R. Bry diflers 
from most anguinid nematodes in that hoth eges and 
125 (aldhoueh 2s were cousidered to be J ts at the 
lime) dre repored ly survive anhydrobiotically 
within senescent walls (Fuweett 1938) Our 
examination Of (he contents of walls formed by A. 
microlaenae, revealed that adults also. survived 
desiccation. Given (his observation und [he limited 
details of the survival of eges provided by Paweett 
(1958). further investigation OF the survival of a. 
microlaenae wes undertaken. The investigation 
included examination of (1) revival of adults and 


Juveniles O04. wrerelaenae following rehydration of 


API UH MIAIeCILiE Reatigy Withe Cyirtin ble LI ee sity 
ob Adak. Glen Osi SA SHEL usta, 
Tlarrespondiie elhiow. Benue anoethey fb aehelaieles eeliaie 


the contents of naturally desicedted walls, (2) 
hatching of rehydrated cygs over to range of 
temperatiires and of different development stages 
aid (3) ce production and deposition in rehydrated 
adulls. 


Materials and Methods 


Source of walls 

Galls formed by al mileralyenae inn Mo stipoiedes 
were obuiined from two sourcess (1) a field 
population collected in September 1999) fron 
Toowoomba, Queensland (27" 34° S$ 157° 57° Vy an 
sfored al room temperature until used for this study 
(hebruary—May, 2001), (2) a cultured) population 
collected in Vebruary 200) from infeeted Az, 
Siipaides groyn in a shade house al the Waite 
Cumpus. Adelaide, South Australia (34° 38° 5 138° 
38" bj. Phe cultured population was established ite 
June 1999 from galls colleeted at the same site in 
Toowoomba thus the two populations were of the 
sime provenance, 


Cantems af galls and. revival of adults and juveniles 


fallawing relweration 


‘Twenty valls each from the field and cultured 
populations were disseeted under water with the aid 
ofa steree microscope. Following incubation lor 24 
Hat 20° C, counts were made of the adult fenmale ane 
male nematodes. eves and 2s aod the viability of 
adulis and juveniles was assessed. Adults and 
juveniles were scored as alive they were turaid and 
emibited movement: some viable but stationary 
individuals may have been exchided andse the count 
Was fH conservative estinnate, 


Meet of temperature and eae development stage on 
huaching of relwdrited ees 
Cyes comamning clearly developed juveniles were 


Ib MDE SUNA & FT, RILEY 


incubated in shallow water (about 2.5 mm deep, 
Adjusted for evaporation diuly) it covered glass 
iiishes at various temperatures and examined daily 
for hatehing over 7 duys. Pive replicates from both 
field aid culties populations of about 25° capes 
per dish were incubated al &, 16, 20, 25, 
31 CL respectively, Observaiions were nol continued 
heyond 7 days due to fungal colomsation. a problem 
iso noled by Faweett (1938), Also, five replioutes of 
about 25 immature ewes were incubated at 20°C for 
7 days and observed cuily, 


hae prodictton dnl deposition Ay rehvelrated cued 
femenles 

Adulis from) freshly dissected galls were placed 
alone Orin pairs in shallow water in covered gloss 
dishes as lollows: (1) a female froma wall without 
males, unpaired (2) a female from a gall with no 
males. paired with w mile, (3) a yout adult lemale 
lroma gall with males. paired with a male und (4) an 
older adult female froma gall with males, paired 
with @ male, Young adult females were relatively 
more active and only slightly curved in comparison 
io older females, which were more obuse, less active 
und spirally coiled, Each combination was repliculed 
ten times und incubated at 20° C for 21 days. The 
females were exupined every 2 days tor ese 
development and depesition, In a separate 
experiment, a further (adult females from galls 
with Inales were incubated alone m Petri dishes on 
LS water agar at 20° C and examined daily for egy 
production and deposition over 10 days. 


Sratiaticed cnalusis 

GENSTAT 5 (Lawes Agricultural Trust, Rothamsted 
Experimental Station) was used for statistical 
analyses. 


Results 


(he contents of the galls from. the two populations 
ape summarised iy fable b. The populations did nat 
differ siatistically U/ lest, 38 db a0 any ateibule other 
than the wumber of juveniles per gall As the 
dist butions ofthe ees. juveniles und tolul progeny 
counts were Abtoorimal, these were transformed (lag 
vt) for analysis, The number of juveniles per pall in 
the field population was less dur iy the cultured 
populition Grieg logierbl) of O70 vo 1.208, 
2.16, 38 dh p > 0,037), indicating that these galls 
may have heen collected at a slightly curlier stage of 
development bul the contents ofthe galls of the two 
populations were otherwise equivalent in 
(ANiLative Lerints. 

Fight galls from the two populations (20% of galls) 
contihed females bul no males (lable 2). There 
were no ces or J2s i these galls, Pxuluding these 


Vann 1 Caen uf leas wally formed shy Amzuina 
nievolacnae i Microlaenmstipwides frome nye sedeees Or 20) 


Coltired Population 
Meanesb (Range) 


Meld Populanon 
Meant SE (Rone) 


females 2,140.29 (1-6) 2400.29 (1-H 


Mules 1.240.235 (+4) 20.37 (0-6) 
Adults 3.21042 (1-8) 4 As().62 (1-12) 
Pemale:Male (720.05 (05-1) O.6-40L0eE (IL-1) 
Bugs ISH (UT 3d) T4418 (1-234) 
hhiveriles 1514.3 (0-58) & |= 321) 


lotal Progeny 30167 (4-792) 185237(0-355) 


Vann 2. Nearer af lve) galls in Miemlaena stipuides willy 
varios combinations af iddtA\rguma microlens 


Females Mules per galt 
per eull a) i a 7 5 6 Holat 
4] + OM - = = 3 
2 ni a 7 - - - \S 
a za - a U | - - 7 
4 - - 3 - = 
35 - ] : | 
hy i] - - | 2 
Totul K 16 9 J 4 | | HM) 


— CONDI aan tat lend 


wally with only females, the proportion ot lemales 
per gall was 0.54 across. both populations, which did 
not diller statistically from an expected ratio of | t. 
Not only was the mean ratio close to 1:1, bul alse all 
galls with both females and niales contuned 
combuinbions close to that ratio (Table 2). 

Regression analysis (exchiding galls wilh only 
femules) did not reveal any signifeant relationships 
between the number of progeny per wall and the 
number of females, mules or total adalis in the gall. 
llowever, sivnificunt negative relationships were 
found belween the number of progeny per adult 
(male, female or total) and the number of females, 
males and adults in the gall. Correlation coelicents 
ranwed Prom -0.408—0.279 and probabilities of the 
regression coefficients from 0.02-0.03. Although 
only explainitig a small proportion of the observed 
variation, these analyses indicate that the fecundity 
of the nematode wis limmled inere by the reaourees 
available within the gall than the number of adults 
present because as the number of adults increased in 
the gall, the number of progeny did not increase 
proportionally, 

All adults dissected from the galls recovered when 
rehydrated, were turgid and moved, albeit relatively 
Jillle, Suintlirly, after tehydration if was estimated 
that 65-75" of the juveniles were alive, exhibiting 
the vigorous movement typical ol invasive stage 
unguiiid juveniles, 


INCHING ATICROL dN SERV IVATL ANT REPRODUCTION 4) 


Percentage hatch 


Time (days) 


fie), Metin accumubitive percentage hatety ol olyeetier 
niprelaenae eees (lite deyelopment-stage) incubated at 
VATOUS temperatures over seven day period (nS), A. 


fues trom leah walls from oa fel) popalatron oof 


Microlawre  Stipejdes collected at Toowoanoa, 
Queensland ( 155). Day 7 = 18). By leeyes trom Ad 
stipeidey grown ih pots ab the Winte Campus. South 
Australne wilh inpeuliin Hom the same Topwoonba site 
(LSD, Day 7 > 12.0). 


100 


80 


60 


40 


Percentage hatch 


20 


5 10 15 20 25 30. 35 
Temperature (° C) 


bie 2.) Mean total percentage Tate of olaeliiies 
AdeVokiGhde Oger (hte development se) meubated at 
yarious temperatures for 7 days (n= 10, LSD, ~ 7b) 


hus cohydrated ata late stage af developoent 
were shown to have the ability to survive 
finhydrobiotically with 60-70% hatehing when 


incubated al 76 ar 20°C. Accumulative hatehime of 


eves over Lhe seven day period lor eaelr temperature 
and population is shown in Mig, | and total hatehing 
mt eavh temperature in Fig, 2. There was no 
statistically sigmifieant diference between ie lwo 
populations abany temperature or tine. Temperature, 
however, had ow marked effect on hatehing. 
Masiiouin meat hatching of 69% occurred ab 20° ¢ 


but this did not differ significantly from the mean 
hatching rate of 62% at 16° C (LSPs, 7.5), At 
higher temperatures hatching was either significantly 
suppressed (24° C) or did not oveur (S12 CY At Re € 
hatching was not observed until the 6th and 7th days 
and then only approached 10%, 

Only limited hatching was observed trom ceys 


rehydrated at an early development stage 
(embryonic) when incubated at 20" C. Hatching 


began aller 5 days and was seen in four ol the live 
replicates reaching 7 1O!% in 7 days, 

Allempts lo observe ca2 development und 
deposition in-adulis removed [ram the galls were 
lurgely unsuccessful. About half the adults were 
colomsed by Hiner (unidentified) over the three week 
period. Only two females, both from the group of 
older females taken from alls with males 
(combination 4), were found to hive deposited: any 
eyes) one a single eve and the other two ees, 
Females incubated on agar were more prone to 
fungal colontsulion with most females being 
colonised within a week. 


Discussion 


Phe study has shown thar adults. eggs and J2s oF 
Lomierafience arable to survive anhydrobiotically. 
Although all adults survived desiccation, survival of 
eyws was only aboul 70% lor those approuching 
maturity and considerably less for mmmature epes 
However, given that some immiiture eges developed 
dnd hatched upon rehydration, itis possible that the 
lower observed survival rate of eres refleeted the 
incubation conditions and fungal colonisation raiher 
than the murinsie survival rate. Similarly. only about 
70% of juveniles appeared to survive desiveation. 
The survival rate might have been vreater iF intaet 
valls had been soaked betore dissection to ellect a 
slower uptake of water. 

Our linding that adults survive desiccation dilers 
trom Faweett’s (1938) observation that adults died 
rupidly when galls became dry. Chis inconsistency 
may be due to the condition of ihe adults at the ime 
of desiccation, If the reserves of the females had 
heen largely exhausted by production of eves, these 
adults might not have been ahle to survive 
desiccation. However Faweert records that the adult 
females hud deposited 150 to 400 cugs each, whieh 
is consistent with our material su this explanation 
seems unlikely. Also, we found tn our material that 
al adults had survived, including those from galls 
containing large numbers of proveny, An alternative 
explanation may lie in ditferences in the rate of 
dryiny: or the storage conditions. Our material either 
dried nathrally as the host plant senesced or wis an 
dried indoors before being stored for up te 15 months 
undet laboratory conditions, This treatment would 


Ie B DE STIIVA KT) S RILEY 


nol purteularty Hevour {he survival of the adults or 
explain the different findings. There is no obvious 
esplanation for these contradictory findings. 

dnenina microlaenae, With its capacity for adults, 
ches and juveniles to survive anhydroblotically, is 
umisnal amongst angzuinid nematodes as most have 
only aosingle survival stage. However muliiple 
survival stages are reported (with Limited detiils) for 
Mesoazuina wasinckia (Steiner & Seow (935) 
Chizhov & Subhotin 1985 and Suhangtin 
radieloole (Greel L872) Paramonoy 1967. 
Womersley (1987) indicates that ll stages of AL 
amisinekia survive anhydrobiotiealy, but cites the 
report oF Pantone & Womersley (1968) which makes 
Ho mention of this behaviour All stages of 4. 
radicidoled are said Lo hibernate’ over wihter in root 
walls (Krall 1991), although this dormaney muy pot 
be anhydrobioss, Mois eoncelyable (hat multiple 
survival stages, as mn wh adcrefaenae, represent a 
trmsitional pattern between  speeies with 
anhydrabiotie juveniles and those with 
wilivdrobiotic adults, However, a recent molecular 
phylogeny of the Anguinidae (8S, Subbotin pers. 
vomm. 2001) provides no support for such o 
proposition, 

‘To be of selective advantage, the survival of adult 
females, which have no further opportunity lo teed, 
should tacilitate comtinucd reproductive activity, Our 
luiure ta demonsiriie any significant cee production 
in adult females removed fron galls and rehydrated 
may he a resull of unsulible experimental 
vonditions, Reproductive activity is more likely (o 
continue in relrydrated intaet galls, as oeeurs in. 
runny (Riley et af 2001). However, given that 
eyes and juveniles are present i highly variable 
numbers belore desiccation, t would be diffieull to 
demonstrate further ege produation. in rehydrated 
ititet wally, 

Similarly, the survival of males points to the 
likelihood of further (asemipation atier revival frou 
anhyvdrobiosis, a behaviour Khow te @eeur in 4 
thinthontoe (Mageenti et af. 1979). Given the 
moderately darge number of eges depusited hy 
anguinid fentales. multiple mating events are likely 
as malonenatades are known ta produce relatively 


Jow nuinbers of sperm (Maggent) 1981). As noted by 
Maweelt (1938), and confirmed by our observation 
hat females in galls without males did nut 
reproduce. 4 aiferolaenae only reproduces sexuilly, 
so Survival OF fates is consistentywith the possibility 
of reproductive activily lollowing dormaney, 

the other notable tinding is the range af 
femperiture for hatehing. Although if appears that 
hatchifi is favoured by temperatures of about 
16 20° Cy halehing of do anorelacne occurred 
Gutside (his range. I contrast, reproduction aff. 
enistalls only occurred at abort 20° C although the 
lemperalure requirements for hatch were hot 
separately determined (Riley ef ef 2001), deine 
microlucnae has been found tn sites fram. the 
lemperate uliinate ol Vietoria Wvaweell 1938) wilt 
Winter-dominant rainfall, through to the subtropical 
climate of southern Queensland with summer- 
dominant raintall (tis shudy), Tn contrast, 4, 
australis. is only known trom the Mediterranean 
climatic vone ol Western Australia. The brouder 
lemperature response Olt, microfacnae 18 Consistent 
with ils wider distribution, 

The oeeurrence of galls containing only females 
and the absence of galls contaiminy only tales dues 
hat appear to have been due solely to ehanee, as the 
sume was lound for ve caeadrediy (Riley 2001), Th ts 
possible that this oveurrence is indivative af a 
invehanism such us pally only being inituted by 
venctigully female J2s or an environmental ser 
determination, 

Further study ofthe survival of immature egos and 
post-dormaney reprudueuve behave abl. 
mieroldende is needed, bul given he constennts 
imposed by fubeus associated with the galls, 
progress in Unis urea may require using a fungicide 
with mo tox or physivlowiwal effeet un the 
nematode. 


Acknowledgments 


Drs K. Owen and Po Wallimson are Unaimked for 
collecting infected plant material front booweombu, 
The salary of 1. 1 Riley is in part finde by the 
Graitis Researeh and Develupment Corperaution. 


References 


ASCHOSINIL NE ChO88) Arrested levelopment ra plat 
parasitic numuloades, Helovnaihal, thy, (Ser 2) SR, 1-19, 

Cran YN, de Shinponin, S.A. (L987) Revision uf the 
nenmiode sublmily Anguininag (Nematoda, lylen- 
chida) on the bastssof ther bivlogical charaeterisne pp. 
SIR da Fortunen Ry 
sulveied Whenimic papers in nenmtology Valo 
(Cahfarmu Gepariment of Mood ond Agricul. 
Sacraiienita CA, LSA). 


(Hdl) “Hwthish tratislanians. of 


Pawer i S.C Mo (19348) A disease al the \isryilian aross 
MWeraluengy stipaides Ro Ue cnused hy a nemo 
Jliuiiolind iniondaentene 1. sp. £ Hedminl. Vo. 197-32, 


Kisii bb C8) Wheal and peass nenutodess olgidi, 
Subungiina, aid veld gener pp. 721-760 Jy Nickel, 
WOR (hak) *Maneal ar Agriculinetl Merits” 
(Mureel Dekker New York. USA), 


ANGUINA MICROLAENAE SURVIVAL AND REPRODUCTION 49 


LimBer, D. P. (1973) Notes on the longevity of Anguina 
tritici (Steinbuch, 1799) Filipjev, 1936, and its ability to 


invade wheat seedlings after thirty two years of 


dormancy. Proc. Helminthol. Soc. Wash. 40, 272-274. 

Macaenti, A. R., HART, W. H. & PAxman, G. A. (1973) A 
new genus of gall forming nematode from Danthonia 
californica with a discussion of its life history. 
Nematologica 19, 491-477. 

—s (1981) “General Nematology” 
Verlag, New York, USA). 

PANTONE D. J. & WOMERSLEY C. (1986) The distribution of 
flower galls caused by Anguina amsinckiae on the weed, 
common fiddleneck, Amsinckia intermedia. Revue 
Nématol, 9, 185-189. 


(Springer- 


Ritky, 1. T., SHEDLEY, D. & SIVASITHAMPARAM, K. (2001) 
anhydrobiosis and reproduction in Anguina australis, 
Australas. Pl. Path. 30, 361-364. 


TRIANTAPHYLLOU, A. C. & HIRSCHMANN, H. (1966) 
gametogenesis and reproduction in the wheat nematode, 
Anguina tritici. Nematalogica, 12, 437-442. 


WoMERSLEY, C. (1987) A reevaulation of strategies 
employed by nematode anhydrobiotes in relation to their 
natural environment pp. 165-173 Jn Veech, J. A. & 
Dickson, D. W. (Eds) “Vistas on nematology: A 
commemoration of the twenty-five anniversary of the 
Society of Nematologists” (Society of Nematologists, 
Hyattsville, Maryland, USA). 


VOL. 126, PART 2 
29 NOVEMBER, 2002 


Transactions of the 
Royal Society of South 
Australia 


Incorporated 


Contents 


Spratt, D. M. & Nicholas, W. L. Morphological evidence for the systematic 
position of the Order Muspiceida (Nematoda). - - -—- 
Shattuck, S. O. & McArthur, A. J. A taxonomic revision of the Camponotus 
wiederkehri and perjurus species-groups (Hymenoptera: 
Formicidae)— -— — - - 
Dutkiewicz, A. & von der Borch, C. C. Stratigraphy of the Lake Malata Playa 
Basin, South Australia = _ 
Dutkiewicz, A., von der Borch, C. C. & Prescott, J. R. Geomorphology of the 
Lake Malata-Lake Greenly complex, South Australia, and 
its implications for late Quaternary palaeoclimate— —- — 
Styan, C. A. & Strzelecki, J. Small scale spatial distribution patterns and 
monitoring strategies for the introduced marine worm, 
Sabella spallanzanii (Polychaeta: Sabellidae)- - - — 


Brief Communication 
Taylor, D. J. First records of two families of freshwater Amphipoda 
(Corophiidae, Perthiidae) from South Australia~ -— -— — 


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


51 


63 


91 


103 


117 


125 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 126, PART 2 


MORPHOLOGICAL EVIDENCE FOR THE SYSTEMATIC 
POSITION OF THE ORDER MUSPICEIDA (NEMATODA) 


By DAVID M. SPRATT* & WARWICK L. NICHOLAST 


Summary 


Spratt, D. M. & Nicholas, W. L. (2002). Morphological evidence for the systematic 
position of the Order Muspiceida (Nematoda). Trans. R. Soc. S. Aust. 126(2), 51-62, 
29 November, 2002. 

Muspiceida are tiny, highly specialised nematodes parasitic as adults in the 
connective and organ tissues of vertebrates. Nine genera, six monotypic, are 
recognised in two familes. Life cycles are unknown but modes of transmission have 
been widely discussed in the literature and postulated as occurring by cannibalism, 
cutaneous penetration or during lactation or grooming. The phylogenetic affinities and 
systematic rank of the Muspiceida have long been in doubt, some morphological 
features suggesting similarities with the Secernentea, others suggesting similarities 
with the Adenophorea. 

Key Words: Muspiceida, Muspicea borreli, morphology, SEM, TEM, Dorylaimia, 
Adenophorea, Enoplea. 


Transactions af the Reval Soeiedy ofS. Aast (2002) 12602) 54-02, 


MORPHOLOGICAL EVIDENCE FOR THE SYSTEMATIC POSITION 
OF THE ORDER MUSPICEIDA (NEMATODA) 


by Davin M. Sprart & Warwick L. NICHOLAS’ 


Summary 


Spray DL M. & Maicnotas WoL, (2002) Morpholowical evidence lor the systemuue position of the Order 
Muspiceida (Nematoda), Tras, Ro Sue, 8) Ass, 126(2). 51-62. 29 November 2002, 

Muspicelda are tiny, highly specialised nematodes parasitic as adulis iy the connective and organ tissues al 
vertebrates. Nine genens, six moneryple, are recownised in two Hinilies. Life eyeles are Unknawe but modes of 
transmission have been widely disvussed) in (he Hleratore and postulated as occurring by cannibalism, culineous 
pencimabon or during lactation or grooming. The phylogenetic allinivies and systematic rank of the Muspieeida 
have lone been in doubl, sone morphological leanires suggesting similarities with the Secernentea, others 
suggesting similarities with the Adenophorea, Morphological siidies usiqy light iiierosenpy. SIEM ail TEM at 
Masnece bore’ Samben 1925 from wild Was domevticns Sehwarte. & Seliwarte 1943 (see Figueroa ev a. 
1986) in Australia sirengthen the killer proposition Although an ofichiostyle Was Holabserved in larval ar adult 
AL barrel, evidence indicues that plasmids are absent in bis species. What had previously been interpreted as 
“phasmidoul cells” in this and twe other speetes wm avhich both adult and larval forms were stidied are in fact 
erudal wands, terminal in adults and sub-ternmiinal in third-shige larvae, with no-associated nervous tissue This 
Hadi. togerher with previous evidence from ather dithors strengthens the view (hat the Muspicerda provide a 
link between the Mermithida and the Trichweephalida ( triebinetlda), dnd on morphological grounds are 
adenophorean, not secermentean. This conclusion aceords with resent studies of phylogenetie relationships 
williithe Nemiutoda and a re-elassifearion of the phylum based on morphological and Jif) cyele Know ledge are 
maleeular data. in particular SSU rDNA sequences [rom animal and plant parusitie, and fFee-living waa, I 
geeonls With dhe placement of the vertebrae parasilie Muspiecida. Droctophymatida and ‘Trichinellida 
(-Trvhocephalida) alonesie the jaseet parasive Mermithida and) Marimernvthida, the phunt-parasitie 
Dorviwimide and (he free-lying Mononehida tithe subeluss Dorylainia. oF ie cliss Choplea, 


Key Words: Muspiveida. Wiynieca horned. morphnlow. SEM, TEM. Dorylaimia, Adenophorea, Pnoptea. 


Introduction 


Phe nematode order Muspiceida Bain & Chabaud, 
1959 18.00 entivimatic eroup of parasites occurring 10 
the skin, eyes, brains or vascular system of 
vertebrates (Sprae et af 1999). Its phylogenetic 
posttion within the phylunt Nematoda has always 
been uncertain. a sitiation which has been further 
exposed by significant advances in understanding of 
petitlode phyloweny, 

Classification within the phyliim Nematoda has 
been, until recently, based upon morphological and 
ccolouical features associated with phenotypic 
characters of free-living or parasigne nemutodes 
(Chitwood 1933. 1950: Chabaud 1974: Anderson 
1984, 2000: Anderson & Bai 1982: Anderson e/ a/. 
1974; Invirs 1083), Ushie these eriteria, the 
overwhelming majorny (estimated 92%) of 
nenatode parasites of vertebrates belony fo the class 
Sucermented und are thought ro have arisen from 
rhabditel, free-living. micrabivoraus. sui nematodes 


Lmitih lase spraiviresind au 

CSIBG!) Suabimible Beosystent CPO Boy 244 ¢ uoherea ACT 
THO ANttsteatid 

“Departmental Lokiny sind Anoloyy. aud Me Bleetian Mieco cpus 
\Jrt Auatiatiait Nuttonal Lenrvurstay, PA Phas yt ntreer ACT 
ONO AN teite ti 


(Anderson 1984). With very few exceptions, the 
third stawe larva is inlective ta the detiniive host 
(Anderson dace. cl). Under untavourable conditions, 
free-living chabditids produce “dauer” larvae. third- 
Stage larvae ensheathed in the cuuele af the second- 
stage. These can be indu¢ed to exsheath und resume 
development in the presenee of Ipod. 

A oiminority of nenatede parasites af vertebrates 
belongs to the elass Adenophorea, However the 
evolunonary origins of the Adenophorea, i 
contrast to the Secernentea, remdin controversval 
(Mayyenti 19835 Bain & Chabaud 1968, 19749: 


Inglis T9832 Anderson 1984: Adamson [O4A) 
beouuse many wuts reconnise its lack of 


monophyly or ils homoaxeneity campared with the 
Secernentea. The adenophoreans parasitic in 
Vertebrates. ure believed to have arisen from a 
dorylalin liteane (seas Chitwood L980) with a 
speur-like stylet in the buccal eayity Conehiosty le). 
most of which are predaceaus or plant-parasitie and 
adapted to piereing and feeding on lissues (Anderson 
J984), This may have given them a capacity to cross 
fissuc barriers by pencuaton and my possibly have 
pre-udapted them toa parasitic way of file (Pitlleborn 
1929) Pour parasitic groups (reeoenised as 
subonlers or suiperlamilles) are tiodeht to be derived 
irom the darylaims, three in vertebrates, Triehinellina 
(-Trichinelloidea), Dioclophymatina (- Didctophymataides) 


§1 OM SPRATT & W I 


and Muspievina (=Muspieeoidea), and one in 
inverlebrales, primarily ansects. Mermithing 
( Mernithoidea) (Anderson 1984, 2000), Wilt the 
exception of the Muspiceina. (he affinities of the 
parasite genera found in these suborders are 
exemplified by the presence. as m free-living 
dorviaims, of an onchiostyle ty first-stage larviie. In 
conirasl fo the Secernenten,  adenophorean 
nematodes do nol havea “daver™ larva aod infeet the 
host inthe Grst as well as the third or fourth larval 
stive. Trond the oaytset of parasitisn, the 
Adenophorea were probably adapted ta teed on 
tissues. hence ihe hesi's intestinal environment was 
probably never a trophic necessity (Pulleborn 1074: 
Anderson 184: Adamson 1986), Althouvly the 
alimentary tract may provide an outlet Tor 
transmission stages im species of some geners 
(Tnehinellina: Capilfarioa, Teehuriy), this linetion 
may be filled hy the urinary system (Triehinelina: 
Trickesomegdes, Dioctophy mating Dioctaplvin) ar 
possibly the skin (Muspicema: MWispicea) in orher 
wenen, ‘Lhere are no lamen-dwelling. titestinmal. 
parasitic Enoplea (Adamson L986). Wright (1989) 
demonstrated that aistrojmtestinal jnhabitants among 
the Trichinellina are in fact inlimately associated 
with epithelial lissues. Modes of transmission and 
postemmbryanic development of these nenarades 
raise the possibility that the entire grovp arose as 
aaspeliles ol earthworms and terrestrial especially 
Jurval. Toseets. Ik was Ihese early assbetates which 
diversified and pave rise to the parusites i both 
invertebrates and vertebrates (Anderson 1984). 
Recently, — phylovenctic 
udenophorean (13) and secernentean (13) nematodes 
have heey assessed using, marpholovical datu snd 
SSU rDNA sequenee data (Kampler ez ad (998). 
Both dala seis strongly supported the classic split 
into Adenopharea and Seeernented, recognizing cach 
as monophyletic, However. subsequent analyses ala 
datthake of SSU TDNA sequences trom 
representatives (53) of enimal parasitic. phint 
parasitic and {vee living dasa resulted ia markedty 
differene vonclustan (Blaster ed a/. 1998). The fatter 
iuntalyses indyated that vonvergent morpholowcal 
evoligion was commen and that the Adenophoren 
tity hé paraphyleue becuuse a includes the aneestors 
ol the Secvernentea, live clades were recoynised. all 
ofowhiel ineluded parasitic species. Dorris ur at 
(1499) stipeested thar aninal parasitism) arose 
independently atleast six Himes and plank parasitism 
dl least three times. Two esclosively udenophoredn 
clades were strongly supported. In particular, Clade | 
urouped the vertebrate parasitic order “Trighye 
cephialida (= Tichimellina oF Andersen 2000) with 
the inseci-parasifie Mermithida, plant-parasitie 
Dorvlatmida ane free-living Mononchida. 
Subsequently. De Ley Ae Blaater (2002) presenied 


relationships oot 


NICHOGLAS 


4 comprehensive yel appropriately conservative 
Treatment of the systeniahe position and pliylogeny 
of the Nematodu based om the overall congruenee 
between morphological amd moleeulir phyloweneuic 
analyses, notwithstanding the fact that maleeular 
sampling oof (asxonomie diversity within the 
Nematoda remains limited. especally within 
Adenophorea relative ta Secerventen, 

This seminal work incorporates a tajor shila 
balance in an effort to combine parasitic and non- 
parasitic iw within a single phylogenetic lnerarclys: 
They argue that sueh a balanee stands on 
inorplological evidenee alone. following inesitably 
from the eombipaion af two varies hypotheses. 
propased on the basis of morphology (Lorenaen 
sa. 1994) and dite eyele data (Inwlis 1983: 
Anderson IS4). These hypotheses. were the 
pataphyly of the Adenophorea with respeet to 
Sceernentes and the assuiyptloy thst all parasitic 
nematode taxa derived from free-living ancestors. 
Both hypotheses ure now supported stroayly hy 
DNA sequence analysis, placing the origin af the 
Nematoda sarmewhere between chiomadorids, 
enoplids and donylatmuls (De Ley & Blaster 2002), 

The phylogenetic affinities and systemarie pink of 
the Muspiceida have long bees in doubt (Anderson 
21100). The presenee of “phasmid-like™ strictres vy 
(hree of the genera ih whieh the larval stage has been 
deseribed (Bain & Chabaud (979) would place the 
Muspiveida inthe Secermentea, Other morpholovics! 
feulures would place the Muspieeida in the 
Adenophorea, a systematic group whose valatity 
itsell has been questioned, a8 outlined above, As in 
adult Mermithida. the intestine is replaced by a 
Trophosome bul pharynageal ghind structures suggest 
aflimiies with the Trehocephulida (=Trehinel fide). 
The amphids have not beet deseribed. |t) cotrust Wy 
secernemteans, udenophoreans usually have caudal 
and epidenval wands. lack phasmids. have a sinyle- 
cull secrelory-exerciury System Usually with a bon- 
curtcularised terminal duet have well-developed, 
usually post-libial amphids. commonty fave 
cephalic and somatic setae anitare found mostly ir 
aquatic enyirohinents (Bird & Bird 1991). ‘The 
epidermal glands are unicellular siruetures locales in 
the talerul cords and open through pores in the 
culicle. Lach is associated witha bipolur nerve cell 
and constitures @ Neuroseeretory wit of unkpows 
funvoon, They are highly susceptible Lo ietomeily 
(OMS, unpub.) and so they may have an imporunt 
role in watey-electralyte balariee, The caticlat 
seeretory glands are usually Uiree in number snd may 
Qpen through a spinneret, a valve siruectune ql the lip 
othe tail thought to be used for attachment to the 
Substhituun in-aquane forms. Placement of the arder 
Muspieeida gear the Trichoeephatida was 
strengthened with the finding by Spratt er af (1v99y 


SYSTEMALIC POSTION OF MUSPICFIDA (NEMATODA} ne 


of Jaterul bacilary bands in immature femele 
Hayeowknenue perplexun Spratt, Beveridge. 
Andrews & Dennell, 1999 In this character. the 
Muspiceida appears to provide a link belween Lhe 
Merinithida and the Tnichocephalida. tts position 
would be strenethened further if a true oneliostyle 
(i.e, it alleredt sub-venteal tooth atid nota formation 
of the ventral wall of the buceal cavity - see 
Lorenzen 1983) were present in the adulls or larval 


slaves of at lest Some members of the group and if 


clear vyidenee of the absence of phasmids was 
found. 

Muypivea barred’ Sambon. 1925 was deseribed by 
Sambar (1925) based on “la (iluire de Borrel” first 
reported by Borrell (1910). The parasite occurred 
malurally in the subcutaneous connective tissues, 
inguinal wid mediastinal lymphatics and in and 
around canverous tumours of mammary glands of 
wild populations of Wis #useu/is Linnaeus, 1758 
from Strasbourg, Vrance, Hewas also found in mice 
from London and Germany (see Brumpt 1939). 
Surprisingly, no new reeords of the purasite occurred 
from the work Gf Brampt (1930). who erected the 
fatnily Muspieejdae for the nematode, until it was 
reported in weld) Touse duce, re deatenticws, m1 
Australia (Sitvleton, L983. Singleton and Redhead 
POM). 

This report provides further merphological 
evidenee fp the systematic posiien of the 
Muspicwida derived tron) hyht, scanoing and 


(raisnussion clectron microscopical studies af 


Maspieea borrede trom wilh Muy domeséers 
Australia. 


Materials and Methods 


Wild Mey domeviions Were trapped at Canberra 
Mice were anaesthetised with ether vapour and killed 
by cervical disloalion. Carcases were skinned and 
skins aid) carcuses washed in Llank’s Balanced Salt 
Soliton (HBSS )an a Peter dish. venthy scraped with 
a blunt razay blade and rewashed) The pelt was cut 
{ite 5 pieces and suspended in HBSS nna Bacrinann 
apparatus (Thtenpont era. 1979) at 37°C for 3h. The 
immersed shin was auitiled approximately every 15 
min Wilh a glass stitver or forceps, Fluid) was 
collveted from the Baermann finde! inte 4 test ribe 
at 15-30 minute intervals. the tube allowed to stand 
for several minutes, Lhe supernatanr renroved und the 
sediienn was thet removed and examined for AL 
borreti ina Petri dish, In mice ktowar to be iifected 
in the subcutaneous lssues, the liver, spleen, 
lymphatics, lungs, heart, brain, mammary glands, 
eetiitals, congue and lips were teased apart in BBS. 
alluwed to stand ina test the st 47°C! for} band the 
sediment re-esuinined for vematudes. 

Por hight microscopy, same nematudes were lixed 


in 5% formal saline then Wwansferred to Si. aqueons 
glycerol which was ullowed to evaporate to 
anhydrous vlycerol in a desiccator. They were then 
mounted on vlass slides i anhydrous. glycerol with 
the cover slip supported by glass beads oF the 
apprupnate sive. and the cover slip ringed with 
Glycvel (Gurr, LR). Other specimens were fixed 
overpight in Berland’s fixative (95% by volume 
Mlacial acetic acid, 5% concentrated formalin), 
transferred to 70% ethanol and cleared in 
laclophenol for microscopic examination. Live 
species were stained with L% toluidine blue and 
with Uy acetic orcei prior to microscopic 
examination, 

For Scanning Eleetron Microscopy (SEM), 
lormalin fixed specimens were Washed in saline then 
post-lixed in)" osmium tetroxide for | he washed in 
distilled water. freeze-dtied, mounted on metal stubs 
using nail varmsh as glue and coated wath yok 
palladium ander vacuurn, 

For Transmission Electron Microscopy Clb), 
specimens were fixed overnight in eold 2.5! 
Wuteraldehyde in phosphate butler, pH 71 
conning 3) sucrose, then post-lived in tty 
gainiuny tetroxide for 1 oh, Specimens were 
progressively trunslerred through wraded ethanols 
and ¢paxypropane to Spurr epoxy resin. Alter 
hardening the resin at 60° Co for 48h. thin seetious 
Were cut, mounted on finnyvar cwated slot gids, and 
stained wilh 6"q aqueous urahyl aectare and 
Revnolds bead eptrale, Two specimens were used lor 
transverse sections ond nyo tor longitudinal seetions 


Results 


Morphology of cedide (Pits 1-9, | 2-24) 

AIL M. hoeredé Fron wilbiniee were recovered from 
subcimmeous tisstles. Adults were 1200-2300 jum 
long and 23-270 fn wide witha blunt cephalic end 
tapering postenorly to a bilid caudal end (Migs 1-5, 
8-9), The ota) opening was small (1.25 jam in 
chameter). sub-terminal, disphiced ventrally and 
surrounded by 6 Small sensory eephalte papillae 
Niere interpreted as 6 inner labial sensilla (Figs 2. 14. 
16-17). Sixteen Jong cells with farge nuecler wand 
nucleoli extended fron the cephalic end to the verve 
ring (Figs 3, 4). The 6 inner tabral sensilla pissed 
through the citicle to the exterion However, external 
openings wee not abserved forthe outer lateral and 
cephalic sensilla. A pair of povket-lke averal 
amphids which stun deeply with vital dyes was 
observed under light microscopy but appeared, under 
Jight und scanuing electron micrascapy, to have no 
cuticular ducts opentm ta the exteror (bis 4. 10) 
A yvaniable namber of cuticular cephatic nodules was 
present wr some specimens (Fiz. 17). Conspicuous 
transverse cuticular sttiae or unnalations were 


ut 
+ 


D.M. SPRAPT & W. L. NICHOLAS 


co 


= 


wn 
on 
peaeBanis 3 


<x! 


Son eR20e © 


| | 


Figs le UL. Muspicea borreli trom tssues of Muy domesticus. Vig. 1. Anterior end showing pharynx (ph), thin sand (st) of 
connechve Lissue al commencement of trophosome (ts) which contains large nuclei and numerous lipid globules, nerve 
ting (nr). lwo pairs of latero-venteal pharyngeal glands (vg) and ovi-testis (ot) containing unfertilised eggs, latero-ventral 
view. Fig. 2. Cephalic end showing cutivular (cu) lining of distal end of pharynx (ph) leading to yentral oral opening, 
lateral view (dorsal on right side). Pig. 3. Cephalic cad showing pharynx (ph), suspected sheath cells (se) of eight cephalic 
papillae and amphidial pouch (ap), lateral view. Fig. 4. Optical section of same showing pharynx (ph), suspected sheath 
cells (se) of other eight cephalic papillae and other amphidial poueh (ap), lateral view. Fig. 5. Optical section of cephalic 
end showing amphidial pouches (ap) and conspicuous nuclei and nucleoli of cells of lateral cord (co), ventral view. 
Fiz. 6. Nuclear pattern af lateral cord (ca) in mid-body region, lateral view. Fig. 7. Nuclear pattern of lateral cord (co) at 
postertor end. Fig, &. Caudal end showing bifid form, pair of large terminal caudal glands (cg) and ussociated nuctei (nu), 
ventral view, Fig. 9. Postenor end showing termination of waphosome (ts) in thin strand (st) of connective tissue and pair 
of caudal glands (eg). Pig. 10. Anterior end third-stage larva showing sub-terminal, ventral oral opening and lone 
cuticular lining of pharynx (ph), lateral view, Fig. 11, Caudal end of third-stage larva showing conspicuous caudal glands 
(cg) emptying laterally, yentral view, Seale bars: Figs 1-10 = 50 pum; Fig, TE = 20 pm, 


Ww 
Ww 


SYSTEMATIC POSITION OF MUSPICEIDA (NEMATODA) 


Pigs 12-17. Adult Muspicea borreli - Nomarski interference micrographs and scanning electron micrographs. Fig. 12. Ovi 
testis (ot) containing densely packed nuclei of developing oocytes and uterus (ut), lateral view. Fig. 13. Optical section 
of same showing uterus containing sperm (s), lateral view, Fig. 14. Anterior end showing pharynx (ph), trophosome (ts), 
one of anterior ventral glands (vg) with single nucleus (arrowhead) and duet (d) from gland emptying into pharynx, lateral 
view. Fig, 15, Caudal extremity showing one of two large caudal glands (cg) and opening to exterior (arrow) in one lobe 
of bilobed tail. Fig. 16. Cephalic region illustrating oral orifice (0) located ventrally and six sub-median papillae 
(arrowheads), en face view, Fig. 17, Cephalic region showing oral orifice (o), conspicuous cuticular nodules (n) and 
transverse cuticular annuli, dorsal view. Scale bars: Figs 12, 13 = 150 pm; Fig, 14 = 100 jum; Pig. 15 = 50 pum; Fig, 16 

LO wim; Fig. 17 = 20 pm. 


a BoM RPRATT & WoL NICHIONAS 


18 


1 ec 


SYSTEMATIC POSTION OF MUSPICEIDA (NEM ATODIAS al 


present throuwhout the length of the adult nematode, 
and occasionally fused with one another (Pigs: 1-2, 
$-4. 15-17). The lateral cords were wider ancl more 
conspicuous that the dorsal and ventral cords und 
had many nuclei with conspicuous nucleoli arranged 
resularly ina simvle row throuyhout the length al the 
adull (Figs 5-7). The cuticle comprised a narrow 
epledticle and thieker coral median and bitsul 
voles (Fis Th, 19). The ventral epidermis was in 
eontuct willl the endo-cuyticle and formed a diffuse 
band of electron-dense membranes arranged radially 
in-shicks, greatly increasing the surface area of the 
epidermal cell membrane in associlion with the 
endo-etiticle (Fig. 18). The cytoplasm of the ventral 
epidermis also contained ghindular tissue with 
vranitles oF protein and lipid (Figs 18-19). 


A tre onchiostle was not observed in adult or 


lunvill forms OF AL horreli The vestigial pharyns was 
short, Fibrous anid hal a narrow lumen lined with 
cuticle for only a short distance (Pigs 1-3, 22) The 
pharyns contained gland cells (Pig. 21), extending as 
anuvow strat oF tissue from the cephalic end wbour 
150.200 jim inte the pseudocdele and terminating 
close 16 the attachment of the ventral wlands, neer the 
level oF Ihe nerve ring (Pigs 1, 3. 4, 14). The nerve 
ring oevurred near the junction of the veorral 
pharyviigesd! vlonds and the pharynx (Pies 1. 3.4). .4 
pair of larae, dense, distinetly bilobed, latero-ventrdl 
pharyngeul ghandswas suspended trom the posterior 
portion of the pharyny by two fine ducts and «4 
smaller pairlay tomiediately posteriorly (bres bl. 14). 
Large slellale coelomocytes each with 2 voluminous 
Wueleus, lirge surface area, convoluted membranes 
and invagindled lamellae occurred throughout the 
pseudoceele (Vig, 22), The trophoseme extendec 
almost the entire length of the pscudoeoele ay uo 
suing of large cells with prominent nuclei and 
yucheoll (Fis |. )4 23, 24). Havas conneeted to the 
pharyns at the anterior end and the bady wall al the 
posterior end by thin strands of conneetive tissues 


(Migs 1,9), The charaeterstic granular appearanve of 


the: (ophasome was due to (he presence of [pid 
yaetoles (hig, 24), The yulvaowas atrophied and no 
conection was detected belween the non-linelonal 
plwevesx and the uferus, The tail of the adult wees 
bifiireate and each tip terminated in a caudal elena 
witha pare open externally (Fis & O15). The 
wlinds stated deeply with both. 1% toluidine blue 
and woth }o dectic oreein. Five prominent nucle! 


with conspicuous nucleolt were associated with the 
vlinds (Fig. &). They lacked the bere cell 
commections associaled wilh phasmids and they 
vpened ‘to the exterior Weaspicea hurreli was a 
protandrous hermaphrodite: — spermatovenesis 
vevurred ji the wall of the genital pouch and 


spermalozna were released from the ovi-tesles into 


(he ulertis which ucted as a receptacle for sell 
fertilization (Fie. 13), Spermatogenesis terminated 
When the distal cells of the genital cord conunenced 
division for ovovenesis (Vi, 12), Larvae developed 
simultaneously inthe uterine pouch (Fig. 24). 


Morphology of Vived-staue larva d Figs W-|), 25-30) 

Larvae developed io the third-staye inthe ulertus of 
ihe female, at which time they were 295310 Lin 
lony and 16-17 wim wide (Fips 10, 11). Three body 
eunicles were detectable in living larvae escaping 
Irom female worms (Piz, 25): the inner and outer 
were smooth, the middle cuticle possessed 
conspicuous tansverse striae. The onal openite wits 
subterminal and ventral (Fig. 10) A conspicuous 
cuticular pharyngeal [iiihe extended posteriorly 
1QU-T100 jim owhere ib appeared to become 
surmounded by an intestinal-like  strueture (° 
trophosome) (Pig. 10) whieh extended to within 20- 
25 pin of the caudal end. A slender lumen and 
secyelory ehuiules were evident in this slracture 
(Pies 26, 27). An arius wits mol delecied, A pair of 
conspicuous caudal glands consisting af highly 
convoluted slandular nissue which stated deeply 
with vital dyes, emptied laterally about 20 jum fear 
the caudal end, whieh terniitated i two bippets 
(Migs 1], 26-30), 


Discussion 


The Muspiceida are tiny, highly speciulised 
nematodes parasitic as adults in the Hssues of 
vertebrates, most of them inhabinny the 
subcunmeous tissues of ther hosts GAndersoan & 
Bain JY82), Nine genera, Sik Mmonotypre, are 
regornised un yer amtlies, One, lhe Muspicendae 
Sdmbon. 1925. contains the genera Miapieeu 
Sanmibon, 1925 from the subcutineous issues of mice 
(Mus spp.oand Rinwsevolvee Bain & Chabaud. 
LOGS. Lokenente Chabaud & Bain, LO74. Penitinta 
Bain & Chabaud, 1979 and Maser Rausel & 
Rausel. TYSS from the subeutyneois tissues af the 


Hines P8230 Transdvissian electron ficromruphs af adoll Mispicee Avews) big be. Lonuitudipal section Hhrouah cobele. 
epideniis dhe miiscke, Fite 14. Transverse section through sume Mssues. biz. 20, Tansverse seetian thiauel pharynues| 


thins. fig. 21 Transverse section thraugh pharym, Vig. 22. Caelomacyte, Pik. 23. Tenphosoiie, Seale bars 


I tite bre 


cuticle bristil owe cl doehamocvte. ee cuticle eorticul Yobe. ve epieuliele. em embrye, hd hypedermis. len lonitadinal 
Husele, Ip Typed melusien. We cubele Med zune nu kaw, pe phuryngeslgtund, ph phuryax. pl pharyngeal lumen. 


Jyh HONE Ut Liters 


58 D.M, SPRATT & W, L. NICHOLAS 


Vigs 24 - 27. Transmission electron micrographs of adult and larval Muspicea borreli. Vig. 24. Transverse section through 
adult female, Fig. 25. Anterior end of juvenile from subcutaneous tissues of mouse. Scale bar = 5 jim, Fig. 26. Embryo 
in-utero. Svale bar = | pum. Pig. 27. Secretory granules in embryonic intestine, Scale bars: Fig. 24 = 20 jim; Fig. 25 = 5 
pm; Figs 26, 27 = 1 pum. cl coelomocyte, Cl outer cuticle, C2 middle and inner cuticle, em embryo, il intestinal lumen, 
Im longitudinal musele, pg pharyngeal gland, ph pharynx, ps pscudocoel, sg secretory granules in embryonic intestine, 
ts trophosome, ut uterus. 


wings and feet of bats. The second family, the 
Robertdollfusidae Chabaud & Campana, 1950, 
contains the genera Robertdollfusa Chabaud & 
Campana, 1950 from the eye of corvids and the brain 
of falconids, Durikainema Spratt & Speare, 1982 
from the portal and intracardiac veins and epicardial 
lymphatics of kangaroos and wallabies and the 
pulmonary arteries of koalas and brush-tail possums, 


Lappnema Bain & Nikander, 1982 from the 
subcutaneous capillaries of the ears of reindeer and 
Haycocknema Spratt, Beveridge, Andrews & 
Dennett, 1999 from the myofibres of humans. In 
addition, larvae of a presumed muspiceid are known 
from white-tailed deer (Beaver & Burgdorfer 1984, 
1987) and infective larvae of a presumed new 
species of Robertdollfusidae are known from the gut 


SYSTEMATIC POSITION OF MUSPICEIDA (NEMATODA) 39 


Figs 28 - 30. Transmission electron micrographs of larval Muspicea borreli inatero showing caudal glands, nuclei of caudal 
glands and terminal lappets. Fig. 28. Longitudinal section through caudal end . lig. 29. Transverse section through caudal 
glands and showing, pore opening of one gland (right side). Fig. 30, Longitudinal section through caudal glands. Seale 
bars = 5 um. cg caudal gland, nu nueleus, Ul terminal Jappet, ut uterus. 


ice LL M.SPRATT & W. 1 


of Studied demos i Cameroon (Bain & Rere 
1993), One of us (DMS) has observed harval 
Durikeinemea sp. in the abdomen of Culicaides 
virlaviae Mache. 1941 (Diptera: Ceratopozonidac) 
near Atherton, Queenshuid where the iree kangaroo. 
Dendrohiun lamholiz’ Callett, [884 is known to 
harbour Durtkainenra macropt Sprau & Speare, 182. 

Out evidence indicates that Attispleca horreli 
passesses a nuinber of the morphological features 
which link ab wilh the Adenophoren rather than the 
Secernentea (Bird & Bird 1991), Bunn and Chabaud 
(1979) reported that larvae of Likoneme, Aduapined 
and Riawsvalvene had comparable phasmids 
(“cellules phasmmidoides”), ic. a pair of large lateral 
cells each conneeted to an opening much Purther 
posterior, Our evidence fram fight and transmission 
electron micrascapy indicates that in third-stage 7 
borreli at least, these large lateral cells represent 
caudal glands with short ducts which open laterally 


fo the exterior (Pigs 11. 28-30) rather than further 


posteriarly. They contain highly  conweluted 
glandular Gssue wilh no sugwestion of ferveus 
lssue, as OCCU [I phasmids (Coomuns & De Grisse 
1¥81), Bain & Chabaud (/oe eit) moted thet 
phusmids were barely pereeptible inadult Muspriecer 
and Riniswedvuie but were particularly pronaunded 
in the genera Cokunena and Pennisi. possibly 
{unehoning as orans of absorption, Ruther than 
ibsorphve struclunes, phasmids' are sematio sensi tht 
with a sensillun’ consisting of one or more sensory 
HWeurons and eseurt cells (Coomans & De Grsse. lax 
em). One oF the latter commonly ecalled tre 
pinemidial wlandt (Chibwood & Chitwood 1950), 
Consequently. these conspicuous leatures ol 
Lukaneie and Peanuivia are more likely 1 represent 
modified eaudul olands than phasnvids: 

We helleve (he srateen tune cells with large nuclei 
Wie nucleoli extending from the cephalic cnd to the 
nerve rig represent sheath cells cach surrounding a 
semaller gonal slistally (see Cuomans & be Cirisse 
Wh). Phe See inet dabial-sensilla passed through 
the euuiele tu the exterior, However, we were Unable 
(Oo obtain) appropriate TEM svetions () contin that 
the outer lateral and cephalic sensilla did nol open to 
the cslenion Similirly, well developed, post-labial 
amphidtal pouches are present in Ad herred? bur these 
did not appear in SEM. lo open to the exterion, 
possibly because openinus were cvbated in wold 
palladium 

Peas af the ventral epidermis ie clectrot 
dense Membres arranged radially in stacks anu 
waidulve tissue with eranules oF protein and lipid 
Sugeesi achive assimilalive, secretory or excretory 
finchons and resemble the bavillary bands found in 


the later, dorsal and/or ventral lypadermal curds al 


THichovephulida t-— Trebinelliduy of vertebrates 
However, true baellary hands consist of elandular 


NIHOLAS 


and non-glandular cells. the Tormer openings to the 
exterior and thought to have @ role dn asic ot 
ionic regulation, the latter hol opening to the exterior 
and believed to function mm cutiele formation and 
maintepanee. and in storage of food materials 
(Sheffield 1963; Wright 1963, 1468), Similarly, the 
internal structure of the large stellate coelomocyles 
suseested an osmo-regulatory or phagoeyiic 
function, We were unable to detect the single dorsal 
gland cell exeretory apparatus with pon. 
culiculanized terminal duct (bow features of 
Adenophorea) described by Buin & Chahaued (1979) 
und emplyime into (he lumen of the pharyirs. As in 
adull Mermithida, the intestine in WW burredi is 
replaced by a trophosome. The traphosonte was 
formed by a series of large cells with prominent 
nucle) and nucieol) rather than a mulbi-nuclvated 
syneyuniim as deseribed by Bain & Chabaud (1974). 
Aldmugh the caudal glands resemble those observ ad 
iW utlier adenophoreans (Maggenti }981). the lattey 
normally oceur as three single eelled wlands- rather 
than Ovo mullicellular ones. We have found ne 
evidence ol a stylet jircitherthe adult or larval stages 
of Me horrelé, 

Detailed cellular ultrastructure was not achieved ia 
this study because by the time specimens had beer 
extracted lront the sub-dermal tissues and fixed ton 
Transmission electron microscopy. substantial 
degenerative changes had occurred. However, oan 
sirudture was satisfactorily preserved and 
observations by light and scanning electnin 
Misrascapy were nat impaired, We suspeel that these 
wematades vontain substantial amieunts at 
cudonucleuses (ind protedses, and that autolysis 
ocet's duit (he post mortem: examination oF the 
host 

Sonm species of Muspieeidi ure dioeerius 
(Ponisi interven) Buin & Chubaud, [274, 
Darkeinema spp, Mayoockoeina perplex) (Bain 
& Chahaiid 1974 Spratt & Speare 1982; Spratt de 
Gill J998), others: are protandtous lermaphroudles 
(Muspicva bortelic Riewveolvimia spp. Lakenena 
luihorchise Chabaud & Baing L974 Mayerter 
veyperations Rausely & Rauseh. 1983, possihly 
appnena ay Bar & Nikander, 1982) (rump 
I¥ath Bain & Chabaud 196k, 197 Chabad & Rain 
O74: Rausch & Rauseh |983: Bain & Nikender 
JO82) In some species (Rebertelalliiver purindtarc 
Chabaud & Cavipana, 50) the vidya ts Functional 
and larvae piss through al (Chabaud & Campana 
1950), In other species, the vulva is fioetiondl suid 
larvae pass through ip but then migrute herween wwe 
layers ul body cuticle und emerge from Ue heute 
resion af the wdult worn (Areiirgelvetaico rhiiioloph 
Bam & Chabuud, 968 (Bain & Chybatid |Vax), ya 
third prop ef speeies (Qurikuinene spp. 
Havcockneme porplesane), the vulva is alrophied ane 


SYSTEMATIC POSITION OF MUSPIOLIDA (NEMAPODAY fi 


ewes batch inside females, develop to thiru-stape 
linvae and burst from the head region killing the 
adult (Spratt & Speare |9S2: Spratt & Gill 190K: 
Sprallelal 1999). developmental feature known as 
wilotakia oenricida: (irschimim 1960) which 
offers an efficient mechanisnr tor auto-re-infeclion 
al’ the host, Larvae eseaping: from the ruptured 
cephalic region Of 7 perplex inta human mascle 
wits illustrated by Spratt er gf (1999). Semelparity. 
the death of adulls upon expulsion of young, 1s rure 
in parasiie nematodes bur aceurs also in the 
Mernmithida. 

he venus Mayerie is distinguished from other 
genera of Muspiccida bya number of morpholovical 
features. the most characteristic being the presence 
ofa Demanian system (Chahaud caf. 1983; Rausch 
& Rausch (983) The genus Paycoekneme Vs 
distiiwuished from other racmnliers of the order by the 
presence of a large amorphous “cell” supportirys = 
granule-fitled ask or gourd-shaped reservoir in the 
rectal revion of mature und vravid Tenrales and the 
presence of lateral bacilaury bunds comprised of a 
sinule row of cpidertnal glands of pore cells spaced 


irregularly und.extending posteriorly in the region of 


the vulvitin tmiature females (Spratt eral, 1999). 
The life cyeles of muspiceid nentatodes are 
unknown, but the modes of transmission have heen 
widely discussed in the fiterarure and postulated as 
occurring by cannibalism. cutaneous penelration, or 
diiring lactation or grooming (Sambon 1925: Brunmpr 


1930) Roman 1965: Bain & Chabaud 1968, 1979, 
Chubaud & Bain 1974: Bain & Sikander L982, 


Spran & Speare [982° Anderson 19% Adanison 
1986). [hus been suggested that these parasites have 
monoxenous life cycles and have evolved in their 
hosts direetly from soil-dwelling ancestors 
(Adamson 1986), The most primitive muspiceid ile 
cveles are though to invalye Hille tissue migration, 
Larvae are beleved to penetrite the skin. dey clap in 
(he subeutaneois. tissues und infective stages. are 


thouwht to leave through askin lesion ind seek a new 
host. A tissue nifation becames necessary: when 
perculancous transmission is repliced by oral 
transmission, as dhay oecur in Muspieed and possibly 
Japprema. 104s presumed that parasites oF ihe deeper 
lisshes, ag. Durikaieme spp. and (L perplextiin ate 
derived from these forms, 

In conclusion. although an onehiostyle was nol 
observed in-adultor larval A, Aeere/i. morphological 
evidence presented here indicates that phasmids are 
absent in this species and whal have previously been 
interpreted as phasmidvid cells are in fact cuudil 
glands with no assoviaied nervous tissue. This 
finding. together with the previous findings by Bain 
and Chabaud (1979) and by Spratt ev e/, (1999) 
strengthen the view that the Muspiceida provide a 
link between the Mermithidi and the 
Trichocephalida, and on morphological grounds are 
adenophoreans, hot sccerenicans, This conclusion 
is inaccord with Clade | or Blaxter ef af. (1948) and 
Doris ev a (1999) whieh grouped the vertebrate 
parasitic, order “Lriehovephatidu with the insect 
parasitic Mermithida, the plant  -parasitic 
Dorylainida aid the free-living Mononchida, I also 
accords with the tentative classification of the 
Nemutoda by De Ley & Blaster (2002) placing the 
vertebrate parssilic Muspiceida, Dictophymauda and 
lrichinellida( Trichovephalida) alongside the inseet 
parasitic Mermilhidivand Manmernmthida. the plant 
parasitic Derylaimida and the — lree-living 
Mononehida in the subclass Dorylunia. of the class 
Cnoplea. 


Acknowledgments 


ly is a pleasure to acknowledye the lechmeal 
assistanee of Dr Amand Stewart and Mr James 
Bovden. The late Dr Ruy C. Anderson and Dr lan 
Beveridge offered valuuble crilicism of an earlier 
dratt of rhe manuseripe. 


References 


Anmwaisos, ME (U8) Phyloneneticandlysis ofthe higher 
vlasstlicalion wf the “emalada, Gay. J Anal @3, 14 7s- 
1482, 

Ayprersoy, RO. (i984) The aes a) cuoparasiie 
nematodes. Cam A Anel, 62, 317 

(2000) “Nemitlode 
~theirdes Sinponent Sune) Passion, 
Imernadonal, Wallingford 4. 

& Bain, O. 1/982) “C14 Reys to the Nematode 

~ Paniles OF Verlebriles. Neo 4 Keys ne Gener of the 

Superfambes  Rhabdrtoides,  Dioctophyiatodes, 

Prichinetlorden dad) Muspiceotdea (Conmonwealll 

Agricultural Bureaux, Farnham Royal) 


raraxilos ol yertehrayles, 
Second de (CAB 


, SCADA. ALO Wipe at Soe tycn 
~ Keys w the: Nenwiode Pardsites of Vertehres, No. 1." 
(Coommonwealth Avricultural Bultcuus. Parvin Royal), 

Bary, OF & CHapsub, AL GL ()46s) Deseriptien ile 
Rioweolan viiiwlophi ian spy Nematode parastle 
de Rhinolophe, montrant les alfinués entre 
Muspreeowen et Mermithuidea Ann. Parasitol ben, 
coun AAS 45-0 

& (1979) Sur les Mispreenhie 
(Nematoda-Dorylannina). dhiel S34, 2007-259. 

a A NIKANDIR. So (TUS) Un ndemitode 
aphastmidicn duns les qapilladies de Poreile cu penne, 
Lupprena aris a. fen. asp. (Roberidolllisihic). Thjed 
SR, A8a-39Eb 


62 DOM SPRATT & WL. NICHOLAS 


& Rise AL (1093) Inteative lurvae of o tow 
species of Robertdolfusidae (Adenopborca, Benmalodi) 
‘ thhe gut ot Sila damnosum in Cameroon, Shak 68, 

2-184, 

Braver. PoC. & Boraboriiag, Wo CLORd) A microliiri af 
exceptional size from the ixedid tick. Aveley damnal, 
frum Shelter (shind, New York, .. Puravizal Wh 963- 
vIitee 


avas a (1987) Creal Comment: 
“Microfilacia wl exceptional size is larva ol! muspiceoid 
hematade.” Jhref 73. 389. 

Hin, A EL & Bien fo (199E) Te Structure al 
Nermuodes.” Sceond bul (Academie Press, Sun Diego, 
Cyditornia). 

Biaxger, Mob. Dr biy, PL Civrpy oh Ro Line Le xX. 
Scie brats, Po Vikksimarih. AL WAND bbb. KK. 
Machry, bY. Denes, Ma. bP aissh, be Me Wibay dh Tk 
THOMAS, WL OK (1998) A inoleeitar ewolulimary 
framework for the phylum Nematoda, Mule 392, 71-78, 

Borkii. AS C1920) Parasilisme ef tuimeurs. clan dus 
Ruste Phavis. 24. 778-784 | 

Become. 6. (1980) AMaspiees herreli Sambon 1825 et 
waneers de souris. Dan Pareisitel, hii comp. 8, 400 
343, 

CUMRALIA A. Gi (197A) MCUEE Rews: to the Nempode 
Parasites i! Vertebrales. Ne. 1, Class Nematoda, Keys li 
suhelasses. Orders and supertamilies,” (Commonwealth 
Agricultural Bureaus, Farnham Royal). 

& BA, OL (1974) Données nolivelles suv ta 
highowe des nértodes Muspiceides fours par Pete 
Wun parasite de Chiroptires: Lukeuema hieaschest 1, 
gen nespy tan Parasia. lane comp: 48. 819-854, 

& Campana, Yo (M050) Nouvenu parasite 
remnarquible par Vatraphie de ses. arganes. 
Raleridoliinsa pataderu (Nematoda, daeeetie wells}. 
Thi 25, 325-444, 

bars, ©. Vincion J-R, Rausacie RL. & 
Racserk VOR. (1983) Organe de Mansi de Man it 
inscomation tunmalique. Reve Nemata, t. (27-131 

Chirpwoon. BG. (19383) A revised clissification of the 
Nematoda 2 Memesttal 20, 141 

_ (1950) An outline classification ul the 
Nemuloda pp, 12-25 Ja Chitwood, 13. Gh & Chinwaud, 
(Mets) “Intrndietion le Nemanilogy” (Universiny Park 
Press, Baltimare). 


COOMANS. A, A Dr GRIssSk, A. CIYXT) Setisnrey: strictness 


op. 127-174 fa Zuckerman, By M.. Rohde, B.A, (keds) 
“Plant Parasite Nematodes, Volume I" (Acaemic 
Iruss, Londen und Sew York). 

De bier, Pak Blasio, M, (2002), Systematic pasiven and 
phylogeny pp, 1-30 de bee Db, (Edy The iiolowy al 
Nemutades” (Laylurd& Prunes, Lorin). 

orgie, My Dre bby ThoX Biante, Me Le (199) 
Moleeular anilysis of nemitode diversity and the 
evelution of panisitisin Parasind Toeay VS 1488-193, 

TG ROAS TE TICTY, Dh MECREPNZUT SL, PRSNEM ETE bie, U1 af 
KEEPIN TC PYRO) Polyroorphism of lyinphocyle antiaens- 
encoding loei in wild mice Curk Toyn Mierohled 
Immunol, 127, 224235, 

Pi ebnOes FCP 29) On the Jarval migration al) seme 
palusitic nematodes. i the Aadly af phe: host aun) 13 
siumifiaince. A Hesminthal 7, 15-20, 

Hes cuiMiash. Ph (1960) Reproduction af Nematodes. pp, 
1H0-167 die Sasser, do A enkids, Wo Ro (Babs) 
“Sermitotogy, Pundanentiels ancl Recent Advunees will 
Trophasicon Ploat Parasitic und Sor borms” (Lonwersity 
ol North Caroling Pross, Sorth Carob), 


Incite, W.-C, (1983) An outline classification a the 
phylum Nematoda, lain. Ane, 3a, 243-255 

KAMIFER, oS. Sipitananrn, Co 8& Ort 2 (ays) 
Vhylogenetie analysis oF FONA fron udenopharean 
nematodes and impheations Cor the Adgnopharea 
Seoernemled controversy, favert Biol, 117. 2030, 

Lorravis, S. (1983) Phylogenetic systemities: problems 
achievements ind its Upplication to the Nematoda pp. Tl 
23 Jn Stone, ALR, Phat, UM. & Khalil, b, T. Unds) 
“Concepls in nematode systematics” (Acudemis Press, 
London & New York), 

—_2 (1994) The phylogenetic systematies ab tee 
Iwi dematodes”. (The Ray Society, London), 

Vasnorek Al (M981) “General Nematolony" (Springer 
Verlag. New York, Heidelberg, Bertin). 

— (1983) Nematode tigher classification as 
TiMuenced hy species und Kimily concepts pp, 25-40 da 
Stones AOR. Pal, Hh M, & Khalil bP (bd “Coneepts 
UT nematode systemutres” (Agademic Press, Lemdon & 
New York). 

KANSCH, ROT, & RAUSCH, WOR. ClO83) Maserre 
vuspertilonis 1. 20M Spy (Dorylaimine Muspiceldie), a 
netioiade from Neurelic bats (Vesperutionidue), .ln 
Parasitol. hum. comp S58, 301-370, 

Rowan, Lo (19865) Superdamalle des Muspiccides 
(Musprewodea) pp, 721-726 he Grasse, PM CR 
“Thé de Zoologie, Anatomic, Systemulique, Bialasie” 
Vol 4+ Pr 2 Némathelminthes (Masson, Paris). 

SaAMBON, LW. (1925) Researches on the enidemiolopy al 
cancer made in feelund aud taly (luly-ODetober }824), J, 
trop, Med Hv, 28, 34-71, 

SHEED. H.C. (1963) Electron mieroseapy af the 
bacillary hand One stichuaeiie of Zeabaeiy vais and 7 
eulpis dF Porgsiial, 49, 908-1009 

Siseil dom. fe Re (1985) Population dynamics ol Veey 
mavcidus and its parasites in Mallee wheutlands on 
Victoria during and after a drought tak Wildlife Res 
12) 447-445. 

= & REDHEAD. TT DOC 880) Strucnire and btoliyry 
of house mouse populations that plague irregulirlys av 
vvalutioniny perspective, Brat. Flinn See. 41, 285-300 

SeeATL, 1, M, & Serve, Ry (1982) Barikoineme macy 
gen, elosp, nov, (Muspiceorder) Rebertdollfiisidacy, 
remurkable nematode Fron Maeropudidae (Marsuptilia). 
An. Parastiol inim, conp. 3753-62, 

& GALL, PAL L998) Derikuinenne ple olure tt 
hap. (Nematoda: Muspreeoden: Roberttollfisiduey 
from the pulmonary dteries OF the koala Phascedentey 
eiiepens Nt associated patholowien! ehummen. Ayye 
Petrosdtal 39 VOT-10t, 

ce — Bevtreiner 1. ANDREWS, JOR DD SODtAE Ey, 
SN PIYUY) THepeorcheina perplesay ey lbhap 
(Nematode Robertdollisidveds an intrumyothbre 
parasite in min, fhe, 43, 123-144, 

Tiierronn Da Ree uerrn. Bode Vaseariis, OP cory) 
“Diagnusing helinintiiasis through  cuprelugieut 
cxumination” Glingsen Research boundation, Beersc, 
Belen) 

With, ReoA. (1963) Cytology al the bavilliry bangs al 
Capithiriay hepatica do Marphiak Wz, 243-259, 

(1968) Stracture ol the bacillary hands af 
Trrehans mvocaitory, 2 Pareisitel, $4. 1100-1110 

(2989) Paurssites in peril - the Urichuroid 
nemilodes pp. GS-80 fie a, Ro Cha) Current Coneepts 
in Minteiology™ (Phing Kong University: Press, Pyne 
Kanye), 


A TAXONOMIC REVISION OF THE CAMPONOTUS 
WIEDERKEHRI AND PERJURUS SPECIES-GROUPS 
(HYMENOPTERA: FORMICIDAE) 


By S. O. SHATTUCK*® & A. J. MCARTHUR** 


Summary 


Shattuck, S. O. & McArthur, A. J. (2002) A taxonomic revision of the Camponotus 
wiederkehri and perjurus species-groups (Hymenoptera: Formicidae). Transactions of 
the Royal Society of S. Aust. (2002), 126(2), 63-90, 29 November, 2002. 

The Camponotus wiederkehri and perjurus species groups are defined for the first 
time and revised at species level. Thirteen species are included in the wiederkehn 
species group, six of which are newly described while four previously valid species 
are synonymised. These species include arenatus sp. nov., aurocinctus (Smith) (and its 
new synonym midas Froggatt), ceriseipes Clark, donnellani sp. nov., gouldianus 
Forel, owenae sp. nov., postcornutus Clark, prosseri sp. nov., rufonigrus sp. nov., 
setosus sp. nov., terebrans (Lowne) (including its synonyms testaceipes Smith, 
latrunculus victoriensis Santscht and myoporus Clark), versicolor Clark and 
wiederkehri Forel (with its new synonyms denticulatus Kirby, latrunculus Wheeler 
and wiederkehri lucidior Forel). The perjurus species group contains the single rare 
Species perjurus sp. nov. 

Key Words: Hymenoptera, Formicidae, Formicinae, species-group, Camponotus. 


Transeclions of the Raval Society af 8. Aust. (2002). 126 (2), 63-90) 


A TAXONOMIC REVISION OF THE CAMPONOTUS WIEDERKEHRI] AND PERJURUS 
SPECIES-GROUPS (HYMENOPTERA: FORMICIDAE) 


by 8. O. Suarruck’ & A, McCARTHUR™ 


Summary 


SyAactiick, 8. O. & McA, A, 1. (2002) A axonbmie revision of the Ceaporioms wederkelrs and perprus 
ypecics-groups (ELymenopten: bormicidae). Trenseetany of te Raval Sach ab det (2002), 12 (2), 63- 


90, 29 Navermber, 2002 


The Coniponatis wiederkehed and perjurns species groups are defined forthe first time ind revised al species 
level Thirteen species are (nelided inthe wiedeekehey species group, six of whieh are newly described while 
four previously yal species are synonymiscd These species include aendii sp. noy.. amocincnis: (Simh) 
(and is new synonym vidas Prognil), cermeipes Clark, dannellanisp tow. geteiainis Forel owende sp. Woy, 
posieoraitns Chirk, PrINSerE SP. NOV. PuTOnigris Sp. MOY. setae: sp. nov, deriGrany (Lowe) (including its 
synonyins resfaceipes Seth, fatrumediis victorivasis Santschi and prveporyy Clark), veryicolar Clark ana 
nicderkehe’ Vorel Owith its new synonyms dentioutamns Kirby, dairmnentus Wheeler and wiederkedrt lucidior 
Norel), The perfirus species avoup containg the single rare species perjuray sp, Woy. 


Key Warns: Hymenoptera, Pormividue, Formicinae, speuies-sriup, Covnpenidtin. 


Introduction 


Iithis paper we revise species oF ants in the newly 


delined wiederkehri and perjuray species groups of 


the wonus Camponotus. Fourleen species ure 
recupnised, seven of which are deseribed for the first 
lime; four previously valid species are treated as 
synonynis. These groups are restricted to Australia 
and contain species which range Irom common to 
rare amd fram widespread to restricted in 
distribution, They are most abundant and species 
rich ain semi-arid regions and all are apparently 
ground nesting, taxonomically, the species weated 
here were previously placed in the subgenera 
Murmopliuma, Mvemusanlas, Myrnoturha and 
Tangenyrmex, placements whieh were made when 
ihe species were orizially deseribed and have not 
been discussed since, During, this study it has 
beeome quite clear that the current subgeneric 
classificarion witht) Cumporanis is ehaotio and 
neatworthless, Species here placed in’ the 
wiederhehr? Spectres group shure similarities in 
overall body shape and size including the placement 
of the compound eyes and We configuration of the 
musosome and petiole. In addition. all share a cluster 
ol vlongate hairs on the base of the mentum. This 
cluster is unique in the Benus and strongly suggests 
they are monophyletic. At present. the higherlevel 
classification within Campanas is poorly under- 
stood and unul (he eptire genus is examined more 
closely, ik iS inappropriate to speculate on 


CSIRO Entomuloay, GO) Box P700. Canberra, ACT 28h 
Avintralin 

~Sauth AUS UTE WES 
Adomtalin S000, Auistea tier 


North ‘Rorawe, leldes Souel 


relationships among species, Canipouertiy genidetians 
is associated wilh a lealhopper and C. ferebrans is 
assoviated with a butterfly, Por an overview of Lhe 
subfamily (Pormicinae) and genus (Cemponotis) in 
Australia see Shattuck (1999), 


Methods 


Mevsurements 

Size and shape characters were quantified and: are 
reported as lengths or indices. Measurenients sere 
made with a stereo Microscope using, 2 dual-axis 
siige micrometer wired to digital readouts. The 
following Measurements and indices are reported. 

Cl Cephalic fadex: HW/HL, 

HL Maximum head Jeneth in lull face view, 
measured from the anteriommost point of the 
elypeal margin to the midpommt ofa line drawn 
across the posterior margin of the head, 

HW Maximum head width in full face view 
excliding the eyes, 

ML. Mesosonral length measured from the anterior 

murgin of the pronatal collar to the posterior 

extension of the propodeun lobes. 

Maximum length of mid tibia, excluding the 

proximal part of the articulation which is 

received into (he distal end of the femur 

St Seape index: SL/HW, 

SL Lenwth of the seape (first antennal seyment) 
excluding the basal neck and condyle, 


MTL 


Location of material examined 

AMSA, Australian Museum, Sydney, New South 
Wales: ANIC, Australian National Insect Collection. 
Cunberra, ACT, BMNH, The Natural [History 
Museum. Londen, UK; MCZC. Museum ol 


Od S. CLAN ATTUOCR & ALL MCR HELE 


Coniparative Zoology, Harvard — University, 
Cambridge, Massachusetts. USA: MING, Muséuin 
(Histoire Naturelle, Geneva, Switzerland: MYMA, 
Museum af Vietoria, Abbotsford. Victoria; SAMA, 
South Australian Museum, Adelaide, South 
Australias WAMP, Western Australian Museum, 
Perth, Western Australia. 

Mast of the non-type material is itt ANIC and 
SAMA, 


Callectors uf marenial exemirtedt 

WAS, A.A Sompson: ACK, ALC, Kistner) AHL, 
A. VL Burbidge; AJM, A, J. MeArthur: AIO, 
A, Johospn: AIKN, A. IK, Nousala; ALY, ALL Yens 
AMD. A. M. Douglas: AML.A. M. Lede .VMM., 
A. M. Morguny ARP Ao RL Petlie: AWE, 
A.W Forbes; AZK, A, Zeita: BBL. BB. Lowery: 
BHO, Bo Holldobler; BPI, B. Pike: BRIL 
BR. Hutchins: CBA. CL Barrett: CHW. Cb Wattsc 
CNIL C. Nilsons COTM, Co T. Mereaviels CWA, 
(. Warner; DCT, B.C. I Rente: DCO, DB, Cox, 
HDA, D. Davidson: DHE 1. Hirst) DHE 
1), FL Ristiner: DSC. BD. Schultz: PRB. E. B. Britton. 
EBR. EB. Broomheud: REDE. I BD. Edwards FER, 
Pb Riek) EGM. Foc. Mathews; PLO, EF. Lockie, 
PTROU, Troughton EXP, South Australian Museum 
Lapedition. PYF, BR. Yeatmany FAC, FA. Cudmore: 
Pse. FO Sehaeters PSH, Fo Shepherds COA, 
(i. Campbell, GCM. G, ©. Maedling GPa, 
C1 Gross: GPR, G, Friend) GIM. G. J. Mulan 
GLH, GL. Howie; GPB, G. PB) Brownings GRILL 
Ciriffith Collection South Australian Museun HRW. 
HB. Whites HES. Horn Centenuy Survey NVMA: 
LWR, WW. Protny HAE. Ho Ueatwoles HMC, 
(LM, Cane; HOF, A, O. Fleteher; HOW, 1. Owens, 
HIRE, He Reynolds; WWE, He Wesselman: TAR, 
I, Archibald: IPB. 1, PB. Common; IGE. 1. Gee; 
WA. |. Valentine: JAR A. Vorresr JAH, 
JA. Heridge; JAR, J. Archibald: JBA, | Balderson 
IBS. J. Bo Stuckey ICG. I CL Cioudies JCM, 
10. Myers: JDL. 1 0, Erskine; JDL I tf. Dixons 
1M. J. Db, Majer; JED, J, &. Dowse: J 
Jb. Feehan: JPR, F Field; JFL J. Findley: 10. 
10.0. Tepper JHA J, Hawkins: ILA, Lawrenee, 
IMC. Jd. MeuAreavey; JRB, I Ro Bo Law, JRE, 
A, Mead: JR 1 Ruble: JS. Shaws ISM, J, Sith: 
JTW. J. Thurmers JWI, J, Wilkinson: ICCA, 
Ie Caspurson, RIVAL KK, Daveys RMA. KO Mairing 
KMC', K. Mekelson; KRO, K, Roth: KRP 
kK. BR, Pullen; ROR, &. T. Richards; LHI, L, PMitele 
LPK, LL. P. Kelsey; LOU, L, OQueale MAA. 
MOA, Adums; MDA, M. Davies; MIT, Mitchell: 
MIDI M. J. Douglass MLS, M.-L. Simpson; MMA. 
MI. Mulpitail: MPE.M. Peterson, MSLL MOS. Upton: 
NBII, NN. BL Tindale: NOS. Nature Conservation 
Soviety of South Australia ies NEA SN. Lawrence: 
PAL PL Aitken, POCO, P, Copley; PGE. P. Gees PGR. 


I), Greensladey PHU, P. Hodson, PIF) PJ, Paruher, 
PIN, Po M, Greenslade: PPL, P. Plym: PRE, 
PR. Birks; PSW, PS, Ward; RBI. R.A Valliday: 
RBR. R. Brande, RCC. R. Co Chandler: RON, 
RD. Nutting, REL RB, Elder: REO, R. Poster, REC, 
R.H.Croviery REIM, RT Mew: RIB, Roh Bartell: 
RIK, Ro. Kohout, RIW R. J, White RRA, B 
Raven; RSE R. Smith; RSM. RS. Melnnes: RVS. 
RV. Southeott, RW R. Wo Tuvior SAI. 
S.A. Harringion: SANPGLS, S. Aust, Narianal 
Parks and Wildlife, Goydery Lagoon Survey 
SANPNOPS, South Australian National Parks and 
Wildlils, North Olary Plains Survey: SANPNS, 
South Austealian National Parks and Wildlife, 
Nullarbor Surveys SASPPITJ, South Australian 
National Parks and Wildlife, Pitantjatiwa Lands 
Survey: SANPSDS. South Australian National Parks 
and Wildlite. Stoney Desett Survey: SANPSOPS, 
South Australian National Parks.and Wakdhye, South 
Olary Plains Survey; SANPVS. South Australian 
National Parks and Wildlife. Vertebrale Survey: 
SANPWERS, South Australian National Parks inal 
Wildlife, Western Flinders Ranges Survey; 
SANPYS, South Australian National Parks and 
Wildlife, Yellabinoa Surveys SBA, S. Burkers SPO 
S. Donaellan; SLEDS. Lewer, SMOOS) Morrison: 
SOS. S. Of Shattuck: SRM, 5. Ro Maren, TAW, 
TOA, Weir VGR, TL Greaves: TOW, TG. Wool: 
TRO, tL. Robinsons WALLS W. AY Lows WT 
WB, Hilchvoeks WCCO. WC. Crowley: WDD. 
W. D. Dodd: WHC) Waterhouse Club, South 
Australian, Museum: WKH, Wo kK. Plead: WER, 
W. L. Browns WEN, WoE Nutting: WMC, Wester) 
Mining & Royal Geographical Soviely Expeditions 
WMW. WW. ML Wheeler YOO YC. Crozier. 


Genus Calnpanatiy Mayr 186 | 


Definition af the C. wiederkuhri species gramp 
Members of the €° yieederdcAr?y species group part 
be separated from other Australian Campananw by 
the presence of a cluster of four or more distier 
elongate curved or “J-shaped hairs on the hase ot 
the mentun) (near the posterior region of the baces! 
cavity) in all worker castes (Fig. 1). Ina few speeies 
related te OC) epAippiaes similar hairs are present bul 
(hese are scattered along the Jeneth of the mentum 
rather [han being present as a pusterior cluster. 


Complexes withtt the C. wiederkehrt species group 

The C. wederkehri species group ean be divided 
into four complexes as follows. While it is likely that 
these complexes represent monophyletic groups lane 
there is no eVidenee that they do not) synapormorphies 
supporting [hese wroupings have wot been souvht in 
this study. Ibis more appropriate for these studies tu 
be developed asa holisue study of the wes. 


CUMEONGTOS WIEDERRETREAND PERFURE SS SPECTS GROUPS hd 


airecinems complex: Includes CL arerertts, 
diracihetus, Gwensae, xetosuy and vervienlor. 
This complex is defined by the presence of a 
distinct and angular netanolal groove in ino: 
workers whieh is depressed (sometimes only 
slightly) below the anterior region of the 
propodeudin (Pigs 3.8, 9). 

ceriveiper complex: Includes C. cerivepes, 
danmellan, prasseri and rufonigrus. In this 
coinplex the posterior section of the mesonotum 
is Weakly bul distinctly convex immediately 
anteriog of the metanotal groove (more so iy 
minors, less So in majors) and the metanotal 
sivove 1 minors varies Cram a distinct angle go 
i shallow concavity (Migs 12, 14, 18, 34. 30). 
posteurnulns complex, Inelydes © postearin ts. 
In this complex (he entire mesosoma in minor 
waurkers 1s stronly arehed. laeks a metanotal 
eroave and the pasterior face af the propodeam 
isomy weakly differentiated from the dorsal! face 
(fig. 31): mn mayor workers the posterior carners 
of dhe head taper rearwards inte blunt 
protuberimees (Figs 28. 29). 

fereheans complex: Includes Co gaefdianis, 
ferebrons and wiederkeliri, In (his complex the 
posierior section. of the mesonotum is. fat (ar 
nearly su) intmediately anterior of ihe meranotal 
groave and the ametunotal vroove in minor 
workers absent or weakly developed (Figs 23. 
47, 58), 


Definition of le C. perurus specie grou 


This species group rs reeovuised by having the 


head produced upwards so that ifs attachment to the 
pronoturn iswell below ifs upper muruin (Fig. Of). tH 
Nias 4 redueed number of hors on the menturr 
compired lo species oF the wiederkelinr group. 
upproachitie the arrangement totind in relatives of C! 
ephippiiin, THS group contains a single species. C) 
perjuenss deseribed below. 


Key to workers of the Camponotus wiederkehrt 
sprees group 


{ 


2 


breet hairs present on dll surlices OF tibide 2 
[rect hairs absent front outer surfaces of tibiae. 
inner surface with aw double-row (Gallhouple 
appressed pubescence may be present) evnncet 
Metinotil broove im mibor workera distinel but 
sometimes shallow trough (Pig. 42); knawn only 
from the Kimberley region of northern Wester 
Ais traltee (Big AES) cic suai trece seers ereer MO RUSES 
Melanolal groove in miner worker wenkly 
developed (Fig. 23) or absent (Fig, 47% koowa 
ouly from Souther Australia (Pies 24. 48)....-3 
Number of erect hairs on propadeum preater 
(han 40: pubescence on head und aster abundant 


6. 


and with individual hairs overlapping! siummit ol 
petiolar node in protile rounded in miter 
workers (igs. 23), a blunt Seta in major workers 
(Figs, 21)-.. Agere coy lethiny otis 
Sumber af erect hairs on  pronodetun less than 
25: pubescence on head and yaster sparse and 
with individual hairs generally non-overlapping 
or at most only shehtly overlapping; summit of 
petiglar node in profile angular ia both minorand 
mayor workers (FIPS 4S, 47 yore hres 
Entire mesasomm in lateral view strongly arched, 
lacking & metanotal groave and with the 
posterior face of the propadeum only weakly 
differentiated Irom the dorsal face (Pigs 20. 30): 
posterior corners of head of major warker 
tapering rearward into blunt protuberances (Figs 
28, 24) oy ubeceviyuavay eae vevesyaserees cop ANTCOLTLAS 
Mesosoma in taleral view Hatborat most with the 
pronotum and mesooaium weakly arehed and 
separated trom the propadeum. by aw weak angle 
(hig. V4) or a distinet. angular or concave 
melanolal groove (Pig. 9), The posterior lee oF 
the propodeum always separated [rom the dorsal 
face by a romued angle (Fiz. 14): posterolateral 
corners OF head rounded tn tmajors (Vis 33, 34), 
Melanolal grouve in minors depressed below; the 
anterior region of propodeum (Pies SY): 
melanglal wroave in majors anyular (Pig. @): 
dosti of petiolar nade in minors broadly or 
weakly convex. (Mat or weakly colcave. the 
anterior face much shorter thang the posterior lage 
(Pigs &. 8); petiolar node in majors broadly 
rounded ubove (Pig. 6).. mallee vsebe'ee wi 
Metanhotal groove i miners s AbSOHt (Pia. "s8) or 
angular (Fiv, 14) and always even with the 
antertar revion of propodeum: metanolal groove 
in Majors a broad, shallow angle (big. 12), 
dorsum of petiilir node in minors anwular or 
browdly rounded, the anterior face at mest only 
slightly shorter than the posterior lice (Pig. 14%; 
petiolar nade io mars AUBAIAE above ({ig. 12) 

saittaaatias mu 
Darsal ‘and anterior regions “of pronntuim ‘dath 
red-bluck. distinetly darker than the yellow-red 
Mesonotum ane propodeupy oo reManuy 
Prlire mesosoit wallop i calor varying 
from dark red-black to blaek on. 7 
Flongale (overlapping) and dense pubescence 
presenr on dorsum of head. mesusoma, gusier 
und tibiae... us bien eee te WES 


1G OO UNIO COG Derren Cre p Orne ee tep ee 


vuapescancnwconari er t 


Short (non-overt cerlapping) ‘and scaute red puluseerite 


present on dorsum uf head, miesosoima. gastcy 


ANE (TDTAC  ayicicu caeerereesiae of 


Anterior region ol “first ‘gasiral “werwile “dark 
reddish black or black, similar im colour to 
propadeum: matpetat groove in minors distinc) 


ae 5. O. SHATTUCK & AW. MCARTITUR 


and depressed well below We anterior region of 


propodeun (Tips 8, Q) eee ee tPCT MES 
First and second gastral tergires red, distinetly 
lighter in colour than the reddish black 
propodeunl, Metanotal yroove in minurs weakly 
lo moderately depressed rail the anterior region 
of the propodeum (Figs 52, 53) 0.0... vervicolor 
9. Posterior section of mesonotum Hat (or nearly 
so) fmiediately anterior of metinotal yroove, 
metanotal groove absent or weakly developed in 
minors (Pig. S&8): anterior clypeal margin in 
majors prajeching with a straight central region 
separated from lateral regions by distinct angles 
(Fig, S)sscsecseetesrece wwiederkehpi 
Posterior section ol mesonutam weakly but 
distinctly convex immediately 
Thelanolal groove (more so in Minors, less so mn 
majors); melanotal groove varying from oa 
disunet angle to a shallow concavity in mimors 
(Pigs I4, 18, 36); anterior clypeal margin 
it majors broadly convex aeross entire width 
Tiedt WO oiedscotetnerparijem pare too gtntarmaat 10 
LO. Propodeum with at most 4 clongate erect hairs 
which are limited to the angle between the dorsal 
and posterior faces.. — _ cannelfan 
Propodewin generally with more ‘than 10 ervet 
hairs which are always seatlered along the entire 
PSH] RUBIA Sensi sc onjeneadeonieliasinopensloeneleosllelde es I] 
1. Metanotal groove well defined and angular (Fig 
39): blavk head ‘Conte g with red mesonotuni 
spoocenben, PUPOMELTUS 
Metanutal vroove a weakly defined concavity 
(hips 14. 36): head same colour as mesonotum 
(both either red or black)... eee 12 
2. Seapes relatively short (i miners. SI 5) (E Tee 
15); peltolir node of minors ercrfis more 
upright and narrower (Fig, 14)... acertselpes 
Seupes relatively long (in minors, St > 1.4) (liv. 
15): petiolar node of minors generally lower anc 
broader (Fiz, 36)... SPLOS8ehs 


NPeles of the ©, wiederkerhi Spemley Srp 
Camponotus arenatus sp. voy 
(FIGS 2-4) 


Neiternil Examined 

Halonpe, Minor worker from South Austratias 
ILumbidee [labelled as Hambridge| National Park, 
17 Weeember 1970, 1 B. Britlon (ANIC) 
Pevainyes Iwo munor Workers. same diate as 
holotype (ANIC, SAMA), 


Offer material exam/rived 
Northern Territory; |5km 5 Alice Springs (PJM), 
South Australia; Cowell (BL): Maralinga (GNC): 


anterior of 


1 


Pie. t, Underside of the bead showing distinelive cluster ot 
elanwate curved or "shaped hairs Gindieated by arrow) 
On die mentum. 


7 


ligs 33 ¢ Head of sitar 


adrenals workers. Pig. 2 


worker Pros 3) Mesowome and petiole of iiinoe worker 


Yumburra CP, 26km NS Inila Roel Waters (TOW) 
Western Australia; 20mi. W Sandstone on Mt 
Magnet Rad. (AMD & MID). 


Worker diagnosis (niner worker) 

Tibiae ond seapes lacking ereet hairs. Uy minor 
Workers mekinolal grooye depressed below level ol 
the anterior region of the propodeum; dorsal surfiice 
of node broadly convex, its anterior tace much 
shorter than the posterior five (Fig. 3). Dorsal and 
anterior regions of the pronoun dark red-black, 
distinetly darker than the yellow-red mesonetumn and 


CAMPONOTUS HIKDERKILBE ABD PERLORES SPECIES GROUPS oT 


fig. 4. Distribition af Co areas material examined 
during thes study, 


propodeum. This species ts superficially similar to 
C, donncHani in overall colour pattern but differs in 
the larger sive of the minor worker and the depressed 
menmotal wrooyve. 


Deseriptian (minor worker) 

Anterior elypeal margin’ broadly convex (Fig. 2). 
Dorsal surface of pronotum weakly conver and 
sepurated fram the sweakly convex mesonolum by a 
shallow angle: metanotal groove slizhtly but distinetly 
depressed below the level of the anterior propodeuni; 
propodeuin uniformly and weakly conves and without 
a distinc! angle, ratio of dorsum to dectivily about 1.5 
(Fig. 3). Petiolir node with a short anterior face whieh 
is weakly differentiated from the broadly convex 
upper surface, the rear fice indistinguishable from the 
upper surface (Fiv, 3). Ereet hairs moderately 
abundant on all surfaees of the head and dorsal 
surlages of the tnesosoma, petiolar node and vaster. 
absent from seapes and tibiae, Head and anterior 
regions Of pronolum black, posterolateral pronotum 
(immediuely above the fore conte). mesonotum, 
propodeun. petiole and legs yellow-red. gaster 


varying from-entirely yellow-red toa combination of 


the yellow-red anteriorly and red-blick posteriorly. 


Maasurenioalts 

Minor worker (n=). CLOT? O795 TILL Y4nim 
220mm: HW 180mm — 1.740: ML 345mm —- 
3.8hnm: MTL 226mm -— 247mm: St LAW) S92 
SL 2.38mi 2.59mm, 


Comments 

This uncommon species S$ known from a lintited 
number of miner workers, Lt ranges from soutlt 
central South Australia, north to southern Northern 


Territory and west-central Western Australia (Pig. 4). 
The only biological uformation is provided by the 
single worker collected by BoB. Lowery. Ib was 
swept fron mallee on red send. 


Kiyntilogy 
From orcha, alluding to the sandy nature of the 
known collection sites of this species. 


Componots airocinciis (i Smith) 
(FIGS 5-10) 


Formica curoeincta SMI. L838. 39, 

Camponalus auraeinetus Mayr, 1886: 355, 

Caniponolus imdas Froggan, 18962 390; Clark, 
1930; 22 (queen described. worker redescribed), 
New synonymy. 

Camponaitis-sp. & — Imai eral. 1Y77) 469. 


Material examined 

Camponotus unrocinns. Worker holotype or 
yyntypes from Adehude, South Australia. Ao single 
specimen (minor worker) in BMNIL is labelled as the 
type of this species. However. this specimen was 
acquired im 1870, several years after the original 
deseriplion was published. [tis currently not known 
Whether the acquisition date is in error or the type 
specimen is lost. For the purposes of this study this 
specimen is Considered a ype specimen for (his name. 

Cumponolis midus. Syntypes trom Hlamurta, 
Norther Terrttory (1 worker (missing from point) and 
1 queen in AMSA; 7 workers, | queen and | male in 
MCZC: | worker In MVYMA: 3 workers in BMNH 
(with an additional 6 workers labelled as °C’, Australia. 
Horn Coll, 96-37" and bearing a Type label), 


Other metterial examine. 

New South Wales: J2kin S Coombah (PSW), 
45km N Balranald (SOS); Ascot Vale (RSM): Black 
Mill Creek (RHM): Broken Hil (FSH): Beaken Lill 
Airport (RSM): Matakana RS (BBL): Mount Gipps 
(RIIM): Mundi Mundi, op, Broken Hill (PIM & 
IVA): Pinnacles. 12mi. W Broken Hill (BBL): 
Poonearic. W. Smith property (RHC & YCC & 
AKN): Silverton (PIM). Northern Territory: |Skar 
s Alice Sprimgs (PIM): 23mi. SN Narwietooma TS 
(RSM & JED): 33kin E Ayers Rock (EF): 7km W 
Curtin Springs (SOS); Andado (HOF): Kings Creck 
Stn (SDQO): nr Ayers Rock (BBL): Old Andado, 
e(5km Eby Andado HS (ib) Uluns NP OLS kin 
ESE (HCS), Queensland: Muncoonte Lake (RRA): 
Cunnamulla (BBL): Foxes Ch. (GiCA); Sandringham 
(PIM), South Australia: 10km NW Emu Junction 
(JAH): 1Okim WSW Mt. Playford Murnpeawie 
(IRE): iim. S Loxton (BBL): | km ENE Arabanu 
Hill, Murnpeowie (RE 14 km SW Taplan 
(SANPVS): L4km SbhyW Beltana GER: l4km 


8 S$. OOSHATTUCK & A. J, MCARTHUR 


/ 


Fies 5-9, C. enracinetus workers. Fig. 5. Head of major worker. Pig. 6. Mesosoma and petiole of major worker, Fig. 7. Head 
of minor worker. Figs 8-9. Mesosoma and petiole of minor worker. 


Fig. 10. Distribution of CL anruciactus material examined 
during this study. 


WNW Renmark (KRP); [km N Vokes Hill junction 
(JAF), [km W Emu Camp, Victoria Desert (PJM); 
2.5km N Limestone dam (SANPSOPS); 26km SSE 
Mlintjitja (SANPPITJ); 30mi E Farina, Mt. 
Lyndhurst (ETR); 3lkm NW Renmark (KRP); 3kin 
W Emu Camp, Victoria Desert (PJM); 4.8km SE 
Coongie, Coongie Lakes Study site 1OE (JRE): 
40km W Vokes Hill Junet, (JAF): 40km WNW Emu, 
Victoria Desert (PJM); 40mi. SW lron Knob (JRE); 
45km WNW Emu, Victoria Desert (P.M): 4km NE 
Marroo Hill, Cowarie (PRB); 5 km SW Farina 
(SANPSOPS); 60kim E. Vokes Hill, Victoria Desert 
(PJM); 6km W Koonchera, Birdsville Track (PIM & 
JAF); 70km E Emu, Victoria Desert (PJM); 9km 
ESE Wapalanchie Tank, Cowarie (TRO); Adelaide 
(GRI); Adelaide (JGO); Alton Downs old HS, 
¢.48km SW Birdsville (JEP); Ampeinna Hills 10.5 
km E (SANPPITJ); Andamooka Ranges (MIT & 
GPG); Approdinna Attora Knolls 86.3 km SW 
(SANPSDS); Barton Siding (AML): Beda Hill 
(JAF); Bimbowrie 2 km NE (SANPNOPS):; 
Brookfield Conservation Park (Site No. 1) (SOS); 


CYMPONOTUS WIEDERKREHREAND PR RJURES SPLCIES GROL PS 4 


c.lSkor SSE Poochera (RWT & RIB): G22tkar NS 
Beltana (JERK): Calperumt NE Boundary (AIM) 
Cambray (PJM): Cheesman Peak 13.2 kov NW 
(SANPPIV); Clifton Hills Outstation AP & DED: 
Coongee Lakes (JRE); Coongie Lake (DMI), 
Coongie Lakes (JRE): Cordilla Downs Stn 
(SANPSDS): Cordillo Downs Sta (SANPSOPS). 
Corrobiome Hill, Eyre Peni. (RCA): Dangyali CP. 
Red ‘Tank Dam (AJM); Darke Reske, Fyre Pen 
(BBLA: & Purni Bore at junction of French Track anu 
Rig Rd Simpson Deserl JAP): Emu Camp, Vietoria 
Desert (PIM): bin Junetion LO km NW (JAN), 
Fladunna Sto, ITH): Varina 5 km SW USANPSDA). 
Gammon Ra. NPL Baleanoona area (AdM); Crawler 
Runpes (PIM): Glenelg (WBA): Gum Lagoon (EGM 
& JANE): Uamilten Ck. (RBR); Hamilion Si 
(WEKIT): Hincks NP (EBB): IHhingita 23 kin WS 
(SANPPET): Tron Knob 40 miles SW (BPR): Kendal 
(AWE): Killiparu CP (SLE), Kimba (PIM & IVA): 
Kamba, ede of Pinkawillinie CLP. (PSC): 
Koonamore (PIM): Koonamore 9 km & 
(SANPNOPS): NKoouumore, Nillingheo (PIM): 
Koouehers Waterhole 6.25 ki S (SANPGLS); 
foonchera. Birdsville Track (PIM & DAF): Kopi. 
Fyre Pen (PJM): Kamytjanu 25) km WW 
(SANPPITD: Lb, Meraniangye. Vietorla Deser 
(PIM): L. ‘Torrens. nr Beda Hilt (JAR): lake 
Appadare 2 kim S (WHO): Lake Callabonna (AZE): 
luke Gilles CP (BRL): Lake Palunkarinia (0b). 
Lite Pine HE G. 32m. SW Whyulla (EBD; Mabel 
Crack (PCR), Marpoo Waterhole (PGE & IGE); 
Marsethy Will 3.6 kine SE (SANPSDS); Maryiona Hill 
Js kin ESE CSANPPITS: May PAT 93 ton WNW 
PANPSDS); Mantevatlina ore (ISH), 
Mocaivale. Daneali CP UAIM): Mount Lindsay 
31 km WNW (SANPPIED): Mt, Guason SC 
Woomera (PJM), Mt. Sturt, tin salt lake. NN. Levre 
Pern. JAP): Munvaroa CR, Thin SSW Moonable HS, 
A7ha fr Whyalla WKH), NW Yaninee, Eyre Penin. 
(KOA): Olympre Dam (CGM & CWA); Paney, nr. 
Pitk Lake. Gawler Ranges (WHC) Pinkawillinie 
CP Lyne Pen. (AES: Pinnacles Mine (RLIM); 
Pipalyitjan 27.5 km NE (SANPPITJ), Poochera 
(BHO): Purni Bore 77 kin To (SANPSDS): Parni 
Bor, SW Simpson Desert (PIM): Radiam Hill 
(PAD. end of L, Windabout (BBL): 5 Koonchera. 
Birdsville Track (PIM & JAI: 5 of Mano Ra. & Ske 
NW Mi. Kintore (SANPPITY, Serpentine L,, Creat 
Victoria Desert (PIM): Sementine Lakes (AT): 
Srmpsot) Desert (DSC); Sinelain Cap CPHL): 
Stockyard Plain GAIM): ‘Taplan 14 Kin SW 
(SANPVS), Thirty Thousand Tank (Ct M); 
‘Voralwwk Varn (AR Trinity Well (as Trinity) 
(EXP); Unusrnou Roekhole (SANPPILI), Vokes 
Hill To km N WAT): Wallatinoa 160 kin W 
(SANPPLED Yelpawaralinna Warerhale 76 kon 
NNW (SANPGLS). Vietovias Yhin ESE Hatroh 


(ALY): Bannerton (COND: Hattah (ALY): Lake 
Mournpall, Hatlah-Kulkyne Wal. Park (SOS): 
Millewa South Bore (ALY): Halls Creek (RMA): 
Mungilli Claypan (KDA), Western Australia: {| km 
W Terhan W-H (PIM & HHE): Hini, NS Mt Aloysius 
(RSM & JED): 163km Shbyk Broome (LPB): 16km 
W ML. Aloysius JER): Loker W Mi. Aloysius JEP & 
TWE): LOmi SN Mi Aloysius (RSM & JED): 20mi, 
W Suridstone of Mt Magnet Rd (AM & MID): 
22m, WSW ML. Porrest (RSM & JED); 24k SSW 
Turee Creek HS (MPE): 28mi. NE Carnegie HS 
(RSM & JED): 66h SWhyW Docker River, 
Northern ferritory WE & TWE): Canning Stock 
Roule (EXP): Cavenazh Ra, (KTR Koonuldy Cave 
(WHO): Mevkatharra-ilhluna Pool Cunning Stock 
Renite (EXP)o Norseman (BBL): Norseman Area 
(AM & MJD): Sir Fredrick Ra. (KTR), 


Harker ciewnasis 

Tiblae Jacking erect hairs, In minors, metanotal 
wroove depressed below the level of the ameror 
revion of the propodeum: dorsal surface of petiatar 
node relatively long and (al, ils anterior lace much 
shorter than the posterior face (Pigs 8,9) Mesosoma 
uniform in colour, varying from durk red-black to 
black, anterior région. of first eastral tergite similar ip 
colour lo propedenm. gastral tergites offer With the 
irailing edge volden yellaw. the golden colour (when 
present) varying in width trem a narrew band toa 
invalving niost al the tengite. 


Dexeription prajar worker) 

Anteriong clypeal margin weakly conver (Fie. 3}. 
Dorsul surfaces ol pranotuin aod mesonalum conves 
und separated by # shallow angle: propadeum 
uniformly convex and without av distinet arte; 
petiolar node with distinet gnterior and posterior 
faces, its upper suridee varying froma broad. blunt 
angle to uniformly conves and sumenuines with the 
medial section iearly Mat (Fig. 6), Ercet hairs absent 
from scapes. petidle anc tibiae, absent or a lew 
seauicred buirs ov the ourline af head and dorsum of 
mesosomy ond faster: underside of head wih tone 
to about 30. Body varying from dark red ta red- 
black, the head and dorsal Surfaces of pronotum and 
mesonucum sometimes darker than the lateral 
mesonatum. propodeurn. legs atid petiole: gaster 
reddish black with yellow-gold banding alone the 
posterior edge of each segment which varies lrom 
being absent ¢o involving the entire visible portion af 
ihe segment. 


Proxevipaueay (ine wether) 

Anterior clypeal margin convex to broadly dngular 
(Wig. 7) Dorsal surfiees of pronotum and 
Mesonolum convex and separaled by wa shillow, 
broad angle. the posterior metanouuim ending tr the 


7 SO), SITATTUCK & A, MCARTHUR 


INnclanotal Lroove, metunoral groove distmel, 
separated from the anterior propadeum by a short 
face which varies from steep (Pig, S$) lo gentle (Mie, 
4): dorsal and posterior faces of propodeum Mat to 
weakly concave und separated by a& broud, gentle 
angle. Anterior face ol petiolar node short and 
separated from the dorsal face by a sharp angle, 
dorsal face elongate and Mat ta weakly coneave and 
sceparated from the posterior face by a broad, 
rounded angle, posterior face lat (Pips &, 9), Lreet 
hairs absent trom seapes and leys, absent or with a 
lew seallered hairs on the outline of head. 
mesesoma, pebole ated guster; underside of head 
with Up ta about 30 hairs, Body varying fram red to 
ted-black, head and sometines propodeun, petrole 
and middle and hind legs usually slightly lehter 
than the pronotum: gaster dark reddish black and 
sometintes with yellow-gold banding along the 


posterior margin of ench segment whieh varies. 


trom narrow lo iivelving the entire visible 
seymenl. in whieh case the gasler ts completely 
yellaw-pold, 


Measurements 

Workery (nm 20), CL ORO (minors) — 
(majors: FL 204mm —-4.05mm: UW Lasmin - 
4Odnun ML LGésnm — S.l4mm, MTL 2.58mm — 
Vldmm St 1L63 (majors) - 1.53 Oninorsy, SL 
2.5Umnay = 3.00mi. 


b.22 


Capunents 

Componatus tides. established by Progpatt 
(1896). is hele considered oa 
C gurecimenes i his deseription of Co mitdas and i 
is Unclear i fe was aware OF amrocine my, und il se. 
how it differed from fis speeles, Clark (1930a) 
redeseribed ©. yaiifes und sepurated il trom 
© weracineius “by the shape af the thorasa aud 
nede, and the calour ofthe gaster TO aaaedrery 
the posterior margin of the segments is tarrowly 
yellow. In nidas the whole of the Sexntents, except 
the base OF rhe frst, are entirely bright golden 
Yellow” Linropuinately, the currently available 
aleral shows that all of these chardeters ure 
highly variable, Many show an east-west clinal 
pallern. with Several changing rapidly aeross 
cential South Australia, For example, 
Co oaracieins specimens fron Western Australia 
are generally darker and lairter (especially on the 
underside of Ure head) compared to those (ron 
eastern South Australia. eastward. The western 


populations also lend lo have broader bands of 


golden-yellow on (he guster with completely black 
vasters essentially unknown. In contrast. eastern 
populations atten have ourrow bands ar hiek 
banding§ completely, the gasters being uiiforinly 


synonym of 
Coumedeniedes. Froggatl made no mention of 


black. Other characters, sch as the depth af the 
etanolal vroave and the relative leneth af the 
petiolar node. vary considerably within loeal areas 
or within single nest series. This variation sulaests 
that a single widespread and variable species is 
involved rather than bye (or more) separule species, 

Cumponolus aurecinctus 1s known from south 
cenwal Queensland, westemm New South Wales anu 
northewestem Victoria west throiigh South Australia 
and Souther Serthern Terntory to west-central 
Western Australia (Fie, 10) 1b is ground nesting, 
shows a strong preferenee ftorsandy soils and is most 
oflen found us foragers during daylight hours, One ol 
1s (AJM) has observed this species at Stockyard Plain 
and Dangeil) Conservation Park, South Australi, 
foraging in the vicinity of Cumponatiy tereteines 
The karyotype of this species was discussed by linal 
etal (1977) (us Chomponatus sps). 


Cumporoliy ceriveipes Clark 
(PIGS |) -1é) 


Camponatis (Vurnoplvinas ceriseites Clark, 
IGGK: ATR, 


Marerial examined 

Sinipes. Six workers from N, end af Reeveshy 
Island, South Australia, December, 1936, J, Clark 
(Fin ANIC, 3 in MYMA). 


Other nievial examines 

Northern Territory; ISkm S Alive Springs 
(PIM; NW Alice Springs. Arartinga (PJM), South 
Australia; (Okm WSW Lamerog (PIM): Gkim SW 
ML, Pleasant (PIM): Bamlf, Courony (PIM), Belair 
(PIM): Bridgewater (PIM): Calea (BBLy Calea. 
J0KIN SE Streaky Bay (BBL); Cape Bauer (RWT & 
RIB & BBL): Cliflon Hills Outstation (IAP & 
DHE: Coorong. Coolatoo (PIM); Coarony. Skim 
WNW Pitlochry HIS (PIM). Eyre Pen. okim W 
Wanilla (PIM): Innes Natl Pk. York Peninsula 
(PIM): Kangaroo Is.. [kn N Breakneek Cle. (PIM): 
Kanuaroo Is. N Breakneek R&R. (PIM): Mi Compass 
(BBL): Mt. Lolly (BBL), Mt Rescue CP. Jimmycs 
Well (JAP); Port Parham (BBL); Sandy Creek, MIL. 
Lolly Ranges (EYRE): Poochera (PSW) Streaky 
Bay (BRL): Vietor Harbour (PIM). Western 
Atestralias 20km 5S Candingup (SOS); Simi. Bhs 
Ruvensthorpe (RW): Cape Arid NP Yokinup Bay 
(ALB): Coalmineg Beach, Walpole-Nornaglip Natl. 
Pk (ILA & NLA); Esperance area (BBL): Greets 
Pool, William Bay Nath Pk (SOS) Junana Rock, 
Thin NW ML Ragged (RWT): Ocean Beach, 
Denmark (BBL): Redgate Beaelt,  lLecuwin- 
Naturaliste Natl Pk (SOS): Waterfall Beach 
William Bay Natl Pk (SOS) Willtum Bay Rd. 
Denmark (BBL): William Bay, Denmark (BEL) 


CAMPONOTUS WIEDERKEHRI AND PERJURUS SPECIES GROUPS 7I 


Figs 11-14. C. ceriseipes workers. Pig. 11. Head of major worker. Fig. 12. Mesosoma and petiole of major worker, Fig. 13- 
Head of minor worker. Fig. 14. Mesosoma and petiole of minor worker, 


15 


Scope Length (mm) _ 


Head Width (mm) 
Fig. 15. Distribution of scape length versus head width for 
C) ceriseipes and C. prossert minor workers. 


Worker diagnosis 

Scapes relatively short (in minors, SE < 1.5) (Fig. 
15). Posterior section of mesonotum weakly but 
distinctly convex immediately anterior of the 
metanotal groove (more so in minors, less so in 
majors); metanotal groove a shallow, weakly defined 
concavity in minors (Figs 12, 14), Petiolar node 


Fig. 16. Distribution of C. cerise/pes material examined 
during this study. 


angular or broadly rounded above, the anterior face 
at most only slightly shorter than the posterior face 
(Figs 12, 14). Tibiae and scapes lacking erect hairs, 
propodeum with more than 10> erect hairs 
(occasionally with fewer) which are scattered along 
the entire dorsal surface (never limited to near the 
propodeal angle as in C. donnellani). Anterior 


12 SO) SHATTUCK & AJ. MCARTIHOR 


clypeal margin in majors broadly convex across its 
entire width. Head same colour as mesonotum (both 
either red or black). 

This spevies is most often contised with the 
morphologically similar © proyweri, The surest way 
io separate these species is based on seape length, I 
larger minor workers of C° ceriseipey the scupe is 
relatively short conipured with similar sized 
C. prassert Workers (Pig. 15), Note, however, (hat 
this difference ts minimal or non-existent in smaller 
workers cue to allometry in this character. Other 
characters uselul in separating: minor workers of 
these taxa are the generally higher atid narrower 
peliolar node (Fiz. 14) and shiny integument in 
C. ceriveipes compared to the lower and broader 
node (Fig, 36) and duller integument in C. prossery 
The shape of the nade works well for the majority of 
minor workers while the shininess of the integument 
js more problematic due to the highky qualitative 
Walle OF and poreater variation in, this character, 


Deseription (najar worker) 

Pronolum) and mesonolum genily convex, 
metanotum distinet. propodeal dorsum weakly 
convex, sometinies a litle stronger near metanotuine: 
angle well Younded and indistinet, anterior face of 
peliolar node stright, summit narrowly rounded, 
posterior fave straight, Teebly concave near summil 
(Mv. 12)..Nnterior margin of elypeus weukly conver. 
sdiircely projecting, with a weak carina (Pig. 1) 
Posterior markin OF head, winderside of head. 
mesosoma. node and gasier with scattered long setae 
tihiae und svapes lacking ereet hairs. beued red le 
black, scape red to black, funiculus dark brown: 
provetuin ced to dark brow; Hesonotuna Fed ta-dark 
brown; petiole red to black, gaster very dark brown 
to blacks legs red to black, 


Meseription paititagk worker) 

Anterior elypeal margin convex, carina distine 
(Vig. 13). Pronotum und mesonotuni on even, broad 
convexity: Mettnotiin indistinet; anterior eewioy at 
propodeum feebly concave, posterior region straight, 
angle distinet and widely rounded. ratio of dorsum ta 
deelivity wear 2 (Pig. 14), Anterior faee of petiolar 
ode straiwht. ingined forward. sufi rounded, 
posterior face siaight (iv. 14). Posterior margin of 
head, underside of head, mesosoma, petiole anid 
waster Will) scattered Jone setac. Gbine and seapes 
lacking erect harrs, Head red to black, scape red to 
black, fiimeulus clark brown; pronolum, mesonolurn, 
propodeum and petiole cach red to black; waster-very 
duirk brow to black: legs red to blavk, 


Meas nrenrtents 
Workers (ne 94) CL O82 (minors 
HL | 42mm 3.3hmm:s WW L.2Snam 


1.24 (major), 
4.0emnan MI, 


236mm - 428mm. MPL L.S9mm — 258mm: PW 
107mm = 245mm. SE 0.68 (rajair) — 1.42 (minar), 
SLL 75mm 2 Simm. 


Remarks 


The speciinens here treated as belonging 16 this 
species show considerable variation in body colout, 
The head und mesusoma range from uniform red to 
uniform black with essentially all intermediate 
combinations displayed among the available 
material. ‘here is a weak trend for the Western 
Australian speermens to be darker and a distinel 
Irend for the Northern Territory specimens to be 
liphter. However, numeraus specimens bridge the 
gaps between these colour farms, especially within 
Westerts Australia, avid specimens nearly identical by 
(hose from the Northern Territory occur in South 
Australia along with more typical workers. 

Camponoins eeriseipes ranges from eastern South 
Australia west along the coast through Western 
Australia with two known collections from southern 
Northern Territory. [thas been found in coastal sand- 
plain heath, coastal serub, dinrestune mille. low 
scrub on a dry ride and on vegetated eoastal sand 
dines, Nests have been found ander rocks and i 
open sand and workers have been volleeted from 
pital fraps and while beating vegetation. The 
speeies has been found with nmiyrniccoplilides 
(Orthoptera) at Mount Conipass, South Austratig, by 
B, B. Lowery. 


Cantponatus daanellani sp, wv. 
(FIGS | ?-{9) 


Murer evened 

Tfolaiype. Minor worker from Kings Creek Stadion. 
Northern Territory, 23 August, 1999, S. Donnell 
sandhill (ANIC), 

Same dale us 


Papaivpes, “MWe ior workers, 


holotype tANIC, SAMA), 


Oren minerial exanincel 

Northern Verritury: 29km ESP Uluru, Uluru 
Kata ‘Vjuta (JWA): [Ske ESE Uloru. Ulori-Kuts 
Tjuta (JWA) South Australia: 3. llan WNW Mt 
Lindsay (SANPPLIJS), LE shore Serpentie Lakes 
GAP), 


Worker diagnosis 

Propodeum with at most 4 elongate erect hairs neur 
the angle between the dorsal and postenor faces. 
Pronoun and mesenotinr Mathy convex. metanotul 
grove indistinet, anterior region af prapodeal dorsum 
feehly concave. sitaight postertor Petiolar node 
browdly rounded above, is anterior face ab most only 


CAMPONOTUS WIEDERKEHRI AND PERIURUS SPECIES GROUPS 3 


1 8 at \ ‘ Pe 
a ae Ne Pe, 7 
YW aN . WK 
\ a dh 4 } 
~~ 


Figs 17-18. 0. donmmellané worker. Pig. 17, Head al anor 
Worker, Mig. 18. Mesesoma and petiole of minor worker, 


19 5, 


42d, 
te . ae 
a 3 y 
oy a4 
PB : = 
a a 
\ 
“A . 
is 
—— 
\ 
] ae 
ne mer Fal 


Vat 


Fin, 19, Distribution of CL donvellani material examined 
tiring this study. 


shebtly shorter than the posterior face (Fig. 18). 
Tibiue and seapes lacking erect setae. Anterior 
clypeal margin feebly projecting, broadly convex 
across its whole width. 

Camponotus dannellani is similar le C. arenetus in 
overall colour pattern but differs in the smaller sive 


of the minors and the Matter mesosomal dorsum with 
u less distinct metanotal groove. It may also be 
confused with smaller. paler workers of C. 
ceriseipes, but differs in having fewer erect hairs on 
the propodeal dorsum. 


Description (minor worker) 

Pronotum and mesonotum vently conver, 
metanotal grove indistinct: anterior region of 
propodeum feebly concave then straight, lacking an 
angle between the dorsal and posterior faces, ratio 
dorsum to declivity about 3 (Fig. 18), Anterior face 
of petiolar node about as long as dorsal face and 
separated from it by a moderate convexity; dorsal 
face weakly conyex and separated from the posterior 
face by a broad, rounded angle; posterior face flat 
(Fig. 18). Elongate ereci hairs scatiered on all 
surfaces of head (including underside), mesosoma, 
node and gaster, absent from scapes and tibiae. 
Anterior clypeal margin convex broadly angular 
(Piz. 17). Head, mesosoma and petiole red with 
upper surfaces of head, pronotum and sometimes 
Mesonotum infuseated with dark red-black, legs red- 
black basally, red distally; gaster dark red-black, 


Measurements 
Holotype, CLO.S89: HL 1.S8inm: HW 140mm; ML 
2.58mm; MUL 1.78mm; SI 1,32; SL 185mm. 


Remarks 


Camponotus donnellant has been encountered a 
limited number of umes in north-western South 
Australia and south-western Northern Territory. [thas 
been collected from a sand hill in association with 
Triedia spp. in the Great Victorian Desert of southem 
Norther Territory, Little else is known of its biology. 


Envinology 
Named after Dr Steve Donnellan of the South 
Australian Museum, the collector of this species. 


Camponotus gouldianys Forel 
(PIGS 21-24) 


Cumpononis gouldiants Forel, (922: (00, 


Marerial examined 
Svntpes. Two medi workers from Sea Lake. 
Victoria, both badly damaged (MHNG), 


Other material examined 

New South Wales: Balranald (JWI): ¢ 26km & 
Euston (RIK). Northern Territory: Iamurta Spr 
CP (JAF & DHI). South Australia: 10km NE 
Chilpuddic, Gawler Ranges (PJM), LOknt NW 
Ceduna (RPO); likin FE Poochera (RWT & RIB & 


74 8.0, SHATTUCK & A. J, MCARTHUR 


22 


\ 


Pigs 20-23. C. gouldicnus workers. Fig. 20. Head of major 
worker, Pig. 21. Mesosoma and petiole of major worker, 
Fig. 22. Head of minor worker. Fig. 23. Mesosoma and 
petiole of minor worker. 


Fig. 24. Distribution of CL goilelianus material examined 
during this study. 


ELO); Ilmi. E Kimba (PJM); 12km E Ceduna 
(REO); 12km E Warramboo, Eyre Pen. (PJM): 13km 
E Ooldea (JAF); I3mi. SE Streaky Bay (TGR): 
15km NW Renmark (SOS): [8km E Ceduna (RFO): 
20km E Ceduna (JAF); 20km E Paney HS, Gawler 
Ranges (PJM); 20km E Ulooloo (PJM); 20km ENE 
Umberatana (PJM); 20km NW = Minnipa (AJM): 
23km NbyW Renmark (SOS); 32km N Renmark 
(SOS); 3mi. W Penong (TGR); 41km EbyN 
Nullarbor (RWT); 45km WNW Emu. Victoria Desert 
(PJM): 4km W Wirrula (JAF); 4mi. E Oraparinna 
(GFG); 53km FE Vokes Hill, Victoria Desert (PJM); 
53km NbyW Renmark (SOS); 58km E Vokes Hill, 
Victoria Desert (PIM): 5km N Poochera (RWT & 
RJB & ELO); G0km N Colona (EXP): 60kni NNE 
Ceduna (JAF & PJM): 6km W Nundroo (RFQ); 
7.4km SW Poochera on Port Kenny Rd (RWT & 
RJB & ELO); 7.5km NW Venus Bay (SANPNS): 
79km NNW Renmark (AJM); 7kin NE Purnong 
(SANPYS): 7km SE Belah (SANPSOPS); 7km SSW 
Munyaroo CP (WKH); 7km W Inila Rock Waters 
(SANPYS); 9km N Atkindale HS (SANPSOPS); 
Aldinga Scrub (SMO); Allendale HS 9 km N 
(SANPSOPS);, Baratia 6 km NW (SANPSOPS): 
Belah 7 km SE (SANPSOPS); Blyth (BBL): 
Brookfield Conservation Park, 0.Skm S Camp area 
(SOS): Brookfield Conservation Park, Camp area 
(SOS): Buckleboo (EBR); Calpatanna CP, Eyre Pen, 
(JAF); Calpatanna Waterhole (JAF); Calperum 
Amalia (AJM); Calperum Murphys (AJM); 
Calperum NE corner (AJM); Cambrai (PJM): 
Canopus Dam (AJM): Canopus HS, Danggali CP 
(TWE & KRP); Ceduna (KCA); Ceduna 10 kin NW 
(RFO); Ceduna 18 km E (RFO); Chadee (LQU); 
Chowilla (TGW & PJM); Clements Gap CP (DHI); 
Colona 60 km N (EXP); Cooltong (GLE): Coultong 


CAMPONOTUS HIEDERKEUREAND PERIURES SPECHES GROUPS N 


CATA & MAA) Cowell (BRH): Dangygall Tipperary 
Dam (AIM); Danguali, NE eonet (AJM): Plash Jack 
Dum (SANPSOPS): Gawler Ra Lake Eyerard St 
(GPG): Gawler Ra Serubby Peak (IAM): Gawler 
Rumues (TUM): Hideaway Hut (SANPSOPS): Inila 
Rock Waters 7 kin W(SANPYS): Katarapko Creek 
(AIM), Kimba (PAL): Kokalha. Gawler Ranges 
(AIM): Kooma, Fyre Penisula (PJM): Keogamore 
(PIM): Koonamnore HS (JAP): Kyehering Soak 
(RCC): Luke Everard Stn, Gulwer Ranges (GPG): 
Lake Gilles GARY Lock (AIM), Loxton Paynes 
Varn (AJM); Loxton Snoderass Farm (AJM): 
Mambray Creek. Port Augusta (PJM): Middle Daim 
(SANPSOPS): Middleback Str. (AJO): Minnipa 20 
kin NW (AJM): Mitcherie Roekhole (SANPYS); 
Monmolata (SANPSOPS): Moorowie Plain (PJM): 
Morganvale. Damgeali CP (ASM), Mount Aroona 
(SANPNWPRS) Mount Ive (AJA & PIF): Ment 
Rescue CP (IAP) Mundoota NT (PIM Mugyaroe 
CP oY kin SSW (CW. Head): N.S.W, Coombah 
(PSW): Nundmoo (AJA & SBA); Nundroo 6 kin W 
(REO). Nundroo Roudhouse (REO) Oak Bore 
(GUM): Ooldea (AMI Doldea 13 kin E (JAM: 
Orparmnmy 4+ om F (GPG), Oraparinna. Pindebs 


Ranges (PIM Orroroo (G1G);  Pandappit 
(SANPSOPS): Paringa (SANPYVS): Poochera 


(BIO): Poochera (GPG): Poochera (RW & KIB & 
BLO): Poochera (AJM): Poochera area (RWT): 
Pooehers area (RWT & PSW): Poochera Cemetery 
(AJM & CHW): Poochera Hotel (SOS): Pooehen, 
“Freihtting site” just Sot villave (RWT & RIB) 
Pooginoak Flat (GL: Port Kenny (SANPYS) 
Purriong 7 kU NE CSANPYVS): Rockwater Rockhole 
(SANPVS); Sall Lake (PHU): Serubhy Peak: Galwer 
Ranves (IAP & WKH) Stoekyard Plain (OLE): 
Streaky Bay (BBL); Streaky Bay (IMC): Streaky 
Bay (POR), Thirty Thousand Tank (GCM y: Tinda 
Catch (SANPSOPS), Tipperary Dam. Dangesli 
CP (AIM: Venus Bay (SEG): Waikerie (BBL): 
Weding Well. Calpatanma CPL fiyre Pen. (LAI) 
Weebubbie (PAL); Whyalla (PJM & RBET): Windsor 
(HEW), Wingoohe Hill (SANPSOPS). Wirrula 4 kin 
W LAB): Wirulla ({8C Ad: VYalata (SANPNS); 
Yaris (OWA): Yelpawandinna Creek UAH & 
DD: Yookumurra (WHO): Yurbarrn CP (ALD: 
Yumburra dow fence (IAP): Yumbarra Rockhole 
(SANPYS), Vietorias 3. 3km N Millewsa South Bore 
(ALY): Hattah 6.3 kin NW CALY): Lake Huttah (IDI); 
Mildura ICM): Millawa South Bare 3.3 km N 
(ALY Sea Lake (ICG), Western Australias 10- 
T5kin N lune Rock, on Balludonia Rd (RWI): 
Hk NE Pewk Charles. Peak Charles Nath [tk 
(SOS); (0k S Balladonia (SOS\: (Omi. SP Rarone 
(RW), (thin Se Mt Ragged. Cape Arid Nath Pk 


(SOs): L60km ENE Esperance (PS W):23ki EST al 


Cocklebiddy (RWT): 23m) W Fraser Ree. TIS 
(RW): 2Smi. NbyW Balladonia HS (RW): Joni, 


SE by F Zonthus (RWT); Skin SW Mt Ragged, Cape 
Avid Natl Pk (SOS): Skin S Balladonia (SOS): GOrmi 
E Balladonia Stn. (TGR): Akim S Norseman (IED): 
Balladome &0 kin FE (AIM & SBA), Border Village 
(KMA); Cape Arid National Payk (AIM & SBA); 
Cape Arikl NP (RPP); Esperance (BBL; Bucla 
(SOS): Gora [as Goora} Hill (TGR): Jarrahsead 
UAIM & WMA): Junana Roek, Senay NW ML. Rueyed 
(RWT): Kambalda3 1.308 1IS.41F (JOM) Madura 
(AJM); Madura (IPB & MSU). Mt Raped (BBE. 
Mundrabilla Motel (AJM & SBA) Weebubbie 
(PAI): Worsley (IDM). 


Wiener iterrosis 

breet hares present on Uibiae arid scapes. Melanoial 
groove absent in miner workers, Propadeum with 
more than 40 erect short and long setae. Pubeseence 
on head and paster ubundant. wilh) widbyidual hires 
Overlapping, In profile, dorsum of peliolar node 
rounded in minor workers, a blunt angle in major 
workers, The relutively clongate bouly with abundant 
erect Huirs wall separate this species fram clase 
relatives, 


Destviption (majur workers 

Anterior clypeal imarin with aw nearhy straight but 
crenulate medidl prajcetion with abgular coruers 
(Pig, 20), Pronotum weakly convex: postenur 
mesonotum, mictanotum and dorsum of propedeum 
Nat and lone: propedal anele rounded, declivity 
straight, rina dorsum to declivity about 2 (Fig. 27). 
Anterior fee of petiolar node convex, summit blunt, 
posterior lace mostly convex (Mie 21). Ryeept for 
finiculus. entire body covered with plentiful ereer 
selac, Head red to dark brown, scape dark browe to 
black. funsculus dark brown, pronotunt red-brown, 
propodeum red-brown; guster black: Jews Hahter thar 
Mesosma. 


Deseripuien (miior Workey) 

Anterior elypeal tianin leebly convex. stronely 
pryecting, crenulale. anierior corners with wide 
ungles; medial cuotna blunt (Viv. 22). Pronerm 
fecbly convex; mesonolunr and dorsum af 
propadewm fat and long. sometimes leebly coneave. 
angle rounded. posterior faee straizht. rauo al 
dorsany ia deelivity about 3 (Fig. 24). Anterior faee 
of peuolar node convex, summit bluntly eounded, 
posterior fice Convex (Fras 23), Except lor funeutus, 
entire body covered will plentiil ercet setae. Mead 
red lo dark brown, scape dark brown te black. 
funticulus dark brown. mesesona. node. and pastes 
darker legs Hahter Uni mesosomi. 


Mecasarennits 
Morkers (fh 20), CL O86 (ninor) = LT (major) 
HL 83mm © 4.24rm; LEW LSYimm 47 linn ML 


76 8.0. SHATTUCK & A, J. MCARTHUR 


287mm — 4.9lmm; MTL 2.22mm — 3,04mm; PnW 
118mm — 2.66mm; ST 0.65 (major) — 1.60 (minor); 
SL 2.46mm — 3.08mm, 


Remarks 


This is one of the most commonly encountered 
species in this group. It occurs from western New 
South Wales and Victoria west to south-central 
Western Australia and can be found in a range of 
habitats including mallee on a number of soil types. 
In sandy soils nest entrances are at ground level 
generally close to the trunks of mallee or other tall 
vegetation. In heavier soils nest entrances are 
constructed of soil formed into a column about 30 
mm diameter and 100 mm tall with an entrance hole 
in the side near the rounded summit. The purpose of 
this turret is not known but is likely to be related to 
predator avoidance and/or lo prevent water entering 
the nest during flooding. A nuptial flight was 
observed at Waikerie, South Australia on 15 May 
1998 at 3 pm when the temperature was 25°C. This 
antis known to be the host for an unusual group of 
leafhoppers, members of the Eurymelidae 
(Hentpera). These leafhoppers live in the ants’ nests 
and forage nocturnally along with the ants (Day & 
Pullety 1999), 


Camponotus owensae sp. nov. 
(FIGS 25-27) 


Material examined 

Holotype. Minor worker from 32km NNE_ Inila 
Rock Waters, Yumbarra Conseryation Park, 31° 44' 
OL’ S 133° 26' 59" E, South Australia, 20-24 March, 
1995, TH. Owens (SAMA). 


Paratypes. Three minor workers. same data as 
holotype (1 in SAMA, 2 in ANIC), 


Worker diagnasty 

Tibiae with abundant suberect hairs. In minors, 
metanotal groove depressed below the level of the 
anterior region of the propodeum; dorsal surface of 
petiolar node relatively long and flat. its anterior 
face much shorter than the posterior face. Elongate 
(overlapping) and dense pubescence present on 
head, mesosoma, gaster and tibiae. Body colour 
black. The configuration of the metanotal groove 
and the abundant pilosity will separate this species 
from others in this species group, 


Description (miner worker) 

Anterior clypeal margin projecting, median 
portion nearly straight and feebly crenulate with 
rounded angles laterally (Fig. 25). Pronotum, 
mesonotum, metanotum and the anterior one-filth of 


25 


Figs 25-26, C. awensue workers. Fig. 25. Head of minor 
worker. Fig, 26. Mesosoma and petiole of minor worker. 


Fig. 27. Distribution of C. ewersae material examined 
during this study. 


CAMPONOTUS WIEDERKEHRI AND PERJURUS SPECIES GROUPS rz 


propodeum # strong. even domed convexity 
distorted only by the two feeble. well separated 


sutures of the metanotum, the posterior four-fifths of 


propodeum rise [rom a wide concavily to a posterior 
hump which includes the rounded angle and the 
mostly straight posterior propodeal face (Fig. 26). 
Anterior face of petiolar node straight. shorter than 
posterior face, summit narrowing upwards to a 
rounded angle (Fig, 26). Entire body black and 
covered with plentiful erect and flat lying white 
setae except antennae where setae are flat lying to 
subereet. 


Measurements 
Minar worker (n=2). CI 0.80 — 0.83: HL 2.04mm 
2.35mm: HW 1.63mm — |.95mm; ML 3.33mm 
3.89mm; MTL 2.98mm — 3.08mm; PnW 142mm — 
1.60mm: SE LSO— 1.71; SL 2.79mm — 2.92min. 


Enmology 

Named afler Helen Owens of the South Australian 
Department of Environment, Heritage and 
Aboriginal AlYairs, who found this species during a 
faunal survey. 


Remarks 


This rare species has been collected only once 
from south-western South Australia (Fig. 27). 
Specimens were collected in pitfall traps in mallee. 
Nothing else is known of its biology. 


Camponotus posicornutus Clark 
(FIGS 28-32) 


Camponotus (Tanmaemyrmex) postcornutus Clark, 
1930. 121, 


Material examined 

Syagpes, 10 workers from Bungulla and Tammin, 
Western Australia (1 in AMSA, 5 in MCZC, 4 in 
MVMA). 


Onher material examined 

South Australias Blythe (BBL). Western 
Australia: 26mi. NWbyW Norseman (RWT); 32km 
W Salmon Gums (GPB); 35km $8 Kambalda (JAF): 
38.8kin ex Murchism R-Billabong (DHK & ACK & 
WLN & RDN); S53mi SSW Coolgardie (RWT); 
71km S Payne's Find (GPB); 9mi SW Grass Pateh 
(RWT): Binneringie Road,  6km ESE 
Widgiemooltha (JAF); Bungulla (TGR); Frenchman 
Bay. S Albany (LPK); Kalbarri Natl Pk (BBL): 
Mullewa (WMW): Norseman Area (AMD & MJD): 
Parker Ra. [as Parkers] (TGR); Salmon Gums. 
70mi. N Esperance (BBL): Tammin (TGR): Tardun 
(CIM). 


28 


29 


30 


31 


Pigs 28-31, C. postcornutus workers. Fig. 2%. Head of 
major worker, Fig, 29. Mesusoma and petiole of major 
worker, Fig. 30. Head of minor worker. Fig. 31 
Mesosoma and petiole of minor worker. 


78 SQ. SHATTUCK & A. MCARTHIIR 


a 4 A 
ose hess % ~ 
aed oe ae | 
we I* 2 \ 
—_ ae 
ue \ 
f 2%, 
Wo ? SS " 
es - a 
\ | 
} - “"; a | } 
i Be i 
+" . a | 
) i 
ho 
a ‘ 
<4 
Lx 


Pig, 32. Distributton of C. pesteornmiis maternal examined 
durings Unis study, 


Worker dliegnosiy 

In minor workers, the pronotuim, Mesonotund and 
dorsum of propodeum form a strong, even convexity. 
(he metanolul groove is absent and the posterior face 
of the propodeum is only weakly diferentiated from 
the dorsal fuec. The posterior corners of the head in 
major workers taper rearward into blunt 
protuberances, The shape of the mesosoma and the 
cephalic protuberanees i major workers will 
sepurale this species from close relatives. 


Description (major warker) 

Medial section of anterior clypeal margin weakly 
projecting anteriorly with broad lateral angles anda 
fechle media! concavity: cara distinct (Fig. 28), 
Posterior corners Of head produecd as blunt homs in 
Major und medium workers (Pigs 28, 29), Pronotum, 
Mesomtolum and metanotum form au even convexity, 
propodeal dorsum and posterior fee form a separate 
even conyesliy without angle (Fig. 29), Anterior face 
of petiokiy node convea, summit moderately Sharp, 
posterior Faee straight (Fig. 29). Dorsal and 
undersides of head, Mesosoma. petiole, waster und 
coxa with sparse reddish, lone erect setae. Mntire 
body dark red-brown with the gaster darker, 


Deseripion Gnitur worker) 

Anterior clypeal margin projecting weakly, carina 
Sharp (Fig. 30), Pronotum, mesonowim and dorsuni 
OF propodeum form a reasonably even convexity, 
propodeal angle broadly rounded, posterior lace 
straight ratio oF dorsum to declivity ahoul 2 (Fig. 
31), Anterior face of petiolar node convex, summit 
hluntly rounded. posterior face convex (Mig. 31). 
Dorsal and undersides of head, mesosoma, petiole, 
gaster und doxd with sparse reddish long ereet setae. 
Entire body dark red-brown with the gaster darker, 


Measurements 

Workers (w=8) CT 1.06 — LASe HL 195mm - 
4.lomm, HW 2.06mm — 4.84mm: ML 3.28mm — 
490mm; MTL 2.16mm — 2.84mm; PrW b.7} ram 
3.13mm; $10.57 ~ 1.14: SL 235mm 2.77mm, 


Remarks 


This species is ground nesting with a simple 
entrance hole, His most common in south-western 
Western Australia with a single eollecion trom 
South Australia which is lighter in colour than those 
from Western Australia, Material is mostly trom 
relatively dry areas such as mallee. 


Camponotus prosseri sp. nov. 
(PIGS 5, 33-37) 


Material examined 

Holonwpe, Minor worker Irom Streaky Bay, South 
Australia, 30 August 1974. B. B. Lowery, mallee, it 
sund (ANIC), 


Paratypes, 25 workers. [A gueens and | male. same 
daia as holotype (2 workers und | male in SAMA, 
remainder in ANIC). 


Other material examiied 

New South Wales: Imi. S Hillston (BBL): 4m, 
N Condobolin (BBL): 628k N Coonabarabian 
(LPR): 7mi, S Hillston (BBL): Berrigan SP 
(BBL); Poonearie (RHC & YCC & AKN). South 
Australia; 20km EF Uloalao (PIM): 32kin WN 
Renmark (RRP): Thm SE Baluh (SANPSOPS): 
Aldinga (BBL): Innes Natl Pk, York Peninsula 
(PIM): tines Nath Pk. York Peninsula (PIM): 
Koonamore (PIM); Loxton Payne's Farm (AMA); 
Loxton Snodgrass (AMA): Mution Bay. Yorke 
Pen. (RSI); Poocheru (PSW); Poochera (RWT & 
RIB), Port Lincoln, 2k N Cape Tournefort 
(PIM); Port Lincoln, Eyre Pen. FP Horse Rock 
(PJM); Port Lincoln, Horse Reek (PIM); Port 
Lincoln, Spalding Cove (PIM): Port Parhani, 
Sdn. N Adelaide (BBI); Streaky Bay (BBL); 
Surcaky Bay (BBL); Yurmbarra C'P, 6m NNE toila 
Rock Waters (1OW). Western Australia: 28k 
WSW Israelite Bay, Cape Arid Natl Pk (SOS): 
30kin W Israelite Bay (GPB.& GJM): 54imi SSW 
Coolgardie (RWT): Sami. SSW Coolsardie 
(RWT); 62k NE Albany, Llassell Nall Pk (SOS); 
72km SW Norseman (SOS): 8Okm. West Talbot 
Rd. Beverley (AMD & MID): Albany (LOR); 
Balladonia and Madura (BBL): Eucla (SOS): Gora 
fas Goora] Rock (TOR): Kings Park (BBL): Mt 
Rugeed. Cape Arid NP (AHB); Norseman (BBL): 
Salmo Gums (BBL). Stirling Ra, (GPR): Stirling 
Ru. NP (GPB) 


(\AMPONOTUS WIEDERKEHRI AND PERJURUS SPECIES GROUPS 7 


35 


Pigs 33-36. C. prosseri workers. Fig. 33. Head of inajor 
worker. Fig. 34, Mesosoma and petiole of major worker 
Fig. 35, Head of minor worker. Fig. 36. Mesosoma and 
petiole of minor worker. 


37 


#a 
= 
2 
. 
o 
da, 
. 


Fig. 37. Distribution of Co prosseri material examined 
during this study. 


Worker diagnosis 

Anterior clypeal margin in major workers 
broadly convex across its entire width (Fig. 33). 
Scapes relatively long (in minor workers, SI > 1.4) 
(Fig. 15). Tibiae lacking erect hairs, propodeum 
with more than 10 erect hairs which are scattered 
along the entire dorsal surface. Posterior section of 
mesonotum weakly but distinctly convex 
immediately anterior of the metanotal groove 
(more so in minors, less so in majors): metanotal 
groove a shallow. weakly defined concavity in 
minors (Figs 34, 36), Petiolar node angular or 
broadly rounded above, the anterior face at most 
only slightly shorter than the posterior face (Figs 
34, 36). Head same colour as mesonotum (both 
either red or black). 

This species is morphologically similar to 
C. ceriseipes and is easily confused with it. The 
difference is outlined under C, ceriseipes above. 


Description (major werker) 

Anterior clypeal margin weakly convex, scarcely 
projecting, with a weak carina (Fig. 33). Pronotum 
and mesonotum gently convex, metanotum 
distinct, dorsal propodeal face weakly convex, 
sometimes a little stronger near metanotum: angle 
well rounded (Fig. 34). Anterior face of petiolar 
node straight, summit rounded, posterior face 
straight, often feebly concave near summit in 
dorsal view (Fig. 34), Posterior margin and 
underside of head, mesosoma, petiole and gaster 
with scattered long setae, tibiae and scapes lacking 
erect setae. Head red to black, scape red to black. 
funiculus dark brown: pronotum red to dark brown; 
mesonotum red to dark brown: petiole red to black; 
gaster very dark brown to black: legs red to black. 


8D SO. SMATTOCK & AL MCARTIIUR 


Description (minor werker) 

Anterior clypeal margin convey. carina distinet 
(Fig, 35). Pronotunr and mesonotum an even, wide 
convexity, metanolum iWdistinet, propodeal dorsum 
lvebly concave anteriorly, slraight posteriorly, angle 
widely rounded. ratio of dorsum to declivily near 2 
(Fig. 36) Anterior fee of petioliur nade short. Mat, 
iichined forward, sunt rounded. about as heh us 
long, posterior face short, flat (Pig. 36). Posterior 
margin und underside of heacl. mesosamu. petiole 
idl waster with seuttered long setae, ube und 
scapes lacking ereel bairs. Head and mesesoma 
clothed a fine flat-lying pubescence sufficiently 
dense in places lo hide the integument Head red to 
Klack, stape red to black, funiculus. dark brown, 
pronotum, MesenGtlun, propadeum and petiole each 
red to black? waster very dark browi to blacks leps 
red to black, 


Ves ure ens 

Workers (n=94). CL 0.72 (ntinar) -- 12) (major) 
HL 1.50mi 3.2 hii AW 1 OS8min— 4.88 Mb 
24trm -— 443m: MTL 2. betrim — 2.66mi: Pr'w 
O0%m — 2420: SLO.70 (major) — 1.76 (minor): 
SL | Yn - 27 inn, 


Hivoralay 
Named afier Dr tan Prosser. Canberra, Australia, 


Romarks 


Hoe specimens considered here as belonging to 
Ins Species Show consistency in overall head, 
inesosamal and peuolur shape as well as overall size. 
The lenuth of the scape varies but this variation is 
Melly correlated with head width (Fig. 15) as would 
be expected for a single tixon, However, these 
specimens do show considerable variation in colour 
und to a desser extent pilosity, Allowing fora lew 
upparently callow or faded individuals, all specimens 
have the head und gaster black. ‘he imesosena, 
petiole and legs, however, vary from black to vellaw- 
red, These colours show considerable variation in 
intensity with essentially all shades between Ure 
extremes present. In general most nest series. are 
fairly consistent in colour patiern with the exception 
of the petiale and dees. which can vary amony 
individuals. Clawever, the variation between series 
shows @ more interesting, pattern, The pronotin is 
generally black but is partially to completely red jria 
few collections from Western Australia, he 
mesosema and propodeum yary from black to reed 
bur this variation ocew's throughour the range of the 
species und the lighter colour fs much more 
common, especially for the propodeum where red is 
more common than black, It should be noted that the 
development of te ted colour follows a distines 


pallern, ‘The propedeum must be red for the 
mesonoium to be red. and the mesonotum must be 
red for the pronotum to be red. ‘This means that the 
most common colour pattern is black with a ved 
propodeurn followed hy black pronotum with red 
mesonotum and propodeum and finally individuals 
with a completely red mesosoma. The colours of the 
petiole und legs vary independently of the 
mesvsumia. 

The variation in pilosity is substantial bur 
generally less obvious than that found in’ body 
colour Both the erect hairs nd appressed 
pubescence vary from sparse to abundant on all 
major body regions. And as with) colour most 
Variation veeurs hetveen nest series mither thin 
Within nest series, However, mo significant 
weoeraphic pattern was detected regarding the 
development of pilosity, and there was Ho obvious 
correlation between colour palterms und pilosity, The 
only exception to this isa set of specimens trom 
south-western Western Australia whieh had abundant 
lone erect setae, I spite af this ohe group, it proved 
diMficull to sevrevate the available material ino 
subsets for which diagnoses could be deyelaped. 
There were distinct sets ol individuals which shured 
colour or pilosity patterns but there remained a 
Huniber of specinens which were either imermediate 
heiween these sets or whieh could not be pluced 
comloraibly within these sets, Asa result, all ofthese 
speeimens are here treated as helonvitiy too single, 
wide-ranging taken which shows considerable 
variation ina number of characters, with a core Urat 
some of these may well represent distinel species 
which are not diagnosable with the material currently 
available. 

Biologically. these ants have been found in 
mallee, Ce//iriy woodlands and coastal scrub. They 
are known to nest under stones as well as ih open 
soil without Covering. especially in sand. and they 
hive been taken in pitfall traps. Ihey are knowl to 
loraze on low vexetation including mallee anil 
yellow box. 


Camponotus rufanigrus sp. nov, 
(FIGS 38-40) 


Material examined 

Holatype Minor worker fron Cambrar South 
Nustrolia, 4-7 Mebruary 1972, PLM. Greenslade, 
dune Hb. (ANIC), 


FPuranpes. 8 workers, sanie dala as hololype except: 
| collected 21-25 February. 1972. dune Il, 2 
elected 7-10 Febriiiry. 1972 dune Hh | eotleeted 
25-24 February, 1972. dune thi 2 culleeted 2s 
January, 1972, dines 2 collected 18-2) Pebruary, 
)O72, dune PLE CANIC), 


CAMPONOTUS WIEDERKEHRI AND PERJURUS SPECIES GROUPS sl 


38 


39 


Figs 38-39, (! rufonigrus workers, Pig. 38. Head of minor 
worker, Fig. 39. Mesosomiu and petiole of minor worker 


40 3 


*. 
a 
" Nw. 
A a 
w jt 
\ -— 
\ penn 


Piv. 40. Distribution of CL rufonigras material examined 
during this study. 


Other material examined 
South Australia; Gawler Ra. (PJM), Yumbarra 
CP, 23.5 km NW Inila Rock Waters (HOW), 


Worker diagnosis 

Anterior clypeal margin broadly convex across its 
entire width (Fig. 38). Tibiae and scapes lacking 
erect hairs; propodeum with more than 10 ereet hairs 


which are scattered along the entire dorsal surlace. 
Petiolar node angular or broadly rounded above, the 
anterior face at most only slightly shorter than the 
posterior face (Pig. 39). Black head contrasting with 
red mesonotum, 


Description (minor worker) 

Anterior clypeal margin evenly conyex, carina 
strong (Fig. 38). Pronotum and mesonotum forming 
an even convexity, metanotum indistinet, propodeal 
dorsum concave anteriorly and flat posteriorly, 
angle rounded, declivity straight, ratio of dorsum to 
declivity about 1.5 (Fig. 39). Anterior face of 
petiolar node flat, short, summit widely rounded. 
posterior face convex (Fig. 39), Dorsal and under 
surfaces of head, mesosoma, petiole, gaster and 
coxa with sparse long erect setae. Entire body 
clothed in fine short indistinct Mat lying 
pubescence. Head, anterior of mesosoma, Most of 
node and gaster dark brown to black, otherwise red- 
brown. 


Measurements 

Miner worker (n=3). CLO.85 — 0.86; HL t.37mm 
1.60mm; HW 1.l6mm — 138mm: ML 2.19mm — 
259mm; MTL 1.53mm — 1.96mm; PnW 0.98mm — 
1.20mm: SI 1.44 — 1.55: SL 1-7S5mm — 2.14mm. 


Enmalogy 
Named afler its red and black body colour. 


Remarks 


This species is known from three localities in 
southern South Australia (Pig. 40). Two collections 
consists of single minor workers, while one (rom 
Cambrai) contains nine minor workers collected at 
six different times during January and February. 
1972. Thus this species has been rarely collected and 
then generally in small numbers. The limited 
biological information suggests that this species 
oecurs on sand. 


Camponotus setosus sp. nov. 
(FIGS 41-43) 


Material examined 

Holotype. Minor worker from Manning River Gorge, 
16°39'S 125°55'R, Western Australia, | June 1992, 
S.©. Shattuck (ANIC). 


Puratypes. 2) minor workers, same data us holotype 
(3 in SAMA. 18 1 ANIC). 


Other inaterial examined 
Western Australia} |.5km W King Edward River 
crossing (SOS). 


a2 S.O. SHATTUCK & A.J, MCARTHUR 


42 


Fis 41-42, -C. sefosus workers. Fig. 41. Head of minor 
worker. Fig. 42. Mesosoma and petiole of minor worker. 


43 ; 


“2% fn te dt 
« gf i 4 
wet . % Voy, 
eed ke Nye i 
ag* * ye \, 
f \ 
ee mh 
ey 
b 
4 r 
) - | 
a 
1 ae 
\, . { 
i ~ =e , ,' 
id — * 4 i 
~ 0 
| 4 
ie “ 
‘ant 


hig. 43. Distribution of CL yetesuy material examined 
during this study, 


Worker diagnosis 

Erect hairs present on tibiae. Metanotal groove a 
distinct, shallow trough. These two characters will 
separate this distinctive species from others in this 
group. 


Description (minor worker) 

Pronotum and mesonotum form together an even, 
raised convexity followed by the angular trough of 
the metanotum, the weakly convex dorsal surface of 
the propodeum, a widely rounded angle and the 
straight posterior face (Fig. 42). Entire body covered 
with dense flat lying pubescence, erect setae absent 
from antennae. Pubescence on posterior of gaster 
yellow, elsewhere white. Gaster black, most of head. 
mesosoma and node black, the remainder with red 
patches; antennae dark brown: coxa and femora red. 
tibiae and tarsi brown, 


Measurements 

Workers (n=4). CI 0.85 — 0.88; HL LS8mm 
|. 96mm: HW 164mm — 1.69mm; ML 3.08mm — 
3.20mm; MTL 2.34mm — 2.54mm: Paw 1.50mm — 
1.54mm:; SI 1.45 — 1.57: SL 2.45mm — 2.62mm. 


Fiypmology 
Named alier the abundant long setae present on 
most regions of its body. 


Remarks 


This apparently uncommon species is restricted to 
the Kimberley region of Western Australia (Fig. 43). 
All known collections consist of ground-foraging 
workers in open Eucalypris woodlands. 


Camponotus terebrans (Lowne) 
(FIGS 44-48) 


Formica testaceipes Smith, 1858: 39 (preoccupied 
by Leach, 1825: 290), 

Camponotus testaceipes — Mayr, 1862: 662. 

Formica terebruns Lowne, 1865: 278 (first 
available replacement name for Formica testaceipes 
Smith) — Mayr, 876: 65. 

Camponotus (Myrmoturba) latranenlis 
vicioriensis Santsehi, 1928: 479 — MeArthur ev al. 
L998; 587. 

Camponotus (Tanaemvrmex) myoporus Clark 
1938:379 — McArthur et al., 1998; 587. 


Material examined 

Formica testaceipes: Syntype workers from King 
George Sound, Western Australia (BMNH - see 
McArthur ef al. (1998)). 

Formica terebrans: Syntype workers and queens 
from Sydney. New South Wales (see McArthur ev al, 
(1998)), 

Camponotus (Myrmotirba) latrunculus 
victoriensis; Syntype workers and males from 
Elsternwick and Belgrave, Victoria (see McArthur er 
al. (1998)). 

Camponotus (Tanuemyrmex) myoporus: Syntype 


C4AMPONOTUS WIEDERKEHRI AND PERJURUS SPECIES GROUPS 83 


ary, 


46 


47 


Pigs 44-47, C. terebrans workers. Pig. 44. Head of major 
worker. Pig. 45, Mesosoma and petiole of major worker, 
Fig. 46, Head of minor worker. Fig. 47. Mesosoma and 
pefiole of minor worker 


Pig. 48. Distribution of CL ferebrans material examined 
during this study. For additional material see MeArthur 
ef th. (199K). 


workers from Reevesby Island, South Australia (3 in 
MYVMA, 611 ANIC - see MeArthur ev af. (1998)). 


Other material examined 
See McArthur ef a/. (1998). 


Worker diagnosis 

Erect hairs present on scapes and tibiae. Metanotal 
groove weakly developed and essentially absent 
(Figs 45, 47), Propodeum with 10 to 25 erect hairs. 
Pubescence on head and gaster sparse, with 
individual hairs generally oon-overlapping or at 
most only slightly overlapping. In profile, dorsum of 
petiolar node angular in both minor and major 
workers (Fig. 45. 47). These characters will separate 
this taxon from close relatives. especially the 
morphologically similar C. gouldicnus. 


Description (major worker) 

Medial section of anierior clypeal margin straight, 
projecting anteriorly with rectangular lateral corners, 
crenulate; carina indistinet (Fig. 44), Pronotum and 
mesonotum weakly convex: metanotum distinct as 
two parallel, transverse grooves; dorsal surface of 
propodeum straight, angle well rounded. posterior 
face mostly straight, length of dorsal and declining 
faces about equal (Fig. 45). Anterior face of petiolar 
node convex, summit sharp, posterior face mosily 
straight (Fig. 45). Entire body with plentiful long 
erect setae tending to suberect on tibiae and scape, 
absent from funiculi. Head red-brown to black. 
funiculi lighter, mesosoma and node yellow to 
brown, gaster darker than mesosoma. legs lighter. 


Description (miner worker) 
Anterior clypeal margin with median section 


Ka $.O. SHATTUCK & A, J. MCARTHUR 


convex and strongly projecting, carina distinct (Fig. 
46), Pronotum and mesonotum mostly weakly 
convex: the smallest workers without a metanotal 
groove; dorsal propodeal surface straight, angle well 
rounded, posterior face straight, ratio dorsum to 
declivity exceeds 2 in smallest workers (Fig. 47), 
Anterior and posterior faces of petiolar node 
generally parallel, summit bluntly convex (Fig. 47). 
Entire body with plentiful long and short erect setae 
iending to suberect on tibiae and scape, absent from 
funiculi. Head brown, funicuh lighter, mesosoma 
and node yellow to brown, gaster darker than 
mesosoma, limbs lighter. 


Measurements 

Workers (n=20). CL 0.85 (minors) - 1.11 (majors): 
HL |.36mm — 3.28mm: HW 1.15mm— 3.64mm; ML 
2.07mm — 3.64mm; MTL 1.56mm — 2.39mm: PnW 
0.91 min—2,02mm: SI 0.66 (majors) — 1.54 (minors); 
SL 1.77mm = 2.39m1n. 


Remarks 


Camponotus terebrans is common in sandy soil or 
disturbed sites across much of southern Australia 
(Fig. 48). Nests are sometimes located adjacent to 
the trunks of trees or shrubs with abundant excavated 
soil deposited around the numerous entrances. In 
some cases excavations have been observed to 
apparently damage or kill nearby shrubs. In other 
cases nests and their entrances are in open areas and 
lack mounds, Colonies may be very large and 
sometimes have “highways” leading to trees and 
other colonies. This species is often found in 
association with Oevris spp. butterflies (Braby 2000). 
For additional details see McArthur ef er. (1998). 


Camponotus versicolor Clark 
(FIGS 49-54) 


Camponotus (Myrinosaulus) versicolor Clark, 
1930h: 122. 


Material examined 

Svanvpes, Workers from Emu Rocks, east of 
Ongerup, Western Australia (6 in ANIC, 3 in MCZC, 
3 in WAMP, 5 in MVMA, 3 in BMNRA). 


Other material examined 

Western Australia: 33mi. SbyE Karonie (RWT); 
Oni. E Newdegate (TGR); Bungulla (TGR); Emu 
Rock (HRE); Newdegate (HMC & TGR): Norseman 
(BBL), 


Worker diagnosis 
Tibiue and seapes Jacking erect hairs. In minor 
workers, metanotal groove angular to. slightly 


49 
0 


I 


50 


51 


J 


52 


53 


Pigs 49-53. C. versicn/or workers. Fig. 49. Head of major 
worker. Fig. 50. Mesosomiat and petiole of major warker, 
Fig. SL. Head of minor worker. Pigs 52-53, Mesosoma 
and petiole of minor worker, 


CAMPONOTOS WILDERKETREAND PERIORUS SPLCLES GROUPS a5 


hig Sd. Distribution of Co versiralor omterial exaniqeal 
during (hrs study, 


depressed below the amerior region of the 
propodeum (Figs 52, 53); dorsal surface of petiolar 
node i iinors pelatively lone and flat lo weakly 
convex, Us unterior face much shorter than the 
posterior face (Figs 52, 53). Mesosoma black and 
wilh at least the first two gastral tergiies red and 
distinedly lighter i colour than the propadeum, 
gastral tergites never will) zolden-yellow bands, The 
configuration of the metanotal groove combined 
with the distinctively colouted gaster will separate 
this species Irom close relatives, 


Description (major werker) 

Dorsal surfaees of pronotum and mesonatim 
convex and separated by a shallow angle, 
propodeum uniforinly conver without a distinet 
angle; peltiolur node with parallel anterior ancl 
posterior faces, its upper surface slightly elongated 
Nut to Weakly conver (Pig. 50). Erect hans sparse ov 
outline af head including tnderside, scattered on 
mesosomi, petiole, coxa and waster, absent trom 
libiae und scapes, Anterior clypeal margin weakly 
convex (Fiz, 49), Body red-black, head and petiole 
slightly lighter than mesosonia; gaster wilh the first 
two tergites red. the remainder red-black. 


Description (minor worker) 

Anterior elypeal margin convex (Pig. 51). Dorsal 
surfaces Of pronotum and mesonotum convex and 
separated by a shallow, broad angle; metanutial 
eroove either a broad angle (Fig. 53) or a shullow 


trough (Pig, 52); dorsal and posterior laces of 


propodeum Mat to weakly convex and separated by 
itmosta gentle dnyle, Anterior lace of petiolarnode 
short dnd separated from the dorsal face by a distinet 
angle. dorsal face clongate and flat to weakly 
convex and separated from ihe posterior lace by i 


broad, rounded angle. posterior face (Mal (Pigs 52, 
53). Erect hairs abundant on outline and underside 
of head. mesosoma, petiole, coxa and waster: erect 
hairs absent from scapes and tibiae, Body dark red- 
black or black with the head sometimes slightly 
lighter: gaster with at least the first two tergites red 
and the remainder dark red-black, or sometimes 
entirely fed, 


Mleesurements 

Workers (9-7). CLOUB2 (minors) — 1.06 (majors): 
HL 223mm 3.20mm: HWW 1 s83mm  3.42mm; ML 
306mm — 486mm; MEL 2,729 — 3.00mi Sd 
145 (majors) - 1.60 (minors); SL 2,93mm 
4.9570. 


Remarks 


Camponotus versicolor 18 ad uncommon species 
which is limited to a narrow band across southern 
Western Australia (Pig, 54), It is most similar to 
C) aurocinetus and can be separaled from it by the 
darker body colour and red pasiral tergites, Minor 
workers of Co aurocinetus also have larger numbers 
of erect hairs on the head and mesosoma compared 
lo this Species, Essentially nothing is) known 
concerning the biology of C. versicalur, 


Camponotus wiederkeliy Forel 
(FIGS 55-54) 


Cumpanons wiederkelw) Povel, 1894; 232, 

Campononny dentioulats Kirby, 1896: 204 - 
Clark, !930@ 19 (worker redeseribed), New 
synonymy. 

Camponotus (Mivemoarurba) tearunculus Wheeler. 
1915: B14. New synonymy. 

Camponanis wicderkelui lucidion Vorel, W100: 81 

Crawley, T9715; 136 (queen description), New 
synonymy. 


Adeiterial enemineed 

Campanas wiederkelrr Syntype workers from 
Charters ‘lowers, Queensland (MHNG). 

Camponotus denticulatus: Syntype workers from 
MacDonell (as MeDonell) Ranges. Northert) 
‘Territory (2 in MCZC, 1 in MVMA), 

Camponotus (Myrmoturba) fairunculus: Synlype 
workers from ‘fodmorden, South Australia (1 in 
SAMA). 

Camponotus wiederkehri ducidiars Symype 
workers and males from Tennant Creek, Northern 
Territory (3) workers in MCZC. 2 workers in 
MHNG). 


Other material examined 
New South Wales: Waukerou (RIIM). 10 mi N 


8G 5. O. SHATTUCK & A. J. MCARTHUR 


Z 


KY 


56 


57 


58 


Migs 55-58. C wiederkehrt workers. Pig. 55. Head ol major 
worker, Piz. 56. Mesosama and petiole of major worker. 
Fig. 57, Head of minor worker. Vig. 58. Mesosoma and 
petiole of minor worker, 


59 


Fig. 59, Distribution of CL wiederkehri material examined 
during this study. 


Broken Hill (RHM). Northern Territory: |.Skm N 
Alice Springs (PIM & RJW): 12km SW Katherine 
(PJM); |5km S Tea Tree (MMA & JHA); 20m1. SE 
Anthonys Lagoon (TGR): 25km S$ Andado Stn 
Rodinga Ra (JAP & DHT); 35km S Darwin (LHD): 
37km E Wallara Ranch (SOS); 3km E Serpentine 
Gorge (SOS); 50km WNW Hermannsburg (SOS): 
7km W Timber Creck (MMA); Alice Springs (CBA): 
Alice Springs (WLB); Alice Springs (WCC); Alice 
Springs (LHI); Alice Springs (PPL); Alice Springs 
(RRO); Batten Ck., 30km WSW Borroloola (JEF): 
Bing Bong HS (JEF): Bitter Springs Creek (JAF & 
DH1); Bullita Outstation (MMA): Camfield (LAR): 
Colyer Creek, Skim N Alice Springs (SOS): 
Corroboree Rock, 2(km E Alice Springs (SOS); 
Darwin (SMO); Darwin (HWE); Doyles Ridge nr. 
Birdum (TGR); Flying Fox Creek (SMO); Glen 
Helen (SOS); Helen's Ck., Banka Banka Rd. (TGR): 
Wlamurta Spring (JAF & DUI); Jasper Gorge (TAR): 
Katherine (RVS); Kings Canyon Nat, Pk. (SOS): 
Kings Creek Caravan Park (SDQ): Kulgera (JBS): 
Kunoth Paddock, 30km NW Alice Springs (WAL): 
Kunoth Park nr. Alice Springs (PIM & WLO): 
Macdonnel Downs (SAMA Exped.); McArthur R,. 
4skm SWbyS Borroloola (JEF),; Narwictooma 
(AWF): NW Brunette Downs (TGR); Phillip’s River 
(TGR); Port Darwin (WDD): Rimbija Is,, Wessel 
Islands (EDE): Rimbija Is.. Wessel Islands (TAW): 
Roderick Creek (JAR); Ruby Gap Gorge (JAT & 
DIH1); Tennant Creek (JFF); Trephina Gorge Nature 
Park (JBS); Trephina Gorge, 55km ENE Alice 
Springs (SOS); Turnall into Ormiston Gorge (SOS): 
Umbrawarra Gorge (JAR & TAR); Valley of Winds, 
The Olgas IEF & TAW); Victoria River (BRH); 
Yulara, campground (SOS). Queensland: |.5kim 
WNW Riversleigh HS, nr. Gregory R. (JAF); 106m: 


CUMPUNOTES WILDERRE THRE ASD PERIURUS SPECIES GROUPS #7 


NW MIL Isa (PGR): WOmi WoMt. Garnet (BBL), 
Lom), ESE Gilbert R, Crassiig, k of Croydon (IED): 
Ih, ESE Emerald (FED): Jom. & Carpentaria 
Downs HS.OSE Einasteigh (ED): Imi, SE Lacraine 
HS (ED) 2km S Woodstock (PJM), 2s8ini, N 
Thorntonia HS, NE of Cayooweal (JED): 2mi. SE 
Camel (hk, Its. W of Ingham (IED): 2ou, SE Mary 
Kathleen (IFD); 4mi. WE Oorind? JED): SUmi. N 
Julia Creek (REL). s2kin S. Woodstock (PIM): Simi. 
W Lotus Vole TIS, N oof Normanton (JED): 7kin E 
Charters Towers (PJM). 9mi. NE Cantooweal (IED): 
Barcalchne (Gb: Blackall (IBS): Carpentaria 
Downs (JED): Charters Towers: Clermont (ABL): 
Cookuwwn (MO) Dalonally, nr Cloncurry 
RK, WED), Doomadgee Mission Station (PAL & 
NEP) lonerakl (AC): Emerald (JHA); Emerald 
Distriet (SAM): Greenvale (IED): Greenvale Station 
area (SAID): Helenslee (TOR): Homestead (FHI): 
Jergho (WAC! Mareeba (BRL): Morniigton 
Misstou (PAT & NBII) ML Isa GRO) an, Dibaba 
(RWT & JEP) Quilpie GSM) St. George (BBL): 
Set oR. Crossing CSALDe Surbiton (PAC) 
fowasville Charters Tenvers Ree CUGRo: Edita LS, 
NE ot Carmoowen! (IED); Winton (FAC), South 
Austeatias (Oka) Wo Mabel Ck. (PIM: [km NN 
Maryinna Hill (SANPPITI): [55km N Cook JAF): 
Jkin BNE Pipalyatara (SANPPITI): 26nn cS 
Kunyiinnd (SANPPIT): S3km FE Vokes Hill, 
Vietoria Desert (PIM). GOkm S Pimba (MAA): Tin. 
Ly Wilzenu (TOR): 8Okin E Ramu Junelion. Vietoria 
Desert (PIM), Andumooka SAPD: Antunes (DCO); 
Argona Damn (AJM & IDE: Belah (SANPSOPS); 
Birthday Hill, N Tareoola (PIM); Blood Ch. (CBA); 
Hox Creek (AIM & JDEQ: 22km S Belt (JEN); 
Cliflon Hills Outstation IAP & DET: Coober Pedy 
(BBL). Copper Hil (HIFRI: Curdimurka, L. Pyre 
(BBL); Davenport Range (AJM & MAA): Douglas 
Creck (MAA); Dulkaviina (PCO): Eraabells 
Mission (NBT) Ermubella Mission Stn. (RBL): 
Everard Park (IFT): Farina (PIM): Gawler Ranges 
(PIM): Unileaway Hut (SANPSOPS): Lake Eyre 
(BBL): Luke Cairdner (AAS & MLS): Mabel Ck 
CPC), Minilli (SANPPIEA): Mitehell Nab 
CSANPPTVT) Mt, Coopering (SANPPITA); Mr. 
Finke (PIM & JAR) Muserive Ranges (BBL); 
Neurutard (SANPPIT); Ooldea (AML: River 
Diamenting (AMM): Rebertstown (SANPSOPS): 
Ronald Well (SANPPLEI) S end of L. Windabout 
(BBL): Sereevh Owl Creek (WMC): Che Twins KS 
(RSM): Vokes Li GIAT): Vokes In (GPO); Vokes 
Hill, Vietoria Desen (PIM Wonitkita Bole, 
Mussraye Ru, (SANEPETS): Woocalla (RSM); 
Yarndea (AJM & PT), Western Australia: 10kin 9 
Souther Cross (MIMj)c LOGkin SEbyE Broome 
(1B) Tikin N Wiluna (PDA & SRM): 1o3kin 
SUhyl Broome (FB) 45ni cS Onslow (GCA): 
s0km ® Kalgoorlie (PIM) ASmi SSW. Coolsardic 


(RWT) 7km FE Kalgoorlie (JER), Thm W 
Kununuira, Balidicoot Ra. (DCE & JBA) Ashburton 
Royer (RUM & GCA}: Baleo Mission (ARP): 
Balladonia (BBL): Black Stove Range (KTR): 
Caneyrass, NNE Kalvoorlie (JED): Derby (WDD): 
Jigalong (IHD; Kalgoorlie (PAL: Kaluoorlic [as 
Kalgoohe| (TGR); Kalurmburu Mission (MDA): 
Kimberley area ne Kalumburu Mission (<3) ni) 
(WLE): Kununurra beat ramp (REM & CCA): 
Latrrange Mission, 120km S Broome (KMC); 
Lydon RK. Carnarvon (RHM), Lyndon River. 
Carvarvon (RAHM): Meekatharra. Mt. Newman mid: 
Gascoyne R. (PIM): Mitehell Plateau (miming camp) 
(DCE & IBA): Moolit Bulla (NBT); Onslow (REM); 
Ord R. (SAH): Pilgancoor Mining Centre (SBY); 
Pindar (CTM): Port George iy (JRB); Roehourne 
(WDD): Windjana Gorge NP ¢PSW). 


Whar ker flereorsis 


Anterior clypeal margin in major wearers 
projecting. the central region stranght with 


rectangular sides joining the lateral regions (Pig. 54). 
Posterior section of mesonotum tat (or nearly so) 
Womediavely anterior of the metanotal “roove, 
metanolal vroove essentially absent or weakly 
developed in miners (Fig. 58), 4 broad. shallow 
angle in majors (Fre. 56), Pesolar node angtilir ey 
broadly reyinded above, the unteriar face al ost 
only slightly shorter than the posterior face (Figs 5h, 
48). Tihiae and seapes lacking erect hairs. 


Deneeiptian Gndjor worker) 

Medial section of anterior clypeus stronuly 
projectine, is margin strareht and lateral corners 
broadly angular carina weak (Fig. 35). Pronoun 
and imesunotun) a slightly raised even convexity: 
melanolum with two distinel groves, the anterior 
seclion oF the propodeal dorsum leebly concave 
anteriorly und feebly convey posteriorly. propodeal 
angle widely rounded, posterar face mostly straight 
ratia of dorsunt to deelivity about 1 (Fig. Sa) 
Anrerior and posteyior faces af peolar node strtaleht 
simi Plat, narrow and -shalp, sometimes bide) tate, 
WS posterior imarpll feebly coneuve (Piz. 56). 
Dorsum and underside of lead, mesosoma, petiole, 
coxa and vasterwith plentiful sentteree erect setacy 
reduced wumbers on propodeal angele and deelivity. 
absent front scapes, Hat byte Gn obiac, Head yellow- 
red to dark brown. antenmae red 4 red-brown, 
mesosoma and node vellow-red to brows waster 
daiker legs Hhter, 


Deseriphan (nider worker! 

Medial section of anterior clypeus strongly 
proyecuing, its margin convex, ctenulate: earmna 
distinet (Fig, 57). Pronotum weakly conves, anterior 
scetion of mesonotum weakly conves, the remainder 


SS SO. STIATTLOR & AJL MCARTIIUR 


joins with propodeal dorsum to form a long flat 
surface ending ina widely rounded propodeal angle 
and short posterior face, ratio of dorsum (o declivity 
ubout 3 (Piz. 58). Anterior face of petiolar node 
Inostly convex. summit sharp (in front view pointed), 
posterior face niostly fat (Pig. 5%). Dorsum and 
underside of head, mesosama, petiole, coxa and 
waster with scattered long setae; reduced numbers on 
propodeal angle and declivity; absent trom Hhiae and 
seapes. Late body clothed with fine pubeseence. 
Mesosoma yellow-red to dark red-brown, sometimes 
with darker ov lighter patches: head, node and gaster 
venerally darker, les lighter 


Neus POMiOady 

Harker (i 20) CL ORO (minors) 
HL U.5limm 333mm. IW 1 .2tmm- 3.6)mm: ML 
25huin- 323mmeMTL 192mm 2-62mm: PnWw 
(97mm — 2.1 3mm; S1.0.68 (majors) — 1.60 Gniturs)- 
St 144mm 245mm. 


1,08 (majors): 


Remurks 


This is one ofthe most commonly encountered und 
widespread species ip this uroup (Pig, 59). In 
southern Australia nests ate generally mounds 
approximately [50 to 200mm in diameter wath 
steeply sloping sides and a fat summit with the 
entrance ina Slight depression in the centre. These 
mounds are ofien decarared with small stones, Nests 
are offen in heavy soil in open areas undoare less 
eommon ar are absent from areas of hit rainfall. 
Oflen several inounds may be seen withih a few 
metres Of cach other. 

Morphologically, this species (as conceived fete) 
shows minimal varialion in body shape and pilosily 
(uther than that expected for a polymorphic taxon) 
hut dows show considerable variauion th colour, The 
eolour ranges from clear yellow-red to black with 
essentiolly all grades of colour in between. In most 
cases the colour is uniform within an jadividual bu 
various degrees of i fuseation oi the Mesesema are 
common. Also, most variation occurs belween rather 
than within pest series although the development of 
infleution does vary within nest series, Finally, this 
colour variation shows Dtthe geographic pattern with 
essentially all colour forms being found in all 
regions, (he only eXception being northern regions of 
the Northern ‘lerritory where fight lors 
predominate. 

‘The wpes of © wreederkefrd and ©) wiederkehel 
Jnéldier copresent the more lihly coloured forms af 
this taxon, ‘These lwo taxa were separated based ut 
wivial and won-signiticant differences in sive. 
sculpruring and the shape of ihe anterior clypeal 
margin (Forel 1910) and they clearly represent the 
sume Gaon Cumpanons Jadrumeulis represents an 


intermediately coloured form and compares well 
with the types of ©. wlederkehd, Wheeler (1915) 
Was apparently unaware of Co wieelerkedied as he 
mide fo mention of it in his deseription of ©, 
latrunenius and this is likely the cause of this 
synonymy. The final previously proposed mame. 
dewticularns. represents the dark form of this taxon, 
However, it is morphologically very similar to the 
other forms placed here and no justification could be 
found for treating if as a separate layer. 


Species of the C. perjurus species erony 


Camponotus perjurus sp, Nov. 
(HIGS 40-62) 


Material cvuntiied 

Halaivpe Minor worker from 74 km E by N Cosme 
Newberry. Westert, Australia, 13 November 1977, 
Jf. Feehan (ANIC). 


her material examined, South Australias kok 
NNF Ceduna WAP): Emu Camp, Victoria Deserl 
(PJM): Mt. Gunson, SEE Woamera (PIM). Westera 
Australia: 40kni SE Ravensthorpe (RW), Borden 
(EFR), 


Worker didgniasis 

Noud of minor worker produced upwards so urat ils 
Hitadhment to the prondtum is. well below tts uppet 
mari (Fig, OL). Often with Weak purple or green 
indescent hue on head and body. The altuchment of 
the head is unique to this speeies group. if rot the 
venus, and will readily separate (his speeis fran 
others, 


Deseription fainar warker) 

Anterior elypeal margin wide. projecting, evenly 
conver und feebly cremilate, with a Feeble mieiial 
carina (Fig. 60). Pronotuim and mesonatum i raised 
convexity which smoothly joing the feebly concave 
dorsal surface of the propodeium, the propodeal ingle 
rounded, its posterior face short and straight, the 
ratio of dorsum ty declivily aboul 4 (lig. 61), 
Metanotal spiracles high. near the dorsal resosoniil 
surfice. Petiolar node leaning forward. parallel 
anteriorly and posteriorly, with a Tony, weakly 
conver summit (Fig, 61). Body red-brown except for 
vaster und parts of legs which are darker, sometimes 
wilh a@ weak purple or green iridescent fue, Entire 
hody elothed in fine while indistincl pubescence with 
sparse Jong seme on the anterior and posterior of 
head. mesosoma, petiolur node andl pester, absent on 
the underside of head 


Measurements 
Minor worker (nS) CLO7Y YS. TL Lam 


CAMPONOTUS WIEDERKEHRE AND PERJURUS SPECIES GROUPS 


60 62 


61 


So 


\ 


Figs 60-61. C\ perjirus workers. Fig. 60. Head of minor 
worker, Fig. 61. Mesosoma and petiole of minor worker. 


2.3lmm: HW 172mm — L-&4mm: ML 2.44mm — 
3..imm; MTL 2.32mm — 2.43mm; PoW L.4timm — 
1.54mm: SI 1.22- 1,28; SL 2.14mm — 230mm. 


Etnmoalogy 
From perjurns, to lie about one’s true nature, 


Remarks 


‘This species appears to be a mimic of members of 


a] 


Fig. 62. Distribution of Cy perjirns material examined 
during this study, 


the /ridomvrmex purpureus species group (subfamily 
Dolichoderinae), This is based on the purple or green 
iridescent colour which ts similar to /rieomvrmex 
viridiaeneus Viehmeyer (Shattuck 1993). Also, only 
single foragers have been found and most of these 
have been collected in assoctation with /ridomyrimex 
spodipilus Shattuck and Camponotus prosseri 
Shattuck and MeArthur. They have been found from 
central South Australia west into south-central 
Wester Australia (Fig. 62). 


Acknowledgments 


We would like to thank the following for providing 
comments on earlier versions of this manuseript: A. 
Andersen, B. Halliday and two anonymous 
reviewers. The illustrations were prepared by N. 
Barnett, Financial support was provided by the 
Australian Biological Resources Study, CSIRO 
Entomology and the South Australian Museum, 


References 


(2000) “Butterflies of Australia, their 
Melbourne: 


Brauy, M. F 
identification, biology and distribution”. 
CSIRO Publishing, 1008 pp. 

Crark, J. (1930a) New Pormicidac, with notes on some 
little-known species. Proc. R. Sac. Vie, 43, 2-25, 

= — (1930b). Some new Australian Formicidae. 
Proc. R. Soe. Vie. 42, 116-128, 

(1938) Reports of the McCoy Society for Field 
fnvestigation, and Research. No. 2. Sir Joseph Bank 
Islands. Part |. Formicidae (Hymenoptera), Prac, h. Soe. 
Ke. 50, 356-382, 

Crawney, W.-C. (1915) Ants trom north gnc central 
Australia, collected by G. PF, Hill. Part bo timn Mag. Nev 
Hist. (8)T5, 130-136, 


Day, M. F, and Pullen. K. R. (1999) Leathappers in ant 
nests! some aspects of the behaviour of Pogonoscopini 
(Hemiptera: Eurymelidae). The Victorian Natiralist 116, 
12-15, 

Forer, A, (1894) Quelques lounnis de Madavascar 
(récaliées par M. le Dr. Valtzkow): de Nouvelle 7élande 
(récoltées par M. W. W. Smith); de Nouvelle Calédonie 
(récoltées par M, Sommer); de Queensland (Australie) 
récollées par M. Wiederkehr: et de Perth (Australie 
occidentale) récaltées par M. Chase. dim, Soc. Entamal. 
Belg, 38, 226-237. 

_ (1910) Formicides australiens regus de M. M. 
Froggatt et Rowland Turner. Rev Suisse Zoal, 18, 1-94 


OU) S.O. SHATTUCK & A. J. MCARTHUR 


(1922) Glanures myrmécologiques en 1922, 
Rev, Siiisse Zool. 30, 87-102. 

Frogcatr, W, W, (1896) Honey ants. pp, 385-392 In 
Spencer, B. (Ed.) “Report on the work of the Horn 
Scientific Expedition to Central Australia, Pt. 2 
Zoology”, Melville, Mullen & Slade, Melbourne. 

IMAl, H. T., Crozier, R. H. and Taylor, R. W. (1977) 
Karyotype evolution in Australian ants. Chromosoma 59, 
341-393, 

Kiray, W. F. (1896) Hymenoptera. pp 203-209 /n Spencer, 
B, (ed.) “Report on the work of the Horn Scientific 
Expedition to Central Australia, Pt. | supplement”, 
Melville, Mullen & Slade, Melbourne. 


Leach, W. E. (1825) Descriptions of thirteen species of 


Formica and three species of Culex, found in the 
environs of Nice. Zoological Journal 2, 289-293, 


LOWNE, B. T. (1865) Contributions to the natural history of 


Australian ants. Entomologist 2, 275-280, 
Mayr. G. L. (1862) Myrmecologische Studien. Verh. Zool.- 
Bot. Ges. Wien 12, 649-776. 
(1876) Die australischen Formiciden. Jowrnal 
des Museum Godeffroy 5, 56-115, 


(1886) Notizen tiber die Formiciden-sammlung 
des British Museum in London. Verh. Zool,-Bot. Ges. 
Wien 36, 353-368. 

McArtuur, A. J., ADAMS, M. & Suarruck, 8, O, (1998) A 
morphological and molecular review of Camponotus 
terehrans (Lowne) (Hymenoptera: Formicidae). Aust. J. 
Zool. 45, 579-598. 

SANTSCHI, F. (1928) Nouvelles fourmis d’Australic, Bull, 
Soc. Vaudoise Sci. Nat. 56, 465-483. 

Sharruck, S. O. (1993) Revision of the /ridomyprmex 
purpureus species group (Hymenoptera: Formicidae). 
Invertebr. Taxon. 7, 113-149. 

(1999) Australian ants: their biology and 


identification, Monographs on Invertehrate Taxonomy 3, 


226. 

Sarr, F. (1858) “Catalogue of hymenopterous insects in 
the collection of the British Museum. Part 6. 
Formicidae”, London: British Museum. 216 pp. 

WHEELER, W. M. (1915) Hymenoptera. Trams. R. Soc. S. 
Aust, 39, 805-823, 


STRATIGRAPHY OF THE LAKE MALATA PLAYA BASIN, 
SOUTH AUSTRALIA 


By A. DUTKIEWICZ* & C. C. VON DER BORCHT 


Summary 


Dutkiewicz, A. & von der Borch, C. C. (2002). Stratigraphy of the Lake Malata Playa 
Basin, South Australia. Trans. R. Soc. S. Aust. 126(2), 91-102, 29 November, 2002. 
The 19 m-thick Late Quaternary stratigraphic sequence within Lake Malata, Eyre 
Peninsula is dominated by autochthonous gypsum, present as relatively mud-free 
gypsarenites and gypsum-clay laminae overlying a skeletal peloidal grainstone of the 
Bridgewater Formation near the base of the lacustrine succession. Calcite and 
dolomite mud are minor components of the column and several metres of these 
deposits appear to have been deflated into marginal lunettes. The skeletal peloidal 
grainstone has been severely modified by dissolution and formation of phreatic 
calcite, dolomite and gypsum cements under alternating pluvial and arid conditions. 
Discrete units are separated by disconformities and attest to rapid changes in climatic 
and hydrologic conditions over the lower Eyre Peninsula, commencing with 
emplacement of the Bridgewater Formation ca. 400 ka. 

Key Words: Quaternary palaecoclimate, salt lakes, Lake Malata, Bridgewater 
Formation, carbonate mud, gypsum, dolomite, Eyre Peninsula. 


Transactions of the Royal Society of S. Aust. (2002), 126(2), 91-102. 


STRATIGRAPHY OF THE LAKE MALATA PLAYA BASIN, SOUTH AUSTRALIA 


by A. DutKiEwicz* & C. C. Von DER Borcu* 


Summary 


DurKiewicz, A. & YON DER BorcH, C. C., (2002). Stratigraphy of the Lake Malata Playa Basin, South Australia. 
Trans. R. Soc. S. Aust. 126(2), 91-102, 29 November, 2002. 

The 19 m-thick Late Quaternary stratigraphic sequence within Lake Malata, Eyre Peninsula is dominated by 
autochthonous gypsum, present as relatively mud-free gypsarenites and gypsum-clay laminae overlying a 
skeletetal peloidal grainstone of the Bridgewater Formation near the base of the lacustrine succession. Calcite 
and dolomite mud are minor components of the column and several metres of these deposits appear to have been 
deflated into marginal lunettes. The skeletal peloidal grainstone has been severely modified by dissolution and 
formation of phreatic calcite, dolomite and gypsum cements under alternating pluvial and arid conditions. 
Discrete units are separated by disconformities and attest to rapid changes in climatic and hydrologic conditions 
over the lower Eyre Peninsula, commencing with emplacement of the Bridgewater Formation ca. 400 ka. 


Kry Worps: Quaternary palaeoclimate, salt lakes, Lake Malata, Bridgewater Formation, carbonate mud, 


gypsum, dolomite, Eyre Peninsula. 


Introduction 


Lake Malata is an ephemeral salt lake situated 33 
m above mean sea level in a mid-latitude region on 
lower Eyre Peninsula, South Australia (Fig. 1). It 
covers a total surface area of around 21 km?, which 
excludes numerous small deflationary playa lakes to 
the east of the main basin. Lake Greenly, 10 km 
south-west of Lake Malata, forms another major 
playa lake in the region but appears not to have been 
connected to Lake Malata in the relatively recent 
past (Dutkiewicz 1996)! and indeed has a different 
stratigraphic sequence (Dutkiewicz & von der Borch 
1995), Notably, Lake Malata is dominated by 
autochtonous evaporite deposits which are 
interbedded with carbonate mud, whereas Lake 
Greenly is dominated by carbonate muds 
interbedded with minor evaporites. Lake levels in 
Lake Malata fluctuate rapidly and seasonally as a 
consequence of surficial hydrological closure and 
rapid changes in the inflow-evaporation balance, 
which relies heavily on regional rainfall. During the 
wet winter season the lake retains < 0.5 m water, 
which evaporates in summer leaving behind a cm- 
thick halite crust. Although there is little direct 
evidence for the origin of the lake, 
geomorphologically its formation appears to have 
coincided with the emplacement of the Bridgewater 


“School of Geosciences, Building FOS, The University of Sydney 
NSW 2006, Australia 

‘School of Chemistry, Physics and Earth Sciences, Flinders 
University, GPO Box 2100, SA 5001, Australia 

' Durkinwicz, A. (1996) “Quaternary Palaeoclimate from Lake 
Malata-Lake Greenly Playa Complex, South Australia” PhD 
thesis, The Flinders University of South Australia (Unpubl.). 


Formation sub-parabolic dunes during — late 
Quaternary sea-level high stands. These dunes, 
which consist of skeletal peloidal sands, may have 
effectively dammed the pre-Pleistocene drainage 
channel thus forming local depocentres. Also, as the 
Bridgewater Formation forms the main recharge 
aquifer in the region, groundwater seepage along the 
dune lobes would have invariably enhanced lake 
basin formation within interdunal corridors and in 


=s 
i 
As wy Y \ PORT AUGUSTA @ 
\ / \ | { 
Vl AUSTRALIAN . 
f f i \ | 
\ I | } 4 
x : {| mel 
mT \ WA \ / WHYALLA @ \ 
Yj oH / ° 
{ PORT PIRI 
STUDY e L ¥. 
ARES \ ) TAS. } & \ 


————— / pal 


EYRE 


\ _ 


ys . 
¥ \ 


34°00 


\ PENINSULA, 


_ 


i 


\ 


| 


a) 


SOUTHERN \ ~ “= ZL 


J 


VORKE 


1 PENINSULA 


OCEAN 


kM \M 


Fig. 1. Map of Eyre Peninsula showing Lake Malata and 
location of sediment cores. 


92 A, DUTKIEWIOZ & CC) VON DER BORCTI 


areas of low reliel, Prominent geomorphologicul 
features include clay pellet lunettes, gypsunr lonettes 
and beach deposits along the eastern margins ol must 
playa basins (Dutkiewier ef a/, 2002) some of which 
reach 9 m in height. Apart front the sub-paraboli¢ 
dunes, pisolitic red soils and calerete of possible 
Tertiary age dominate the geamorphology iy the 
south and west of the main basin (Dutkiewiey oy cal. 
2102), 

This paper focuses on the sedimentary successiqn 
withit) Lake Malata, which provides evidence of past 
fluctuations in lake level, groundwater chemistry. 
and Quaternary climates. The carbonate-evaporite 
eveles reflect hydrologic and geomorphologic 
settings of the basin. detrital influx. groundwater 
seepage and recharge, und wind shear, which oflen 
redistribtiles surface water and wet sediment Herdss 
the entire lake surface and deflates dry sediment inte 
marginal lunettes, Post-depositional diagenesis of 
primary und clastic carbonates and evaporites will be 
discussed briefly as these also have been influenced 
by climatic oseiilations. 


Methods 


‘The stratigraphic sequence is based chiefly on five 
diamond drill cores taken from the main basin in 
1987 by Gilfillan and Associates Ply, Ltd. to 
determing the viability of gypsum mining (Fig. 1) 
The eores sampled the lake sequence lo basement 
and are available for viewing at the South Australian 
Department of Mines and Energy core library in 
Glenside, Adelaide, Despite their deteriorated stale. 
compaction ol up 60% and 80% recovery, careful 
sampling and detailed petrographic study ofabout 50 
(hin sections allowed a stratigraphic succession aad 
palaoenviranmental reconstruction to be established 
for Lake Malata, Unfortunately, sediments from the 
drill cores were unsuitable for radioearbon and 
thermoluminescence dating due to contamination, 
exposure to sunlight, and paucity of suitable material 
avuilable for dating, Consequently, a piston coring 
method was used to sample 1.8 m of fresh sediment 
Irom the eenter of Lake Malata (Fig. 1), AMS dating 
al the sequence, however, was unsalisfactory due to 
high concentrations of Na, Mg and K salis and low 
organie carbon contents (Dutkiewies 1996)! 

All cores Were logged and the mineralogy of 
selected horizons analysed in some detail, The colour 
was determined using the Munsell colour chart, 
Uneonsolidated material was wel sieved; the coarse 
lruviion was eXamined under a binnewlar 
microscope, and the composition of the Tine faction 
determined using X-ray dillraction. Consolidated 
material was cut perpendicular to bedding, 
impregnated and used for thin sectioning. The thin 
seclions were partly stained with Alizarin red-S, and 


Lied ASE 24 in dei 4S 


wer vent 


op rei on he rey 
‘ 
A \ 
+ twit 7 
inks Lingl ' 
=| SY EEARE IME 
a sot = 
_ | 1 
oe, : bh 
5 ls =| Mr hae oe! 
| | | Bevan evi ih i i. 
‘ - : ba 
ae Pas! ' 
» (Le ive teks 
eo oa  —— 
uh 
hele TED 
Su PAP IN ITE 
“ 
" 
lu 
Wut 4 
/ SKELETAL 
DRA 
be HAR ATONE 
~ tacorsl 
~ enn wa 
" SE Laminate 
Tee Ceepanan erm . 
“ e+ s 
ee tie 
=r 
\- Jw rahi 
| OSIM sre Fee 
cet 
i" + 
7 
, -*+ 


, unin? = —— 
(6 ENSECeHI - 


=—— ke 


=) ee 


= :o8 


Vig. 2. Correlation af cores throuzh he Lake Matar basin 


studicd with a polarisiig microscope. Textures and 
cements were further exumined usiig Seanning 
Kleetron Microscopy at CEMMSA at Adelaide 
University, 

Ciypsum sumples in band specimen are deseribed 
ising a vrainesize classification scheme of Warren 
(1982) while primary and secolidary gypsum 
pelrofabric descriptions are based on criteria outlined 
by Bowler & ‘Teller (1986) ancl Magee (1991). The 
skeletal peloidal sands and grainstones in Lake 
Malata have been correlited with calcareous 
ycohanites from the Bridgewater Formation using 
detrital, mollusean, foraminiferal, eehinoderm, algal, 
bryozoal and peloidal compositional classes. 


Stratigraphy 


Gypsum constituies at least 7O% of the bulk 
sediment within the Luke Malata basin, Carbonate 
(calcite and dolomite) and detrital clays form a 
relatively minor component und oweeur us fine 
laminations or literbeds rather than diserete lnits. 
llowever, strandline deposits, which inelude several 
phases of carbonate pellet lunette and gypsum 
foredune deposition (Dutkiewiex ef af 2002), 
sugtost that at least 5 m of carbonate mud and at 


SURATIGRAPHY Ob Th LABRLEMALALA PLAYA BASIN, BOL TH AUSTRALIA OS 


least 10) nt oP gypsum sand have been removed from 
the lake basi during periods of deflation and 
lunetie-building: spanning ca. 115-6 ka (Dutkiewics, 
eft af, 202), Individual units comprising the most 
complelely sampled succession from diatijond ceil 
core DH-S, which appears to have been faken from 
the palueo-like center are diseussed in detail, A 
eross-sechon through the Lake Malata basin using all 
available dianiand drill cores js shown in Figure 2, 
Contacts between the individual units are sharp with 
disconturmities between units @, 3 and 4 and 6, 4 
and 3. 


Unit 7 Basement (Weathered Gieyss) 

‘The basement consists of yellowish orey, very soll 
and lughly weathered gneiss which contains 
abundant pebble and sand-sized vrains of clear and 
grey quartz, sericite and iron oxides. The gneiss 15 
exposed around the southem and eastenr margin of 
Lake Malala where il forms a yraben-lype structure. 


Unit 0. G\psant-Rich Seiieite 

This unit coosists of very light grey to light grey, 
heavy and very dense sericite chay containing 
randemly-oriented. displacive pyramidal pypsum, 
The vypsum is lentieular im thin section and displays 
uv diversity of @rain-sive with crystals ranging [rom 
Joos Han Emin ap lo Sayin in length. Vhe-erystals are 
isolated und lack contact with each other, The pour 
sorting af the erystals reflects the variable porosity 
and permeabiluy of the serie matrix. which 
together determine the jo sith growth of the 
pyramidal sypsum, The centres of the crystals 
frequently display polyerystalling overgrowths, seen 
as distinc! crystal voniny under polarised Tight. Iron 
oxides dre eGmmonly incerpordted lang the 
cleavage plines of gypsum. Te sericite matis 
displavs a high birelringence under crossed polars 
and is clearly the weathering product of the 
underlying unit. Unit is approximately 20 en in 
WiekWess i DHS and 3 min thiekness in DH-3, 
reflecting the irregularly in basement and variable 
depth ab he weathering von, 


Unir” Leineted Gyypsurenite 

Unit 3 consists af Mnely laminated vypsarenite. 
which reaches approximately | im in thickness i 
DH-S and disconformably overlies Unit 6 (Figs 2, 
3a), The wait bas not been recognised elsewhere in 
the basin and possibly represents local deposition 
within a deeper. central part ofthe basin, In taet, Unit 
4 direetly averlies Unit @ in all cores with the 
exception of core DH-5. The gypsarenite comprises 
allernaling wavy ninm-thick laminae of very fight 
vey fhe to mediwm-grained, moderately to 
well-sorled Stary BY pPSUN, CoArser vps ThA 
pitty al Clay anal dolowites and fiediuns Mahe pley 


clay which drapes the underlying vypsumerich 
luminae. Mast of the gypsuny erystals are prismatic 
and appear as equant polygons in thin section 
(Piz 3a) The lineryramed gypsum ts 
closely-packed, matrix-tree. with only minor Lo trace 
amounts of line-erained irom oxides. Prismatic 
gypsum comprising Ute coarser layers. on the other 
hand, aeeurs in ad matrix ol nod-ortented clay and 
carhonale, predominantly kaolinite and saccharoidal 
dolomite. and displays concentrations of Tran oxides 
vlony cleavare plunes (Fig. 3b), Matriv-tree, 
cuarse-prained gypsum js alsa common bul 
represents ariding al the litter crystals rather than 
discrete larninge Displacive, lenticular or pyramidal 
vypsium forms are rare but occasionally gecur wilhiyy 
ihe coarser gypsareniie layers. where they dre 
ovented cindomly or sub-vertically to bedding. 
Unlike the clay laminae Comprising i more recent 
and better-preserved Unit 3, the clay in Unit 5 lacks 
oplical onentation, A possible explanation for this ts 
the relative abundance af coarse grains sueh us 
wypsuin, quarte und iron oxides. which are 
incarpoeraed in these laminge and prevent the cliv 
particles lrom Becoutite aligned. Clay obentaton 
may also be disrupled by post depositional zrowlht of 
gypsum within overlying and toderbying keyers. 


Cnt d' Sholeral Peloidal Giaiistone 

Uni 4 consists of a strongly ceptented skeletal 
peloidal grainstone. whieh diseonformably overlies 
Unit 3 in DH-5 and Unt & i DHE to DH-4 The 
pramstone aleains only Sen to thiekness i DH-S 
but reaches a Mantimuy thickness of 3.5 mn ih the 
easternmosk basin core DH=}. where it forms a 
unique and complex seguence of diagenetic 
curbonale-evuporite fabrics, These iwelude moldic 
porodity filled by poikilotopic gypsum and 
dolomicrospar (Pivs 4a, b), dolomierospar and 
microspar cements conwining displacive gypsum 
discoids (Fig. 4c) and dolomicrosparcoated 
allochems with @ late pore-(illing gyspsum centent 
(Fig. 4d). A possible explanition for the difgrenee in 
Udit ThickHess between DH-4 and DH-S is that DH-4 
is rohilively proxinal to the paargii of the lake and is 
betler sulted For the deposition of sandy nearshore 
facies, in particular calvarcous sands derived, from 
surrounding sub-parubolic dunes. In DUS. the 
drdinstone is -essentrally a light alive prey. 
well-lithiied  eypsiterous  wavkestone with 
appronijiately 40-60%. fabyie-sclective (maldie) 
porosity and displacive. poorly-sorted pyramidal 
gypsum within a mrerite main. (Pigs 3¢, ad). The 
vypsunt erystals Geer as ojuled lalls. characterised 
by sharp oerystal faees  indieatine minimal 
dissolution, Althouwwh the gypsum is generally 
randomly ar sub-vertically oriented to bedding, 
individual crystals show a tendeney tor displacive 


U4 A DUPRIPWICA A CO YON DER BORCII 


STRATIOR ATTY OF THE LAKE MALATA PLAYA BASIN, SOUTILAUSTRA LTA "3 


é ihe 
*¥ eee fe AL 
Oee © ve, = 


Migs 4. Photomieroeraphs of the skeletal peloidal grainstone in Unit, Thin section imaves Geb) aiken under plain ight. al 
Moldic porosity partially filled by poikilolopic gypsum, Note mictite coatings (mek mieriised wlochems (mau. eHpey 
lochemical voids (Vv) oeersionally filled by gypsum (2). otergranilar cement consists almost entirely ol poikilotopic 
vypauin (2). b) Alfochemieal void purtially filled by woledrul dotomicrospar (dy and gypsum (oh Ditergranuhir porosity! 
is bled by poikiotopie eypsum and minor unhedral dolomierospan ec) SEM image showing displacive gypsum diseaids 
within dolomicrosparculeite micnospar cement, d) SEM imige showing a dolomicrospar rind (dr) sound a nneritised 
allochern Me rind forms contiel sutures With neighbouring dolomicrospar rinds surramding alluchemies) voids which 
ane parhauly Milled by dolomicrospar (C0. Intereranubir cement consists af poikilotopie 2vpsiim aad sparse dolomicrospar, 
Note the presence of inlergmimulir pyrile (py) 


Fig, 3. Photomicrogriphs wf Units 3, 4atd 5 from the Lake Makai basin teken under plain ight Gu Gypsum-chiy couplers 
1 UNLS, ‘The thicker laminie Consist of fine fe coarse-grained equant (prismatic) gypsum (2) and the thinner laminae 
consists OF nen-oriented cliy (noc) dominated By kaolinite. (A) Prismatic equant eypsam erystals in Unin 5. The mates 
consists al kaolinite and dolomite, Tron oxides are commonly ieorporaied alone the cleavaze planes of gypsum, le) 
Conmipletely leaehed portion oF the skeletal peloidtil grunstone comprisii Unit 4 Allachemical voids Ov} are present 
within a dolomientie mivix. Disphicive discoidal (pyramistili gypsum Gel is cemimen and offen forms clusters, (dy 
Abundant eypaun chseaids in Unit 4+ The discoids ure rindemly oriented 16 bedding and.are rarely. Ta contact wilh een 
other. (e) Pyeinidal 2ypsoin on Unit a. Note aoning within the centre of the ervstal caused by the mefusion of eon axes 
around wv pre-existing detrital eare or gypsum nucleus. 1) Poorly-formed diseoral (pyramidal) gypsum eeystal fron the 
base of Unit a. The centre of the crystal is qccupied hy a queries core (2) Gypsun) diseaids comprising the basal 
eyvpsurenie in Uni 3, The cevstals are oriented sub-verticully ta hedding and show zoning near the crystal edges rehited 
lo chissolotion and re preeipiiition of the gypsum or variable growrh fates of The site erystul. Quarty (2) Cg) cores ure 
oecusimally present. (h) Care secbon showin reeulirly alivenatige laminae ph eypshor iehr and clay (darks eompradny 
the gypstim-ecliy limite Wi Unit 3 in DHRS 


Oty A-DUTKIEWICY & C0, VON DER BORCHI 


: ; : " - — 
—_— 4, = 6 
— i - we ~ | 
O4en 
a . 


big. SVhin scetion bi. oT) and SEM (h) phatemicragraphs showing the gy psamn habirs im Unis 3 and 2. Polarised fight Ga 
phon Hight (oP. ay Ciypsum-chiy couplets comprising the eypsun-elay luminite iq Unit >. The lini consist of highly 
quented elay (ic) dominated by kioligine. overran aid underlain by mnicker layers Of extremely fine-grained, equa 
ofismatle ewpsuny orystily Gr, 1b) Gypsum discoid (pyramidal aypsum) from the upper gypsurenite Gaver in Uni 4 The 
fice OF dhe eryseil tothe (O00) cleavewe plane which shows considerable receysithsation. Note the presenoc ot 
fine-grained Suh-spherular dolomite Girrnw), Ge) Freecharing and minor reworking al at sypsur erystal Gaeowey near the 
base af the upper ’ypsurcorte horizon in Unies. Displugive (pyruinidal) 2ypstin is (he most Gannon erystal dorpholoey 
br this wie The manis consists of a mixture of kvolinile wad dolomite. The wypsut is very poorly serted, fd) Course. 
interenwe bladed (py nannicaly sypsanr near (he base of Unit 2 showing the presence el Hibrous hiolinite (2) along the 
cleavage planes. Main light te) Subevertically to nindonily oriented discondal fpyrstridiul) gypsurt withio. a caleite 
(tolomite THT COMmprising ey psarenite i Ui’. Reery stiller and zoning are parteohiely eonmnon my this ancy OP) 
Abundiol iron oxides forating clusters within Clinit 2, 


STRATICRAPTY OF TUL LAKE MALATA PLAYA BASIN, SOUTIFAUSTRALIA y? 


sub-circulur cluster arrangement (Fig, 3) 
reminiscent of vypsite nodules. Preserved 
allovhemical components in DH-4, us well us the 
veneral shape and size of the voids in DH-5, stzeest 
{hal the porosity has resulted from a coriplete 
dissolution of skeletal and peloidal allochems 
sourced by the Bridgewuter Formation. Detrital 
grains include (ine-wrained quart, plagioclase and 
iron Oxides. whieh have not been affected by 
dissolution, Crum counting and cluster analysis, 


although resivieted to a sinall unleaehed purtion of 


the umt in DI-4+, show “ood correlation with the 
Bridgewater Cormation and the 9 m beach ridge 
dlong the eastern margin of Lake Malgla 
(Ddutkiewies ep af 2002). Although textures and 
Kibries dvseribed for DH-S are consistent with 
pedogenesis, in DH-+ the skeletal pelorul 
grnstone has underuane extensive phreatie 
diwenesis which ts reflected in fabric-selective 
moldic porosity, Sopachous dolomictospar rinuds. 
intergranular and itragranubar — veid-lilliny 
dolomicrospar, and puikilotopie and yeid-illing 
eypoura cement (big. 4). 


Linn 3. Laminated Grpsarenite 

Lint a disconformably overlies Unit 4 ond adains a 
thickness OF 6 mn, Tt consists of a jediuit light grey 
(y Tigh) grey ovpsurenile containing variuble 
dmounts of jnterdispersed dolomite mudd and 
kaolinite. Hine-geumed eypsarenite, and displacive 
wypsite nodules, he anit is interbedded with o finely 
laninated eypsarcujle over the 95-4005 mm and 
11.5-]2 m depth intervals (Fig. 3h). The laminuted 
sequences consist of alternating inm-thick wayy 
Jaminae ot very helt grey, Sugary, fine to 
meditin-wained, Well-sorted zypsuin. finer (- ] mn 
hick) laminae of medium dark urey. aplically 
orivnted kaolinitecand light grey nin-thiek. laminae 
of medium-prained ypsarenite io matrix of clay 
and dolomite mud Clay driping is common, A 
metre-thick layer of fine te wnediumegreinied, 
moderalely-sorled vypsarenite separates the 
linrinated inlervals, 

The eypsun-clay laminae overlie a gypsarenile 
layer which consists af randonily and sub-verivally 
onchted pyrarnidul gypsum crystils witht a iralens: 
OF sdechuroidal dolomite (Pig Se). The gypsum 
crystals ure relatively wale aeruss tle Gans und 
Show variable wranrsize aul degree of sorting, A 
cm-thick layer ol pyramidal crystals oriented parallel 
to bedding, and showing Tittle variabilicy to 
wriesize. is diya present. Zoning, of crystuls os 
common wid may be utiribuited ford) the 
IWcorparation Gt iron oxides along cleavare planes 
und erysial boundaries during a change in fhe srowrs 
nite or during sleeve dissolution of the érystal 
(Fig. Je 2) crystal growth arounel a debris! core 


(Migs. 36 @): 3) the development of gypsum 
overgrowths at the manans of gypsum crystals 
which lack optical contunuity with the rest of the 
crystal. I this sense, the pre-existing wypsuni 
crystals provide a nucleus for subsequent gypsum 
growth: and 4) selective dissolution followed by 
re-precipitation al’ central and marginal parts ol 
crystals, All of these teatures have been observed in 
this part of the unit, 

The repetitious oature of the elay and: eypsun 
laminue hears a striking resembhince lo similarly 
varved sequenees trom Luke Tyrrell (Bowler & 
Teller (989). Prungle Lakes (Magee 1991) and Lake 
Eyre (Magee ef afi 1995). In Lake Malala. the 
individual laotinace eousise of ob) fine ta 
voarme-grained. reversely graded. closely-pucked, 
matrnctree, horizontally oriented prismatic gypsuiny, 
2) rekwively course-grained, frequently reversely 
sraded. horizontally oriented. prismatic gypsum ina 
noutrix of noneoriented clay (ie. 3a), Here. the clay 


contains ubundaunt iven oxides and myipor 
Jiie-grained,  displacive. vertically oriented 


pyramidal gypsura: and 3) optically oriented clay 
(kwolivite) with minor iran oxides and miner 
toarsc-gramed prismatic “vpsum (Pig Sa) The 
laminac are equally spaced and eyele, 

The vypsurenite overlying the gypsunt-clay 
laminae consists of medium co coarse-praincal, 
poorly-sorlwidk pyramidal gypsum im o mauris of 
non-ariented clay and Saccharoidal dolomite with 
abundant wypsite rodules. The gypsum is randomly 
orjentited to bedding and displays perfectly formed 
palyerystalline discoids under the SEM (Wig. Sh). A 
small number of the erystals, however, are prismatic 
and Oriented parallel to beddiig. The eypsite nodules 
are several mm in diameten displacive and consist of 
sth-sized pyramidal gypsum forining imutrix-free. 
cumulus-Shaped clusters. Iron oxides are abundant 
und geeur alone eleavage planes of the pyramidal 
vypsuin erystals, Frielurius and apparent reworking 
ola number of ervstais are evident (Fig. Sel. 


(nie 2> Caypserenive 

Unit 2 vonsists of yellowish grey is light olive 
erey, Slightly muddy, medium to coarse-vrained und 
poorly-serted wypsarenile. The Lit ts approximately 
Som ih thickness wel sharply overhes Wait 3. The 
mud traction coosists of dolomie with rmiinor 
amounts of delrilal kaolinite, heconing inereasingly 
ealcite-rich (law-Mg caleite) towards the tap of the 
uot where dolontie aid Kualinite are present onty in 
dee amounts. Kaolinite is oceasionally incorporated 
within wypsum cleavage planes (Fig. Au), 
Centinpete-thick laiitiations of pelatively oiuddy 
Bypsarei|ie allermatiae with less muddy eypsarenite 
are conmon between 45-5 mound 35-4 m A 
decinretre-thick layer of vrevish vellow ereen 


UN A DUTRIPWICZ & CC YON DER BORCH 


kaolinite is present belween 5 und 5.4 m overlying 
a dm-thick layer of coarse-wrained displacive 
diseoids af pyranvidal eypsum measuring about 
foem a length, White, irrewulin and displacive 
evpsile nodules und gypsile layers are common 
herween 3.5 and @ m. tn thin seetion, the 
pypsarenité consists al pyramidal wrystals oriented 
randomly and sub-vertically to bedding dispersed 
within a carbone (calcite and dolomite) matrix 
(Fig. Se) Gypsum grain size is varmble, with 
mdividual erystils ranging: Feo less than | mm to 
Jem in lenwth, Gypsite nodules alsa cousist of 
raidonmly opiented pyramidal gypsum erystuls 
However the crystals occur as poorly-developed 
discos and form dense. displacive, matriy-(ree 
nodules within slightly muddy yypsarenite. 
Miboweh detrital cores contribiite 16-erystal voning, 
the majority of (he gypsun) taths are zoned dye ta 
dissolution and rapid re-precipilalion of vypsium 
(Fig. Se). Lanse. bladed, occasionally traerured ara 
Hilergrown, prismatic gypsum up to | can it bengal 
is conimon near the base of (he unit, where it shows 
replacement by low-My culene (') alone the 
cleuvupe plaues. Minor amounts of horizontally 
oriented prismatic erystals and clusters of iran 
ose minetals (Fig Shy tire alse assouiited with 
ihts layer. 


Cnit lh: Ciypserentite 

Unit | consists of olive urey, muddy, medium to 
course-praiied, poorly-sorted uypsarenite 
interbedded willy em-thick layers of gypsite anda 
dm-thiek layer al organie-riel, olive black law 
mdenesion calente mud near the base. Phe unit is 
upprosmudcly 70 en thick and sharply overlies Uivit 
2. The gypsum is pyramidal, with ineliwidital lors 
ceaxes oriented randomly or sub-vertically to 
heading. The erystals occasionally show inclusions 
of mud, indicative of Fastagrowih rates withii a mud 
matrix (Kastner 1970), Clusters of tran oxides are 
present locally, Only Unit Tis represented within the 
piston eore (C16; Fig, 1), 


Jnterpretation of depositional environments 


The ostrativruphie sequence refleets Jargely 
uroundwater controlled oscillations in lake levels 
associated with hamid and urid climatic episodes, 
which in a vertical sequcnee are marked by the 
presence of suling lacustcine facies (carbonates ang 
eyapurites) intermittent avoltan deposition of 
skeletal peloidal sand. Junetle-buildins an 
puduvonesis af marginal regions: 


Wire, © 0) (IMO) ) Hieelet af ihe Queteriiry Rrideceaies 
Voom of the Seupavese and (onirst Seuth Aetealia’ The 
thesis Me inders Cineevrsiey, of Seely Atetbe Cl publ a 


Laminated clay-rich gypsarenile (Uinit 5). whieh 
disconformubly overlies wealbered basement urerns 
Woqvhe resion. forms the hase af the lacustrine 
sequence. “The clay was most likely depasited in 
Lopoeraphic lows as channel runoff during u 
relatively pluvial period, willy flow amb erowon 
iWitiated during and alter heavy rains, Deposiiien 
way have oecurred in the early Pleistocene price to 
the initial emplacement of the sub-parabolic duttes 
cH 700 ka (Wilson L299}, whieh buried vast aneas of 
ihe land surlace and played a key role ty the 
formation al the lake basins and the regional aqualer, 
‘This is supported by the absence af skeletal peloidal 
allochemes within the chiy which would otherwise be 
expected ia be tamsported with the flow. sitll 
valeareous sand and eraimstone necurring higher tH 
ihe sihabigruphie sequenec. Lamimation af clays. mn 
DH-S iy pssoeiwOn WILA ypSHren ile. Nuleusts 
inlerimitlenl, possibly annual deposition controlled 
by the duration and frequency of the pluyial 
uprsodes. Regionally, the clay probably represents i 
relatively low How regime, where only ulay-sized 
particles with a very fine sand fruetion are deposed 
in fhe centre af the busin, 

Deposition of skeletal peloidal sands (Limit 4) as 
nist Hhely relited ty Wilson's (POOL) phase | (ea 
400 ka) or curly phase He (ea, 220 Kit) emplacement 
Of the Bridgewater Vormation sub-parabolic duties 
The skeletal peloidal sand diseonformably overlies 
basement elay and shows tntense diagenesis anil 
strong cemenuiion io the ventre of the Luke Matota 
basi, IL is disconformably overlain by lacustrine 
gypsurentios in Lake Miata as indicated by sharp 
ioe! discontinuities and the presenve ol 
indurated horizons ahove und below The geil. whieh 
are the result of subserial exposure and pedoyenesis, 
Deposition of Unit dis closely related to the take 
Malala foreding ridge, deposited during a prolonged 
pluvial phase va. 319 ka (Durkiewior en ué 2002), 
The skeletal peloidal sand has undergone incuraion 
and cementtion partly due to subaerial exposure antl 
partly due to preerpravon of phreave intergranular 
voment, which relleets ollemaiune groundwater 
Aneluations. Phe idurated (pedoeenic) lores 
intheale minor breaks i deposition oF the sane, 
whieh is controlled Targely by sediment supply anal 
the intensity of the westerly winds, Relatively impor 
amounts of licusiine carhonate have heen deposiled 
intermittently owihin the sand, fornine 
discontinuous interbeds, as conditions heeame mane 
pluvial for short-lived periods of time. 

Vast amounts of the mobile skeletal peloidal sand 
would have been transported vito the bitsin prier to 
dine Stohilisation, during, (he landward migrate ol 
the dunefield and durin subsequent episodes of 
dune re-werivation, Phe sand was transported intr tne 
Inke basin imastly by Lhe streng preventing westurtnes 


STRATICR ATHY OF THE LAKE MALATA PLAYA BASIN SOL THAALSTIOALDA uy 


und partly by local vnott which drained the 
stib-parabolic dunes, Emplacement of ihe first phase 
ol skeletal peloidal sand initisted the formation af a 
major unconfined ayuiler and the onset of lacustrine 
carbonate deposition. The lake it this (ite wis 
relatively fresh, atid the recharue rates high, The ea, 
31 ka Lake Malala ridge and abundant dissolution 
feulures in the reohanwe aquiler and within the Luke 
Malta busin indieate evidenve for high wroundwater 
lables, Dassalution of qloehems, partloularhy during 
the freshening techarge episodes, was essential in 
providing sufficient! ions for the subsequent chemical 
precipitation of low-Mp calevte. 

‘The skeletal poloidal sand experienced sone 
reworking within the basin, as indicated by the 
presence of a relatively thin Jayer of the saud 
overlying laminated eypsarenite i DEAS. I this part 
of Lake Maliuta. the sand experienced induration and 
pedoenesis as reflected ji thy presence of a 
eryploerystalline Perle cemen| and displucive 
pyrinidal gvpsunt associated with a Muetuaiiy 
water table. Pedogenesis appears uy have been 
particularly effective in areas of fateral dining of 
the sandivraistone and may be related to the vale of 
the ceuostone asa recharge conduil and preferential 
drying of low recharge parts of the lake. Uhick beds 
af skeletul peloidal wraiistone, on the other hand, 
experienced intense diggencsis i) the Corm oF 
carbonale-eyaporite Habrics related to aserlations: in 
vroundwaler and the phreatic dhayeneue 
environment (e., DHF). In particular, carbonate 
cements formed during periods af mereased 
PHUViTEY, telatively low evaporation/inflow ratios 
assocrled with relatively high Take levels and law 
salinities. The dolonite represents a conbinution of 
teplacive and void-filling cements linked with 
labric-seleetive dissolution of allochems, Gypsunt 
eenichty ure vou-hiling und post-dule eurbonare 
cementalion. They were fonmed during arid periods 
characterised by high evaporution/iiiow ratios: ane 
law lake levels. 

Follows (he first phase of dime migration (and 
fornia Ob thy calcareous recharge aquifer), the 
lakes were qtotindater! with) carbonate-enriched 
eround and surfage waler. wilh solutes derived 
largely through the dissalurion at skeletal and 
peloidal allochoms. This is evident m the first eyele 
Of diigenosrs iy the skeletal peloidal urainstoic 
withi) the busin, whieh is marked by the 
preaipitatian Of carbongle cement, aud im the 
considerable thickness of chermically-precipilatee 
carbonate mud in regions of former lake extent 
overlying the Jirst phase of skeletal peloidal sana 
depasinon (Duickiewles 1996) The thickness and 
the relauively homogeneous nature GF the carbonate 
WHS Th thirgindl dreas (Prtkiowies (996)! indleate 
hat precipitation ocewed under relatively 


long-lived hydrologically and elimatically uniform 
conditions ina low energy. open-lake environment 
Fhe carbonate mud units carrelite with Nininated 
vypsarenitce i Lake Malaita (DIEH-3) trom whieh 
several metres of carbonate huve been removed by 
deflation during the construction of carbonate pellet 
Junettes Over ad period spamiine ca, YH ta 1S ke 
(Dutkiewiey ef a 2002), It is. possible thal the 
indurated horizons separated by — untithified 
carbonate mud within these deposits are related to 
lunelie pedoyenesis associated wath tajor falls in 
groundwater fevels (Duikiewiex et af 2002), 
Possibly due to burial ald moisture content, 
carbonate amd clay pellets assueniled with lunette- 
building have not been deteeted ac depth within the 
mud sequences, 

Deposition of the skeletal peloidal xrainstone in 
Lake Makita was followed hy theonsel of alternating 
Shallow and relatively deep saline conditions 
associated with frequent groundwater Muetuations. 
This ts ustrated by the presence ofa distinc finely 
laminated gypsarenile Sequence (UniL 3) comprising 
wllernaling laminae of clay aod prismatic gypsum, 
which overlies the skeletal peloidal wrainstone 
towards the basi centre. In Lake Maltta, carbonate 
sedimentation was restricted fo Matginal areas. 
proximal (a the recharge aquifer, wilh wypsum 
deposition confined to cental. deeper parts of the 
basin. The repertwve nature and constant thickness of 
the lanvinae iw this Unit suweest an alleenatine 
wet-dry, seasonal depositional eyele. Clays whieh 
comprise the thin (= | mm) laminae and tepresent 
the fine-grained clastic component, were truisparted 
into the basin durifg the wet winter seasoir as tunel 
which drained the eastern and western eliy-tich 
slopes surrounding (he lake. While ihe finer clasties 
Were transported into the deeper, central parts of the 
basin. coarser elastics, including skeletal peloidal 
sam! owhiely is currently eroded from the 
sub-purabolte dunes bordering the southern lake 
murvin, were deposited in the near-shore regions. 
Magee (199!) sugeested that a density difference 
between the dilure inflow and concentrated lake 
brines Would allow the Frosh feodwater carrying a 
suspended ehry da slide over the brine tor 
considerable distances prive to the clay foceulating 
and settling to the lake bollem, Tyapordtion. 
eonbined with reduced inflow inte the lake during 
the dry months, would subsequently coneenttle the 
surluce brine and allow prismatic Lypsum to 
precipitate within the brine bady or at the bring-air 
interiee. 

In Lake Mahta. primary, sibaqueois precipitation 
OF YYpsiT is Supported by: ty) the absence of 
reworktie [atures such as fracturing and roundinw. 
whieh are indicative ol abrasion during: transport. 
lhe consistent with cryskds growing at the 


WW) AD TRIE WIC? & ©. 


sediment-water interlace (Mayee T99}): 2) the 
absence of variuble grain size wilhin a single vypsum 
laminae. whieh ts indivative of diagenetic growth of 
crystals following depasition: and 3) the presenve of 
wavy laminae, sugvestifu the presence shallow 
water. Therefore. gypsum comprising the limine Is 
of dhe “settled” variety of Magee (1991) having 
formed at the brine-air interfiee and then settled to 
ifig lake sediments as described hy Schreiber e7 cl. 
(1982). Coorsening-upwards of the wypeunr crystals 
stiggests that the surface brines became (nerewsinely. 
siling and supersaturated towards the end of the dry 
season, producing larger and fewer erystals 
(Schreiher 197%: Magee 191). Tn order for 
suibuyueous vypsum to precipilale and accumulate in 
siznificane amounts, the basin must be groundwater 
controlled and contin permanent saline water whieh 
is maintained only when the basin receives a 
coustant supply of water and experiences ligh 
evaporation) rates (Roser 1994), The presence of clay 
withing number of the gvpsiin layers bs prelate to 
brief Mooding episodes during the dry phase, which 
apart from supplyine fie-grained vlastigs to the 
luke, are insuffivient to dilute the tering below the 
Jevel of wypsuin saturation. row oxides are supplied 
cither during the flooding Of the basin or are the 
product of sulphate-redueings bacteria oxidising ina 
sulphides. The eyele is repgated with the mest yet 
episode. during which clay drapes the tinderlying 
eypsum, This provides un impervious layer which 
seals the eypsuly and prevents i) from undersorny 
Uissalution. us He brine Freshens by mixing vith the 
dilate in Mow 

Mechamsams aivolved in orientation of olay 
particles aie fot completely understupd A Wumber 
of proposed mevhanisnis Neve been reviewed hy 
Magee (1941), although to date very little work has 
heen done on hizhly oriented elay particles: Most 
noteworthy cantributions by Mead (1964) and 
Sonnenfeld  (JOR4)  supbested Compaction, 
Ue-watering of clays and Moeculsuiun as possible 
controlling faefors in particle alignment Bowler and 
Teller (I9S6) Subpested that formate and 
preservation of oriented clays im salme Mcusiine 
envignments is dependent on salinity und the 
Activity OF benthic riiere-organisnts. They proposed 
thal deep water, aerated, low salimiry enyironments 
would support seavenging organisms which are 
Irkely (odlisturb oriented clay particles. On the other 
hunt, organisms cannot become established under 
conditions of extreme salinity and transported clays 
are able ot flocoulate and sete undisturbed. Sine 
miendaund is relatively rare (or rarely preserved) in 
Luke Vidata, the explanation of Bowler and Teller 
(1986) provides a likely mechanism for cry particle 
alignment in the laninae doeumented here 

Conditions following the seasoial deposition al 


©. VO DER BORCH 


the gypsum-clay luminite changed drumatically 
wilhin the Lake Malata basin as lake levels dropped. 
This was due to un overall Merease in the 
vyaporacian/inflow ratio caused by a decreuse m 
precipitation, whieh as the mam source of recharge 
into the lake, and/or a decrease in the (raeuon of 
eroundwater Jos duc to leakage through an 
increasing impermeable skeletal peloidal erainstane 
(Dutkiewice ef af 2000). Sediments directly 
averlying the laminated sequence are no longer 
yarved. and ave danyinated hy pyranndalrather than 
prismatic vypsum (units 2 and 3). In fact, pyramidal 
gypsum is the most common form oF yypsum willl 
the evaporite beds and conyprises thick units within 
Luke Malata and Lake Greenly. Pyramidal gypsum 
has been found iy raahy coastal settings Such as Hatt 
and Teeman Lagoons in Western Australia (Arake! 
LOS), Tricial Coast (Shearman 1966) and more 
recently my Lake Tyrrell (Bowler & Teller 198o) ane 
Proungle Latkes (Magee (991). Unlike pristmatie 
eypsunt, Which forms within a standing brine beily, 
pyrantidal pypsunt precipitates moterstibally Choi 
saturated pore (waters immediately below the 
sedjnent surfuce Within the capillary zone ander the 
iifluence of high evaporation rates (Bowler & Teller 
1986). In Lake Maluta. ibis commonly fut within 
a carbanate’clay multix, where it either completely 
displaces the surronpding mauix forming mud-free 
vypsuremily, or ulidermoes. diagenetic growlh with 
vypsareniles becoming  coarse-wpaied and 
puorly-sorted while isolated crystals beeen 
massive and eouch several centimetres in lengilr 
Absence of Solid Tnchisions within the ussive 
gypsum indicules slow growl ander Witter 
conditions (Kastner 1970). Pyramidal gypsuny 
comprising, wypsurenites. on the other hand. os 
generally cloudy duc to The incorporation of 
Impurities, sugeesiqie fast wrewth bnder wort 
cnnclitions where the gypsareniles are THaderalely to 
well-sorted, ond non-uniform conditions where the 
eypsarenile 1s paorly-surted, 

Bowler & Teller (P9KH) sugested thal sediment 
layers contaming abundant pyramidal gypsum 
crystals muy be dood mdicdtors oF past Nuctalans 
Woeoundwater, The fact that pyranridal oe discordal 
By psu Conprises LYpstin Hpettes/foredines alane 
the euslern margin of Lake Malate ir itself Sugeest: 
seasonully oscillating hydrological conditions 
(Dutkiewite ef af 2002). Sinee the eypsum 
lunettes/furedunes vontain only traces of carbonate 
or clay pellets, ibis the venerally mud-free thick 
By psarenite beds such as Units 2 and 2 which are the 
most Hikely source of the gypsum. In this seenane. 
vypsunr is reworked by wave avtion during a 
relatively wel episode and deposited at the eastern 
luke margin. where itis subsequently detlaled (ited 
Junete or loredune during the next dry episode, The 


STRATIGRAPHY OF THE LAKE MALATA PLAYA BASIN, SOU TEE AUSTRALIA 101 


semsonul deposition of the gypsum foredune mimes 
the earlier deposition Of the gypsum-clay laminae. 
which are no lonver forming due to-an overall drop 
in lake level and a shill trom a throughflow to a 
relatively closed diseharge basin. As advocated hy 
Bowler (1983), neaesurlace precipitation of 
zypsum and other salts assists in pelletisation of 
lacustrine mud and clay. This process 1§ required for 
delation of mud and clay from ihe lake surface and 
is possible only under a groundwater discharge 
regime The general absence of gypsum within 
carbonate pellet lunettes indicates that thust of the 
eypsuin precipitated as groundwalers rose slizhtly 
and the capillary fringe reached the lake surfiee. 
following a period of deMation and oscillating low 
water tables, 

Units | and 2. which comprise the Lake Malata 
sequenee. corcelate well with the alternating 
carhonale-evaporte beds in Lake  Greenly 
(Ditkiewier & von der Borch 1995), However, 
correlation oF individual beds ts impossible, partly 
due lo deflation of several metres of carbonate and 
its subsequent deposition along the north-eastern 
marvin of Luke Malata (Dutkiewice ef e/. 2002), and 
pamly duc lo local hydrology. geomorphology and 
aquifer charactertsiics which control ihe deposition 
oF eurbonates in one basin and evaporites within the 
other basin. However, withit a single vertical 


sequence. the carbonate beds are associated with 
humid conditions and relatively law 
evaporauon/intiow ratios. whereas the avpsum 1s 
ussuciated wilh and conditions and relatively high 
evaporation/inflow ratios. Thermolumimeseence 
dating of curbonate-pellet lunettes suggest that these 
humid-arid oscillations may have been operating 
sinee ca, 16 ka, which post-dates the tnajority of 
carbonate pellet lunetle deposition and overlaps 
with formation of the gypsum lunette/foredune ex, 
5.6 ka cal BP (Dutkiewier cf al 2007). 


Acknowledgments 


A. Dutkiewiez gratelully acknowledues rhe 
financial support of Flinders University. We thank 
Joe Lorenzin forthe preparation of thin sections and 
staff at the DME eore library and the CEMMSA at 
Adelatde University for their help with aecessimy and 
analysing the Lake Malata sediments, Core C16 wis 
sumpled with the help of fellow Flinders University 
post-graduate and honours students. We are indebted 
to (he Modra Family for their hospitality, and help 
during our prolonged stays in the field. Chris von der 
Boreh acknowledzes the continuing support of the 
School of Physies. Chemistry and Earth Sciences al 
Flinders University. We thank Martin Williams for a 
thorough review ol the manuscript. 


References 


Atantiy A.V, (1880) Crenesis and diiaenesis ol Aplocene 
evuporide sediments in Hull and Leeman lazoons, 
Western Australia, “Sed, Per 50, |305-| 326, 

Bowink, J, M. (i983) Lunettes as indices of hydrologie 
chime: erevew of Australun evidence, Proc, Reval 
Ser, be 98, 147-168, 

ke Tae, LOT CToS6)) Quaternary evaporites 
and hydrslogie changes, Lake Uverrell north west 
Victoria. aw i Barth Sey. 33, 43-023. 

Doratiwit4, A. Herevkgo A. L. & Dighton. J.C. (2000) 
Pas! changes (0 isotapie and soltte bakinees ina 
comtinetal playa: clues from suble msotapes of lienstrine 
carbonates, Chen, Geel. 1b$. 309-329, 

& vow bie Bomcno Coc) (1995) Lake Greenly, 
hyre Penmeula. South Australias sedimentalowy, 
pulucoclimalie and — palucahydrologie 
Polovurene, Pdluenelim, Pulacmenl 3,436, 

— Oo & Perscend, JR, (2002) 
Geomorphology of the Lake Malata-Luke Greenly 
Comples, South Australia, anal ils iraplications for Late 
Ouitlernary Pahieoctiimite, free, AO See, 8. often 12t, 
LO4-1bS- 


evclos: 


Kasinik. M. (P70) An jnelision Hourglass pattern in 
symthetic eypsum, Avi. Mineral. 33. 2128-2130, 


Maarr, J W. (199})) Late Otiaternary licustrne. 
sroundwater aeolhin amd pedogenie gypsum in the 
Prungle Lakes. southeusterm Austraha, Palicogeute, 
alaenchim., Palaeaeeal B4, 4-42 


~ Bowl rR LM Ming Ro Goh A Wirt iam, De 
L. G. (1998) Shativnuphy, sedimentoloy, chranolamy 
and palacohydrology of Quaternary lacustrine deposits at 
Madigan Gill Lake byre, South Australia. Pafworcou 
Polacwelin, Palaenecal W3, 4-42. 


Mi apn. RoE C1969) Removal of water and rearrangement 
Of particles during the compaction of ehiyey sediments - 
review. GS. Geolosical Surver Professtanal Paper. No. 
AQT-13, 23 pp. 


Rosia, MOR. (1909). The Importince of proundwaters in 
plavas: A review al playa classifieations and. the 
sedimentalogy and hydrology of playas, J Rosen. MLR. 
(Ld) “Paleoelimte and Basin Evolution of Phayi 
Systems” Geol. Suc An Special Paper So, 289 


102 A. DUTKIEWICZ & C. C. VON DER BORCH 


SCHREIBER, B. C. (1978) Environments of subaqueous 
gypsum deposition. pp. 43-73 In Dean, W. E. & 
Schreiber, B. C. (Eds) “Marine Evaporites” SEMP Short 
Course No. 4. Soc. Econ. Min. Palaeont., Tulsa, U.S.A. 

, Roto, M. S. & HetmMAN, M. L. (1982) 
Recognition of primary facies characteristics of 
evaporites and the differentiation of those forms from 
diagenetic overprints pp. 1-32 /nm Hanaford, C. R., 
Loucks, R. G. & Davies, G. R. (Eds) “Depositional and 


Diagenetic Spectra of Evaporites - A Core Workshop” 
SEMP Core Workshop 3, Calgary. 

SHEARMAN, D. J. (1966). Origin of marine evaporites by 
diagenesis. /nst. Min. Met. Trans. 75, 208-215. 

SONNENFELD, P. (1984). Brines and Evaporites. (Academic 
Press, New York, U.S.A.). 

WarreN, J. K. (1982) The hydrological setting, occurrence 
and significance of gypsum in late Quaternary salt lakes 
in Australia. Sedimentology, 29, 609-637. 


GEOMORPHOLOGY OF THE LAKE MALATA-LAKE GREENLY 
COMPLEX, SOUTH AUSTRALIA, AND ITS IMPLICATIONS FOR 
LATE QUATERNARY PALAEOCLIMATE 


By A. DUTKIEWICZ*, C. C. VON DER BORCHT & J. R. PRESCOTT 


Summary 


Dutkiewicz, A. von der Borch, C. C. and Prescott, J. R. (2002) Geomorphology of the 
Lake Malata-Lake Greenly complex, South Australia, and its implications for late 
Quaternary palaeoclimate. Trans. R. Soc. S. Aust. 126(2), 103-115, 29 November, 
2002. 

Lunettes, foredunes and beach ridges from the Lake Malata-Lake Greenly playa 
complex on the Eyre Peninsula attest to major changes in lake level and palaeoclimate 
over the last 320,000 years. These have been dated by a combination of 
thermoluminescence and radiocarbon techniques, thus allowing correlation with Late 
Quaternary Oxygen Isotope stages. The lakes experienced a major wet phase ca. 320 
ka followed by multiple arid episodes linked to relatively cool periods and low 
eustatic sea-levels between 115-16 ka. Aeolian activity and aridity wre particularly 
intense during the Last Glacial Maximum with the onset of a dry climate and 
carbonate pellet lunette-building commencing as early as 26 ka. The Holocene 
palaeoclimate is marked by seasonally oscillating wet and dry periods reflected in the 
intermittent deposition of gypsum lunettes, carbonate ridges and quartz foredunes 
around the eastern margins of lakes Malata and Greenly. 

Key Words: Quaternary palaeoclimate, salt lakes, Lake Malata, Lake Greenly, 
lunettes, thermoluminescence dating, Bridgewater Formation, carbonate, gypsum. 


Tranacnons of the Royal Sactety of 8S, Ause (2002), 1242). 103 115, 


GEOMORPHOLOGY OF THE LAKE 
AUSTRALIA, AND TTS IMPLICATIONS F 


ALATA-LAKE GREENLY COMPLEX, SOUTH 
OR LATE QUATERNARY PALAEOCLIMATE 


by A. DurRiewic. C.C. Von Der Borcu! & JR. PRESCOTT! 


Summary 


Durkiewicd, A. von DER Boro, CoC. ASD Parscorl I. BR. (2002). Geomorphology of the Lake Malata-Lake 
Cireenly complex, South Australia, and its implications for ue Quaternary palacoclimate, Trays, R. See, 8 Aust, 


126(2), 103-115, 29 November, 2002, 


Luneties, foredunes dnd beach ridges from the Lake Malata-Lake Greenly playa complex on the Eyre 
Peninsula attest to major changes in lake level and palaeoclimate over the last 320,000 years. These have been 
dited by a combination of thermoluminescence and radiocarbon techniques, this allowing correlation with Late 
Quaternary Oxygen Isotope stages. The lakes experienced a major wet phase ca. 320 ka followed by inulliple 
and episodes linked to relatively cool periods and low eustatic sea-levels between 115-16 ka, Aeolian activily 
and aridity were particularly inlense during the Lust Glacial Maximum with the onset of a dry climate and 
carbonate pellet lunette-building commencing as carly as 26 ka. The Holocene palueoclinnte ix murked by 
seasonally oscillating wet aad dry periods reflected in the intermittent deposition of gypsum lunettes. curbonite 
ridges ind quarty foredunes around the eastern margins Of likes Malata and Greenly, 


Key Worms; Quaterniry palacoelimate, silt likes, Lake Matita, Like Greenly, luneties, thermoliuminescence 


dating, Bridgewater Formation, carbonate, gypsum. 


Introduction 


Like basins are one of the richest archives of 
ferrestrial palacoclimate data (e.g., Williams ev el. 
1998, Mason er a. S994: Rodé et al, 2002), In 
particular, surficially-closed basins such as salt lakes 
are extremely sensitive to changes in climate ind 
respond accordingly by adjusting their lake and 
groundwater levels. ‘They are widespread in 
south-western, south-eastern and northern parts of 
Australia where they often represent the termini of 
large endoreic basins (e.g.. Bowler & Magee 1988; 
Magee ef al, 1995; Macumber 1991: Bowler 1971). 
As salt lakes are susceptible to drying and erosion, 
one of the most challenging aspects associated with 
their study is resolving the problem of discontinuous 
strauyraphie records. This, however. can be achieved 
by examining and dating not only the sedimentary 
succession on the basin itself. but also the 
gcomorphologic features such as beach ridges und 
lunettes, as these invariably formed during major 
changes in lake levels and climate, In this study we 
describe strandline features of the Lake Malatu-Lake 
Greenly Complex (Fig. 1), which contain a rich 
record of major climate change exposed along. its 
eusterm shores, Most of these features have been 


School of Civasciences, Building 5, the Lintversity of Svdney, 
SSW 2006, Australia 

School of Chemisty, Physics and Earth Scienwes. Flinders 
University, GPO Bos 2100, 54 S00] 

Department of Physicn sine Mathematica! Physies. University: of 
Adelie, S05, Australia 


dated by thermoluminescence and radiocarboi 
dating and provide a framework for late Quaternary 
climate change in South Australia. These are 
discussed in detail in this paper. 


\ EYRE 


sae 
dating 
¢ } 
aid 
j 


feb ei 


PENINSULA 


\ 


\ 
‘ vik FARE x e } 
Ni Perrin Ps F, 
Soy / s 
sas Vy ¢ 
Yk 


PPNERSH LAG 
" ) 


; ) | ¥ 
OUBAN ‘ a { / = 
<4 [Tt f 
PUT as / ‘ / & 
tN { ( 4 
: a7 
= ’ \ o 3 
a OR i \., ryvewnyoaTan 
F } 7 \ sykalt 
} , ee \ Fo 
“ — / - 
“WRU ST RA LIAS me iWWeakog a 
{ f —l ina t foLant ' 
: eA SA | | a ! 
srupt we 
AREM iy 


Fig. t. Map of Eyre Peninsulit showing the location af the 
Luke Mulittt-Lake Greenly Complex, 


144 A. DUTKIEWICZ. ©, C, VON DER BORCH & JR. PRESCOTT 


General Setting 


The Lake Malata-Lake Greenly Complex consists 
of y chain of north north-east trending Quaternary 
carborute-cvaporite playa dukes. situated approx- 
imiwtely. 33 mabove mean sea level in a mid-latitude 
reviolh on lower Eyre Peninsula, South Australia 
(Fig. 1). The main basins. Lake Malata and Lake 
Greenly. are separated by an extensive easterly 
trending calcareous, sub-purabolic dine system, 
whieh forms the main regional aquifer. Emplacement 
of these dunes during the late Quaternary sea-level 
highstands (Wilson 1997)! rhost likely caused the 
danuning of a pre-Pleistocene drainage channel, with 
subsequent groundivater seepage along the dune 
lobes facilitating formation of lakes Malata and 
Greenly. Both basins are eroded along their 
SUULH- West margins with adjacent basement rocks up 
ty) S metres above the present-day lake floor 
suguesting L graben-type depression. Numerous 
smaller playas, located exclusively to the east of the 
(nuit basins. appear to have formed much hitler, via 
the interactive processes of deflation and 
droundwater disecharuc. Hydrologic. stratigraphie 
und geomorphologie evidenee collected to date 
indicales thal the main basins. have never been 
surlicidly connected, 

At present. all fakes fn the Lake Matlata-Lake 
Greenly Complex are ephemeral groundwater 
-dischirge playas characterised by a em-thick halite 
erust during the dry summer months, Depths of up ta 
(13 mol water. partly due to direet precipitation and 
partly due to redueed evaponition exist during the 
Wet wibter months, ‘The solutes are derived trom 
murine sult aceession vid aerdsels aml by 
evaporation of inflow (surbice and) groundwater), 
Which delivers chemical weathering produets fron 
surrounding sedimentary and basement rocks and 
syndepositional veeyeling of evaporites. The 
fiydrology afd geachemistry of the main basins have 
heen discussed elsewhere (Dutkiewler ef al, 2000). 
Although detined by the same mineralogical suite, 
basin sediments fram Lake Greenly and Lake Malate 
ure distinctly different. Lake Greenly sediments ave 
dominated by carbonate mud (calcite and dolomite) 
mensurioe severgl metres to decimetres in thiekness, 
with (he uppermost 3 m oof the basin sequenee 
interbedded wilh dim-thick layers of sypsarenite 
(Durkiewier & yon der Borch 1995), In contrist, 
Lake Malata is dominated by gypsum, which occurs 
in the fort of relatively mud-lree., m-thick 
eypsarenites, und maim-thick gypsunm-chiy laminae 
whieh overlie a cemened skeletal peloidal 


1 Winsor, €96  CPY Te “Geolory of the Qualerminy Bridvewater 
Momation of the Seuthwest and Central South Austad Pht 
thesis, Uhe Piiders Criversiry of South Mistral HU npabl ) 


grainstone near the base of the suceession, The 
skeletal peloidal grainstone overlies weathered 
basement. The difference in the relative abundinee 
of carbonate and gypsum over the lake complex. is 
related to the local hydrologic setting of each basin 
and raintalVrecharge distribution over the region, 


Geomorpholigy 


The morphology of the playa lakes depends on the 
nature of the pre-existing surfaee. the angle of ihe 
Jong basin axis to the direction of the prevailing 
wind, the presenee and depth of surtace water, the 
proximily of (he groundwater (9 tle hike surface athe 
playa-groundwater chemistry. Aeolian reworking, 
ground and surfice water fluctuabons and 
interactions play a secondary cole in modifying the 
lake geomorphology. whieh ulumately reflects major 
charges in climate, A number of geomorphologie 
feutures directly associated with the Like 
Malata-Lake Greenly Complex include islands. 
spits. luinetles, inrewular sub-parabolie dunes. beach 
ridges, sandy beaches. marginal seepage-spring 
zones and surface drainage channels (Pig. 2), In this 


1p LETH 
— 


~“onlter® yarn 
\ 


iv 2 Ceamorpholigic ina At the Lake Matatar Litke 
Greenly Comples showing the location af TL wid AMS 
Urfed suiiples. 


V5 


GREENLY COMPLEX 


4 
+ 


IMORPHOLOGY OF THE LAKE MALATA-LAKE 


GEC 


MPLOtl F £96 


PY LL F9T 
YY 61 F 9c 
BY oO + I 
PATE FOOL 


PYBOFE LI 
PY POF LIT 
BAP SF PSL 


TY LOFO'S 


+ 


WATT F TOL 
da P1600 + IRF 


ay 


(266) HOosalg 2 Zormaryind)) isa} nNealejd ay] uo neaieyd ajqnop ap Sursn paurwygo sunpasoy ayp Jo uaie 


IGOWAL SaTBIIPLL ATV JASUNOA Ary - 


(£66 NOdrsalg I? ZIMayNg Vy L/ F STE puke LY LY = TLE) Sade Ow) Jo adwlaav 
(S661) ‘JE 12 JOAINIG PUR (PEG) Jawiey pue sdAIMIg UO paseg LONRIGYED | 
ASE (PaLAq|RauN) [RUONUAUOGD 7 TCL AZO “ON GET ASH) Anauwo.qaads ssew joyesajanor fq parep 


‘piu ayeuoqies Ladue) pur jeraypad Apyaips 

aluoqies AYRYD palranpul 

Ajalpiapou 0) Apyean fo sajqqos &q ULELaav ayuogies [Bla] [ad 
“SIUALUAPL] JAfLUILURIOT PUP apoOrPAso “eyjalVoy lwepungy 
“puxs ZEN pauTess-asRos OF LUNIpaLU ‘paios-AfarRsapojA 


“LL uoysuies [eplojad pRiapays pauless-Lun pau ‘pavwos-|jaa *payuatuad <jayesapoyAl 


AL 
TL 
TL 
TL 
TL 
“LL 


“LL 
TL 


LL 


TL 


LL 


od APO PY (TL CTE ES = ST) OFS 
SINV 


POAT Suneg 


‘amnuogies <ypeya palwanpur Ajaysopow 

oO ApyBaw Jo sarqqos Aq wPpaAo pnw aruogir [Pala 

‘PUPS BIPAINOD SUIAPIAAO pri ayeuOqaes [RIAaTpag 

‘ake] ajeuoqira IBNpoU peyeanpul <jsuENS 

rade] aBUOGgIeD AYpPYS payeanpuT 

Ayaqesapour 2 FurAjsaag oyeuogsea perypjad snorayisdad AyysysS 
Jadny ayeuogmes parwmMpur uv FuUTAPIrg ayeuoqed eyed 
Jade arevogies pawinput ue Aq umpiaao ayruaqira Jade 
‘zuenb paureys-uor jurpungy watey aeuoqura 

JB[NpOU palwANpUl UB SuLApiaao ayeuogued peyaqjad ApYsysS 
‘qweuoques jeiayjad “Kadepo Apysrys “snosaypisd in, 

“auU0 Suess TepLojed 

[Ba[ayS Pauls-as.rod O} LIMIPALU *papos-jfaas “pajuatias APSuoNG 
Jade] ayRuogina payanpul ur Aq 

WIRLI2A0 ayeuOgied Aer [e1ayad puw snosayisdés ApYysyS 
“akRy] areuoqiva painput 

un 4q umpano ayeuogies Aadvyy qriayjad ApYyaysS 

Auiddes aysdds vw gq 

UIP[ISA0 altuaresdAd parros-<jaresapou ‘paurzas-asivo5 


anauny 12jad ayeuogma 
ayauny taijad aivuoqur 


quNpaoy zuene?) 
aunp oyoqrred-qng 


ayauny jatjad ayuoqury 
ayauny jajyjad awuoqieg 
anauny iayjad amuogquen] 


aHaun jayjad alngogirs 
aylauny yayfad aruogins) 
aveauny iayjad aruoqiey 


ayaun] jajjad ayeuoques 
ayeuny jajjad auogiey 


(Sapu RIEL aye) 
Asp aunpaiog 


anauny 1yypad arruogiry 


aneuny jajjad arruogiey 


aunp tunsdArnjanauny unsdin 


uondtissaq 


ainyeay aBo;oydsouloay 


819 
FID 


SUTW 
LUI 


STIN 
rlw 
El 
CI TN 
C/T 
L/TUTIN 


OVIW 
O1WN 


LIW 
IW 
TIW 
HINT 


atunyy ajdureg 


‘Saineay adojoydsowoes Aay fo sasv D,, Puy aouaasaurumjounayy, “| a[oRL 


106 A, DUTKIEWICZ. C. C, VON DER BORCH & J. R. PRESCOTT 


hig. 3. (a) A chain of playa lakes that have previously deflated into a gypsum lunette (Le). View is towards the north. Centre 
of photograph spans approximately 600 m. (b).A 5 to 6 m-high, Wuncated lunette profile at the south western margin of 
Lake Greenly, Basement is unconformably overlain by an indurated pedogenic horizon (P), which is iy turn overlain by 
an altered carbonate pellet lunette (Le), Gypsite layer (Le) overlies Le with a gradational contact. (¢) Lee side of two 
carbonate pellet lunettes (~1.17 ka ML14 and ~17.3 ka ML15) along the eastern margin of a small playa lake north of 
Lake Malata, forming a prograding lunette sequence separated by a samphire-vegetated mud flat. These represent Phase 
LI and Phase J deposition events in Fig. 4. (d) Thin-section photomicrograph showing bimodal carbonate pellets (>) with 
a minor amount of well-rounded carbonate-coated quartz (q) of similar grain-size, Plain light. Scale is 0.25 mm. 


paper we describe some of the key geomorphological 
features ranging in age from 319 + 72 ka to | 0.9 
ka (Dutkiewicz & Prescott 1997; Fig. 2, Table 1) 
which formed as a result of major climate change. 


Lunettes 

Although clay, quartz, and gypsum are the most 
common mincrals comprising lunettes (crescentic 
dunes associated exclusively with playa lakes: e.g., 
Bowler 1983; Warren 1982; Williams er al. 1991; 
Chen eta/, 1991; Macumber 1991), those in the Lake 


Malatu-Lake Greenly Complex consist cither of 


gypsum sand or sand-sized carbonate pellets. In 
general, the gypsum and carbonate pellet lunettes are 
part of a prograding sequence, which rises 2 to 3m 
above the present-day Jake floor, They are 
characterised by at least two disconformities in the 
form of pedogenic layers or erosional scarps and 
younger deflation basins (Figs 3a, b, ¢). The huincttes 
occur along the eastern margins of most playas in the 
complex und provide a partial indication of the 
amount of material that has been deflated from the 


lake basins. Their associated pedogenic horizons 
(disconformities) are potential time-stratigraphic 
markers for strandline-basin correlations. 

Four discrete units representing four major phases 
of lunette deposition have been recognised trom 
exposed sections and dated by TL between 115 + 14 
ka and 1,17 = 1.1 ka (Dutkiewicz & Prescott 1997; 
Pig. 4). The distinction is based largely on the degree 
and style of pedogenic alteration of the indurated 
carbonate layer (disconformity) overlying the soft 
lunette material, and field relationships of lunette 
deposits. Notably, progressive pedogenesis and loss 
of original pelletal texture are a function of 
Increasing age while the composition and colour of 
the lunettes and pedogenic horizons reflect the 
immediate source area, Internal structures, such as 
low-angle planar beds normally expected from 
seasonal accumulation, are very diffuse or 
non-existent and are attributed to the breakdown of 
pellets by moisture and pedogenesis. Individual 
deposits may reflect multiple phases of lunette 
deposition and stabilisation, although the general 


CEOMOREHOLOGY OF THE LAKE MALATA-LAKE GREENLY COMPLEX 107 


( 
5 lef beoka > W. 
SE s4- 0.7 ku =) S35, \y 
A Ss @ see 

W357 08 ko SS a — 
wa- wwe] 8 @ 8 Be - 
36 S 
“ou Phase Ill 
43.4 ~/- 


i@ Phase fl 


=F Bi 
| od 7* >. | Phase | 
<a ST 


ra ; 1 
| uf ~~ _| 
a . ———_ 
—— = 2 5 *™ w po f = — 
a 8 o 7 o an = \ Pal 
nN . a ow aa f hi 
F Bb _— / 
TTT! 
eat St 
a / f f Glasiay Caaelsurite 
/ ar tray La 
‘ / Mar Sealhabal) 


au Shute Hees itl 
OS) Graietony 


Intinnisel Gerrmanets Latent ine Seuquace tea, 
7 
Heals) Panetanval WT Holguin ies Liahoailanrsity 
ew 
.-_ «= Poco Someretioe é Hoepreany 


hl 46 1.2 ka Thermolumineseenve ages 


Fig, 4, ldeslised schematic west-east section of Lake Mulia showing three major phases of curbonure pellet lunmene 
deposition init single onlapping sequence, Thermoluminescence dates are also shown. 


homovenvily of the deposits makes differentiation of 
these difffteult, 

The pelletal fraction consists almost exclusively of 
Jow-My ealeite and minor clay Minerals dominated 
by montmorillonite, The coarse. non-pelletal fraction 
usually contains mipar amounts of abraded gypsum 
discoils OF prisms, quartz, iron oxides, peloids, 
foraminifera, Ostravede carapaces and Corelle 
Irazments, and trace amounts of Uthoclusts, pyrite 
and remnants of algal mats including rare charophyte 
oogonia, In thin-section and under the scanning 
electron microscope (SEM), the carbonate lunettes 
vousist OL birnodal sand-sized Carbonate pellets ana 
minor sancd-sized quartz (Mig. 3d), Quartz grains tend 
tO be well-rounded and earbonate-coated. while 
gypsum is discoidal, poorly-sorted, shows effects of 
dissolution and abrasion and frequently occurs as 
semi-cemented aggregates, Gypsite nodules are 
commen ithe gypsiferous lunettes, 

In general. the degree of pedogenic alteration 
corresponds with the age of the Junette and conforms 
with Netterberg’s (1967) calcrete classification, Por 
example, (he oldest lupeties show intense pedal 
development in the form of cavernous, nodular 
calerete und complete loss of the original pelletial 
structure in the underlying deposit, Younger lunettes, 


on the other hand. are capped by a chalky, powdery 
culerele or more massive and strongly indurated 
hardpan calerete, both of which are nodule-free aod 
requently comprise several undulose sheet liryers 
which themselves reflect multiple phases of 
pedogenesis. These are overlain ether by thin 
veneers of pelletal soil or younger, onlapping lunette 
deposits, The youngest lunettes display a strong 
pelletal fabric with samphire vegetation acting as the 
main post-depositional stabiliser, 

Three major phases of gypsum tunette formation 
hive been identified in Wis study. The best cAamples 
oecur virtually along the entire lengths of the inner 
and outer margins Of eastern Lake Mualata (Fig. 2) 
Where the luhettes form a prograding, cross-bedded 
sequence measuring up to 7 m above the present-day 
lake (Pig. 5a). An organie-rich layer within the core 
of the man linette has been dated by AMS at 5.59 kit 
cul BLP. and is currently being mined for gypsum for 
avricullural purposes. ‘The cliffed sections of the 
Lake Malata lunettes are onlapped by clean, 
well-sorted gypsarenite which forms the present-day 
beach, The finette sequences are stabilised by a 
Weakly indurated 40 em lo 3 in-thick cupping of 
gypsile which ts colomsed by abundant sult-tolerant 
shrubs and samphire yegeration. The relationship 


Luh 4, DUTRIEWIOZ © © VON DER RORCH & LR PRESCOTT 


Pig. 5. e) DWwe phases of gypsum lunelie deposilion lune 
the liner eusiern meetin of Lake Mahila, Reach snd OB, ) 
consists of cauirseegrnined oy paarenite. Younyer Phi U1 


evyesum lnncies ame stahilised carirely by vewelsition. 
Distance Wonk rhe Dake Malini margit re pkiyd luke 
immediitely behind Phase UA dunete is 400 1 (ba 
Gypsum lutente thy) representing Phuse 1 gypsum 
linote depasition overlying a-elayey carbone peller 
lnnette (ho) alone the etter caste marzin of Lake 
Malats. Poorly-sorted, coarse arained bench sam (B.) 
comprising, basement and skelter pelordal graimstone 
livharclusts forrus rhe presently beach, Note presence of 
Lonirse-prained sy hile eypstum sud te) onkppine Le 


belween the gypsum and Curbomule Tinetles ts nol 
always clear as (he gypsum lunetles are generally 
larger and more extensive, completely abseurinmy 
nniderlywing units which essentially become burriers 
for their developnient. Along the outer south-eastern 
Inarein of Lake Malate and the south-western margin 
of Lake Greenly. exposed sections clearly show a 
gypsum dunetie overlying a carboimue pellet wii 
(Pig, 5b). However tarther north and alony the inner 
margin of Lake Malata, the gypsum larettes appear 
to extend ina southerly direction away trom the 
Hanks of the curbonate luneties without directly 
overlying the carbonate peller units th expased 
sections. Field relanouships and & single AMS ilate 
sugeest thal the gypsum linettes are general) 
younger than the carhanate pellet lunetes. 

The gypsum dunettes coosist almost entirely of 
mediim to coarse-sramed, moderately 16 well surteu 
eypsucenite With small amotnts of varbouate (low 
Me calcite) and trace itimounts oF fine-grained quiet? 
and iron oxides. Consequently, the zypstm linettes, 
were unsutkible for TL dating and only in one conse 
contained sufficient orsanie Gurbon for UC dite, 
The carbonate eantent may be attributed either to the 
presence of carbonate pellets. carbonate coalime Ue 
gypsum crystals during their growth in the Take 
basin, or to biogenic components. Klagments of 
astricades, Cogella and the toruminiter /plidrinay 
are Common within the uppermost 65 en) of the mest 
recent gypsum lunetles. The aypsarenile corsests 0 
{4 mm tong. Sightly abraded anhedral lensotels 
marked by dissalunon kinks. ‘The thickness ob the 
Lypsile Cupp varices between 40 on fo 3 nan ts 
W function of the size and the age of the lunelte, Phe 
thieker id the more indurated. the gypsite horizan. 
the older the lunetie. The vy psite consists entuely al 
10 jc) long aciculorarystals under the SEM. Ciypsite 
Wise Qecurs ay em-hiek Trterbeds within He aypsum 
lunetle svyhere if mest likely represents srabiljsetiu 
of individual, possibly annual, aeolian layers, Tn (he 
same manner that the indurated carbonate layers 
represent diseontonmities within the carbonate peller 
lnettés sequence, the eypsite UNS represent periods 
of nor-depesition within the eypsum fibettes, Lavy 
angle aeolnin bedding 1 well-deyelopedt within (he 
gypsur funettes and reflects griin-size viarnilions 
and general sorte of the gypsarenrte willtin the 
individual lanimnac. 


Beach Ridees and kervduntes 

The most distinct veamarpholagie leatire 
asandidted with the Lake Maluté-Lake Creenly 
Comples isa 9 iehtgh, ainewate, indurated skeletal 
peloidal srainstong beach ridge dated by Th at 419 2 
52 ku (Duthiewiey & Prescott 1997: Table 1), avluets 
is present along the eastern marvin of Lake Mutat 
(“Lake Malata Ridge? in Figs 2 and 6). The ridge has 


QUOMORPHOLOGY OFTHE LAKE MALATA-LAKE GREENLY COMPLEX 10) 


LAKE MALATA 


CARBONS TE MUO 
OYPSAREN | TE 


29} SKELETAL PELOIDAL SANDYGRAINSTONE | PLETSTOGENE | 


PZ LeTERITISEN GAGLMENT (PRE ee FS Conley 


His. 6. Sehendatic westeast section across Lake Malar showing the western margin ol the bike busin bordered by 
culererised (C) sub-purabolic dunes (D), and the eastern margin bya major arcuale ridge (R) ancl a series of trunsverse 
eypsunt luneres (1) and purbonnte pellet linerres (Let and small pliyi lakes (P), 


been correlated with asinilar gradostone sampled by 
drilling, which overlies the basement in the Like 
Mahita basin (Ditkiewies 1996)" and represents the 
largest and quile possibly the oldest seomorphologie 
feature associated with the luke complex. The ridge 
is cuvemous and bas anup fo 1 en-thick capping oF 
wilerete, Which is locally overlain by a thin red soil 
contig abundant ferraginous pisoliles, The ridge 
sediments outcrop sporadically along the south- 
western margin of Lake Malate where they ure 
overliin uncontormably by a sypsunt lunette. 
Sedimentary structures were not Observed, possibly 
due to caleretisalion and a general lack of oulerop 
and exposure. ‘The beach ridge consists of a locally 
tufacecous, medium-grained, well-sorted skeletal 
peloidal grainstone comprising sub-ronnded to well: 
rounded micritised peloids, abundant mollisk 
fragments. coralline red algae, foraminifera, minor 
echinoid Traements. rare bryozoa and yarying 
amounts Of Tithoclists and angular quartz graits 
(Fig. 7a). Ws composition correlates well with) the 
Bridgewater Formation, whieh — comprises 
surrounding sub-parabolic dunes dnd spectacular 
clits on theawest coastol the Eyre Peninsula (Wilson 
199] )!. 

More recent but pervasive beavh deposits are 
found around the shorelines of Lake Malata and 
Luke Greenly aind associated pliyu likes. The 
composition of the beach sand depends largely on 
Ihe-souree and is Uekitess on the sediment supply 
and the proaunity of the source tothe lake thargin, 


Hokie AS CH%G) "Quaterminy Pakeochiate fre bathe 
Malia take Creenly Chive Comples, south Ausiralie’ THT 
thesis Phe HTindees Tiniversity et Sanath AUStrilal OF npubb | 


the water depil, and the teteh of the lake. For 
example. poorly-surted. very coarse sands and 
gravels are associated with basement outcrops, Thick 
(up to 4.5 77) localised accumulations of beach sand 
are common along the south-western margin of Lake 
Greenly aind southern Lake Maha. where they 
consist of Very course ingular quartz, Titholasts of 
busement tock, culerete, abd skeletal poloidal 
erainstone. iron oxides. Teldspar, mica, and skeletal 
pelondal allochems derived from surrounding 
sub-parabolie dunes, Fragments of Coxiella sp., 
ostacodes. faraminitert and charophyre ooponia ave 
occasionally jacarporuled, Medium lo coiurse- 
grained. oaderately-soried, skeletal peloidal sand. 
on the other hand. forinsa beach along the southern 
margin of Lake Greenly where the source is a selor 
parabolic dunes proximal to the lake basin. Ih 
contrast, curbonate playa hikes associated) with bake 
Greenly are characierised by shorelines dominated 
by biogenic fragments including ostracode valves, 
CoxieHa sp. Toraminifert and minor charophyte 
oogonia. These are oceusionally very aveakly 
indurated. Beaches associated with Lake Matlab, on 
the other hand, are dominated by coarse-grained 
gypsarenile und fragments of Coxviellasp, 

A 3 m-high foredune along at the south-eastern 
murginof Lake Matata situated less thin 200 ni west 
of the Lake Malita ridve ts-of particuliur imterest (Brg, 
7b) and has a maximum ‘Th age of | + 0.09 ka 
(Dutkiewiey & Prescott 1997: Table 1). Tt also hosts 
wi undisturbed Aborigmal campsite comprise 4 
stone grinding plate and scraping tools, Phe beach 
consists Ol medium to course-griined, moderitely- 
sorted sand dominited by course to fine-grained, 
anwular fo well-rounded quart. moderne amounts of 


aL A, DUTRIEWICZ, CO 


VON BER BORCI & FR. PRESCOTT 


Fie 7 he} h-seerion photunicroaphs alle skelert pelowlal geinstone eompranig the Luke Mahita Plemiacene siilee 
Phe Srioiseoite ds composedal ibundigt Hirimintiiend) 1A) are ines oie caleareenis nad ated Ga pan peloril alloehens 
Quarles alse present he grimstone is stromly cemented by miectie. Polanco taht (hy 2 tie high quart [reli 
Py! sepanued Tam the Like Mitta odee (Rs hy an ephertoc scepage-spring cone 1S) measuring approsiinately 
1st wade. The foredune has cain TL age ak =| ku, Tetes are uppony imately 2 th tall, ie) Serial view at che 
wisten munghinal Lake Greely Qiarmchtriitd by o prigmdme sequence (Jew lie carbonate pellet funeues Heb and 
carbonate Tses Ro Centre oF piatiprph spats TOO (CU TI section phate irayhoat scdimien) carpresiiy: Be 
IH ECE CH pascd OF DUNC sticede Traciietia WHT a iris rials Thine lit 


nihiuneyeaioed pelos, and fragments of pave sla 
sp. userodes wad foruotiniferd, Carbonate pellets 
Welh uppriimanely He) Tine to tics persia 
CAL? Satl ate presen) ata depr of tar below the 
Siti hati sciith, the deposit eridtes ite very 
cog prinicd unl pooslyesqrled beach safal 
WpProvimalely Tq above Tre piesenlaiiay fieke Lew 
The sand i dominared hy fine to tpurse-grained 
ungular und oeedsionally trarestuned quanme un 
fragments OF Cewe/la sp. Moderate ainiunts ot 
lithoelusts derived from skelauil peloncil zetitistiiics 
dod caleretes, (eer amolints ar peters judd felaspu 
iu Trace aunts oF charophyte oogoriuy ured jie, 
ine alse present, Tere, the sundsoverines a ilm-thitete 


humic eoyuinm layer dominuted te Trawenents of 


Covidia spe. anor amounts OF quire. camel brave 
umiuuiits of foeninitera. asteawodes. vluabophyte 
Henin, pelowls mea and reeem vegermin, Thly 
layer a tum. overlies a stronwely indunited skeletal 
pelordal eraeistone whieh euterops tualy alany the 
Hanon al the lake, 

The gevmorphologie Tbatures along the eastern 


Tre Lake Greenty are sttking ly oiterent trem 
those of Lake Mahi This isammibuled minty me the 
Uillennces i basin merpholowes, pardcularly the 
anmeniatiad af the love basin adis ta the dincetionn ad 
the prewar wid, the ceatire at the oases 
sealimnonts, aid the eroufdwatey chemisipny. ‘The 
wssocniled Luneclicerdwe systeuy is low ye ached uid 
dificult hy map het unmisthkuble wher seen Prom vir 
(Fin Teo Wi tack mapping of the individual lanes 
and ridves could only he aehieved using aertil 
phowieraply. Pn uddition. exposed lane sects 
aré rire jnd tiheties. uippear to have formed a 
progiadime sequence, soureed by episodic deflation 
wih Lake Greenly. Unlike the prograding linethe 
sequence al Lake Malata, where individual bietles 
ere senuraled by dellatinnary busin. the Take 
Greenly linertes appear to he sepanired by a series ot 
curbomare cides. The fides are very senrihin ay hand 
Specuinen wed in Hulerap style fo the tiduroted 
carhonute horizons associited With Tinettes which 
thes the distinetion of these leatupes estrenpely 
Heute in the Tele 


ChOMORPHOLOGY OF THE TLAKE MALATA LART GREENLY COMPLEX WW 


Phe mast casily recognised carbonate Ades oceans 
proximal to the present-day northeastern luke 
margin. where four pluges representing four phases 
Ol dake repressian have been reegenisad, The ridges 
ae approxymately 2 to Fm above the present-day 
Hoorn ot Lake Oreenly and Form fractared and rabbly 
carbomile sheels, They lack the smooth and 
CONMHGetS UPcte outerop style eommoan to 
indirated curbenate horizons usseciuted with 
Hinettes, The orists consistaot low-Mep caleite ane dre 
chilky and (friable, with ae min-thiek coating ol 
laminar calerete, Tubular-vaids are abundant more se 
(hun in the indurated carbonate horizons associated 
With carbonate pellet Tinettes ynd appear to be 
relied to phint growth in rekiliyely soll sediment, 
However, no fossil plant remains were Tine, 
Scanbing eleetram micruscape analysis of these 
erusis Shaws the prescuce of shundant, straight or 
mently ourved, decasionally branching, ~ 5 prick 
taledeonerusted oendolithic  trayments, The 
mourphulogy and sive of these filiments are 
CONSISTENT WIL Tongot strictures. described by 
Kluppa (1979) find jrdicate pedoyenesis in the 
subactial yadose environment, tia thinesection, the 
crusts consist ol mierite. whieh oecasionally displays 
a oelohular texture und abundant, geuenidly 
ninidomly oriented, caleitie shell fragments whieh 
coMptise 5 to (5% of the total sediment (Fig. 7h, 
Vhe shell fragments consist oF low-Me caleite am 
wre generally steaight oor gently — eurved. 
upproxsimately 1O pon in diameter and generally Loto 
Tom oh tength and most likely represent 
disurliculited ostricwode valves. Foruminieral 
fragments are rare. "The deposits have not been date 
due lo the patieny of suitable materiits suele as 
Ormunie mader ane quarts 


Discussion 


Vhe Lite Quitumnary veamonmphaligy im the Lake 
Malita Lake Geeenly Complex is represented. hy a 
comiples supe al ridves, luneltes aad) forediuner 
Which have heen chitwu bebyeern 320 Kuand | ka. The 
penod eyvers a tine of dramatic climate oserthalionns 
during whieh the formation ol the lake comples wus 
nitiited and the likes expericneed a nmajor lacustral 
(Wel) plane followed by uw series of drying and 
deflitionary episodes puretiated by periods of 
Pedosehesis and fehuively wior dacustral events, 
Oserlalins hetween these climilie extremes 
culminated in the presenl-dity states ol the likes js 
WrounUwarerdischange plats. 


Wer phase ta, 2b ha 

The tiny licustral phase is represented by the ra 
V0 bul (Ohyeen Isotope Stige 9: Mies 8) Lake Milita 
betel videe fries 2 ghd Ay deposited dni a phase 


of high lake level, [ts morphology is consistent with 
foredune deposition and we-envisawe thal ih formed 
by deflation of sand from wave-nourished lakeshore 
beaches in very mueh the sure niiner thar coustal 
foredunes and foredine ridges are built immediately 
behind cones of beach swash, Prior to stabilisation of 
the surrounding sub-paraubolic dines, a laree yenount 
of the skeletal peloidal sund was blown and wasted 
inte the like basin und subsequently reworked by 
wind-denerated weves diiriige ay plivial climatic 
phase. A combination yf cehitively lower than 
Present evaporation rates, hiehhy  effeenve 
prewipiahion, mereased runadland recharge and high 
willer ables, associated with an interglacial segelevel 
higwhstand. would Haye resullect in the accumulation 
Of relatively fresh water within the lake basin. The 
size of the ridge suggests that ab least | oie oh water 
was present in the lake basin during the wer winter 
Months. Whielh would Mave been -chiatactersed: by 
higher rainfall and lower evaporation relative Wve 
present Such a rehitively bigh luke level. combined 
WITH stron north-westerly winds assocratce wath the 
Winter months. sould have pitied wave-ve erated 
currents capable of moving large volumes ol the 
skeletal peloidal sand as bedload towztrds the custern, 
and partieulirly the south-custern dake marin where 
the ridge attains jis manimun width, Under these 
conditions the sane veecreted on the eastern hikeshore 
mW! Lake Malaha as a heagh deposit and was 
subsequently delhited by strony southwesterly 
Winds nite a foredine immediately behind this high- 
enedgy beh, The henht of the beach ridge attests lo 
the faet that fas period was relitively lone-lived, 
vhoracterised by enhinved seasonality and a large 
and comliniaus sediment supply. The vidge was 
eventially stabilised by pedosenesis and vegetition 
during an extended period af tondleposition, That 
only one such feature is present within the Take 
Malita-Lake Greenly Coniples tiuicates iv unique 
depositional episede, The absenee ol a sinvilar 
foredune midge long the waster margin ol Lake 
Gicenly miy be aliribuled not only to a lick «) 
sediment supply, bul alse to the basin morphology, 
puriicubirly the ortenhition ol the lon wads to the 
direction of the prevailing westerly wind, The Lake 
Greenly basin is oriented approximately at 45" vo the 
direction oF the prevailing westerly wind, while the 
Take Multi ages lesa 907, 

Consequently, Uhe cidee ceprescuts a Pleistocene 
“nieealake or" bacustral” shige (csi Bowler 1980, 
1YSl: De Deckker 1985) i the evelution of The lake 
busin and overlaps wih Wilsan’s (180i! Phase U1 
deposiiign ol Bridgewater Furmacen duoes duriny 
mid to hie Pleisticene WMloervlacial sea-level 
highstands. Meouluke shiarelines. such usa 13.5 1 
high beuch at Leke Tyrrell, contain abundant 
sells of Cruella sp, (Macuimber L980) Bowler de 


12 A. DUTKIFWICZ, CC. VON DER BORCH & J~R- PRESCOTT 


SPEGMAP isotope record 
(stacked and smoothed) 


a18Q (per mil} Depasitional Events 


a 
3 5 
e =8 i 
A ‘ we = 
50 aoaa Sas : | 
nwo th 7 
il | a = TE wy 
S " 12a 
S Ear 
en 
100 gg 
ia 


200 


Lo JA)EM ARSE 


250 - 


\ey) aby 


aBpee LjaBSey 
SEE) 27 


44 Rensopug 


400 


460 


S00 


Fin BR. Ages ob maja geonomhologe feaniies in the bake 
Cireenly-Lake Mulita comples ploticd against the 
SPECMAL conve Chibeie eral TYs¥s Mebatyre er ri 
1989). Aves of Hic isotope stages | 10 6 after Martinson 
ell (IOX7). Ages for the deposition of ihe Bridgewater 
Fororation yiter Wilsent (1991), Renin nunienils denote 
glycitl terminations. The lenath of cael bar indlentes 
Uncertginty (a mepsurement of Me que of litette/Mdge 
(sve Table | for detail), 


Teller 1986), Curmousty, there is a lack of lacustrine 
fauna (fresh and saline water-rolerant) within the 
beach tidve grainstone and equivalent sediments in 
the basin. Possible explanations for this are: 
1) post-deposivonal dissolution of organisms caused 
by achinge in the physical and chemical environ 
menls; 2) destruction of shells during high enersy 
Iransport rendering them unrecognisable in the 
sediment revord: and 3) the high-energy, sandy lake 
basin may bave been dnsuitable for lacustrine 
OV QUNISITIS, 

Formation of the ridge may have commenced 
considerably earlier than 320 ka, which murks the 
wane shines of deposition from the lee side of the 
ridge, Wilson (1991)! proposca that the cmphiecenicnt 
ol the Bridgewater Formation dunes along the west 
coast of the Fyre Peninsula occurred during sea-level 
highstinds as early as 700 ka (onset of Oxygen 
Isolope Stave 17). Therefore, it is likely that ritye 
aceretion alsa aceurred episodically throughout 


Oxygen Isotope Stages 17 to 7 and Stage 3 
interglacials (Fig. 8), until the sediment supply was 
exhausted and recharge rates decreased due ma fall 
in eustatic sea-level, giving rise lo ow new 
depositional regime in the Lake Matata-Lake 
Greenly Complex. A comprehensively dited sequenve 
through the Lake Mulata ridge could potentially 
provide paleoclimatic information prior to 320 ka, 


Palireoclimente va. J20-115 ka 

We have no direet recor) of sedimentation and 
evolution of the lake system for the period 320-115 
ka, As mentioned earlier, the Lake Mihila ridge may 
have continued acereting intermittently during 
pluvial episodes, particularly during Oxygen bsotope 
Stage 7 until ~ L80 ka, when the emplacement of 
Phase (1 dunes alone the west coast of the Byre 
Peninsula was most intense (Wilson 199])!, The 
morpholowy and pedogenic alteration of sub- 
parabolic dunes overlying thick carbonate sequenves 
south of Lake Greenly (Dutkiewies 1996)" are 
consistent with a later episode of Wilson's (1991)! 
Phase LE dune emplacement during the last 
interglucial (Stave 5; Pig. 8). This, suggests that 
deposition of lacusttine carbontes most likely 
occurred prior lo and intermittently diriiy the wari 
interstadials of Oxygen Isotope Shige @ and during 
the warn intervals Of the Tast iiterglicil (Oxygen 
Isotope Subestige Se). Lundiward migrating sub- 
purtbolic dunes would hive buried tegions relatively 
close to (he coash whereas more distal ures, such its 
Lake Malata, would have been subject to continued 
curhonite precipiition. barge volumes of ubis mud 
are likely to have beep deflated into lineties during 
Oxygen tsolope Stage ©. which is browdly similar lo 
staves 4+. Sand 2 during which lunetle deposition vats 
pervasive. Further sempling and dating ts required 
to decipher the palaeoulimare record during his 
period, 


Mulliple Arta Bpinodes cu, 113-10 ka 

Lunetles form by dellation of siatidess Zcd material, 
which commonly includes pelletised clays derived 
from a drying lake floor by uni-direetional wind 
(Bowler 1973; 980, 1983). bactors mvolved im. rhe 
construction of clay pellet limeites (Muctuating 
wroundwater Jevels, UniediWectional wid, aridity) 
have been cliscussed extensively by Bowler (1973) 
und the same explanation can be applied lo the 
carbonate pellet hinentes from the Luke Malati-Luake 
Greenly Complex. ‘The ages of (he linettes indicate 
that seasonally arid climes and intense prevailing 
westerly winds it southern Australia occurred 
several times sinve the last interglacial. Although the 
PLoawes are not sufficiently preeise to dite the exnet 
onset of each arid episode, at the very beast (hey 
indieate the time when Tynette building wis in Call 


GhOMORPHOLOGY OF THE LAKE MALATA-LARE GREENLY COMPLEX Vs 


swily, Dy gweverul, these correlate with periods of 
relatively low eustatic sea-level and oscillations to 
cold intervals. many of which had net previously 
heen assoeiated with cominental aridity and lunette 
building. The oldest lujene bocizon dated ca. 115 ka 
corresponds toa the dast glacial meeplion and 
fermination of the last interglacial (Oxygen tsotope 
Sub-stape 5d). Acoliin activity increased again ated, 
96 ka (Oxveen Lotipe Sub-stage 5c) 85 ka (coll 
Sub-stape Sb), and 75 ki and 70 ka (Sub-siages 
Sad), Significantly, @ strongly-indurated pedogenic 
horizon dated arcu. 75 ka sugges that Ute luvette 
miverial was most likely modified soon after the shitl 
trom stave S to 4 Which elobally marks the main 
ehtaal transition, Similar perrods of deflation and 
pedogenests are estimated fo have oceurred around 
OW ka und before ubout 20 ka tothe Madiwan Gulf at 
Lake Eyre (Magee er o/. 1995), Lunette-building ia 
the Lake Malakebake Greenly comples occurred 
twiee aluring the interstadial of siage 3° with 
(EXNHOM AcUlia uetivily Centred around the aeo 
vold sdadials ca. 54 ku and 43) ka immediately 
tollowite the end ob the main glacial transition, 
These aves likely correspond to the 60-50 ka playa 
defliution phase and done building at Lake Eyre 
(Mawee & Miller 1998). Untike Lake Eyre. however, 
there js no evidence at Lake Malia of Lake Greenhy 
tora mayor lacustial phase in the period S035 kit 
(Magee & Miller 1998) Towever. further exca- 
vation work ts required to fest whether a beaeh 
depisit of thts ae mathe be buried beneath younger 
acohay sedfyeuts, 

Several Tunetles i the Jake systenr dialed at eye. U8 
ft 7 ka aod 16 ke, cluster on Oxygen tsorope Sue 
2. which marks the peak af the last aliciation for the 
Australian continent around 20 and 17 ka teas, 
Bowler 1986; Colhoun L991). The age of the oldest 
luneibe near this cluster dated ui ca. 26 ka 
comesponds ta the casino benveen Oxygen 
Isutope Stage 3 and Stuge 2, indicatine that the onset 
oF ri conditions ancl linet building during the hist 
shicuil maximum commenced as early as 26 ka in 
(his purtol Australia, This corresponils to a general 
(levrease in the pumber of high and intermediase 
lakes in Adstralia after 26 ka (Harrison 19934 aad the 
unserola dry-lake phuse around 30 Ka ut Lake Eyre 
(Mavee & Miller 1998). Aeolian activity appears to 
have peaked coy. 17-S-16 ka, which correlates well 
With the hast Glaciai Maximum ub b7.4° ka 
(Martinson ef a T9871 During this glacial pert 
the sea level was at its lowest and the climate 
experienced intensiticd aridity and high westerly 
with! speeds (Bowler & Wasson Psd: Perit ee el. 
1900) conducive fo pervasive dune-boldme aver 
urtd aml semiarid regions of Australia (Bowler & 
Wassan (984) Wassat) PO86), Luperte-building: was 
ih ts wig Stages ea The TS ha. wath local 


deposttion still oceurrmig locally until ea, 15-6 ka aad 
Was resiricled ta northern parts of the lake complex. 
Based on records rom upproximutely 35 Australia 
lakes. ELarrison and Dodson (1993) suggest w bref 
interval lo weller conditions during 15-13 ka, which 
Is consistent with absence of lunette sequences at 
Lake Malai. These authors further propose that arid 
condivions persisted alter the last glacni) maxinuny 
CUINAt Oh ndariuan aridity at 12 kay by whieh 
time most Australian Jakes were dry. This. wool 
correspand to pedogenesis of Last Glacial Maximum 
luneltes i the lake complex. 


Weare evetey mm fhe Holocene 

Gypsum funettes in the Lake Milttu-Lake Greenly 
Complex have formed in two stages. in a slihtly 
different manner to carbonate pellet linettes, The 
pypsun Cirst preeipitaled within the Take Basin in 
association with groundwater oscilhiions and 
evaporation attire eapillany trite (Teller eval, 1982: 
Bowler & Teller 1986; Magee (949]). Although 
sandesized diseaids eXposed dirjag a dry period 
when the like levels are low are easily deflated by 
prevailing winds, the similarity i grain-size and 
Morphology of gypsum forming present-day beaches 
and the youngest linettes ut Lake Malate stiggests 
thal (he most reveent pypsum luneites most likely 
formed by deflation of reworked material deposited 
at the fake margin during an ection relatively higher 
lake level. Sinve surficial sediments in the Lake 
Mahuta busin ape dominated by hermi-pyraonual 
eypsaremile, a combination oF a thin skin of waler 
amd strom vind wuld) provide an efficient 
techanisny lor trumsporting and depositing the 
eypsune atthe lake margin. Trinsportition by wave 
wetton is farther supported by the presence of ripple 
marks en gypsarenite-dominated playa surtaces amd 
by the abundance wf biogenic fagmenty witha the 
most recent deposits. The gypsunt is subseygently 
deflated and sorted during a more arid period, 
Therefore, the gypsum lunetes most likely represen! 
foredunes deposited under seasenally oseilauog 
relatively high lake levels pind rebitively low lake 
levels in response Ro Chuuging evaponiban/inflow, 
Strong winds daminanted by a westerly component 
ure reyinred rhroughout the entire vycle af depositian 
and reworkitig, A single AMS-dated horiven trom 
Me myiddle of the lunete indicates that this process 
war well underway ea. 5,6 ka cal BP. mosi likely 
coineiding with the Holocene sea-level bishstund ey 
64 ka (Belper ee uf 2002). The omean annual 
preeipiunion ut this time is estimated te have 
increased by 20-50% (Wasson & Dennelly 1091) 
with masximam lake levels. recarded it most sites 1 
Australia (Howley PYS~: Wasson & Donnelly [9U4, 
Harrison 1993; Harrison & Dodsai 1993), 

Although gypsum linettes have formed in the 


ne A. DUTRIEWICZ, © 0. YON DER BORCH & JR PRESCOTT 


relatively recent past at Lake Malar. this has not beer 
the case at Lake Greenly. The reason tor this is Ural at 
hake Greenly gypsum oceurs several decimetres 
below the Jake surface beneath dolomite carbonate 
muds (Dutkiewiee & von der Boreh 1995). In this 
seenario, surficial curbonate would first have to be 
pelletised and deflated before interstitial 2ypsarenites 
wre exposed to underzo reworking. Therefore, while 
gypsum lune(tes were forming inthe relatively recent 
past al Lake Malata, carbonate ridges were more 
likely to form coneurrently al Lake Greenly, The 
complex system of Tuneltes and ridges wt the custern 
margin of Luke Greenly suggests that this may have 
been the ease. The fraemented nature of the ostricode 
valves in the Lake Greenly mdves serves as an 
indieation of reworking by waye action during 
relatively hivher lake levels. The ridges consutute a 
prowcading sequence lormed hy 
lacustrine carbonate mud under sridially regressing 
lake shorelines. The carbonate mut has undergone 
subsequent swbilsauon by vegetation followed by 
pedogenesis ind. induration in the subserial vadose 
environment Consequently, these beh ridges are 
excellent indicators ot the former lake extent aid 
although undated may be concurrent with the 
lormiution of sypsum lunetes at Lake Malata. 

The recent 3 m-hizh beach deposit (MLA, Fig. 2) 
ab Like Mahia the remobilisation or hich has been 
dated at va 1 1.93 ka (Dutkiewiew & Prescott 
1907), represents a foredune formed by aeosian 
reworking oF coarse beach sand. Acoli deposition 


exposure af 


is Sopported hy the finer grain-size und better Sorting 
of the sand compared ty other beach deposits of 
broadly similar composilion, and by the presence | 
carbonate peloids. Tn particular the coyuina layer 
within the deposit is indicauve oF a dicustrid period 
diving Which relatively high lake levels and lower 
salinities caused by increased precipitation and/or 
dvereased evaporalion rales would hive allowed 
large nuinbers of Cenelis gastropods to inhabit Lake 
Mahuta. The orientation of the (eredune along the 
soulleeastert! shoreline is consistent with the 
orientation OF prevailing southwesterly Wilds, 
whieh operate during the dry summer nionths. 
Aeolian activily was generally high atthis time and 
is further suppeited by the most recent linetie 
building episode in the Like Malata region dated ut 
ea. 12 ka (Dutkiewies & Preset) 1997), 


Acknowledgments 


The Flinders University aod) the upchaeometry 
special fund af the Physies Department, University 
of Adelaide, provided financial support lor this 
study. which is based On PhD research carried out by 
A, Dutkiewies. AMS dating wats funded by AINSE 
Grants 93/127 AMS and 94/176. AMS. Chits yon der 
Borch acknowledves the continuing suppert of the 
Schoal of Physics, Chemistry and Earth Sciences al 
Flinders Wniversity. We are grateful to Martin 
Willams for fis constructive comments on the 
LUSCH pt, 


References 


Rel peRIO, AJ BL HARVEY, N- ae BoulkeyAn. ROP. (2002) 
Spaliah sok feniporal variibility in the Holocene sea 
level records of the South Austeaban codstiine. Seah 
Geel 150, (54. 169. 

Bowler, I. M. ()971) Pleistocene salimihtes and clinvatic 
change: evidence from likes and linetles in southeastern 
Austrilia pp, 47-05 47 Mulvaney. PL & Golson, J 
(has) “Aboriginal! Man and Bayironment in Adstealie 
(ANU Press. Canberrit), 

(1974) Clay dunes: theie occurence. formation 

qn onvinouimental sanificanee. Fadi sed Kev 9, 315- 

4AR, 


(1980) Quaternary chronolowy and pues 
~~ hydroloey in the evolution of Mallee landscapes pp. 17 
36, In Storrier, KR. RK, & Stannard, M, &. (Eds) “Avelian 
Landscapes in ihe SemeAnd Zone of South Fastern 
Australia’ Proceedings of the Australian Suciety af seal 
Seicnes Hrearparated Conferenee. Rivertna Rraneh. 
Mililine. 

, oe — WYs) Atistraliin salt hikes: a palioe- 
Hydrologiest approach Hverbielagia 82, 431-444. 

(1983) Lumnettes as indices of Nydralogic 

Ro Nene 


changer a review of Australian evidence Pro 
Vie 95. IAT-10%, 


(JO8G) Spatial varkibriity und Fydrolawe 
evolunion ‘or Australia like basiiis: Anilogie fur 
Pleistocene hydrologic chiihge dnd evaporite lormation 
Melaermenn, Palacor linn, Tilivooeel. S42 l-A], 

a & Mache, 2 W. (988) Luke brome: 
strittigraphy. facies analysis and evalieiion oF a nay 
playa pp. 4-6 In SLEADS Conference 38. “Salt Latkes in 
Ari Austra’ Austridian Narional Lniversiny, 
Canberra. 

& Tercrk 1b (O86) Quiternany evaporites 
~aind TD hydrolugie finieet Lake ‘Tyrrell, north-west 
Victoria. Avan 2 Bain Sel. A8. 43-63, 

Ae Wasson Rood. (TYN4) Glacial age 
environments af inhincl Australie pp. EX3208 fe Vouel 
1G. (EL) whale Catnoveie Paulaeoelimates of the 
Southern Hemisphere” (Balkema, Roulerdan), 

Cries, XY, Bowne. dM. & MAGI J WC E99 DF) Acoli 
landsuaper | me central Austiilias Giypsiterous and quarts 
dime enyironments fring Lake Amadeus, Sediorentuliags 
AK SLY S38, 

Corndgun, BOA. OI98t) Climate during the Last Gkicial 
maximum in Australia und New Guinea. Australian ane 
New Aealand Geomorpholosic Group Spee! 
Poblication No, 2, 


GROMORPHOLOGY OF THE LAKE MALATA-LARE GREENLY COMPLEX 11S 


Dt Deki. PoUTYRS) Late Austadian lakes dori the 
lust 20 million years: sites for petroleum source rock or 
Invlal ore depositivnsor both! pp, 45-54 Jy Kelis, Aad & 
lalhot. MER. (ds) “Laeistrie Petroleum) Source 
Rocks” Geological Society Special Publicition.. No. 40. 

Dorkinwies, AL & PReseori, do Ro (1997) ‘Phere. 
IWMineseenee aves and palacoctimate rom the Lake 
Malhtli-Lake Cireenly Comples. Eyre Peninsiti, South 
Australi, Quan Sei Rey, 16, 367-385, 

4 Henevkric A. bk Diiiton. JC. (2000) Past 
changes Lo isotopie and solute bulances in ud contietil 
pliya: clues from stable potopes of lachstrine 
carbonates. Chem. Ciew! 165, 309.329. 

& YON bh Boker (1995) Lake Cireenly, Eyre 
Moniisiha, South Australias -sedimentalowy, 
puliwochimatie amd palivohydrologie cycles. 
Taldeoseos., Pedaeactin, Pulecwecnt, 113. 43.56. 

HARRISON, S. PB, 1943. Late Quaternary like-level changes 
and climates of Australia, Quart, See Revy 12, 211-23). 

& Bovson, J.(1993) Climates af Australin and 
New Guinica since [A000 ve BP pp. 265-293 
Jn Wright Bl EL. Rutybaeh.o Jo bo dee Webb TIT, 
I, Ruddiman, Wo SweetPerron, FA & Bartlein, 
Pol (Rds) “Global Climates Since the Last Glicil 
Maasimuin” (University: of Minnesota Press, Mrinesota 
Minneapolrs). 

IMukih. L Metpivar, AL & Mix. AL (1989) Ovonic 
Response to Orbittl Poreme m the Late Quarernary: 
Observauonidl sinh Rxperimentil Scrategies. pp. | 21-164 
Jn Berger, A. Selineider, 8S. Hw Duplessy, IC, (lids) 
“Climule qind the Creoscienees” (Klawer Avademic 
Publishers, Boston. Massachusetts) 

Kiarm. © OF C1979) Lichen strommitotites: eritenon for 
sukterkil exposure vind do mechinisin far the lormation of 
laminar caloveres (culiche). J. Sed, Pel, 49 487 400, 

Mctniryke AY Ro boiMaAs, W. i, Rartin, i. & Mix, AOC, 
(1988) Surhice water response of che Equator) Athintic 
Qeean ly orbital Goreing, Paleoeeun, 4. 19-95, 

Mactmben. POG (1980) The inflience of srotadwater 
discharge on the mallee landscape pp. 67-84 fer Storrier, 
ROR A Stenmard, Mob (hds) “Acolian Landscapes in (he 
Semi Arid Zone of South Bastern Australia” Proeeedinges 
of the Austitian Sociery of Soil Science Inevorporated. 
Conterence, Mildura, Riverina Brinch, 

_ YO lareractions Berween Grounu-witer and 
Sulace Systems in Northern Vietoria. Department of 
Conservation sid Environment, Australia, 

Machi, dW, (l90D) dane Quateriury lacustrine. wrounil- 
Waller, aeolian and pedopere pypsum in the Prongle 
Lakes. southenstem Australia. Pa/aeeuiog., Mileoor lia, 
Palittoweal, 84, 4-42. 

———— & Milibe Go Ih (1995) Lake Lyre 
palacohvdrolomy From 60 ka tothe present: beach rides 
Hd wham Wein aridity. Javed. 144. 307 320 

> Bown hk IM. Milirk, G. HL, & Witttams, BD, 
Lo, (1995) Striligniphy, sedimenrology, chronology and 
palicohydrology ob Quaternary Tacusiine cleposits al 
Madigan Gull. Lake Fyre. South Australi. de, 113, 3-12, 


MAK NstiNg. D) OG, Pisias, N,G., Hays, dt. DL Inibing a 
Moorn, ToC. tn & Siaekibtos. No. (19887) Ape 
(hit ancl orbital theary of ee ages: develupment ofa 
High-resolution Q te 300,000) vears ehronostratiorapny 
Queat Nes. 27, 1-29. 

Masons. 1M. Guckowska. ML AL. Rathi. CLG. ae 
SHWE PrRROT FO AL (1994) The respanse of hike 
levels wend areas to clinic change. Oftin, Clainuy 27. 
1ot-197, 

Nerrprahed. EF C967) Some rowdimaking properties of 
South African calereles pp. 77-41 dn Proceedings of 4th 
Regional Conference an Sou Meehanics and 
Foundation Engine rims, 

Prev. Re Mouser. bomen Kokorkeyv inci ¥. 8, 
Roiakoy. Vo 1 de Lorius, C. (1490), Palacoe 
Gimarolagical aod chionotorical implications of the 
Vostok wore dust record. Nafare 343, 56-58, 

Revo, XN Git S.) Baraayens, Fo CaMin, FA, 
Tenorio, Ry G & Jhiis, Ry (2002) High-resolution 
swine lake sedimeurs ais enhaneed tools for relating 
proxy palcolake reeorls to recent Clue Git series. 
Sed. Geel, TAB. 203-220, 

Sroiver, Mo& Reimer. Pb (1993) Extended 'C database 
and revised CALIB aaieearane Culibrsliie prow. 
Rodiocvarbion 35, 215-23 

—_~« eae . Breck, JW. BURR, G. 

Thee 4K. A] KROMER, £. MceCorMaAc, ECs. 

x Poul fo & Srerkk, M (L989) INTCALOS 

Radivcurbon age calibration 24,0000 cal BP. 
Radiocarbon AQ. 1041 1Oa3. 

Tree. Th. Bowrre dh Mo& Macrnner. PG (1982). 


Moers sedimentatuin in Lake Tyerell Vietoriat. 
Australian Geol, Soe Aust 29, (59-79. 
Wacken, JK (982) ‘The hydrological setting. 


occurrence and significunee of gwypsiim in late 
Quaternary salt lakes in Australia. Seeimeniolouy, 29, 
ANO-637. 

Wasson. Rod. (1986) Geomerpholoey and Quialernary 
Nistoryool (he Austratian cantinentel dunefields, Ceo, 
Rev, Jape, 89, 55-67. 

je & Dossenby. TH, (1991) Palieoelimatie 
reconstructions for ibe Last 30,000 yours in Australia = A 
contribution to the prechenon oF Future climate. CSIRO 
Division ot Witter Resources. Technical Memoir No 
91/A, CSIRO, Canberra. 

WiLtiAMs. M. ALT. Br Dre nnn BR, ADAMSON, DLA. & 
Talbot MOR. CMY1) Episodic Muviatile and aeolian 
sedimentation OW a late Quaternary deserh marin 
system, central western New South Wiles pp. 258-287 
Jn Willitms. M.A. J, De Deckker Po& Kershaw. ALP 
(Mls) “The Camozoie of Austraha: A Re-Appraisal af 
the Evidence” Geological Society of Australia Speetal 
Publichtivn Nats, 

Willi AMS, M,, Dinkeriiy, DB De Deckkek. PB, 
Keesiaw, Po& Coa. J. CHY8S) oQuaternary 
Movironments” (Arnold London) 


SMALL SCALE SPATIAL DISTRIBUTION PATTERNS AND 
MONITORING STRATEGIES FOR THE INTRODUCED 
MARINE WORM, SABELLA SPALLANZANII 
(POLYCHAETA: SABELLIDAE) 


By CRAIG A. STYAN*! & JOANNA STRZELECKI* 


Summary 


Styan, C. A. & Strzelecki, J. (2002) Small scale spatial distribution patterns and 
monitoring strategies for the introduced marine worm, Sabella spallanzani 
(Polychaeta: Sabellidae). Trans. R. Soc. S. Aust. 126(2), 117-124, 29 November, 
2002. 

Spatial distribution patterns were determined for the introduced marine worm, Sabella 
spallanzanii (Gmelin), at Largs Bay, South Australia, as part of a study to determine 
the most efficient methods to survey these worms living on soft sediment habitats. 
Worms were patchy and, across a range of spatial scales, more likely to be found 
together than if they were randomly distributed. Average density varied among sites 
between 0 and nearly 10 worms m’. 

Key Words: Sabella spallanzanii, introduced species, marine pest, survey, spatial 
pattern, distribution. 


Trmsar ions af the Reval Soctery af S Aust (2002), W621 17 12a. 


SMALL SCALE SPATIAL DISTRIBUTION PATTERNS AND MONITORING 


STRATEGIES FOR THE INTRODUCED MARINE WORM, SABELLA SPALLANZANIT 


(POLYCHAETA: SABELLIDAE) 


by Craig A. Spyan'! & JOANNA STRZCLECRIA 


Summary 


Srias. CAR Sieebeek J, (2002) Sill seale spatial distribution: patterns and monitoring strtegies foe (he 
introduced marine worm, Sebella spedlancanl (Polychaetu Sabellidaes Craik. Soe. 3. Aue, 1260). 1-124. 
“) November 200) 

Spatial disuibution patlerns were determined forthe introduced marine worth, Satelia ypetloncaais (GM, 
i Lares Bay, Soul) Austratia, as part ola study (o determing the most efficient methods ty survey these warns 
living on saflscdimenr habiars. Worms were putehy und. deross wraige of spatial scales, more likely to be found 
together thin i they were randomly distributed, Average density varied among sites between and tearly 1) 
worms ai” Boue cifftrent sampling wills were trialled to determine the most efficient forangoing monitaring, 
We did not Hind sirong evidence af conntine bias anon divers of different experience, but af some sites all 
divers muty hive underestionted the number ol wormsowhen using 0 25o-° quadrats, Larger sampling wits (245m 
S Hand Sims (or fransects) sampled grea more quickly ona per we basis, bul br? quadrats were the most 
eHigientsanipling Holl to sorvey warns Willi site Are liner (buy bsuale sampling wiitsize had line effect 
ofthe effort needed to reliably detect an inerease in worre ntibers: die naniber of sites sanypled hada much 
wereder WVAen cea pawer once move than abeul (Oto 15 minutes were spent underwater at edelr site. A ast 
efloetive plin for detecting (@=$=0,05) moderate increases (S09) tn The ubundance OF 9 spadfezeniiy al Larries 
Gay involves diver pairs sampling randomly within sites for 2 mhinules per site. using Sin transects (6 per diver) 
or 15m Wiseety (3 per diver). at = [5 randomly located sues cach rine, A similar time tinderwatter, sanmpling 


with Tn! quadraes (3) per diver). would hove only slightly less power (91 1 


KRIY WorkDS Sibel ypotla cite (ieadoecd species. (iri pest survey spulial pater, disteibulon. 


introduction 


Doring mid 1900s. the European [eather duster 
worm, Sabella spatlanzenit, wis detected in a 
number of marine ateas across southern Australia 
Presumably, ils presence is The resull Of a recent 


introduetion of the wornt inte Australian waters., 


perhups madvertently tnportad in the ballast water 
of trans-oeeanic container ships (Andrew & Ward 
1947: Putli & Gamnbi 2007), The hiree. up te 50 crn 
lone, OS, spallanzami® tends to form, dense 
ugerevations anc. at least on sinall scales, van have 
senous. Impuets om subtidal sessile flora und Fauna 
(Holloway & Keough 2002), Few knowin predators 
idl Vite History teats ineluding fast growth, 
extended spawning tine and fit fecundity (Curie 
ef ei, 2000; Giingrande ef af, 2000) make plausible 
the possibility of large inoreases in local population 
sizes whde rapid weagraphic spread of this murine 
pest 

Th South Australi, 8. spollancaii have been 
ound al West Lakes and the North Haven/Largs Bay 


Depa linent ot biiwaronmental Gialagy, Griverdty ol Adelie 
Seth \astrasi SOUS, Avistate 


‘Preset dduosss Conte for Researenh of) Loolegiatl Tipaits oe 


Coastil Cittus. Ciiiversity of Sydney, SSW 2006, Aus ala 
Present vddiess, Present iiddiess, CSIRO: Muatine Rewueh. 
CVO Bos 20. Nort Beael WA O20 Australia 


areas OFF metropolitul Adelaide siiee at least 1995, 
and the worm now seems ta be firmly established al 
these sites (C. Styan pers. obs. They may have 
been in the Port River for several years hanger (N. 
Halimes, Kinhill Engineers, pers, comm). In South 
Australia and efsewhere, the worms are known to 
colonise a range of habitats, from reels and man- 
made structures such as piers and marinis. lo soit 
sediment habituts, meluding seagrass beds. Given 
{he potential threat (hese worms pose, lupe scale 
Monilorigg programmes are necded, together with 
small scale manipulative experiments aeross ay range 
OF habituts. tu understand properly the ecological 
effects of the warm as they spread, Before any 
Monitoring programme can proceed, however, 
methods deed to be developed to estinmte accurutely 
(he abundinee §. spetlemmand? in the field, We fortis 
here on determining efficient methods lor 
Monitoriag worms on soft sediment habitats, These 
hubitals ure the Most common in South Austrian 
wulfs and so. heeause of their extent, could 
potentially harbour Jarge numbers of worms, Other 
work will he needed Lo determine the most elficient 
methods for surveying worms in other habitats such 
as roeky reels or marinas, 

Determining the abtndanee ol organisms 
underwater is Hob alwiys a trival tsk. even for 
relatively large. sessile Organisms (e.g. Inglis & 
Lincoln Smith 1995; Benedeti-Cecchi es el, 19961, 


Ls COA SPYAN & 1h SPRZELECKI 


Workers lave i Wide range of survey techniques and 
sampling lo choose frome bul, unfortumilely, often 
avbitrary choice about which methods are used ean 
commenty laud to wasted effort and resources 
(Dawning & Downing 1992), Obviodsly, planning ts 
necessary Tor any under witer survey lo crasure hit, 
whilst effort and costs are minimised, surveys retain 
silficienruceuraey und stanstical preeision to allow 
confident interpretation of data (Andrew & 
Mupstone 1987, Underyood 1997). Such planning 
requires information about the relative cost anil 
elforr required per sample (and perhaps an 
evilliation of different sampling units), and 
THI eTaLION abOUL the appropriate urbe of samples 
thal need lo be taker to wenerate desired levels «] 
stalisticul precision and power. ‘The tumber ot 
samples required. tn turn, depends on the way 
visanising are disttibuted spatially, 

Information necessary for survey planning can 
sometimes be obtained fram previous work, but oo 
Tifermalion was available about the distribution ofS. 
speadlancanit oy sott sediment habitats, typical of the 
sulls in South Australia, Se. tn 1996 we conducted 
pilot surveys Of S. spollanzaeis in these habitats at a 
nuinber OF sites within Largs Bay, Gull St Vincent. 
Our study had two specific aims. First, we wanted to 
uelermine how worms were distributed, on at rane 
of spatial seales, and how (his) varied with 
abundanee. Second. we wanted to determine the time: 
taken to coUnL worms, the relanve magnitude of 
yaruuice Components between replicate samyples at 
Jilferent spatial scules, and the differences tH these 
found using a range of standard underwater sampling 
unis (O.25m7 and LA quadrats, Si Lin transects 
aod 25% bi transects). 

Counting even arse organisms underwater is nit 
always done equally well by divers with differen! 
enperionee (Inglis & Lincoln Smith 1995), "Thus. there 
isa potential to bias results and/or (urease varkinee 
estimates GF multiple divers ire used un surveys. 
making: Changes in-abundanee even more difhicult to 
detect, We used i range of divers vondueting: these 
surveys, so Wwe needed to determine whether havi: 
different divers involved surveying afleeled counts of 
worms. So betare including dati From eweh diver in 
ihe planing analyses, we ulso tested whether there 
Wwasainy evidence of bids in the total number ol warns 
experienced hd mexperieneed divers eounted ino 
SUPYCY. USTED RINCOMU ZATION Lests, 

The overall ain of our work then, was lo deter 
the most efficient techmaue lo use for resutur 
monitoring of the ubundancve of S. apiece on 
suff sediment habinals. 


Srysitled V2 (OME) Phe ropriduelive coalouy af phe xentlape Chdmyy 
Hiffers (Seu AUstaT PROUT hosts. Bepirunent ar Brharanmentit 
Hioloey, Liniverite oF Adelaide. South Austolia OF fapulil. 


Methods 


Lovano ap surveys 

We sumpled at 6 sites within Lins Bay between 
February aid Noverber 1996, Lares Bay bas 
chosen as our main Field location for this work 
because we knew fronr previous work! what 
reasonable densities of S$. spal/anzanii were present. 
We also conducted some pretiniary searelynig far 8. 
spatlancani’ at other loculions; specifically, the areas 
surrounding jetties at Edithburgh and) Ardrossan 
(Gull St Vincent) and Wallaroo (Spencer Gull. 
Those iveas Were searehed because we considered 
thal they were fikely to be anchorage porns tos 
bowing und shipping from Port Adelaide. and this 
possible destinauions of worms sproud through ull 
fouling, Although a few individuals of Une similar 
ooking mative feather duster worm, Sebelleayierte sp. 
were ound at all lacations, no S$. spelfansanit was 
found in 2x 40 minute dives at any of these other 
ocutions. 

Largs Bay is a-shiullaw bay (4 ta 8m deep), close ta 
the Port River and seven mitrruds, areas Kaew) lo 
also Suppor High numbers af 8 speallancanii on 
pylons anid mooriies (CLA. Stiym pers. obs.) fhe 
SUbstreta in Largs Bay is predominantly salt 
ScUimenl, Consisting of a qlusaie of seaurass 
(Poxidonia spp) meadows, sand palehes and palehes 
of deeraded seagrass, eharacterised by silty 
sediments: Common. lurve invertebrates i the ured 
ie lide razor fish (Pinte Dicalar), scallops (Cf lain's 
bifrons|. shartish (Unieptora sranifera), whretks 
(Memoploacd australis) and sea tulips | Pyare 
sputifernt)’. General underwater Visibility during the 
study wis enod und ranued between J and lane. 


Sreryvey: peed teely 

Six sites within bers Bay were haphazarely 
scleeted benveen SAUm and |S00m offshore ALcael 
Sie. LWO Scls OF lupe measures were sel oul by divers 
to det as survey base lines. The directivum of these 
hase lines was hipbovardly chosen (there was line 
current How), with divers kuyahg out te Hie as hes 
swim in apposite cirections fron the anchor One 
pair of divers worked along one each base line owah 
auch diver independently placing randomly selcuted 
qaadrat or Gainseet starting points ab pre-determince 
places along the He. Square yuadrats ind transects 
had one edge Tying along, and tafseels (an 
perpendicular to, the base line, Dyers used a In 
slick Lo defing [he transeer width and counter! warns 
us they rolled outa Lape easure for fie sed lisiamee, 
Fach diver was based on opposite sides af the byse 
ihe lo nimise viterforenee between divers 

Ow the first day of sampling (14/2/96), Wwe set pul 
oily (Wo SOm base fines at site |, Pour divers eueh 
nensured fen. «0.5m quadrats, then ten din Tm 


DISTRIBIIEION AND MONTTORING OF S. SPALLANA ANY 110 


quadrats, then three (o five Sin x 1m transects andl 
then one to three 25m x Tm transeets, The time taken 
lo measure cach OF these sets of units (Separately) 
wats recorded for each diver, On subsequent days (ut 
sites 2-5), lwo 200m base lines were Jaid oul, and 
quadrats and transects were conducted as divers 
moved along the base dine (ie. sampling swith 
different units was both spatially ani temporally 
interspersed), On these days cach diver counted ten 
0.250 quadrats, ten Fine quadrats. five Sm transects 
und ane 25m transect. The exception to this was on 
the Hina! day of samplins (26/11/96; site 6) when 
only one 200m base ling was lie gnd a single pair 
obutvers conducted surveys, 


testing for diver bias. 

M cuch site olf survey Leams consisted of 3 
expenenced serentific divers (each wath hundreds ol 
hours af underwater work logged) and one 


mexpeneneed scientifie diver (<!5  hoars 
wnidkerwaler Work), Before pooling divers’ data, we 
tested Whether divers met count worns 


(ilereotly, ludividual divers changed between 
days/sites. but the mix (1 inexperienced: 3 
experienced) did not change. Pairs oF divers worked 
on seplirate base lines within sites, principally: to 
prevent divers gelling in each other's airy, bul alse 
lo increase the spatial spread of the sampling within 
ust. As a result. for eaeh site there was the 
potential to mike 2 sarts of dive experience 
colipansons bebween divers who were sampling the 
suine area; belween an experienced und an 
inexperieneed diver on one base line. and between 2 
experienced divers working on the other line (each 
for (25m, 1Oim* quadrats und Sm transects). 
However, we could hot make hese comparisons for 
wrmumber of base lines where worms were absent or 
found i only a few samples. 

We used u series of randomisation rests to compare 
the difference jn total nuinber of worms counted by 
the iyo divers ona base line wath the distribution of 
dilferenees between divers ifthe counts tact insteuel 
been allocared randomly. This. disteibudea was 
constructed with a simple mucro in Bxeel. using the 
IWhwNt random number generaror to allowdte the 
counts tO divers in L000 simulations We tested the 
fiypothesis that if there were a counting bias, one 
diver would counf more worms in folal than the 
olherdiver on the samme base line. wd dilterences: 2s 
lurge us this Cor larger) would be jatregueat when 
county were randomly allocated, We elione to run the 
rindomisution fests rather than Ltesis because the 
uta were very skewed in mest cases. The 
nuldoutisation tests allowed us to make the 
comparison aboul sumpling worms without first 
tramstorming the data or modilying, the hypothesis 
gecordinely 


Spetial dixiributions 

To describe the Spatial distribution of S. 
Villani ala vinge of Scales, and how it changed 
With population density we calculated Marjsita’s 
Indices at cach site, for each of the different sized 
sampling. voils. Morisita’s Index (i) describes hos 
much more likely ras that bwo individuals drawn at 
rindom will have come from the same sampling unit 
than if the population had been randomly dispersed, 
For example, an indes of 1S means that individuals 
are SO% more likely to have come trom the same 
quadrat or transect than if the popalation had been 
randomly dispersed (Hurlburt 1990), 


The formats for bay ts: 


_ (Nj) (Ay (e 
n= (5 Oil He) 
(Morisira 1971) 


Where X is the number of sauiples, (1 is the sample 
neat and ay the swmple yarnince. Because ky is 
essentially a variance jo mean ratio, the null 
hypothesis that ly = 1.0 can be tested, bused on the 
4 disivibtition with (nel) degrees af freedom 
(Mulburt 1990), We calculated Ty for cach of the 
sumple unit sizes al each site. pooling dala fron 
divers und baselines within cach. 


Power caleulanoly & cospelictency 

Power culcubitions were done in Excel (using the 
PiFace itdd-in lo calculate non-eentral T- 
distributions: R. Lenth. University of lowal tora 
simple sampling design where equal Gite was spent 
by iwo divers Working Underwater, before ov afler a 
lime interval over which worm numbers muzht have 
changed. Time (before and otier) was treated as a 
fixed fuctor in Gur analyses. Using measured 
estimates ol the average Lune taken per sample unit 
to dletermMiie the umber of samples that gould be 
taken for a given time underwater, and expected 
Variance assoomted with each saniple unt, we 
eulcotamed either the size of at jaureake in incan 
abundiuniee that pairs of divers could deter. or the 
time taken for pairs to detect a given sized chanpe, 
both with @=P=0.05 (Lindernvend 1997), ‘These 
qulculahons were dene with respect lo moniloriny 
designed i defect changes am 2 spatial scales: 1) 
within uw single. fixed site and 2) aeross rhe entre 
Lares Bay area. We set “=—f=0.05 beeause we were 
equally convened about type foamd UW errors. anil 
wanted a low probability of incurring either 
(Mapstune |49S). 

Within # single fixed site. statistical power to 
detect a chanee tbronah time will be determined by 
(he variinee among sample wots wilhio thal site Gand 
the number of samples taken). Using the variances 


{20 CVACSTYAS & | STRZELECKI 


ussoenited wilh each site, we did the power 
vilenlations for each site and sample unit 
combination separuely, ‘Pimes uceded ala site per 
period to detect a SO% inerease in worn abundance 
(willl @=B=0,05) Were loe-transtormed before 
averaging and back-lrainstormation, Similarky. we 
did the power calculations for cach sample unit using. 
ive other estimates Of the variance within sites, each 
averaged. across sites (but both calculated slightly 
differently), The first mean variance estimate was 
venerated from an hierarchical ANOVA across siles 
2-5 {see below), the second calculated alter 
averaging the coefficients of variation from all 6 
silos, 

Ona larger (hay) scale, statistical power lo detect a 
change whroush ome will be determined by the 
variance in the mean abundance of worms among 
sites, and the number of sites sampled in each time 
period: ie, the lest denominator MS will be a 
combination of variance within sites (among 
sumpling units) und varianee among sites in’ the 
mead density of worms, The magnitude of these 
vilriunee Components can be determined trom the 
inca square estimates in an hierarchical ANOVA 
(Underwood 1997: see also Table 0), Our sites were 
in effect randomly chosen within Largs Bay and, 
despite data not really meeting assumptions (they 
were very skewed), we ran ANOVAS on the rasw ciatie 
lo determine the magnitude of the among and within 
site Mean squares estimates, and henee the size of the 
individual variance Components. We did. the ANOVA. 
only on the 4 sites where we had balanced data and 
equal numbers of samples a each site (sites 2-5). 
Using these varianee component estimates, we were 
able to generate the expevted denominator MS tor 
the test of un inerease between times across Largs 
Bay, under a ringe of scenarios (varying the number 
of sites sampled and time spent sampling per site), 
and for cach of the different size-sanipling units, 


Results 


AL Largs Bay §. spellaizaii’ Were patehy ona 
number of spatial scales, We found quite lange 
differences among ‘sites in the abundance of worns 
(Fiz. 1). Worms were quite common at some sites. 


for example average density was nearly 10 worms m7 
al site 2, bul worms were virtually absent at others 
(c.g. site 3), The highest densities measured in single 
sainple units were 52 und 49 worms m) atsite 2, 10 
4 Sim transeet and a Lint quadrat, respectively. Within 
euch site, and across cach sample tuiit size. the 
distribution of worms within sampling units was 
very skewed, with miiny units not containing any 
worms at all (Pig, 2). 

We found tittle evidence of bias in the numbers of 
worms (hal different divers counted within busclines, 
bor G.25r! quadruts. none of the four randomisation 
ests between experienced divers, nor the test 
between a inexperienced and anexperienced diver 
was significuml al the =0.05 level, Similarly, none 
of the four randomisation tests between experienced 
divers. Gr the one test between an inexperienced and 
in experienced divers was significant for the bm 
quadrats, None of (he four tests between experienced 
divers was significant for the 3m transects, bubone of 
ive Tests between mexperieneed/experieneed divers 
was significant (v=0.035) for the Sm transeets: 
However, We might expect nearly one significant test 
(al 420,05), even if the null hypothesis was true, 
given the overall number (16) of these Lesis we 
vondueted, 


i4 
12 
mw 10 
= 
bo 8 
a 
E 5 
= 4 
2 
Qa 
- is] ao st uy wo 
my w 2 y B 2 
a n v} w a a 


Hig, 1. Mean density (25-E2) ol So spedlencaiy at anehorine 
sles within Largs Ray. AL euch site. density estinumes 
using different sarnpliny units ure showy sepuritely, 
pooling estes [ran divers: 251 Crapsects (rey bars): 
Sit Wunseets (white bars): To? quadrirs (bhick bars): 
(25° queulals (diagonal striped bars), 


TRL) LO Eytiindtes of variance components within and dione aictioriig ses (2-5) af bares Bay followin ANOVA, using 
d f u } MY 


Silifferent sized sanplin wily, 


Sampling unit Sore dt. 
25H tnenisect Among Sites 3 
(4 =70.0) Within Sites 2 
Sm trseer Ariane Sites 3 
(W=l64) Within Sites Th 
re qqutiel rat Ammon Sites : 
(yy =2.Ah) Within Sites 156 


39929 = extn + FOP ne 
31 7Y.3= ef vain 

10302 = cyt + Wai 
936.6 = Cyan 

656-7) = vit Menu: 
30,04 = vite 


Mean Square 


(mone = 91874) 
(Lone = 4683) 


(CPi =15.65. ) 


DISTRIBUTION AND MONITORING OF 8. SPALLANZANT, It) 


TON 
* 
in], “ 
“ ® 7 
Inq . . Py 
y . e + . ‘ . 
1 + r —— 
2 4 5 & 19 12 
Density (worms per m*) 
a4 


Vig. 2. Spatal distribution uf AO spall ata range of 
spatial] scules WIKIA aehering siesund how this yarics 
WOl papntition density, 257 transeets ieirelesi. Sin 
Iransects Cirhimglesi Tin quitdrits (squires); 0.25 im 
yimulrais (diamonds). Morisita’s (M97 DL) index hy) values 
siomilicanily diflecent Crom 1.0 (at y= 0.05) are shown ats 
Hlled syinhala: jhe one non sienimicunt yulue is shown 
unlilleel. 


Im® quadr ats. 


200)- 7a 160 
0,.24m° quadrats 


80 


7 ) 
6 8 10 0 


10 20 30 40 50 


10 ~ — 

5m transects 25m transects 

30 5 
0 Sg ee So 0 -— & = 

ao 
CS RBH GaF cp sgedeg 
- aw MN Kran m 
Worms per sample unit 

Hig, 3. Drequeney Wistoerins of the number ol § 


spallanecaiit per SMNpPHng UNL pooling derass dehorins 
HCH. 


We did not compare estimates of density among 
sites or sampling units because of very unequal 
numbers of samples, very heterogeneous variances 
among sample wnt sizes (and sites). and skewea 
distributions. with large numbers of zeros inthe dita 
(Fig. 2). However. we did note that at fwe sites (2 
uml 6) where worms were relitively abundant, 
estimates of average density made with the 0.25m= 
quad seemed to be less (han those made with the 
ther sampling units (Pig. |), perhaps indicating bias 
(across all divers) For this reason, we did) oor 
consider 1.25 me quadrats inthe power caleulugons. 


Variance component estimates 
ANOVA ate shown in Table 1 

Figure 3 also illustrates the patchiness of worms. 
but ata hanee of (termediate (within site) scales, As 
worms became more abundant, there appeared to be 
some decrease ih patchiness across all ol the Gwithin 
site) seales. bur worms nanetheless were still 
significantly more likely to be found together than if 
they had been ruidomly distributed. ‘Worms were 
quite patchy on larger (within sie) seales. woth fy 
values significantly greater than LO ut the Sim and 
25m transect sewes (Fig. 3). Additional patelipess at 
smaller scales was rellected in the even higher It 
values for (be O.25m" and 1.0m quadrats, whieh 
incorporated varialon ut larger scales and varintion 
al small seales. Only one Ta index (for 0.25 oF 
quadrats, al very low density) was detected ay net 
being significantly greater thin 1.0L At small scales. 
S. spallaicaniy were clearly arranged in distinet 
clumps. with 10s of wotnis sometimes attached to 
the same small piece of (rare) hard substratum such 
iis a Prane shell, Quite offen, however, we use found 
clumps of worms that did not appear to be atiaehed 
lo any hard substran. and whieh appeared to be 
firmly routed in the soli sediments. 

Pig, 4 shows the average lime tiken lo sample cach 
unit at the first site, tor the runge of sampling units 
we tested. As expected, 25:0 transects took much 
longer (nein imesA63 sec) than Sm transects (173.4 
sec), that took longer than Inv (37.4 see) and O.Sny 
quadrats (34 see), Clearly. larger sampling units look 
less Jong to coun! on a per me? basis than snialler 
sampling onits. Error bars on Mig. ob illustrate the 
range of per unil times we found among the divers 
that conduieted our surveys. Foreuch sample unit, he 
slowest diver wis the inexperienced one, but the 
fislest varied among the experigneed divers (age was 
faster at quadrats, another ait transeets). Fiz. 4 also 
iustrates the decrease in the average coelicient of 
Varttion WITH increasing sample unit area. 

Within a site. power calewatons biased un the 
average coetlicient of variition aeross sites found 
that for at given time sampling underwater. line 
quiidrats provided a more powerful ae at 
technique than the other sampling units (Fig. 5), The 
linding that }m? qiadrats wele more ‘ionta (han 
Sim or 25m transects was also found when we used 
the (overall) wilhin sites varianee estimates (i.e. from 
ihe ANOVA of sites 2-5: see Table | und below), 
When the power calculitions were done sepurately 
for Gueh site (Le. across the range of population 
densities illustrated in Fig. 2), using the speeitie 
variance estimate found for each site. we generally 
found the same resull, Whilst the level ane the 
Vilference if power between units varied with site, in 
four out of five cuses, the Tim quadrats were the 
most powerful technique. [In the fifth. Sm trinseets 


following the 


122 C.ALSTYAN & J. STRZELECKI 


c 
4 3 
=) 
a) 
= 
o 
ote 
‘Ss 
yeaa 
& 
1 2 
is) 
Lt i 
= a 
— o 6 
oa 
= 
3 
- 
a 
a 
a 
= 
= 
c 
0 
o 
= 
ro @ £ rs) 
5 3 % g 
iss} a re 
3 3 c = 
io > g = 
tw ma = 
E E = E 
re - tte) Ki 
ra] 


bie. 4 Average of mean time taken to count 8. spallansanii 
per unit (averaged across n=4 divers). contrasted with the 
average coelfielent of variation within anchoring sites 
(across 6 sites). for the 4 different sized sampling units. 
Vertical lines indicate the range of values (maximum to 
minimunt) for both measures. 


14 i me quadrat 
os! —— 
ca | a 
g 06 wl _Smiransect 
5 “ _ = 7 28 mn transent 
a va — 
0.45 yy 
wt i - 
021 “~- 
g ——_-—_ — ——_ 


0) 20 40 60 80 100 120 
Minutes spent sampling 


Fig, 5, Statistical power of difterent sampling unils lo 
measure a 30% increase in the mean abundance of 5, 
speflansanii within an anchoring site between 2 periods, 
given the effort (time) 2 divers spend sampling 
underwater in each period. The calculation of cach power 
curve takes into account the number of samples that cart 
be sampled by 2 divers in each period using a particular 
sample unit. and the mean coefficient of variation using 
that unit, (ce=0.05). 


were more powerful, but Im? quadrats were very 
nearly as powerful for any given effort and the C.V. 
for 5m transects was based on only four samples. 
Across all the locations we sampled. we estimated 
that a pair of divers using Im? quadrats. would, on 
average, need to sample a site for 128 min (range=50 


1 ing eet 
1. (0-8 as” 
fn 
B06 ae” 
a ne 
O04 n?® 
n?® 
s 
0.2 e? 
e. 
0 


2 4 6 8 10 12 14 16 18 
Number of sites sampled 


Fig. 6. Effect of sample unit on the statistical power Lo 
detect a 50% increase in the mean abundance of 4. 
spallonzenii within Largs Bay. between 2 periods, with 
the number of randomly located sites. visited per period. 
A pairof divers spend 20 min sampling in each site using 
either Im? quadrats (circles), Sm transects (squares) or 
25n1 transects (triangles), (4=0,03). 


700% increase 


1 —— 

08 - 
io / 50% increase 
GS 08] / , - 
(-) ~~ 26% increase 
a 04, 

0.2) 

——— —_ 

2 4 6 8 10 12 14 16 18 20 
Number of sites sampled 

Fig, 7. Power to detect varying increases in the mean 


abundance of §. spellanzavi within Largs Bay. between 
2 periods, with the number of randomly located sites 
visited per period. A pair of divers spend 20) min 
sampling in each site usimy Sm transects. (c=0,05), 


- 266 min), at both times, to detect a 50% increase in 
abundance through time (c=fi=0.05) at that site, 
Equally powerful surveys using 5m or 25m transeets 
would, on uwverage, necessitate nearly twice as much 
lime underwater (averages=240 or 248 min 
respectively). 

Atthe scale of across Largs Bay, sampling unil size 
had litte effect on the number of sttes required to 
reliably detect an increase in worm abundance (Fig. 
6). Moreover, the time spent in each site had very 
little effect on power, once 1)-15 minutes had been 
spent in each site (i.e. once the number of sample 
units for each diver was >two). We did not illustrate 
the effect of time per site here, because the results for 
even short dives (e.g. 20 minutes sampling) were 
trivial. Our calculations found that the best way to 
detect a moderate (50%) increase in the abundance 


DISTRIBUTION AND MONITORING GES. SHALLANAANT IG 


aLS. spallansanit across Largs Bay would be to hive 
iver pairs sampling tandomily within sites for a 
ininimal dine grewer than 15 snliutes (say, what 20 
Min per sited, using Sm transeels (6 perdivers or 23 
Iranseets (4 per diver) at = 15 nindomly located sites 
each time. A similar sampling scheme Using inc 
quadrts (AL per diver) would have only slightly less 
power (0,91), The power to detect increases. given 
Ihe number of sites cach sampled by a pair al divers 
for 20 minutes per period is Wustrated jn Fig. 7. 
Inereuses OF 25% or Jess are difficult (a detect 
WiIHhOnL Very Tange numbers al sites bem sampled: 
More moderite increases (>50%) should be much 
sasivr 1 deter. 


Discussion 


Svhella spatlancant diving on sol sediment 
habitus ut Largs Bay were patehy on a range of 
spatial Scales. Worms were found i small, (ht 
clumps, and ther there were patches of these al lureer 
seiles walltin sites: and laree variation im overall 
ubundanee between sites separated by LOOS ta 1000s 
of memes, Whatever the underlying biological or 
physical causes af the distribution of $. spallancenir 
on soft sediment Habitats, ol work has: Tlusteated 
With depending on the spatial scale of monitonug. 
dhis spatial patuliness cam ifluence the decision 
ubuulo which sort of siimpling doitstoutd be used to 
mMeastre chinwes i) worn ubunadiinee, 

We found thal Tine quadrits were clearly the most 
efficient sumpling voit for estimating sworn 
abundinee within t site. "This was despite auc findilys 
that 257 transcels Gand Sm transects) were much 
faster lo conduct per me of seafloor surveyed, 
Essentinlly. the trade-off here betsyeen sampling: 
fewer places wlong ad base line with Rirger sampling 
Lis Chit sampling each of these places more 
preersely ) ute sainpline more places (bul vach less 
preemely) with a smaller unit. favoured the later. 
Whilst [ranseets vaver iene area on a small scale 
relutvely quickly, the extra effort expended counting 
Hone YW fainseel WOuld he bette? Gxpended sampling 
more, randomly determied, positions within wt site 
(10s to 180s OF metres away). with a smaller 
Awnpin wnt “Thus, we conglude that if precise 
estiniites of worm ahundanee are needed an sinuller. 
sile scales. then Tine quidhaits Should he used ty a 
saimpling. wiih TP estimates oF worn ubundiinve are 
needed ao this seule then the use ef Line quadrats aan 
Jead Lo subsuinthil savings in effort and/ge incrases 
IN precision, relative to monitors willl Sa or 25m 
transects. 

Hy conlmist we found (hil when monitoring tor 
chimes al hirger scales, slitistical power will depend 
essentially only oo fiw touny sites are sampled. bor 
ronrloring across Lares Bay the elive ol Sampling 


Unit between quadrats or transeets will be 
unimportant beeause. fora given sampling effort, the 
differences in’ power henveen different sived 
sumpling units were less than the Wwerease (or 
decrease) ih power af yae more (or less) sileavere 
sampled, We also faurid that, provided 4 reasonable 
arnounl of time was spent at cach site lat least 1S 
Minutes). the Uime spenr per site had only @ small 
effect on the overall power of a monitorimne 
progeanme, Thus, the power oof any inonitoring 
programme for these worns cun effectively only 
inerease through sumpling more sires. Consequently, 
we recommend that time spent per site is minimised 
fo about 20 miintites per site (=15 minutes, plus a few 
ext la ensure at leust several counts ave liken 
wilhin a sitet and that effort is put ito sanypling 
more sfles (at Teast 15) rather than sampline sites 
more intensively. 

We did not fing! strong evidence of differences i 
the number of warns counted anion: divers. but 
divers: experience did intluence the time wm iook then 
jo Coun worms. Having more expertenced divers in 
{ strvey feanr jnght speed up monitoring, bu 
Include Mexperienced divers in surveys as well 
Should net seriously bras abundance esuimates or 
make surveys less pawerhil We did uncover some 
evidenee that. at least on soft sediment habitats: wad 
for moderate Worn densities, wll divers may 
ungderestimule warm ahundartee with U.25in' 
quadrats; As aoresull. we would recommend against 
the use of 0.25mi? quadrats as a sampling unit ti 
fulure INONHOrIN prograiines, at lewst on sor 
sediment habitats, OF course, recommendauons 
about nptimal sampling wits and staresies depenad 
on the habimit being surveyed, ane phe spatial 
HSHTBUTON OF Warts ala time ol scales within the 
scale Of interest. So, on hard substrata avhere worms 
are offen found at much bigher densities und perhyps 
not ws palehy, we predict dit smutler (0 25m) 
quadrats jay be more effective, though (hys ail} 
need Hirther testing, 

Our Vinding (hal using o partioular sampling: wait te 
sHMple worms can, io some sitions, lead to moe 
more powerful surveys fora given effort is net 
purticulurly novel, Iidewd- an expectation thot we 
might find ches was the husis for dei this work in 
the Hirst place (ean Andrew & Mapstone [OR7: 
Underwood |997), Nor ys our tinting that broader 
sede osiveys here are more influenced by the 
AUMbSY OF sites sampled, rather than the preeision of 
simpliog within each sile, very surprising: espeeially 
viven the large jntersite variation in sworn 
abundance. We note that these speettic results are, 
hawever, citirely depentent on the spatial 
Uistrbutions OF SY. apedfeuicend at Lares Bay. jue the 
trode OFF between precision dnd effort required: for 
varius sampling Units. Por other species or even 


|24 C,A, STYAN & J. STRZELECKI 


S$. spallanzanii in other habitats such as on piers or 
marinas. the spatial distribution patterns and trade- 
offs may be different and so, consequentially, might 
the recommendations for surveys (Andrew & 
Mapstone 1987; Underwood 1997). The only real 
way to find out how to best survey for other 
circumstances is to conduct a preliminary study 
(similar to this one) for those circumstances; this is 
an often repeated, but apparently seldom heeded, call 
(Underwood 1997). 


Acknowledgments 


We wish to thank Andrew Melville. Emma Cronin, 
Rebecea Fisher and Jan Magraith for help with field 
sampling. Alan Buller, David Williams and Piers 
Brissenden helped sith the logistics of this work. We 
gratefully acknowledge the support of the South 
Australian Department of Environment and Natural 
Resources, through the Marine Environment 
Protection Fund programme. 


References 


ANDRIW, J. & Warp, R. D. (1997) Allozyme variation in 


the marine fanworm Sabella spallanzanit: comparison of 


native European and introduced Australian populations. 
Mar: Ecol. Prog, Ser 152, 131-143, 

ANDREW, N, L. & Mapstone. B. D. (1987) Sampling und 
the description of spatial pattern in marine ecology. 
Oceatiogr. Mar Biol. Ann. Rev. 25, 39-90. 

BenrorrrreCrcen!, L., Arorpt, L,. ABBIALL, M. & CINELLI, 
F. (1996) Estimating abundance of benthic invertebrates: 
a comparison of procedures and variability betwveen 
observers. Max Ecol. Prog. Ser 138. 93-101. 

Currin, D. R., MCARTHUR, M. A. & Conen, B. F (2000) 
Reproduction and distribution of the invasive European 
fanworm Subella spallanzanit (Polychaeta: Sabellidac) 
in Port Phillip Bay, Victoria. Australia, Mar Biol. 136, 
645-056. 

Downing, J. A, & Downinc, Wo L. (1992) Spatial 
ugeregation, precision. and power jn surveys of 
freshwater mussel populations. Can. J. Fivh. Aquat. Set. 
49, 985-991, 

GIANGRANDE, A. LicclAno, M, & PAGLIARA, P. (2000) 
Gametogenesis and larval development in Subella 
spattanzanii (Polychaetas Sabelhdae) from the 
Meditermmnean Sea, Mar Biol, 136. 847-861. 


Hottoway, M. G. & KrouGH M. J. (2002) Effects of an 
intraduced polychaete, Sabella spallansanii on the 
development of epifaunal assemblages, Man Eval. Prog, 
Ser 230, |A7-154. 

HurLpurr, S. H. (1990) Spatial distribution of the montane 
unicorn, Gikes 58, 257-271. 

Incus. G. J, & Linco.n Smith, M, P. (1995) An 
exuminalion of observer bias as a souree of erroc in 
surveys of seagrass shoots. Aust. J. Ecol, 20, 273-281. 

Marstons, B. D. (1995) Scalable decision rules for 
environmental-impact studies - effect size, type-l, and 
typed errors. Ecol, Appl. 5, 401-410, 

Morisrra, M. (1971) Composition of the Lindex. Res. Pop. 
Ecol, 13, 1-27. 

Parr. FP. & Game, MC. (2001) Phylogeography of the 
invasive polychaete Sabelly spallancanit (Sabellidac) 
based on the nucleotide sequence of internal transeribed 
spacer 2 (ITS2) of nuclear DNA. Mar Ecol, Prog. Ser. 
215, 169-177, 

Unperwoon. A. J. (1987) “Experiments in ecology. Their 
logical design and interpretation using analysis of 
variance” (Cambridge University Press, Cambridge, 
UR). 


FIRST RECORDS OF TWO FAMILIES OF FRESHWATER 
AMPHIPODA (COROPHIIDAE, PERTHITDAE) 
FROM SOUTH AUSTRALIA 


BRIEF COMMUNICATION 


Summary 


Since 1994, extensive sampling of streams in South Australia (SA) has occurred as 
part of the Monitoring River Health (MRH) Program, the AusRivAS project and local 
monitoring for the Onkaparinga Catchment Water Management Board and other 
agencies. This work has revealed many macroinvertebrate taxa not previously 
reported from SA, including specimens of the amphipod families Corophiidae and 
Perthiidae. Freshwater corophiids previously have been recorded only from the 
Brisbane River in Queensland’”, and perthiids have been recorded only from Western 
Australia’”. 


Cramvuetions of te Reval Saciety af South Austria (2002), 426(2), 125-120, 


BRIE COMMUNICATION 


FIRST RECORDS OF TWO FAMILIES OF FRESHWATER AMPHIPODA 
(COROPHIIDAK, PERTHIIDAE) FROM SOUTH AUSTRALIA 


Since 199d, extensive sulvipling of strewis in South 
Austria (SA) has oecunred as partol the Monitoring River 
Healith (MBE Program, the AuskivAS preyeer ant Ieeal 
Monloong fob the Olhaproius Catehried) Water Mange: 
mont Bard and other iveneies. This work bas revealeu 
MY TEENIE rlebrile Taxa not previously reported fren 
SA, including speeimens of the amphipod fimilies 
Coropliiicie and Perthiidie  Prestisvler  corophrits 
previnusly have heen recorded analy tram the Brisbine 
River in Quevnslinet! aod perthiids live been recorded 
Onhy leony Westend Australia’ | 

The Coraphiidae is a speciose Guntly ound in marine and 
Frestiwatter Habitars round the werk. iy Austeatia, the anly 
KOOWA Proshiwuler species i Pervecmphiun ever 
found whoo New 4cuhind! Speeinens recently found in 
SA appear ti be Mo eyiivenen. bur as the orginal 
desenplion of the species in Agstrdia is dubious (1A. 
Bradbury. Univ, Adelaide, pers. conn, they ale identitied 
here as Corophiidae SAsp) ‘Phe bocly is stighely Hatened 
dorsoventrilly, dine the rosome ts markedlyso. hens eno 
acecasory Hagelin on the antenniles. The merus ol tie 
second enithapod is dlongate and all pereapods are heavily 
selose: Phe third droped is small witha Onter ranuts Hat 
is bwice us longa he inner ramus, and is partly hidden by: 
wrounded. ene (at clei. Testy felsen. 

‘Sites Where Coraphitdae SAspl oeeurs in SA ure 
widespread fur disginet. They dachide the ‘Tae River on 
Hyre Peninsula (44° 45" BL 85° 53" 8), the Bromer Rives 
ticun Hartley (35 10 BL 1B9° OLS). Gorge Ck 134° 56’ 
b, 139° 00" Sy and Reedy Ck U34? 5), 139° 137 8), hotly 
obo which qre tributaries of Ghe Murray River and Lake 
Bontiey (37° 39° PodOF 19 Sy and the Lake lrome outlet 
drain (37° 3" E14? 079 Sy in the south east Muecues in 
Still and Mowing witter habits with conductivities of 2890- 
T8700 pSicnt. Th is aburlunt ae all ese sites, and atten 
cdliuhiis wal diitehieniaaisiedis (Cermidae) the mast 
common [reshwailer arnphiperl in SA, 

Nhe distabiinon at Corophadie SAsp] in SA sugeests 
Chine HAMOHees. Two révoreks dre fron wattencdueses thal 
evuiply fave the sey Coiake Frome quller, Iud Bevery, and the 
ottivrs drain to the Marniy River or Lake Mlesindring, bath 
of which were connected te the ocean before consirucnion 
Of viver mouth Barfages in M940 Accerdiog to Chilton! 
who drest ieatied 2 avcavenin fron Queenshind 
(Brishine River. this species prefers Mimi waters yea 
the cote Alb New Zeiland records are Torn brackish 
Wallets! 

The unity Pecthiidie previously wos Known only loam 
sautir western, Western Australis (WAde. TE contains a 
single gents. Mera and Iwo species. A aeudielsen and B 
bromehiaiy, The anteniotes ane not smaticantly: longer 
Than fhe antennae. and the aveessury Thawelum af cael 
‘wlennule js 2-seemented, Thoracic segments carry: 
dendritic steraal gill, The gnathapeds are birge atid 
cuntleveres|, Pereopod 6 is longer than percopet 7 tn P 
Minehialhs the toner comes at yropadl aos one yuartor he 


length of the outer ramus: in Po eeaivelsan ibis about pvo 
thirds ts long as the outer Rims ATLOudT specimens fyi 
SA fit the deseriplons of both WA species, the distance 
ovlween fhe Wyo pexToOns sugeests (hat there min be 
tukonomic diflercnees (1 He Rewlbury. pers cout 

Most Specimens at Pevrticd spp. trom SA are Hrom.t small 
drea ol the Mount Lolly Ranwes. in the Onkaparias 
catichment ner Adehiile. but they have-alse been colleetead 
frond the Marniy River at Waods Polar The biner site has a 
ehemica Composition like other sites where Perthia spp. 
have been collected, bul itis a law whund river rather thin an 
Upland sired, SA Waren dises the Onkapatiiwa River as a 
venduit fer Vineray water and at is possible thet 
inuuslocation of species his occurred. 

From the physicochemical elavacleristies af silos in WA 
mul SA Where Peria spp. have been volleeted. it would 
uppear thi the group preters slow tTawiig or scald febiats. 
cool temperatires and fresh oweder -<1500 pm S/em) wvalty 
tow futrivnt levels (total phosphors <Q.4b mg/l. total 
Kyellabl nirroven <tonye/h and meutea) toaciic pH, hey 
senerally occur in cuchments with relatively high raintall 
and hative Vegeninon. The most fkely factors (a pestrie! (he 
Jisuibuion of Perfdd spp. in SA ure conductivity. and 
muotall, 

Higher nif senerilly means green permineney of 
wilter bodies. In the MRI survey at WA (1994-2000), 
Perthia spp. were loud at SSO) sites. All have average 
wnniual radintill vl 600-1400 movye CS. talse, Dept 
Conservation & Lind Maiinement Perth, pers. conin). tn 
SA. all sites other than Woods Point have aenual rainfall 
SOOO nun yr, Only a sn} aed pf SA receiwes paul) tn 
Wis mhywe (her Kangaroo tstaud. Mp Lolty Ranges, MI 
Gambier regiani Woods Point dacs ot receive high 
falrifall, Bit nonetheless as at High deerce al permnaneney. 

The were conductivity of WA ant SA MRH silts 
Where Perida spp. Were found Was arotind SSO) Sic 
CS. Plutse. pers. conn: AWOC. Hapubly. The conduetivity 
of SA MRH sites wes 1S0-(00 O00 yS/erm, und most were 
SIS00 yore Pherelore S50) paver is “Presh> suid 
(uicominen by SA swinidan, Cimdvetivity tray fiat the 
distribution of Meri spp. TSA 

These flew records wid tothe koto biodiversity of SA 
unt my abe contribute to evoluloniry and eeologicul 
stidicn. Amphipods ire potentially uselul is eavironmental 
Hidieators. due to their ecologies! impertines. juumerigg 
ubundance and seasigwity fo toxteanys and pollakanbs'' bal 
Hicir use is linvited to Wie few regions where comprehensive 
Taxonenie ind naticidl History invesienrtions have been 
Underken® These new reorrus pisy extend their pve ta this 
wuly. 

These city are trom programs tunded by Land wad Walter 
Austeulit. Eivaroninent ustatd. the Ravaronnienar 
Protection Ageney of South) Australia, the Onkaparinue 
Canctment Water Manasenient Gourd aad the Murpiye 
Darling Basin Commission. Thanks to Joli Bradbury for 
likenaniie advice, Stuart Hlilse and Iver Grows for 


126 


unpublished data, Beth Hughes and Chris Madden, Paul 
McEvoy, Tracy Venus, Vlad Tsymbal, Peter Schultz, Amber 
Lang and Darren Hicks for collection and identification of 
samples. 


' Chilton, C. (1920) The occurrence in the Brisbane River of the 
New Zealand amphipod, Paracorophium excavatum (G. M. 
Thomson). Memoirs of the Queensland Museum 7, 1-8. 

? Bradbury, J. H. & Williams, W. D. (1999) Key to and checklist 


of the inland aquatic amphipods of Australia. Technical Reports of 


the Australian Museum 14, \-21. 


‘ Straskraba, M. (1964) Perthia n.g. (Amphipoda, Gammaridae) 
from freshwater of Western Australia, with remarks on the genera 
Neoniphargus and Uroctena. Crustaceana 7, 125-139, 

+ Chapman, M. A. & Lewis, M. H. (1976) “An Introduction to the 
Freshwater Crustacea of New Zealand” (William Collins (New 
Zealand) Ltd., Auckland). 

‘Hart, B. H. & Fuller S, L. H. (1979) “Pollution Ecology of 
Estuarine Invertebrates” (Academic Press, New York). 

°*Thomas, J. D. (1993) Biological monitoring and tropical 
biodiversity in marine environments: a critique with recom- 
mendations, and comments on the use of amphipods as 
bioindicators. Journal of Natural History 27, 795-80. 


D. J. TAYLOR, Australian Water Quality Centre, PMB 3 Salisbury, South Australia 5108. E-mail: 


daria.taylor@sawater.com.au 


ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED 


Patron: 


HER EXCELLENCY MARJORIE JACKSON-NELSON, AC, MBE 
GOVERNOR OF SOUTH AUSTRALIA 


OFFICERS FOR 2002-2003 


President: 
O. W. WIEBKIN, BSc, PhD 


Vice-Presidents: 


R. W. FITZPATRICK, Bsc Agric, MSc Agric, PhD 
N. EF ALLEY, BA(Hons), MA, PhD 


Secretary: Treasurer: 
J. H. LOVE, BA, BD J. T. JENNINGS, BSc(Hons), PhD 
Editor: Assistant Editor: 


N. F. MORCOM, BSc(Hons), PhD 


Librarian: Programme Secretary: 
N. J. SOUTER, BSc(Hons) 


Minutes Secretary: Membership Secretary: 
A. J. McARTHUR, BE 
Members of Council: 
J. E. PATTISON, MA, BSc, MSc, Grad Cert Ed R. D. SHARRAD, BSc(Hons), PhD, DipT(Sec) 
P. A. PARSONS, BAgSc, PhD, ScD, FLS M. IQBAL, BSc(Hons), MSc, PhD 
M. A. J. WILLIAMS, BA(Hons), MA, PhD, ScD 


Printed by Graphic Print Group, 10-14 Kingston Avenue, Richmond, S.A, 5033