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)- - - - - -
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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
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
tentacles
stomach
stalk
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
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
¥ =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.
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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
* 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
Pig. 1 Location of the raised gravel beach deposit, Sellicks Beach, South Australia.
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.
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 “
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
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_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,
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
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
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
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
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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
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
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,
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
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
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(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 +
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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
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
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
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.
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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
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.
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)
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
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 &
\
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.
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
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
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
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
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
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
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
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~ 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
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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.
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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
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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
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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
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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
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
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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,
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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
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,
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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
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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
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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
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