Records of the Western Australian Museum Supplement No. 61: 323-333 (2000).

The Chilopoda and Diplopoda of the southern Carnarvon Basin,Western Australia

Mark S. Harveyt, Alison Sampeyl,2, Paul L.J. West1,3 and Julianne M. Waldock1

1Department of Terrestrial Invertebrates, Western Australian Museum,Francis Street, Perth, Western Australia 6000, Australia

2Present address: Lot 1984 Weller Rd, Hovea, Western Australia 6007, Australia3 Present address: Halpern Glick & Maunsell Pty Ltd, John Tonkin Centre,

629 Newcastle St, Leederville, Western Australia 6007, Australia

Abstract - At least 15 species of centipedes and 11 species of millipedes wererecorded from the southern Carnarvon basin. Centipede diversity isdominated by scolopendrids with 13 species, but most species are relativelywidespread both within the survey area and in other regions of WesternAustralia. Centipede communities were found to be strongly correlated withrainfall and soil texture variables. Millipede diversity is dominated bymembers of the paradoxosomatid genus Antichiropus which is represented byseven undescribed species, five of which are restricted to individual surveyareas. The other two species are found at two and three survey areas,respectively. These distribution patterns are seen elsewhere in WesternAustralia with Antichiropus which contains many highly localised species.

INTRODUCTIONThe myriapods are an assemblage of the four

classes Chilopoda (centipedes), Diplopoda(millipedes), Symphyla (symphylans) andPauropoda (pauropods), all of which are known tooccur in Australia (Harvey and Yen, 1989).Symphylans and pauropods are small and generallyrestricted to wetter biotopes, whereas centipedesand millipedes are found in many types ofterrestrial environments. As with so many otherinvertebrate groups, data on the myriapod fauna ofthe semi-arid and arid zones of Australia arefragmentary, mainly being derived from specimenscollected opportunistically and for taxonomicrevisions.This paper attempts to examine patterns in the

composition of the myriapod communities of thesouthern Carnarvon Basin, Western Australiathrough an intensive pitfall trapping program at 63localities (Figure 1) positioned in a stratifiedrandom array to represent the geographical extentand range of major geomorphological units of thestudy area. Species occurrences were analysed forcorrelations with numerous physical attributesmeasured for each site.Our knowledge of the myriapod fauna of the

Carnarvon Basin is somewhat fragmentary, with thescolopendrid centipede fauna being the best studiedelement (e.g. Kraepelin, 1908; Koch, 1982; Koch,1983d; Koch, 1983a; Koch, 1983c) (Tables 1, 2).Other groups of centipedes are poorly known and

one group, the Geophilida, could not be identifiedto species level due to our insufficient knowledge ofspecies-level characters in this diverse and complexgroup. The millipedes are also poorly known withonly a single species having been previouslyreported for the area (Attems, 1911). However, incontrast to the geophilids, we wer,e able tosatisfactorily separate species using the peculiarmorphology of the male gonopods.

MATERIAL AND METHODS

Study AreaThe survey area of the southern Carnarvon Basin

covers some 75 000 km2in the central region ofAustralia's west coast. It is centred on Shark Bayand extends from the Murchison River in the southto the Minilya River in the north, and eastwards tobeyond Gasgoyne Junction (Figure 1). The physicalenvironment is discussed by Wyrwoll, Stoneman,Elliott and Sandercock (2000) and Wyrwoll,Courtney and Sandercock (2000). The climate issemi-arid with temperate weather systems in thesouthern half of the survey area, and tropical andtemperate systems in the northern half.The survey area covers the northern half of the

Irwin District of the Southwestern Province, and thesouthern half of the Carnarvon District of theEremean Province (Beard, 1980). The vegetationalcharacteristics are documented by Keighery et al.(2000).

DOI: 10.18195/issn.0313-122x.61.2000.323-333

324

113°

24°

Bernier 1

25°

DorreI

DirkHartog I

26°

~N

27° 0 25 50

Kilometres

M.S. Harvey, A. Sampey, P.L.}. West, }.M. Waldock

24

1,',-~":r.... <

4:-

27°

Figure 1 Map of the study site showing sites utilised in the study.

Centipedes previously recorded from the southern Carnarvon basin.Table 1

Species

GEOPHILIDALamnonyx tahitiensis WoodTuoba sydneyensis (Pocock)

SCOLOPENDRIDACormocephalus aurantiipes (Newport)Cormocephalus turneri Pocock

Ethmostigmus pachysoma L.E. KochEthmostigmus rubripes (Brandt)

Scolopendra laeta Haase

Scolopendra morsitans Linnaeus

Family

GeophilidaeGeophilidae

ScolopendridaeScolopendridae

ScolopendridaeScolopendridae

Scolopendridae

Scolopendridae

General distribution

Widespread in the Australasian region.Widespread in coastal regions ofAustralia, New Guinea, Solomon Islands.

Widespread across mainland Australia.Widespread across the drier portions ofsouthern Australia.Known only from three WA localitiesWidespread across mainland Australia,as well as parts of Indonesia, NewGuinea, Solomon Islands and China.Widespread across mainland Australia.

Cosmopolitan and widespreadwithin Australia.

Distribution within southernCarnarvon basin

TamalaDenham; Eagle Bluff

Carnarvon; Dorre Island

Dirk Hartog IslandMardathuna Station; TamalaBaba Head; Denham; Dirk HartogIsland; Dorre Island; Edel Land

Denham; Dorre Island; Gee GieOutcamp; 10 km ESE of MeedoHomestead; TamalaBaba Head; Boologooro Station;Calligiddy; Carnarvon; CoburnStation near Hamelin Pool;Denham; Dirk Hartog Island;Gee Gie Outcamp; Shark Bay area;Tamala; 8 km NE. of TamalaHomestead; Useless Loop

Authority

Attems (1911)Attems (1911), Jones (1998)

Koch (1983c)

Kraepelin (1908), Koch (1983c)Koch (1983d)

Kraepelin (1908), Koch (1983d)

Kraepelin (1908), Koch (1982)

Kraepelin (1908), Koch (1983a)

326 M.S. Harvey, A. Sampey, P.L.J. West, J.M. Waldock

Table 2 Millipedes previously recorded from the southern Carnarvon basin.

Species

POLYDESMIDAAntichiropus monocanthusAttems

Family General distribution

Paradoxosomatidae Described from three WAlocalities, including two inthe southern Camarvon Basin;subsequently not recorded inthe literature.

Distribution withinsouthern Carnarvonbasin

Dirk HartogIsland; Tasmania

Authority

Attems (1911)

Field SamplingThe survey was based upon 63 quadrats in 13

groups of two to six (but usually five). Each groupis termed a 'survey area'. The quadrats werepositioned to cover the geographical extent of thestudy area, represent the array of stratigraphic unitspresent and provide some pseudoreplication of itsmain substrate types.Sampling was conducted with the aid of two

different types of pitfall traps. Specimens were alsohand collected during the field program, but to datewe lack the resources to sort and label this material.All specimens are lodged in the Western AustralianMuseum, Perth.Live specimens were removed from vertebrate

pitfall traps (seeMcKenzie et al., 2000) during eachfive-day trapping program (September-October1994, May-June 1995) and preserved in 75% ethylalcohol.Wet pitfall traps consisted of 25 litre buckets (300

x 400 mm) dug into the ground, with the top flushwith the soil surface. Each was fitted with a lid intowhich five 10 cm diameter holes were cut. A pieceof chicken wire (c. 1 cm mesh) was suspendedbelow the lid to allow any vertebrate whichventured into the trap to escape. This may haveenabled larger invertebrates such as beetles andspiders to escape although a 2.5 cm gap was leftbetween the wire and the wall of the bucket. Eachtrap was equipped with approximately 3.5 litres ofpreserving fluid (3 1ethylene glycol, 40ml formalin,350 ml water). The traps were opened in August1994, and the contents were cleared in September-October 1994, January 1995, May-June 1995 andAugust 1995, when they were removed. Sampleswere returned to the laboratory, washed in waterand stored in 75% ethyl alcohol until being sorted.Target organisms were removed, labelled andplaced in separate vials or jars.Five wet pitfall traps were placed in each of the

63 quadrats, the total number of traps utilised being315. Over the 12-month trapping period, we hadthe equivalent of some 115,000 trap nights. The 12-month trapping period allowed for the fullsampling of all seasons, and reduced the suspectedbias which may occur in restricted samplingperiods.

Data AnalysisThe data matrix (Table 3) comprised the presence

and absence of species on each quadrat, rather thantheir relative. abundance. It was analysed using thecomputer package PATN (Belbin, 1993) to procurespecies composition patterns. The associationmeasure 'Two-step' was used to determine thequantitative relationship between each pair ofspecies, and the Czekanowksi measure was used tocompare the quadrats according to their speciessimilarities. The dendrograms derived from theseassociation matrices were derived using a modified'unweighted pair group arithmetic averaging'(UPGMA) hierarchical clustering strategy (Sneathand Sokal, 1973).The biological patterns revealed by these analyses

were investigated in terms of a set of attributesrelated to the physical environments of the quadratspresented by Wyrwoll, Stoneman, Elliott andSandercock (2000) utilising the GSTA module inPATN, and assessed statistically with Kruskal-Wallis K-sample tests.

RESULTSSeveral thousand myriapod specimens in more

than 1080 vials were sorted and identified duringthe current survey, representing two classes,centipedes and millipedes. Of the centipedes,three of the five known orders were recorded inthe survey area, whereas only four of the nineknown Australian orders of millipedes wererecorded.

ChilopodaA total of 15 species were recorded from the 63

quadrats (Table 3), ranging from two species/site attwo sites (MOl and PEl) to 11 species/site at twosites (KE2, KE3).

GeophilidaMembers of the order Geophilida are slender,

long-bodied centipedes which we recorded at 31sites. Any resolution of biogeographical patternswithin this taxon were concealed because we wereunable to identify our specimens past ordinallevel.

Table 3 Centipedes and millipedes collected during the present survey.BB BO CU EL GJ KE MD MR NA NE PE WO ZU1 2 3 4 5 1 2 3 4 5 1 234 5 6 1 2 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 234 5 1 2 3 4 5 1 234 5 1 2 3 4 5 1 234 5

CHILOPODAGeophilida spp. XXXX XXXXX x xx x xx x xxx x x xxx x x XXXXXScolopendrida Scolopendridae Asanada sp. X X X X xx XScolopendrida Scolopendridae Arthrorhabdus mjobergi XScolopendrida Scolopendridae Arthrorhabdus paucispinus X X X X X XXXX xx XXXX xxx XXXXXScolopendrida Scolopendridae Arthrorhabdus sp. XScolopendrida Scolopendridae Cormocephalus aurantilpes X XXXXX xx xxx x XXXXX XXXXX XXXX xx x XXXXXScolopendrida Scolopendridae Cormocephalus strigosus XX XX XScolopendrida Scolopendridae Cormocephalus tumeri XXX XX XXXXX xx x x XXXXX XXXXX XXXX XXXXX XXXXX XXXXX XXXX XXXXX XXXXScolopendrida Scolopendridae Ethmostigmus curlipes X X xxx XXXX xx xxx x xx XXXX x x xx XScolopendrida Scolopendridae Ethmostigmus pachysomll XX X xx x X XScolopendrida Scolopendridae Ethmostigmus parkeri XScolopendrida Scolopendridae Ethmostigmus rubripes XX XX XXXXX XXXXXX XX XXX XXX X XXX XX XXX XX XXX XXXXX XXXXX XXX XXXXXScolopendrida Scolopendridae Scolopendra laeta XXXXXX XXX XXXXX XX XXX XXXX XXXXX XXXXX XXXXX XXXX XXX XXXXXXScolopendrida Scolopendridae Scolopendra morsitans XXXXX XXXXX XXXXXX XXXXX XXXXX XXXXX XXXXX XXXXX XXXXX XXXX XXX XX XXXXXScutigerida Scutigeridae Allothereua sp. XXXX XX X XXXX XXX XXX XXXXX X X XXX XXXX X X XXX XXX XXX

DIPLOPODAPolydesmida Paradoxosomatidae Antichiropus sp. I XXXXX

Polydesmida Paradoxosomatidae Antichiropus sp. 2 X X

Polydesmida Paradoxosomatidae Antichiropus sp. 3 XPolydesmida Paradoxosomatidae Antichiropus sp. 4 X

Polydesmida Paradoxosomatidae Antichiropus sp. 5 X X X

Polydesmida Paradoxosomatidae Antichiropus sp. 6 X

Polydesmida Paradoxosomatidae Antichiropus sp. 7 X XX X XXXPolyxenida Polyxenidae Genus I, sp. I X XX X XPolyxenida Synxenidae Genus I, sp. I XX X XPolyzoniida Siphonotidae Genus I, sp. I X X X X X X XX X X XSoirostreDtida ? Genus 1, so. I X X XXXX

328

ScolopendridaThirteenspeciesofscolopendridcentipedes,

representingfivegenera,werecollectedduringthesurvey.

Asanadasp.wascollectedatsevensites(Table3),buttherelationshipbetweenthispopulationandthoserecordedfromnorthernQueensland (:r<:och,1983b)andWoodstock StationinthePilbara(HarveyandWaldock, 1991)remainstobedetermined. Itnow seemsthatAsanada iswidespreadacrossnorthernWestern Australia.

Only twoAustralian speciesofArthrorhabduswere identifiedinarecentrevisionofthegenus(Koch,1984),andbothwerenewlyrecordedfromthestudyarea(Table3).ArthrorhabdusmjobergiwascollectedonceatKE2, while A. paucispinuswascollectedat23sitesacrossmuchofthestudyarea.Anunidentifiedspecies,A.sp.,wastakenatBOlo

Kraepelin(1908)andKoch(1983c)recordedtwospeciesofCormacephalusfromthestudyarea,C.

aurantiipesandC. turneri(Table1).Both ofthesespecieswere recollectedduringthesurvey,C.

a

M.S. Harvey,A.Sampey,P.L.J.West, J.M.Waldock

aurantiipesfrom34sitesandC. turnerifromallbutsevensites(Table3).Cormacephalusstrigosusisnewlyrecordedfromthearea,althoughKoch(1983c)recordedthespeciesfromsitestothenorth(BarrowIsland)and south (e.g. Mullewa,Northampton)oftheCarnarvonbasin.

Two species of Ethmostigmus have beenpreviouslyrecordedfromthestudyarea,E.rubripesandE. pachysoma(Kraepelin,1908;Koch, 1983d)(Table1).We recordedthesetwospecies,aswellasE. curtipesandE. parkeri.Ethmostigmusrubripesiswidespread overmuch ofmainland Australia,includingseveralsitesintheSharkBay area(Kraepelin,1908;Koch, 1983d)(Table1).Werecordeditfromallbuteightsitesinthestudyarea(Table3).Ethmostigmuscurtipeswas takenfrommostsurveyareas,withtheexceptionofGascoyneJunction,Meedo, Woodleigh andZuytdorp(Table3).ItwaspreviouslyknownfrominlandareasofWestern Australia (Koch,1983d).Ethmostigmuspachysomawas previouslyknownfromonlythreelocalities,allsituatedinWestern Australia (Koch,

,-

k.hllllll ~ ..19...I•••D.I-I••.•.1.h.J~

BDEEDROORBOOAERREREBODOJEOUJJJJBBAAEEUULUAAEUEDEEUUUUEEUOOOLUDE143553341321124532555321411435224233535224544122261231443451511

,-

b

IIII.II"IIII'II".II.I..II...~ III1111111 11IIIII~

BDEEDROORBOOAERREREBODOJEOUJJJJBBAAEEUULUAAEUEDEEUUUUEEUOOOLUDE143553341321124532555321411435224233535224544122261231443451511

Figure2 Attribute valuesfor(a)precipitationinthewettest quarter(PwetQ),and(b)soilpotassium(KHC03)

superimposedashistogramsonthedendrogramstructurederivedfromtheclassificationofthe63quadratsintermsofspeciesbelongingtotheassemblage.Quadratcodesareprintedvertically.NotethatPwetQandKHC0

3wereinverselyinter-correlatedinthestudyarea.

ChilopodaandDiplopoda 329

Table4 Centipededatamatrix,reorderedaccordingtospeciesco-occurrences.

Species Sites

*Arthrorhabdusmjobergi

BMNNMMWWMBWWNKMMKMKBWMBGKBCGGGGIBBNNPPCCECNNNCKMNPCZZZNPZBBBEZIMPBDEEDROORBOOAERREREBODOJEOUJJJJIBBAAEEUULUAAEUEDEEUUUUEEUOOOLUIDE14355334132112453255532141143521242335352245441222612314434515111

I IAsanadasp. * **** * * 1 1

Arthrorhabduspaucispinus * * ******* * * ******* *** I * * 1Ethmostigmuspachysoma * ** * ** ** * 1 ICormocephalusstrigosus * * * 1 * * I

.------------------------------------------------------+------------------------------+--Cormocephalusaurantiipes ** ****** * 1 ************ *********** * * I

Ethmostigmuscurtipes * * ** * 1* * * * * * * * * * * * * * * * * * * * ICormocephalustumeri *******************************1****** ******************* 1

r ~ *******************************1****************************1*Ethmostigmusrubripes ********* ***************1*****************************1**

Scowpendrawem ******************** * 1******************************1Allothereuasp. *************************1********* ** **** * I *Geophilidaspp. * ********** * 1 * ******* **********I

._-----------------------------------------------------+------------------------------+--Arthrorhabdussp. * 1

._-----------------------------------------------------+------------------------------+--Ethmostigmusparkeri * I

._-----------------------------------------------------+------------------------------+--1

1983d).ThepresentstudyrecordeditfromGascoyne Junction, Kennedy Ranges andMardathuna (Table3),allinlandlocalitiesinthenorthernportionofthestudyarea.EthmostigmusparkeriwasonlytakenonceatMRl; elsewhereithasonlybeenfoundonthewesternedgeoftheGreatSandyDesert (Koch,1983d),andtheMardathunarecordisasignificantrangeextension.ScolopendralaetawaspreviouslyrecordedfromseveralsitesinthestudyareabyKraepelin(1908)andKoch(1982)(Table1),andwerecordeditfromallsurveyareas,withtheexceptionofGascoyneJunction.Itisawidespreadspeciesfoundovermuch ofmainland Australia (Koch,1982).ScolopendramorsitansisacosmopolitanspecieswhichisfoundovermuchofthedrierportionsofAustralia.IthasbeenpreviouslyrecordedfromseveralsitesintheCarnarvonbasin(Kraepelin,1908;Koch,1983a);Table1),andwerecordeditfromallstudysitesexceptELl,EL2andPEl.ItseemstobeubiquitousthroughouttheCarnarvonbasin.

ScutigeridaAsingle,unidentifiedspeciesofAllothereuawascollectedduringthesurvey.Itwasfoundinallsurveyareas,withtheexceptionofNerrenNerren,at42ofthe63quadratssampled.

AssemblageCompositionThecompositionofcentipedecommunitieswerefoundtobecorrelatedmoststronglywithrainfallandsoiltexture.Thethree-groupsofsitesdefined

fromtheclassificationshowedsignificantdifferencesinthewettestquarterprecipitation(P<O.OOOl,withKruskal-Wallis=30.0)(Figure2a)andwithsoilpotassium(P<O.OI,withKruskal-Wallis =19.9)(Figure2b)andarangeofothercorrelatesofsoiltexture.Group1consistsofsiteswhichreceivelessrainfallandhavehigherpotassiumlevels,whereasgroup2consistsofsiteswhichreceivehigherrainfallandhavelowerpotassiumlevels.Group3comprisedverydifferentsites:asanddune(MDl)andasalineclaypan(PEl).Theirseparationasadistinctgroupmaybeanartefactoftheirlowspeciesrichness(Table4).However,PwetQandK(HC0

3)arestronglyinter-

correlatedinthestudyarea.

Diplopoda

PolydesmidaOnlyasinglepolydesmidangenus,Antichiropus,wasrepresentedamongstthematerialcollectedduringthesurvey,whichisconsistentwiththeresultspresentedbyAttems (1911)andbyspecimensofmillipedesheldinthecollectionsoftheWestern AustralianMuseum. Antichiropusisabundantinsouth-westernAustraliawhereseveraldescribedandnumerousundescribedspeciesareknown tooccur (Attems, 1911) Harvey,unpublisheddata),andinSouthAustralia,whereA.mammilliferJeekelisknownfromPoochera(Jeekel,1982).Themostnortherlyrecordsareofatroglobiticspecies,A.humphreysiShear,recentlydescribedfromCapeRange(Shear,1992).Theonly

330 M.S. Harvey, A. Sampey, P.L.}. West, }.M. Waldock

--------------M-----------------------------------------------M---------------------------------------

(a) Temperature in the wettest quarter (T wetQ), Kruskal-Wallis =19.3, df 3, P<O.0002.16.00 16.70 17.40 18.10 18.80

GRP +-----------------+-----------------+-----------------+-----------------+1 ---M-2 ---------------------------------------------------M------------------3 ------M-----------4 ----M--------------

(b) Temperature in the warmest quarter (T_WarQ), Kruskal-Wallis =20.7, elf 3, P<O.OOOl.27.10 27.83 28.55 29.28 30.00 I

GRP +-----------------+-----------------+-----------------+-----------------+ 1

1 ----M------2 ----------------------M------------------------------3 ---------M--------4 -------M-----------

(c) Temperature in the coldest quarter (T cldQ), Kruskal-Wallis =18.8, df 3, P<0.OOO3.14.60 15.35 16.10 16.85 17.60

GRP +-----------------+-----------------+-----------------+-----------------+1 ---M--2 -------------------------------------------------M------------------3 ------M---------4 ------M-------------

(d) Precipitation in the wettest quarter (P_wetQ), Kruskal-Wallis =20.8, df 3, P<O.OOOl.100.0 126.3 152.5 178.8 205.0

GRP +-----------------+-----------------+-----------------+-----------------+1 -----M-----2 ---------------M------------3 -------------M-------------------4 ----------------------------M-------------

(e) Soil carbon (C), Kruskal-Wallis =15.9, df 3, P<O.OO12.0.1400 0.3225 0.5050 0.6875 0.8700

GRP +-----------------+-----------------+-----------------+-----------------+1 --------M----------------2 --------M-------------34

Figure 3 Relationships between UPGMA quadrat groups and selected environmental variables (M =mean; GRP =quadrat groups delineated in Table 5).

species previously known from the survey region isAntichiropus monocanthus Attems, recorded fromDirk Hartog Island and Tamala by Attems (1911)(Table 1). This species was not recollected duringthe survey, although seven other species, allundescribed, were taken.Four species were found to be restricted to

individual survey areas: Antichiropus sp. 1 to thefive Zuytdorp quadrats (ZU1-S); A. sp. 2 to twoNanga quadrats (NA1, NAS); A. sp. 3 to a single

Edel Land site (ELl); and A. sp. 4 'to PE4 on PeronPeninsula. Antichiropus sp. S was taken from bothNanga (NA3, NAS) and Zutydorp (ZU3).Antichiropus sp. 6 and 7 are probable sister-species,sharing significant similarities in the morphology ofthe male gonopods. Antichiropus sp. 6 was foundonly at PES, whilst A. sp. 7 was found at Ede1 Land(EL2), Nerren Nerren (NE1, NE2) and Zuytdorp(ZU1, ZU3, ZU4, ZUS), all in the southern portionof the survey area.

Chilopoda and Diplopoda 331

Table 5 Millipede data matrix, reordered into groups by classifying species according to their co-occurrences, andquadrats according to similarities in their species composition.

QuadratsGroups: 1 2 3 4

BBCBC BCCMMNBW EPPNNN ENNZZZZZOOUOU OUURREOO LEEAAA LEEUUUUU1 344 6 22535353 145 1 3 5 21213524

Antichiropus sp. 1 * * * * *Antichiropus sp. 7 * * * * * * *Siphonotidae sp. 1 * * * * * * * * * * *

+ +- +Antichiropus sp. 5 I I * * I *

Spirostreptida sp. 1 I I * * I * * * *+

Antichiropus sp. 2 I * *+ +

Antichiropus sp. 3 I * IAntichiropus sp. 4 I * IAntichiropus sp. 6 I * I

+ +Polyxenidae sp. 1 * * * * * I ISynxenidae sp. 1 * * I * * I

The restricted distributions of each of the speciessuggests that A. monocanthus may not be found atboth Dirk Hartog Island and Tamala as suggestedby Attems (1911). Attems did not provide details ofthe sexes of the material which he examined, and itis possible that one or both of the localities isrepresented by females or juveniles of a secondspecies. The morphology of the male gonopod isthe only morphological feature which separateseach species, whereas females and juveniles areindistinguishable from each other. Reexaminationof Attems specimens is needed to establish whetherA. monocanthus is truly present at both sites.

PolyxenidaTwo species of Polyxenida were recorded during

the survey: one was a member of the Polyxenidaeand the other a member of the Synxenidae,. Theformer was taken at Boolathana (BOl, B03, B04)and Cape Cuvier (CU4, CU6), while the latter wastaken from Boolathana (B04, B05), Cape Cuvier(CU6) and Woodleigh (W03).

PolyzoniidaA single species of polyzoniidan belonging to the

family Siphonotidae was found in several surveyareas (Boolathana, Cape Cuvier, Mardathuna,Nerren Nerren and Zutydorp) (Table 5). Polyzoniidmillipedes are usually found in wetter biotopeswithin Western Australia, where they may becommonly found in the Karri (Eucalyptusdiversicolor) forests of the south-west. TheCarnarvon species most certainly represents anundescribed species, but the relationships of this

species with other Western Australian polyzoniidsis uncertain.

SpirostreptidaA single species of Spirostreptida was taken at

Nanga (NA3, NA5) and Zuytdorp (ZU2-5).

Assemblage CompositionMillipedes were very patchily distributed across

the study area and were found at only 27 of the 63study sites (Table 5), which is less than half of thequadrats sampled. For the analyses, we ignoredthose sites that lacked millipedes. Strongpatterning was detected with significantcorrelations between rainfall and temperature, andother related variables such as soil carbon (Figure3). In general, most millipede communities arerestricted to the wetter, cooler sites, which arelargely situated in the woodland regions of thesouthern portion of the survey area, or are situatedalong the coast.

DISCUSSION

ChilopodaThe strong correlations between centipede

communities and environmental variables, moststrongly with rainfall and soil texture, are consistentwith other groups of organisms examined duringthis study. In general, there is a greater diversity ofcentipede species in the drier regions of the surveyarea, with species such as Asanada sp., Arthrorhabduspaucispinus, Ethmostigmus pachysoma and

332

Cormocephalus strigosus found on relatively dry,sandy sites (Table 4, Figure 2).

DiplopodaAs noted above, millipede community

composition is strongly correlated with climateattributes and soil carbon, with most speciesrestricted to either the areas subject to temperaterainfall patterns or in areas which receive coastalrainfall (Table 5).The relatively small distributions of these seven

species of Antichiropus are probably not an artefact,although the short activity period of arid-zonemillipedes ensures that the sampling has not beencomplete throughout the area. In other. parts ofWestern Australia, Antichiropus species have showstrongly localised patterns of endemism at thespecies level (Harvey, unpublished data), and itseems that the group has radiated extensively insuitable habitats throughout south-westernAustralia, aided by low vagility and a dependenceupon suitable habitat.The Antichiropus species were all taken in the

lower half of the survey area, with the mostnortherly record on Peron Peninsula. The bulk ofthe localities are within the Southwest BotanicalProvince (Beard, 1976; 1980; 1990), and even thesites at Nerren Nerren, which cross the boundarybetween the Southwest and the EremaeanBotanical Provinces, seem to be within thisregion, as A. sp. 7 was found during the periodMay-August 1995 at sites NE1 and NE2, but wasnot found at NE3-5 (Table 3). The presence ofspecies of Antichiropus north of Beard's boundaryat Peron Peninsula (Table 3) and Dirk HartogIsland (Attems, 1911) provide some support forthe hypotheses of Diels (1906) and Burbidge(1960) who included these regions in theSouthwest Botanical Province. However, untilmore is known of the composition andrelationships of the millipede fauna of regions tothe south and north of the study area, littlefurther can be deduced from these minortransgressions.

ACKNOWLEDGEMENTSFunding for this project was provided by the

Commonwealth through the National ReservesSystem Co-operative Program of the AustralianNature Conservation Agency (now EnvironmentAustralia), together with State funds provided bythe Department of Conservation and LandManagement and the Western Australian Museum.We wish to thank those people who ably assisted usin the field program, Mike Lyons for Figure 1,Norm McKenzie for assistance with the numericalanalyses, and Norm McKenzie and Bob Mesibovfor useful comments on the manuscript.

M.S. Harvey, A. Sampey, P.L.J. West, J.M. Waldock

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W. Michaelsen and R. Hartmeyer (eds). Die FaunaSiidwest-Australiens. Vol. 3: 147-204. Gustav Fischer,Jena.

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Beard, J.S. (1980). A new phytogeographic map forWestern Australia. Western Australian HerbariumResearch Notes 3: 37-58.

Beard, JS. (1990). Plant life ofWestern Australia. KangarooPress, Kenthurst.

Belbin, L. (1993). PATN: pattern analysis package. CSIRO,Canberra.

Burbidge, N.T. (1960). The phytogeography of theAustralian region. Australian Journal.pf Botany 8: 75- I

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Manuscript received 28 June 1999; accepted 7 December 1999.