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A new species of Calamoecia (Copepoda: Calanoida) from arid Australia, with comments on the calanoid copepods of the Paroo, northwestern Murray-Darling Basin PDF

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Preview A new species of Calamoecia (Copepoda: Calanoida) from arid Australia, with comments on the calanoid copepods of the Paroo, northwestern Murray-Darling Basin

. ANEW SPECIES OF CALAMOECIA (COPEPODA: CALANOIDA) FROM ARID AUSTRALIA. WITH COMMENTS ON THE CALANOID COPEPODS OF THE PAROO, NORTHWESTERN MURRAY-DARLING BASIN BRIAN T1MMS V. Timms, B.V. 2001 06 30: Anew speciesofCalamoecia (Copepoda: Calanoida) from arid Australia, with comments on the calanoid copepods of the Paroo, northwestern Murray-Darling Basin. Memoirs ofthe Queensland Museum 46(2): 783-790. Brisbane. ISSN 0079-8835. Calamoecia bayiyi sp.nov. is described from claypans in the Paroo catchment in southwestern Qld and northwestern NSW. It also occurs in WA where it was previously knownastheCueformofC. lucasi. TenspeciesofcalanoidcopepodoccurintheParoowith Boeckella triarticulata, C. Canberra and C. lucasi common, B. fluvialis and C. zeidleri presentandB. robustarobusta, B. timmsi, C. bayiyi, Diaptomuslumhottztand Gladioferens spinosus uncommon. Co-occurrences are common, especially in claypans and riverine waterholes. & New species, Calamoecia, Calanoid copepods, environmental ecology?, hiogeography Brian I' Timms,SchoolojCcosciences, UniversityofNewcastle, Callaghan2308,Australia (e-mail: [email protected]); 28 March200} Calamoecia Brady contains 6 species Bayly, ecological data are available. Ecological and m1o99s2t,re1c9e9n8t; aHdadlisteio&ns,McCR.azee,idl21e0r0i1B)a.yTlwyo(aonfdthCe cbioop<eipeoodssrap(hBiacylys,tud1i9e9s6;onMaAluys,tra1l9i8a4n; cMaallaynoi&d halseiBayly liveinwatersinremotepartsofarid Bavlv, 1991; Maly & Maly, 1997; Maly et al., Australia. Also occupying some ofthese waters 1997; Timms & Morton, 1988; Walsh & Tyler, is a form similar to C. lucasi Brady but different 1998) have not included data from the eastern enough tobe recognisedasthe Cue form (Bayly, arid zone. 11998844))., InteahrasPobretenHerdelpaonrdted(TfirmommsC&ueM(oBratyolny,, foTrhmeoafiCmasloafmotheicsipaapluecraasries.tlo. daensdcrtiobecothmemeCnute 1988), and tNheWGascoyne-Murchison (Bayly, on ecological biogeography of calanoid cope- 1998), all in WA. Although Bayly (1998) pods in the Paroo catchment ofthe NW Murray- thoughtthiswestern form ofC. lucasiwas anew Darling Basin. species, he was reticient to proceed partly because it isallopatric with C. lucasis. s. (l.A.E. METHODS Bayly, pers. comm.). C, lucasis.s. is widespread in eastern and central Australia (Maly & Bayly, Specimens were measured under a Wild M3C 1991), but records ofC. lucasi in WA in Maly & stereomicroscope fitted with an eyepiece wBeasytleyr(n19f9or1m)a(nadliiansMCauleyfeotraml.)((19M9a7l)yr&efeBratyoliyt,s dmiicsrsoemcetteedr,wistthainteudngwsittehnCnheoerdalzeosl BilnacDke,PaenXd 1991, and S. Halse, pers. comm.). However C. mountanl on a microslide under the same lucasis.s. occurs at Lake Gregory, S Kimberley microscope. Appendages were studied and (Halse et al., 1998) and another variant of C. drawn using a Olympus BHA microscope fitted lucasi, the 'Lake Grace form' lives near Lake with Olympusdrawingattachment. Terminology Grace in SW WA. and abbreviations used in describing the appendages follow Bayly (1992). Discovery of populations similar to the Cue NW NSW SW Ecological data on the copepods ofthe Paroo mCsbfuyouatmrenpmyatfhtoierdnriimtfCchfiueesweriaPetanhsfrceooepCrsoa.m,rbalehtutaecaswssepiemecansi.iseni.stt.caaaNoinnonnddfetidCor.nmiltelsyudQcadlatrisdhcsai,tttihsna.tecnshrt.dee-, (war1rcae9eent9ngl7gteablrsn,eedadfn1rsoe9iondt9me8s1f;C4rou4ovur°menr2rpm5au'1wyb3Eil,n.myyeo2aand8rai°tNst5aao0)(tr.Ti,iSoinT)nmghatmelosstLPuaa1dar9ikre9ekeas3,iPconeof1evt9r1eh90yer70eaiN+d,n iveness despite chances for interbreeeding. theOverflow lakesarea(143°37,E, 30°45'S)and The Parooareacontains9othercalanoidcope- encompassing about two-thirds of the Paroo's pods includingmany inland forms forwhich few catchment of73, 600km2 . 3 ) 784 MEMOIRS OF THE QUEENSLAND MUSEUM SYSTEMATICS two-thirds along outer margin and lip expanded CRUSTACEA into rounded spade-like structure with denticles Class COPEPODA on and towards its margin. Base ofthis structure Order with short spine on a protuberance on the inner Family CENTROPAGIDAE Giesbrecht side and longer spine next to it. Left endopodite Calamoecia Brady (Ri)l-segmented, a little longer than the first 2 Calamoecia baylyi sp. nov. segments of the right exopodite (i.e. exopodite (Figs 1,2) withoutitsclaw), withalmostparallel sides. This segmentwith5 spines(2terminal,3 subterminal) Calpaompouelactiiaon,lufciags.t4AB-rDa)d.y; Bayly, 1^84 (in pari, the Cue decreasingalmost uniformally in length from the inner spine to outermost one. ETYMOLOGY. InhonourofDrIanA.E. Bayly,towhom so much is owed on the taxonomy and ecology of Female. On first legs (Fig. 1A), spine on outer centropagidcopepods. edge ofproximal segment (Rel ) and 2 spines on outer edge ofdistal segment (Re3) ofexopodite 20MmW0o.2Au57Tn947Ets8meR4mQd,IMoApWnlrLe2onm.s5giotc4mhrH8eo,Os5Ll,ILi1d0OQ.emusT1em9iYenmnPmDpsEerloPtaosent6oXadlm.,e,MQ,AuPMsLA1WeL.Ru25OAm52T.T4mYY8mCPPo3lEtE.loSteaclV3tl,ee0dnbQgo5bttMshyh. 2ws6-ie4t)g..hmTOahenbennstoefirfo(mtuahRtleelllrey)gspl(iawFnriigegtes.hasIbeDasc)neo,nntdpoarourontyxeeirsxmpaoislpnpouiedlxnseoespoo(fwadlsiiettgihesn t1h5ea9usthoQrMonW1725M4a8y7,,19T9u6r.kOeTyHPEanR,M29ACT3E3R,SI,AII4..43°1419''ES,, (sReec2o)ndaalrsyo swpiitnhulseusc.hMaidsdplineeeaxnodpoadiltaergesecgumrevnetd NW Bloodwood Station, 130km of Bourke, slender outgrowth on its distal innercorner. This NSW.colIected by the author, 19 May, 1998; 5 c?s, 3Vs, outgrowth with 2 rows of spinules. Distal exo- Melaleuca Pan, 29°33'S, 144°48'E, Bloodwood Station. podite segment (Re3) with 5 or 6 spines, outer- NSW.collectedbyauthor 19May 1998; aclaypan nearS most with secondary spinules, next inner one end ofLake Wyara, Currawinya National Park, 28°4S1S. 144°157H, Old, collected by author 14 May, 19%; longest but still a little shorterthan the segment. dissected 6 WAM C24994 and vial of10 6s and 10 ?s Curved outgrowth on Re2 subequal to Re3 plus WAM C24996 from an unnamed clavpan, 24°47'35"S. its spines. Endopodite (Ri) 1-segmented, with 7 H1a1l4s°e0921514"JuEly(C1B99558;eodfisIsleacltseedet9al.,W20A00M),Cco2l4l9ec9t5edfSr.oAm. aornd8 2loonng soeuttaeer(e4dgoer.)5.onPrionsnoermesidtee,rm1intaetrimningali,n TtrigieClaypan,24°38,34"S, 11 59'29"E,collectedbyS. elongated lobes on either side of prosome- Halse 17 August, 1994. urosome junction (Fig. 1E,F). Urosome (Fig. TYPE LOCALITY. A claypan 1.5km NE of 1L,F)withgenitalsegmentabout 1.5 x as longas Coomburra Waterhole, Currawinya National wide. Genital operculum with a distinct nipple SW and on a large mound. Left side of genital Park, 28°47'E, 144°22'E, Queensland. segment with a bulge almost halfway along its DESCRIPTION. Male, On first legs, outer edge length,butnoindicationofanassociated furrow. spineson exopoditepromixal segment(Rel and ) distal segment (Rc3) with abnormally large REMARKS. Variation between populations. WA secondary spinules (same in females [Fig. 1A|). Specimens from the Carnarvon region of are On fifth leg (Fig. 1B,C) distal basidopodite slightly different from the type material in a segment (B2) with a distinct space between number offeatures. On the 6 fifth leg, the spine insertion ofexopodite and endopodite, which in of the outer distal corner of the first exopodite the right limb is wider and a little indented. segment (Rel) is much longer (Fig. 2B). The Proximal segment of right exopodite (Rel) claw of the right exopodite (Re3) is markedly somewhatcuboidal withaspineon itsouterdistal bent furtheralong its length than in type material comerandasmall roundedprotrusionon itsinner (ca one-third to halfway along its length). The distal corner. Middle distal segment (Re2) with a right endopodite is curved inwards and the first prominent rounded projection midway on its and second segment (Ril, Ri2) even more inner surface. Claw (Re3) strongly bent about bulbous so that especially the second segment one-quarter way along its length, and lacking an presents a semicireular outer edge (Fig 2B). In inner spine. Right endopodite (Ri) 3-segmenled, the 9 the most noticeable difference is on the proximal (Ril) and middle segment (Ri2 third segment of the exopodite (Re3) where round-ovalshaped,butdistal segment(Ri3)more spines (6 instead of5) are longerthan in the type elongatedandbearing2 longsetaesubterminally. material, so that they exceed the length of the Left exopod (Re)1-segmented with asmall spine segment and ofthe eurved inner proeess ofthe NLW CALAMOEC/A FROM THE MURRAY-DARLING BASIN 785 secondsegment(Re2)(Fig. 2A). Bayly (1998) thought different material he examined from the Carnarvon region 'agreed exactly with the Cue population', i.e. was the form now known as C. bqylyi. The Cue population oiC. lucasis.i studied by Bayly ( 1984)isalmostidenticalto the type material from the Paroo. Like the Carnarvon population the second segment of the right endopodite (Ri2) in 8 is enlarged and with a semicircular outer edge, and in 9 the proportions and lengths ofthe spineson thethirdexopoditesegment are the same.(Fig.2C; Bayly, 1984, fig. 4). The Cuepopulation is similarto the Paroo type material in other features, including the short spine at the outer distal corner of the first right exopodite segment (Rel ) and the marked bend in the right exopod claw t(oRea3)thbieridnaglaobnogutitas qlueanrgtthe.r FIaGs.pec1t.sC,alreasmpoeectciivaelbya,yloyfis8p.fnifotvh.;leAg,s;VDf,irs9tlfeigf;thBl&eg;C,H,anvteenrtiroarlaansdppeocstteorfio9r Both these western genitalsegmentwithlobesoflastprosomalsegment; F, leftlateralaspectofV populations ofC. lucasis.I. genital segmentwith lobesoflast prosomal segment. B and C from holotype, areconsideredasbelonging A, D, E, F from allotype. Scale bars = 0.1mm. to C. baylyi. The Lake Grace population of C. lucasi s.i the projection midway alongthe innersurface of studied by Bayly (1984) exhibits most of the the middle right exopodite (Re2) is hardly features ofC. baylyi including mostofthe minor developed)(Fig.2D),ormoredeveloped(e.g.the modifications seen in the Carnarvon and Cue outgrowth on the left side ofthe genital segment material. It has however some curious features is more pronounced) (Bayly, 1984, fig. 5E,F). unique to it, including an outgrowth on the inner ThisLakeGracematerialshouldbeconsideredas distal corneroftheproximal segmentofthe right C. baylyi,thoughalittlemoreabberantthanother exopodite (Rel). This is much more prominent western populations. thantheusualsmallroundedprotrusionintypical For 5 populations of C. baylyi, prosomal C. baylyi, but not as long as in C. elongata, the lengthsare a little lessthan 1mm, with 8 lengths only other species with an outgrowth in this ca 10% less than 9(Table 1). position. Also unique to this population is the Differential Diagnosis. C. baylyi is easily variablesetationonthedistalsegmentoftheright recognisable by the rounded projection on the endopodite(Ri3) so thatgenerally the2 setaeare middle inner surface of the second right of very unequal lengths and usually much exopoditesegment(Re2)ofthemale(Fig.1B,C). reduced. In addition some ofthe features typical The only other species with a remotely similar ofC. baylyiareeithernotas well developed(e.g. structure is C. australica, in whichtheprojection / MHMOIRS OF THK QUKKNSLAND MUSI LM bp dsi rise, this feature is shared only by C. ampulla and i hansi . s $,. but the latterhas no terminal denticles. In addition, C. lucasi lacks an inner subterminal spine that is present in C. haylyi and overall the length of the left exopoditc isshorterrelativetothe left endopodite (a little greater than half in C hicmI $.$. « three-quarters in ('. baylyl)> c 5)Oilthe iTfthleg,thecur-,o_l outgrowth on the inner distal corner of the middle c.xopodile segment (Re2) is relatively long in C haylyi reaching about to the ltd. 2. Cahtmut ... . . i . i .. -.p. \-\o\. from \V\; A. second aiid third end o\~ the distal exopoditc (sseegcmeMnatteurifaelxofpourddueetaiolfs);-' Bf.iftphoslteger,iosrpeacsipmecetnsforfom6Carringhatrvfoifnthbalseiun, sInegmmaennyt poltuhseritsspsepeiineess,(ifnicgl.ud1 iD)n.g specimen from C4roarven basfn; C, posterioraspeel ol : fifth legs, C ue C. lucasi s.s.. this outgrowth pleogpsu,laLtaikoenG(rafatceer Bpaoypull\a.ii19o8n4,(afftieg.r 4BAa)y;lyI,X1p9o8s4t,erlilogr.5aAs)p»ecStcaolfe 3barfsift=h terminateswell short ot thedistal 1mm. segment plus us spines (Bayly, i). 1961 Only C halsei exnibi j. similar condition (Bayly. 1998)* isabouta thirdalongfheinneredgeandmoreover oi Again on the ¥ fifth legs, some ofthe 5 or6 sbaip pointed. spines of the distal exopod segment (Re3) are Other notable features are variously shared only a httle longer than the length ofthesegment with a limited range of other species. These inC />mVi/ (rig. ID), as forC kahet(5 spines) include: and C, salina also (but C. salinu has only 2 1 )Outeredgesoftheexopodsofthe first pair erf terminal spines instead of6), By contrast in most legs in both sexes have 3 spines each with a few otherspeciesofCalamoecia, including C. tut asi. largestoutsecondary spinules (Fig. IA), Ofother (he spines are more than twice the length ofthe ( alamoecia species examined for these spines, segment (Bayly. 1961) and sigmlicantly in C (. . hdkei hasthem andsignificantly C. fUi i lucasi s.s. there are only 5 spines, though some does not (Bayly. 1998). speeies have 6 (e.g. C. trifida). Onl> C. zeidleri 2)Theclaw ofthe rightexopoditc ol the r fifth has Shorter tenninal spines (which number 5) leg(Rc_i l. lacksaninnerspine(Fig. iB.C)as inC (Bayly; 19X4). atistfaiicdi C, Canberra^ C. Itaisti and (.' zeniieri In the dichotomous key provided by Bayly (bBuatyalhy,ot1h9e9r8)s,peCcieIsnrhaastves.\it.(aBlasyolyl.ack1s9%th)i.s spine (1,I2992N).9< CandhuyIlIvitwoo(u\ldluckaesyi,outb:uttharsoupgohinctoeudpleotust 3)Theleftendnpoditeofthe i fifth leg(Ri I lis above is easily distinguished from this species. I-segmented, very elongated, exceeding the Also, as shown by Bayly ( 19S4) the Cue form of length ofthe left exopod and hears 5 stfpeeeraetmiuirenesalliaiststtdsedhsauarbbetodevremw,iintbahulttsuhpseiunaeslsal.ymeInhoi4ts po:puIlai tiIo.nsProOfSOCmabluylhein.gt*hsOnilnym6mJ'osfan10d 4'.•>?ainmdeaIstu)r8e'dS.I'rom 5 with C lurasi v.v.. which although ol 1 Mean Range ..i \r.-.i, I.' i Souro . similar construction, has 2-5, usually Icngjci length ditlii 41t%crminal/subtenntnalspines.(Bavlv, (-'NTMVi 0.0] a$H l i"i:.,i,il 1,1992, 1998). PBlaraoIwood, g |1..! ,'m>: ; h 1 r-'imil le4g) Tishe1l-esfteegxmoepondtietcd* oafntdhewi5tfhiftah WA• &.fty ')>-( \3Mf I on 1 ;_-inal spade-like terminal profusion Cta.WA }iu\i, I''- i bearing denticles (Fig. 1B,C). hi a , , , ., t;WA ,. .i , on Bayly. 1984 NEW CALAMOECIA FROM THE MURRAY-DARLING BASIN 787 C. lucasi, i.e. now C. baylyi, is markedly larger COMMENTS ON OTHER CALANOID than C. lucasis.s.. The same applies in the Paroo COPEPODS OF THE PAROO where the 2 speeies are sympatric. In Lake Numalla(CurrawinyaNational Park)whichhasa Boeckella fluvialis lives mainly in waterholes associated with the Paroo R, and occasionally in perennial population of C. lucasi s.s. a mean lakes (e.g. L. Numalla) and pans fed by Paroo female prosomal length of 0.68mm has been floodwater, all in the Currawinya National Park recorded from measurement of 30 individuals (Timms, 1997a). It also occurs in waterholes in (author unpCublished) compared to 0.94mm the lower Paroo area. In all the waterholes it was (Table I ) of baylyi in a nearby claypan. most common followingmajorHows,reachingc. 5-12% of the total crustacean zooplankton. In Eficeoslhowgayt.ersC,. pbaratyilcyuilalrilvyesinivnertyurtbuirdbidtecmlpaoyrpaarnys LakeNumalla it was restrictedtothe Coomburra WA Waterhole that receives Paroo floodwater and (as in six sites in the Carnarvon region of soon disappeared when the lake returned to (thHealPsaeroeot)a.l.,In20t0h0e),Paarnodotihteofcocuurrspriensetnhtesseitpesanisn pneorrsmiaslt.edIfnorthoenl2y pandsayrseadcehspeidtebytheflpoaondswhaatveirnigt soon after filling, passes through one generation water for many we1eks. Though B. fluvialis lives and disappears alter a few weeks (Fig. 3). in ponds and lakes elsewhere in its distribution In the Carnarvon region it co-occurs with C (Shiel, 1986; Timms, 1989; Maly et al., 1997) in Canberra or B. triarticulata, but not with the the Paroo it prefers riverine habitats. similarly sized C. halsei(female prosome length Boeckella robusta robusta lives only in turbid 0.81 - 1.00mm(Bayly, 1998)) which also lives in claypans. but there are <10 records in the Paroo turbid claypans (S. Flalse, pers. com.). Further probably because it is short-lived and is not work is needed to determine whether C. baylyi common when present (typically 1-3% oftotal and C halsei are competitors or have slightly crustacean zooplankton) (Fig. 3). Its presence in differenthabitatpreferences.Thecollection from NW NSW and SW Qld (QM W25486) adds NSW near Lake Grace also had B. opaqua and B. considerably to the 3 localities in and robusta maxima (Bayly, 1984) while Halse's Tasmania listed in Bayly (1964). Bayly's (1964) collections from the same locality containedjust contention that the Tasmanian record was B. robusta maxima as well as C. baylyi (S.A. possibly a misidentifieation is supported by the Halse, pers. comm.) In the Paroo other lack oi' any records in a state-wide s&tudy of Tasmanian caianoid copepods (Walsh Tyler, co-occurring copepods include B. robusta zrueopibdutlsoetrai4,aBsn.pdetcCriieasCratcincbouelmramrtaao,,nBw.i(tHthiamncmocs-oioc,ckcCua&rlraeTmniocmecmscsio,af bTr1ohy9be9Sur8asert)f.saoFr(use1rh8tot9hhu6eel)rdkmpornboreobewamtnbohldedyiiSfhsyitadersidnnbeuotytowiloionbnceacoellfniutdByue.rrboeranconlobiyrusdseedtW.da unpubl. data). In all ofthese co-occurrences C. NSW and SW Qld. baylyiisamedium sized Calamoccia, largerthan Boeckella timmsialso inhabits turbid claypans C. Canberra, but smaller than C. zeidleri and all and has been collected even fewer times t—han B. tlie Boeckella species. robusta. It is known from only 2 claypans the Dairisdtrainbdutsieomni.-aCraildazmoonce.ciIanmboaryelydietoaiclc,uirtsisifnoutnhde tPyapnealtocBalloitoydwinooCdNPSta(tBiaoynly(,H'a1n9c9o8c)kan&dTTiumrmkse,y widely in the western third of WA from Port unpubl. data). In the latter it matured earlier and Hedland in the north (Timms & Morton, 1988), its existence was briefer than other copepods through the Carnarvon basin (Maly et al.. 1997), (Fig. 3). Itmaybemorecommonandwidespread, to Cue (Bayly, 1984) andnear Lake Grace in the but its short life and early appearance in pans sinouetashtweersntW(ABa,ylpyo,ss1i9b8l4y).thIrtohuagshnloatcbkeoefnhadbeitteactteodr msoeoannsaftiterisfirlalrienlgywchoellnecttheedy.aredifficult toaccess collectingeffort, norinnorthern SAand southern Boeckella triarticulata occurs in almost all NT which has been searched extensively by W. wetlands sampled in the Paroo, making it Zeidler (in Bayly, 1984). In the Paroo in common and widespread, as it is in much of NSW & northwest and southwest Qld, which has Australia (Bayly Williams, 1973; Maly, 1984; been intensively studied by the author for 13 Shiel, 1986; Maly et al., 1997). It even lives in years, it occurs very spasmodically in time and hyposaline lakes to e.18 g/L. Overall it is most space. common insitesthatarepartly turbidandwithout 788 MEMOIRS OF THE QUEENSLAND MUSEUM fish. Although not a characteristic species of turbid claypans (Timms & Boulton, unpubl. data), if they persist for many months, it maybe the only species present, and certainlytheonly species of Boeckella (Fig. 3) (Timms, in press). C. Canberra is also common and widespread throughout the Paroo, as might be expected from the distribution map given by Bayly (1984). if prefers turbid waters and is particularly common in claypans, even if they Dayssincefilling persist for months (Fig. 3). Though present in other FIG Fluctuations in abundance of calanoid copepods in Turkey Pan, lakes and river pools, it Bloodvvood Station during a "normal' fill in April-May 1998 and another disappears in favour of C. prolonged fill in 2000 (mid to late stages only). lucasi when waters clear and does not survive should and Killanbirdie Waterholes 4 times over 13 salinity increase above c. 5 g/L. years of observation. All records pertain to C. lucasi in the Paroo prefers larger lakes and summer floods andnumbers werealways<0.1% riverine waterholes. often with less turbid water, of total crustacean zooplankton. These e.g. L. Numalla, Corni Paroo Waterhole, L. occurrences are well S ofits normal distribution Yantabadgee. Where fish arepresent it is usually (Timms & Morton, 1988) and must be due to the dominantcalanoid copepod, as Maly & Maly some individuals being washed down from the (1997) have observed. It also occurs in smaller northand surviving fora limitedperiod. Halse et intermittent creek pools that persist for months, al. (2000)alsohave foundthis species somewhat but not in turbid claypans, less persistent small S of its known distribution across N Australia poolsorhyposalinelakes.Thesedataconcurwith (Timms & Morton, 1988). those on this species elsewhere in eastern and cceontmrmalonAusitnralliaargewrhepreermitaniesnwtidweaslperrebadodiaensd ParGoloa,diwoiftehrejnusstspoinneosruescorisd einvetnherasruermmienrthoef (Bayly, 1978; Maly & Bayly, 1991). 1995-6fromOurimpereeWaterholeinCNP.This is the most northerly inland record for a species C. zeidleri occurs throughout the Paroo, but that normally occurs in some coastal freshwater only in turbid claypans, and then mainly in the lakes (Timms, 1973) and estuaries in SE early stages offilling. Its presence in the Paroo AustraliaandNNZ (Maly& Bayly, 1991)andin extends its distribution from NE SA (Bayly, floodplain rivers of the Murray-Darling Basin 1984), and a collection from a claypan near (Shiel, 1978; Shiel et al., 1982). Windorah, Qld (author, unpubl.) suggests it is Co-occurrences are common among calanoids distributedeven furtherafieldto include much of intheParoo. Themostnumerousarebetweenthe the central and eastern aridzone. Its presence in larger B. triarticulata and the smaller C. suitableintermittentwaters,alongwithC. baylyi, Canberra or C. lucasi, as noted elsewhere in C. Canberra and occasionally C. lucasi, provide inland Australia (e.g. Maly & Maly, 1997). further exceptions to the observation that Claypans regularly have multispecies Calamoeciagenerally inhabitspermanentwaters (Bayly, 1978; Maly & Bayly, 1991). complexes, with the relatively large B. robusta, then in decreasing order ofsize, B. triarticulata, Diaptomus lumholtzi occurs rarely in occasionally B. timmsi, C. ziedleru occasionally waterholes in the CNP, specifically Ourimperee C. baylyU and C. canberri Bayly (1984) noted 1 ' MA\ CALAWQECtA I-KOM THE MURRAY-DARLlNd B thatC, zicdlcn co-occurs with eitherorbothoi'B. lakes in south-westcm Australia Journal iriartiailuia and Q Canberra] they are woil Royal Society ol Western Australia 84; 5-11, separated by size. Riverine waterholes also halsh.Va .sum ,R i STOREY aav..ldwAS occasionally have many co-occuring species D.ti.D..l.A\SrU RVI.A \I.l..DJ.&H\KVIA. which may include uny comhmalion qf B. M.S., -000 Aquatic invertebrates and I , irutr/imlala, D. iu/nhullzu G spinosu.s, B oi wetlai ! ani 1 1 cfs ol the southern Catoai - fluvialis, C Canberra and C liuasi. In both Austirna,liWaensiWeurncAuumstrSluipa.plReemceonrtdNsoo.f6t1he2W1e7s-2t6e5r.n hhaabvietatbseeunp steoe5n SCpoeecxiies?tinagn.d Sofetmene 3a-r4es<p1ee.i3es•,- HALSL\,i|iSi..Aii.M,-KJi.nMvIeirl.tIe.btrYa.te\sMioMfAVLIaSk.e WG.:rDe.got9r9y», other speeies in si/e and hence in possible bwcsieni Australia, in relation iu salfnit> and competition(lluichinson, 1951 >,butthesituation ionic composition. Ilydrohioloeaa 381: 15-29 is not persistent, as fl tuwvsi develops earlier HUTCHINSON, Gil 1551. Copcpodologj lor the than its eongeners in pans, and other Oimihologisi Ecology 32: 571-577. co-occurrences of Similar!J sized speeies (/'. MAI Y, 1'..!. 1984, Dispersal abilhy and relai,,. friorticutata ID, lumhoUzi />' fluvialis and (' abundance of Bozcketla and Calamo canbzrra '(.'. lucaxQ are linked with temporary (('opepoda: Calanoida> in Australian and New disturbance by flood want Zealand waters. Qecologia (Berlin! (>2: 173-181 ACKNOWLt IX,3EMENTS mm VhJ.cV:B\vi.Y. i.a.i 1991.FactorsMucn fcn blogeogcaphic pattern-, ol Au.slrala ian 1 thank Ian Baylv tor encouragement in this c1e8n:t4r5o5p-a4g6r1d.copepods. Joi ral fi :: ognipli> psrpoejceicmtenasndfrStoumarWtfelsitaeitsne Afuorstrmaalkiiangandavafiolrabhlies MAIMpetiKt.i.Io.n,&,anMdAeLoY-oeeMur1r'eu1e9e9s7o!I'rBetdw.icnku&i,i,fa an|d comments on the manuscript. Caiamoecia ((. opepoda; I nlnnoida) in Wp HUD AusKalia. lydrobiologia 554: 41-50. LITERATI :RI M\I\, J. I1I.M.SI, S.\. & MALY VI.R !4*)7. I BAY!gVe,nu\sJi£C.ulu1r9i6h1,Ku'AiilevKisojopncpo.iultah:e in'laalnadnoulaat i I >iarihnlion anadndineiiidcemniehefKp-actktceIrinsuo(fCBoopccckpcotdlo, t ). i / i . > . iliari i al ij '-.i, r* l ; ; i ihwatcr ('alariividaj in 'A-stern Australia. Marin.' i I Research 12:54-41. i i ,hwatei Rcfiearch 48: 615-621. 196(4C.-o.IpAcrp.eoivdeiiasigoenCnaoelira'anthoBciodAeauc)s.kterUAaaulsuatsiriadalnhiasnpeJcoiuersnLailt'thoel SHIMRMuu.yrRar.laJS.yuc1TI9e7t8j.I oZtoVoipSelytasontrkeitmao,'n)()P:croIomVc3me-ue2nJdiJi2tn.igess oofl tdhiee MarineandFreshvi i1 I >searchl5: 180-2 -x F9K6 /.otiplanklon orthe Murrav-Uarlmij - ]('?.N..ii.V.a.rii'aiitjioncionpseepxuoadlsdimaonrdphiistsm iencnoolnorngaircianle oIl'*ArlitveeriesvysItVeRm.scV.:(WJaulnkke:r.Dokr.dhre'c(hetd)s.>Thece. , i I22i4r-f1fc2a&n.cc Limnology and Oceanography 23: SHIM R I WALlvHR. K4. & WILLIAMS, W.D 198i4.alaAnonideaw) fsipecieSsouotih CVutattartmtoljetiinind[Ccoopmempeondtas: F.1Ar9ue8ss2th,rwalaPitalc.ainkARtueo-snaeraarolicibain1h3c:l3oI0uolrw!m-ed3i2 oMfurMr.aiym,,,Souritcil ''otSon''ialt"1harneoeu'lieaiSM)ioeueotnl.ehi!r:I-lnU.le.aierITtnrcwlanointrasralocdp1ta0ig8o(:inGdsuaIc4io7dri-etTts5ho4epIedpRoodytaah;le I AIMLlS.o,iwneureHaAnl.lSu.onc.t.i.1ee9i.i8v9V.aollleftNs,.S' W'Nu;SdWi1v,1..mt,hOPiw2rj9oti7chk-ee-3er0dh6Cio.ndMgn1',Mo-li ,Llhi ciodcnnitiinleneiaal *waotfersCheofiiiubcerowinovrelrdtebNtor.ate2s),oifSPtIhle 1993. Saline lafo al tl Paroo, inland New South u ademic Publishing i he i fague)* W'ales. Australia I ^ydrobiologja Z67: 269 R^. 1996. [niaod-watci ci Did copepods oi [997a '. Stud (kinds of the Currawfnya Kangaroo, kiiir and Minders !>!; National Park. Lnpuhl. report to theOueensland Biogeograpty iiit| ! colog) transactions of| tIhe Dcpflftmc \\ OfL.iuironment. K>ow(Kimha 199:8'. New nspeeJie.-sI SoofuCtuhLAauusotrcacliiua.a1n2dU:B"[i-6. [99\7\be*ll\ancdosmpoannsBolnOboedtWwOiOdisSatTairlnioui,idtfhreeslPiawnater I fre--.liv.alerCopepoda' Calanoida) 1mm Western '.' ii.li.i. with special relerence lo iheir use h\ ...lii.i ,\\m.\ Queensland Journal ii! th< Ro '! [crbirds bnernational loumaJ od Sail Lake Society of WtesternAustralia81 177-182. Research5: 287-313. i BAYLY, I.A.I & WILLIAMS, W.D., 1973. Wand 199^. Ysiud> ofl akeWyam an o Hi aliy filled KAI SI H. s! anraheMirceReoAlIog} i(1Mang2r0n0a1n:MCealiboaumrcne). s[anlliinnenllakme \i\n ISO"U!ith|wilcsill Osu-<elienLsa|lkaend,ResAenasilraha .'. ht,'!a u-sp '' &pi poda; I alanolda) fforti tall 790 MEMOIRS OF THE QUEENSLAND MUSEUM In Press. Limnology of claypans of the Paroo, WALSH,R.G.J.&TYLER,P.A. 1998.Reconnaissance arid-zone Australia. Verhandlungen Inter- limnology ofTasmania IV. The distribution and nationale Vereinigung Limnologie 29. TIMMS, B.V. & MORTON, D.W. 1988. Crustacean ecological preferences of tasmanian species of freshwater calanoid copepods (Crustacea: zooplankton assemblages in freshwaters of tropical Australia. Australian Journal ofMarine Centropagidae). Archiv fur Hydrobiologie 141: and FreshwaterResearch 21: 11-33. 403-420.

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