Branch, Shine. Harlow & w€bb 89

Or i gi naI con t r i bu t ions of UerpctologJ

Sexualdimorphism, diet and aspects of reproductionof thewcstern keeledsnake, P1lft onodipsas carinal.r (Serpentes: Colubridae)

W.R.BRANCH*

Departthenlaf Herpetolog/,Part ElizahethNl|se nt, P.O. BoxI 3 117, Hune\rco.]601 3, LasternCapc, tbutll Atr icd

R. SHINE, P.S ITARLOW and J.K. UBBB

SchoolaIBialogi.ll Scienc.esACa, The Universiy ofsytlnev,N.S.|Y. 2aa6,Austmlio

To rrlrod corespondcnccshould bc addrcss€d

BRANCIT WR., R SHINE,P.S. HARLOW and lK WEBB. 1997.Se\1lal dimorplrisn. dicl ard aspe.tsof rcproductionof thc w€stemkeeled snakc, Plrr o adipsas.arinata (Seqle. es:Colubri&e). Afr. J. Herpelol.46(2): 89-97.

Ewntaiiot atul dittectiM oI hnls.wt spcinas prc\,idal onginl data o" noryhalog', s.tul rlintaryllisn\ f.e.ln1!! IabiLt ah.l reptudrctiw biobgt of k. lare wskn k?e!.d s4ka elnonodjpsrs 6inalallo,, lla,ibid drd 4ngalq. lhis snrall, sle,t1l.r hake is ,@tut tal. FqaLs Mhre at a larAet size (47-51 m tuu|wit ldsth. SI/I-) tlnn nnl6 Q9-31 cht SrL), d1,1 dtbh c$idembly laryer winlnt sies (to 77 c SrL). /lt tJle M e bdt lensth. Mlc: Ituw siqnifi@r 1/ldger toik l@rfenales. Reldlite 10 St/L, Mlysis oJ cMtiance detect.d rc sianili@tt dillmcT betn@ the sM h h@d laStJt, h@d vidh, bd, \'i.Ih. or [email protected] shaPe 0..., widlh relativ. to lot811t, d ftlaht ete siz). Atl prcy irds wre tenebrtles. with liads conyisnts 67 dz aI nt diel. tujents 25 '/q bids 4 %q ar,l a/a) unidennf..l vnebtute bd14 4 %- Andta tih'd pfer, aak6 (39 % of o ptey) atd sknts (t8 ye.l@'vrated. Noctumt 86kas f/froedua Chondrodaclylus PahEtog€ko ard Pachldaclyl$) as NEll 4 diannl otps fa\opltopB url Naru.Lsia, wre cate". All skink prc.v wre Mdttya s7. ManMlian pfty wre qte,' pri"nnly by lar4er sndk s. nE ur1&lal fnsnlnted ltea.{ shieds ak.l a'la,Ead paldtirc leeh of\a\6dJjilr9s.eii?,ia na.y be adaplatias nt rymtit shepr\ dtu.sl ptq 10 be.fltact.rlIftnt ret@ls.

Introduction Bmnch 1988:Broadley 1990) or rcSionalsuncys of hcrpc(ofaun(Bauer el al. I 99:l). llcrc arc lt\t rclcrencesto Thc \\'cslcmkc.lcd snake.I'rthono.lipsas cafi1ata, is a r,jre dnss?eci€s ir) tlle lilcrdhrrc. back-fangedsnak€ restricled 10 gmvcl plains and and Girnficr's (1868) 1]?e specimcn ot Plhonadip&s "found savainai ftom llrc Bctlunic Dislicl in southemNamibia lo cdrirdra $as allcgedly b] Mr J. Chapnan on drc extremesoudr-w€stem Angola (Broadlcy1990). Thc genus Zllnb€ze', but Boulenger(1888) noted a sccondspc.imen is nonotwic and of wcertain la-\onomicaflinities. It is llom Damandand. The sp€cies is nov kno$n to be unusualamong colubrids (serl.q! lato) in possessinggready rcnricted1o the *e(em arid regionof soudEm Alric4 ard fragment€dll€ad shiclds,a pcculiar na\illary dentition(3rd to bc absenlfrom ttrc Zmbezi drainage area (Bro.ldley to 6th la4es! posleriormostloodl large and gooved), a 1990). Ar Chapnan travelled fionl Wai\rs Bay on his reduced nMbcr of paladne tl-dl (including a gready joumev (o $e Zambezi.Broadley (l9ll) reslrictedthe 9p€ enlarged"fang"), and hFapophysesdeveloped throughoul localil) to Darnaraland.Lilltc is kno*n of tfie biolog/ of lhe venebralcolumn O4ani €t al. 1982;Sch:i(ti & Mccarthy Pythonodipsascarinatd exc€pl for anecdolrl obsefvatrons 1987). Thescfcatrnes have stimulatcd invesligations into irs (I-ankesl9l4: Falk l92l; Stuan1976). Perhapsconnlsed pos.rblcqaors as a prolo\ipcr"ti e. rc{mblng a putau\c by tlrc original q,pc loc2lry. and ir uencedby thc unusual transidorl]rlfonn betwsrn colubridsand e"?rryvip.6 Ma^ largc. up$ardl]drcclcd noshls of tltc s?ccics.Slcmleld cl al. 1982).Exccpl in phylog€nelicin\esligalions and i (19i0) rnistakenlysuggestcd dn1 d)e sp€cieslives relr natoml (e9., Mc(ens 1955, l97l; Fitzsimonsl%2:

Alr. J. H.rpetol. 1997.46(2r:a9-9j 90

Previous souces of infonMtion on th€ habits of South A.fiican Musem (Capc Torur), Naftib Des€n Pytlnnodirys carinata ltave come prnnarily ftom Reserch Shtion (C,obabeb),Slale Muserun of Namibia obsenations ofcaprivos?ecinens. l$kes(1914) notedtlar (Windhock) and Glifomia Acadcmy of Scienccs(Sar) a capUvespecimen of,3,/r onodipsasc.vinata \\"d tocawffIat Fmncisco,USA). and that during daylight lDurs it often hid under bark or Prcservd sp€cimeNwerc nreasurcdand diss€.rcdas sloneson sandin rhe ter.fiunl Another caDti\c sra c $iA folloNs. We recordcdsnour-ven( lengrh (SVL). lait l€ngttl fed a srnall. lnparric skrnt- r6r|,ely Mab;ra otu abt^. hcadl€ng1h (from posicriormargin of th€ towcrjaw 10tllc which it bit two or ttrire times and shookbefore releasing tip of Ure snour), head ividth (at ttrc widesl point). cyc (FaI( l92lr. Ti'c prq \!"s subsequcny ingesle4tcrdrng dramct€r.body widlh (diamclcrar midbody)and bodYnrass Falk ly2ll I lo inlcr tlLlr ljle snaleposscss a \r€ak venorn tlle sp-lmcnsr\crc opcncd $rrh , ILidlcirrrdtrncrsron, ,Id Ho*ever, Marx er al. (1982) recordedihat l€se small any pr€y itcrns in ttle alimenrat, crnal (including drc snakesconslrict lizzrd prey, wtich continuesto struggtcfor hindgu0 ritrc rcmov€d lor later idcDtficarion ald sometimc after bcing sciTrd. Tlrcse obseNations$ggcsl measur€rned.Sizes of rclatively udigcslcd prey (tcnglh. Unl encnonution plays lirdc parr in prcy imnobilizarion. $idtl\ nuss) \€rc dclcnnincd direcdy, aod reconsliturc.l OllrcrobseNations(Branch pers.obs., Haackc p€rs.co m., s7ls of morc fully digcslcdprcy ilcn$ $€re dctcnnincdbI May 1996)on feeding in alplive specimcnscor irn thoseof comparisonwi(h snDilar-sizc4 inract specimensir thc Marx (1982). et al. The oniy rcccnl reponon lhe biolog/ of coll€ctionof the Pon Elizibe& Mus€ur Relalir€ prcy wild s?ecimens is dul of Sr an (1976), for l}ree snakes na-rs(RPM ofa gut ilcm \l"s calcutarcdby crpressinSjls from thc Namib park. Des€n Two of rhesecamc ftom nass as a percentaScof the snakc's mass (minus rhc prey spaisclyvegetate4 arid areas,whitst the third $€s found in riverinevegetalion nerr s€mi-pem)rncntpools in the Kuiset) S€x and rEproducrivc s1atus of the snakes $erc Rivcr T\ro sp€cimens had eare4 one containingan adutr determincdby visual ineection of tlre Ilblcs rl€re gs:ko (Pachydactylus : p. Sonads. bibronii taqigatus) and rhe mnsidercdnuture ifthq had enlarge4 turgid reslesand/or olh€rtwo unidcntified small rodenb rvhile. thickcmd effercnl duqs (ndicating lhe prcsenc€of Mar\ (1982) ct al. drcw allenlion Io drc slrikinS slrrm). Fcmalcsrverc classificd as Duturc iflhey tradthick- morphological convergcnce bctB€cn ttthonottipsas, musolar oviducls. vitellogenic ovariaJr folliclcs and/or Spalercsophis (a Nodr Afiican colubrid also luving oudrrc€l eggs. Somemature snakes \irh regresscdgonads fragmented llc€d shictds) and vip.rs. Th€se aurnors rnaybe misrakenlyctasscd as inunalure. Additjonal daraon conjecturd dlat lhc unusualfraginentalion of he.adshields diet and reproduction$€re oblainedfrom the lirenturc. and in vipen, boids and Ihe co|ubi'd g.nen Spdkrosaphisand from personalcomrnunication may promolc rykanodipsas rhe wide gapc tqlufto ro Slatislicalanzlsis follows ttut ouUincdfu Shineel al. engrnf hrge prey. 'Ihey also srr8gesteddral the edargcd (19%a- c), afld involvcdt$o-iailcd unpairedt-tesrs rc assess h^F;s in Wlaline Ptthonodipsasmigh fircrion to r€strain the signficancc of se-\ difiereocesin mejn trair \,ztu€s. geckosdlal are prone to escapecapture by rcaringrhcir skin. Mea$rements of mass werc loA-transfon€d prior to \,farx et al. (1982) made ptea a for more ecotogicrj analysisto aclieve nonnrlity of dislributioflsafld €quatilyof inloinatio on Pythonodipsasto addressthese problems. vanances. To cMrnine dillerences in body proponions Subs€4rndy,Scfiard & Mccadry (1987)denorEtiared dlar bcM€en drc se\eq we us€d aialysis of co\.ffiance and the va.rious morphological similaitics be$ccn helemgencityof slopcstes1s. Contingency-table (chi-square) Sllktosophis afi Ptrhonodipst reprcs€ntconvergence. aralyseslrere usedto test the signifcance of obseNedsa\ Wemer(1985) also proposed an alrernarivch!,pothesis for differencesin dietarycomposition. An atpha te\€l of 0.0j h€ad shield fragmenlation Sparercsophis and 11?sused dnoughout. Ptthonodipras, srggesnng $al, atons wirh dorMl colouration and ot]rcr fea(urEs.they fonned a su(c of aalapulion5for Balcs€rr n rtucD of y,rp3rn, \apc6 Results Resolutionot thescconnicUng h)polhescs is harnpc,€db) tle d€a(h of information on the specics,biotogy_ In this Scxual dimoqrhism: We c\arr{ned 55 sp€cimcnsof contexqwc pr€sen( pJ,thokadipsa, here biological data tot Pfthonodipsascainota l,l:,fiuseumcollections Ofdrcsq ll carinata d.nyeJ ffom del2iled aralysis of pres€n-ed r€re Ju\€niles too ffull or poorlv preseNeo ro sex accuratel). The remaindercomprised 15 adult mates,3 juvenilc males,15 adult felrr1lq ar)dgjuvenitc fcmales. nris spcci€sdisplays considemble sexual dinlorphism Materials and methods in bod), size and proportions. AduI( fcnules atain sig ficandylarg€r sizes tlun adult nratcs. For nlosrof thc Informadonon rhc disrnburion Inrphotos/, d,er and trails ir€ measured(snoul-venr lcngrb he"d lengl\ head Rproducf.r\€ brofos/ pr,thonadfsa, of was bascd o \vidrll body widdl body mars). rhe ditrcrcncc i mcan e\aiulrauon ofpre.scncd specimeltIin the co ectionsof the \alucs bcllve€r adult nr.llcs and adull fcrnales N,as Transvazl Mr$eum (prcloria). pon Etiabedr Museurn, slali9icallJ.,sigoifican{ di.lpir€ our relatively low samplc Bmnch, Shine, Harlow & Webb 9t

Tabfe l. Body si?4sand so\1ralsize dirnorphisrn in adultPl,lro nodipsas carinatd. Meanvalues for €achtrail. with slandard deviation in brackels,are slrcwr. The tlro right columnsshow rcsultsof unpan€dflo-tailcd t-resls and slatislical significanceof lle direrencesbetivccn (he sexes. For mass.untransform€d dara sivcn in tablc. but t€sr are bascdor In-lnnsformedvalucs,

TEit t (28dn t5 l5 Snout-venllerelir (on) 38.49(6 99) 54.5r(10.43) 160:l < 0.0001 29.0- 5l 5 3A5-7?\) Tail lererh (m) 6.60(0.99) 6.40(1.r5) 020 0.62 Rmge of lail lo€Irs 5.0-9.2 4.9-84 t8 ?A(2.51) 25.09(2 88) 625 <0000r Rarge of fi@d ldgdrs 15.0,230 19.6-294 FIod !$ddr (mn) r0.32(20t) t4 64(2.71) 412 < 0 0001 Range in hod lvidLirs 0- l0 0-5 Mars (s) 246(r93) 55 I (29r) 346 <0.003 8-65 19,t0 Bodywidlh (m) r0.25(2.80) 13.48(2 6l) 323 Rll.lqe in bodv wiClhs 0-i0 0-5

T^ble 2, Dier ln Pythonodipsas.drraad. Perc€nrageocclrrcnce of identifiedprcy shorn in brackets.N = 28

Prey q."e No Vertebmtebone (3.6 %) Reptilia Lizad bone(1.6 %) Gekkonidae(39.3 %) Arwdutu alia@ 2 I Bouleuad, I 888 Ch dd niacty tus d,su Iife I I I I I Pacb.daclytu q. I I Pdclrdactyl6 bibrc i (= P. ladisat8) I I Stun, 1976 Palnatoaeckofrngei I 1 I l Maprropa bipotts I l 2 I S.incidae(17.9 %) Mabura a.utilahns I Fdt, t92l Mabroa st,iata whlbo\i I lr,labUa sd ab po1. tati ssi,n I I Lac€rtdae(3.6 %o) Pediopla,risatunl6i I I Asmidae (3.6 %) I l Sl'isl- 1980 Av6(3.6 %) i I Mttnrli^ Q5.a %) 6 5 t ctJqt fin (Ta1 e tu Id. %6 | er) I I 5 '7 Bnpty t9 Ah J. He.p.tot. 1997.46(2):89-97 92

0.09; inlcrceprsFt,29 = o.a2, P = 0.3?), or body $,id r relalivelo S\,L (slopesF1,28 = 2.48,P = 0.13:inrerceptl F1,29=0.41,P=0.53).The rateof incrcasein bod) nriss wilh incrc"sing SVL \r"s higher in maleslhan in femiles (slopesF1,28 = 7.03,P < 0 015)but thispaoem is diffortr 1ointcrprcl, as tlle two regressiollines crossat intcnncdial€ SVIj Cigure lb). We also exanrincd possiblescx diflercnccs in head dimcnsronsrn fvthono,lip&s .nnnatn. by ca].rtnE oul o ,na\ses of covariance\vifi hcad lc glh (ralher ilan SVL) o females . as thc co\"riale. Theseamlyscs revcrlcd no significint se\ males dillercoc€sin headslup€ (widdr relati\e to lcngrh: slopcs F1,28= 0.03,P = 0.87;inlcrcepls Fl.29 = 0.02,P = 0.89)or in relalivc eyesi7, (e)c diamclerversN hcadlengdr: slopcs 20 30 40 50 60 70 80 FI,28=0.75,P=0.39;inlerceptsFl.29= 1.16, P=0.29). Onc of rlrc specimeflstlnl $c cxanJnedmNidc.ab\ snout-vent length (cm) exlends rc nma\inlnunmbody si,a kJ.o\st [ot Pythono.lipMs .arir?ald. The prcvioosrecord \ras f.om Fitzsinons (1962). lvbo gavea m.ixirll]jn sizeof 550 S\{- + 75 tajl = 62j nxD lolal for a fem.d€specimen. Thjs is increasedconsiderably by a femalcftom Goarkontes,S\r€kop Riv€r. SNakopmund (IM 41632)widt measuremen.sof 770 + 8l = 851 ln]n. The largeslmalc fIM 69260i unkrc$n locatiry) me.:tsurc: 5t5 + 92= 607mnr E Dict: Of58 snak€sexamined. 33 (56.970) had gul contcnts. of which 20 had identifable prey items. Five mnrarned snull insect fragments (pr€sunably secondarilyinge(€d with vert*rnte prey). and nine had ing€stedfangs and/or unidcntifiablematerial. T\ro prcy it€ s Makrya stiata punctatissina)in th. san|.egut nust Iu\€ beenacceptcd in €"pnvily, 20 30 40 50 60 70 80 be.ausethis sxbspeciesof skink da€snot occu ir sympaay wilh Pythorcdipsas cad,ara. Tlrc cobabcb snoul-vent lenglh (cm) colleclionyielded an addidon l rccor4 ailhough tlrc snake itselfir"s not availablefor e\"mination. The snake.$,lich @ntzjn!,Ja \ery l^ee Palntatogeckorangei in i\s gnt. \\as Figurer. Sqtterplotofse\lul dimorphis in $e q€st@ keled colle.ledat the Hom€bsills in 199l, and found ftesNy dead *\ake,Frkd@dipw en,ara. Mal6 (dors)have retarively on lhe groundduring the day. De€ils ofa funher fi\€ prE, lo'sq tails(a, upper gmph) ed hieler in(rc onbody Ina-\ itcnrs *€re availablefrom other sources(Boutcnger 1888: wi1h ir@sing SVL O, lolq gnph) rha do fenals Stuan1976. l98q Fa* l92l). (circlcs).S@ ten for st2tislicalrBts. Combn:ingdrcse records wilh tllc infonnauonuul rc oboincd from our dissections, il is appare dxlr hlhonadipsas cdinata lakesa la\onotuically di\qs€ arml. siza Clable l). This r€sui( refle.ts the magnitudeof rlrc of prey ilcrns. Table 2 lists rhc items rlur havc bcen dilTcrencqeg. adult fcnul€s weigho4 on av€rage.more identiliedfrorn alinenary facls of rhis spocies; rc nunrber than toice as much as adut( nrales(Iabte l). Tail lerglh ofguts involv€dis shownin brackels. With the exceprionof provid€sa notableexception ro this generalpatlerq being iniecls, which were probablysecondzrily inges0ed (5), att \€ry similar in absolutelength in adult mal€sand fernates preyitems in the gutr e\amincd1l€re vertebrates (20). Wirlr (Iable l). This appar€ntaionuly is due to a sexdiference the inclusionof additionalrecords (8, including thosetaken in proportions;males have longer rails rhan femal€sof ihe in captivitJ),liards compri$d 67 % ofdrc diet, wilh geckos samesnout-vcnt lenglh (Figure la). A hererog€neityof (ll-39 % of all prey) and skinks (5.18 %o)predominaring slopes test confinns fie slatislical significance of this (Figure2). TI|e forner includedbolh noctumal(,4y'oedra diferencc (with se\ as the frcror, S\tL as lhe covariatqand Chondrcdacvus, PahMtogecko ad PacJrydactylus)ain lail lenglh as lhc = p dependentvariable F1,28 9.1?, < dinnAl (Rhoptrops and.Narudasia) Senera. All skinls 0.006). Amlysis of co!"riance det€ded no sisdficanl $€re refcrableto ,4rarrra, aldrcughdu€€ of dleseilelns had dille(-,,< trel$cer.:.trc s(\iis in h€adtcngrh ,ela!i;lo S\t bccr a..-"ptedin captivity. Rodentscor prised 25 % of rlrc = (slopesF1,28 2.26,P = 0.14:inrercepts F1,29 = 3.49,p = dielary items (7), bul prEsrunablycomprised a larger 0.0?).head width relarivero SVL (slopesFI,28 - 3 05,p = pmponionof drc toial diet in tennsof bionrassand ener6/ BrancL Shine,Harlow & webb 93

Vertebrate(3.57%) Bird(3.57%)

Rodent(25.00%) Geckos(39.29%)

Lizard (3.57o/o) Agama(3.57%) Lacertid(3.57%)- Skinks(17.86%)

Figurc2. Diet oflhc wesldr keled sMliq,ryro,&?arar.arr,/z.. Pic chd ndiatinS drcpccilagc @nrlDsilionoi prevclas$s ol,ll recor.lst food i{ens is sbolh 0r = 28). cuts @nr,n ,g god{i\'inecnd nr$cls tord uridentiliiblc dtlcr uc nor nrcluded. Gekos o,qrie thc ldgest proporlionof$e diet con$nnption(because they tend 10bc much larger thrn llrc lhereforeprobably nraturEbetrvccn 475 - 510 nlnt SVL. liTardsLlDl aJc consurned). Tlrc onl] mamtnalianprq ilc'n Non+r€€dingnulure fcmal€s$€re colleclcdin January0), tllat could bc specifically identified $ a bush\€:d gerbil April (l), May (5).luly (l) and Aug sl (l). Thescfcmales (Tatetule cogastet l). Odrerprey items included a drumal conlaincdno olarian folliclcs largerthan 3 ,nm in diameter. lacF'n.tdlizaldeeAoplanls gae,'deJ; l), a diufnd agamrd Thc smallestscn|ally maturc malo (llick cfel€nt ducts liza (Agana anchietae;l), lizard bonesand cla$s (t), a and/orspcrm pre.sent) had a SvL of 292 tnm (SMW 2310). down feafter from an urudendficdbird fl). and \€(eb.atc A male ot S\4- 310 mln CIM 55700) llad thin \lalled bones(l). Nine addrtional hindguts containeddi( shed ellerent ducls and riry tesles and nlay luve slill b€€n fangsanauor other unidentifiable materiat- imnralure. Thesc obscNatiolB indicalc thal male se\:ual RelativePrey Ntass@JN4) could bc deleflninedfor seven nlalurity is rcachedat SVL 290 - 310 mrn Mature nrzlcs intactpltry items (all lizards),and rangcdfrom 0.5 - 42.8o/o widr spennin llrc efierentducls rverecollccred in Mar-'(l), Excluding the unusual rccord of a hatchling ii?rudana June(l). ,nd October(l). trtvd (SvL 13rnm, mass0. I g) in an adult (370 rnn SVL, The smallestspecimens measul€d (PEM R8325 and rnass19.8 g) nalc snakc(RPM 0.5 7o), ihe a\€rageRPM SMW 2309) bodi had S\4-s of 160 nun. On€ of thesc tl9s 28.8 %. Four of drc six snakesconlaining rcdent srRkesri€s collect€don I June 19651tbe date for th€ ofier rcmarns $€re females. Snakes eating rcdents had an wff not recorded. Other srnall jwcniles (165 - 180 rnm a\€rageS\aL of 497 mm (Enge 310 - ??0; n = 6). TIl€ S\,L), all bearingprominent umbilical scars,were collectcd s akes conlaining gecko r€nuins (dehils of llle @abeb inMarch(l), April (l), May(1) andJuly (t). sn ke containing ^ Polnatagecko fauaet are unknown) includedsix nl.dcs,t\lo femalesand an unsexedju\enilc, and lud an averageS\4- of 330 nlm (range 176 - 394 lYlrn: Discussion n = 9). Aialysis of all wild lrlhona.lips^- .artlata r:har consum€dlizards increas4sthe al,cragc S\4- lo 371 lrrnr Pykalo.lipsas corinata ap@rs lo be a Senurnelyrare (rangel?6 - 617 fiun; n = l5), ]vilh eigllt nulei si\ fenulcs snake,ard knowledgeof ils biolog/ and drstnbulion ll?s andajuvenile. There\l?s no significantdillerence bet$,€€n been infrcquendy .evielved (FiDsinons 1962; Broadley rc so\esin the frequencyof ectothennicveNus endothermic 1990). C€rtaiflly,i1 is \€ry rarely encountenrdby hulnans. prey taken (chi-squzrewifi I df = 0.60 alter continuity rvith lessdun 80 s?ecimenscollccted for museumsduring conectioi\ P = 0.44), but there \r"s a significantdiffercncc the lan 130 years. The ertrcne s.2rcrty of Plthonodipsct's in meanSVL betr€en snakeseating lizards and drose€aling ear",raras?ecimcns in mus€umcolleclions enPlrsi.es the mdents(unpaired two-tailed t-tes1 (t with l9 df= 2.272.P = us€firlnessof &csc colleclionsas repositoriesof ecological 0.0349). da14and noljost ta-\onomicinfofinalon. Givcn the amount of time it would take to locateeven a single specimenof "conventional" R€prtrduction: Tlrc smrllesl female with (hick n scular Ptthonodip.ds carinala ia tlltr wil4 oviductsCIM 69260),indjcating rcproductvcmarurity, h2d ecologicalrncthods (e.9.. dclailen autecologicalstudies of a SVL of477 mm. Other fe|rales $'ith SVLSof 460 mln marked aninrals) clearly cannol b€ applied (o such rare (rM 56973)and 510 nun (fM 4611?)slill hadtl n-r?lled speaiesfor logislrcalrcasoils. Nonedleless,our cxarnination oviductsindicaling they nuy not ha.!r yet bnd. Ferlral€s of mt|sqtm sp€cincnscollecaed from a widc are4 over a Ah J. Herpetol. 1997.4614: a9-97 94

long pedod time and by nlalty di{Ierentpeoplq provid€s arl ScIUal dimoryhism: Tlrc rario of Inean S\tL of adutl oppommityto docurnent someaspeat of the bosicbioloB/ of fenule lo adult DtnaleDlrorodipsas carinata is t.4t6. ta lhis enigln ic serpenl recentmmpilation of dala on dris r|ni( anoDg smk€s irr PrrhohoApsascarinata is acrr/ known to be widely gencral(Shirc 1994)rcvealed rhal rhis vatucw"s exce.ded dislibuted in Namibi4 wilh a $nalt e(ension into lhe by or y 4 olt|er colubrid spccies(of 229 speciesfor wbich coastald€sert of southemAngola An updar.ddisfibution data arr available)and by oniy ninc odrcr sn*e spccic.s mapis sho$n in Figwe 3. Therc arc Gw r€.ordstom dune sludiedto date(ofa rolat of375 sp€cics).Hencc,lhcdegcc seas,and the slecies lus been mainly collecledon gavel of sexud size dimorphism in Plthakadip.tas cdinokl plains and in rockf outcropsin the D€sertand Nan]a-Karoo rcprEsenban c(rcmc casecolnpared lo lc Usud si(ualioD biomes(Irish 1994). It also hrs a timited exteruioninto add in snakes. savafiu in the r€stem rcgioN offie EtoshaGame R€serve. The probableevolutionary pressfcs ilvoived in sxch The scatter€dsouthem rccords indicate that it may oe more caseshave aflracted considerable spcculation from biologisls common in the r€gio4 which renrainspoorly collecr€d (King 1989;Shine 1993. 1994). Madscn & Shnrc(199s) (Bnnch 1994). Thereare few dcails of microhabitat.Sturrt have dcmonslratedthal fcnalc Europc.rn addctr (yipero (1976) r€co.ds specimeN from spa$ety vegetalc4 racky ,errr) evcncnce high coslsa-ssociatcd {ilh rcproduc{ion. arcas llom dcrchct buitdrngs. and fiom ncar semi_ includinB: lo\lcr surviwl and lowcr gro\\1ll rarcs of permanent pools in dry rivcr b€ds. The prey of reproduclr€ fetnaics: and reduccd rcprcductive rarcs Pythonodipsascarinata is fomd in a variety of tubitats, consequenton Oe postponenrcnlof subseqtrenlreproduction indicating that the snzke itsclf rnay atso bc a habitat due to dre liSh encrg/ denundsof o\'xtation and SeslatioD. Due to thesecosls, female gro!\di aier maruradonslo*s dramaticallyand thereis .I)usselecrivc pressure lor a fenalc lo alelaynuturalion u iI shc reaclrcsa siTr where hcr first (pertaps,only) clutch is largc. Sucha s.cnarjo nlay evtain fhc la€e bodysi/a in feDEIe4rhanodipsd.' coinata The hugc dillerence ir mcrn ad tt body sizes nriglrr alxo have be€n afecl€d by, or have consequercesfor, e.ologicalditrerences betrve€n lie sc\c.s In scvcralhighly dimorphic sn ko specier ro sa\es have been reportcdlo feed upon difi€renl prey D?es and/or foragc in disererl habitats(e.g., Zinner 1985;Shine t993). Our dieral-adaLl suSgesrdla| the saine kind of phcnomenonnuy occur in Pltho@dipsascarinata also,widr the largcr sex (femrtcs) tal.jng a disgoportionatenumber of largc marunalianprery. HoilEver, our sample siz€s arc too small lo enablc a quantitalivctest of this proposilion. Th€ e{reme fcnrale+iascd djNorptism rn Prthohodipsascarinata alsn allows us to make some inferencesabout mating slsrerns in this laxon. Conprau\c ana.Dseshave identified drc evolutionor lossof rule-Inale combatas a major inllucncc on rhe dcgreeof sexll size drtnorplsnr rn eul6 and no sp€ci6kno$n lo engagci[ male-nale combal displays as crlreme a fcrliaie-biascd dinorphism as that sccn in ltthona.ltptas cdrinala (St]|!rc f994). Hencc,we doubltltat trJcP) prothp&s.otr,a!,t enSagcin combatbehaviour during the maringseason. In conu:as1to tlrcir c\te'ne dispariry ir absotulebody $zq m2l€and f€maleP//ronodipsas cainata are sirilar in body proportions(relative head leng \ etc.)_ The only morphological\,€riable dut sborved dillerent allometric trajecloriesin malesand females\r"s rait lengrtrrelativc to S\|L (Figure la). Sex differences of this kind arc wdes?rEad(bu( not ubiquitous)among snakes,and might r€sult either lrom scleclion for longer tails in nales Co accorunodale le hemjpenesor to enhancemrle successin counship) or for shoner tails in females (postcrior Figure 3. Dislriburion nup of the s€slern *€elert srake, displacemcntoflhe \,cn( lnay providemore abdominat space Iwdodip% @nMLa.. l](l.s r?res€nr quar€.{€gre for eggt (King i9irt; Shine l99l). The significant squrs tom $tich speciM h'!e ben eoded. di\€rgencein refativelail lengtts in ryrhonodips8 @tinata Branch,Shine, Harlow & Webb 95

is consistentwith a gcncnl trcnd docrlment€dby King wiillin llrc Colubrida€,and drc unu$al cnnial f€atrrcsmusl (1989)and Shnc (1994),namely tlat se\ divergcnccin lail thcreforebe convergcni. lengtl relative lo SVL is mosl pronounc€din snakelaM Of tie thrcc s{'.{,l€s ot LtcoPhidion rccorded fronr with slrongsexual sizt dirnorphismin badysize. Namibia (Broadleyl99l), or y drc Namib i{olf snakc (L nonibidnunt) signfica ly or€rlaps the .al8e of 'Ihe Dica: prcy ot Pythonodipsatcarinota includesbolh Pythonodipst .ainata. Tlrcse t\vo s?c.iesoccur in *'ide diuinal and noctumalspccies. tllat rrl2yalso be lerreslrialor S,mpatryin no rcm Namibia (Fig re 3; Broadlev l99l) rupicolous.Of rhe latre\ AIroeduru dficana, Nandcsia and appcrr to inlubil simjlar rocky silLr.'tlions.Anong uolf SVL Iesth'a An.dRhoptropus biporcs$ arc slridly rupicolous, snakes.L nanibianum is of avcragesiz! Cnir\ilnufi whilsl R al/ is rupicolous,but preferssheer b€drack and 460 nrm), and femalesSrow larger dun males(Broadrey ofien retreatsunder low rock slabsat night. Among Jre l99l)- It is possiblelhz( dre unusual€xlrenre e7e enl2l incr€a$d tentTeslrial$ecies, rl.€ g€kes Chondrcdacllls ongutifet dimorplrisn in 4,li?orod?sd cadnatd, znd the ojld Pal atoge.ko rarg€i a-rercslriclcd to sandr habilats. Ircq cncvof mamm.{ianprq ca(cnby largcrspecimcns nlils( lho lac€nid Pedioplanis goerdesi prcfcc gmvel help6to rcduccdielary olctlap bentecn dlc l$o slo ic< plains. Thc geckoPa.ry./zctylus bibrcnii (= P. laevigatus) MaDict al (1982)sxggcscd lhal drecnlarged palalrnc a^d $e skink Mabuyastiala r?r/ba{g/ havewidcr habilat f^ntis of I'vthoho.li?saseinall? arc uscd to olerco'nc a prefcrcnces,and arc oRcnco netuad(Branch 1988). Tbc spccificdcfcnsile slra{egye'nploycd by sone geckos(r'e. "s€lf-nuhlatron" bushveldge6il is noclumal ard occursin opcnsavanna ofl by skin loss whcn seized). AllhouSh sandysoils (Dc Graatr 198I ). gcckos do fonn lhe najor conponenl of the diet of Whal docsihis lell us abou(tlrc foragirg taclicsadoplcd hlhonadipMs .afinta (39 % ot all prcy). \rc discountUlo q (1982) sfategY W Ptlhonodipsas, or the aalapdvesiglrficxncc of ihe lrlpothcsisof Ma.^ a1. as s1lcha defcnsile unuslral cranral features,narnely fmgnenled head-shields is uscd by only a few souficm African g€ckos,none ol and eolarged palatine teelh? Experiencerith capti\€ *\vhichhav€ bccn rccordedin thc dter of Pythanodipsas specimensconfinns tl)c nocrnnal habirsof Pylrcnodipsas , aa),ara Il,i5 dcfenscoEcld srnrs nlosrmn$icuor.a itl (Bmnch carinata Menerld 1955; Wemer 1985). Hencc, llrc t\e Pnhyhctylut nanaqwnsi! cotnplc\ cl al. numerouspr€y ile N bclonginS to diunully-actrve lj:.ard 1996),bur nonc of !)es€ spccicsoccur signifcandy within high Aenem (RhoptropB, Natu.kia, MabuJv, Pediaplatlis, ne rzl,geof Pyhonadipsascari,ata. Moreover,{rc AeoM\ ate {'rdtkely 10 have be€n captucd fm|n ambush proponionof gcckoswillin tlrc diei may no( rellectdietary durine daylight hous. They were probablyloe.ted by llrc qxcialisation on lhis Sroup as drc rveslemarid region of snakein their retr€atsat night. This iffers aclivese?rching soulhemAfnca is a radiationcentre of geckos,pairculaJly by the snake. of Pachtdactylus3dJtd its rclatives(Bauer 1990). They fonn Discussionsof snako ahetoften suessfomging theory drc domirumcomponent of tlrc noclu al vertebralcfauna nl and predator-preysiz. opdmizalion (see Amold 1993for 1Jreftrye of Prrhonodipsascarrol4 and can bc ery€ctedto rcaeotrenew). Horrcvet in sxch discussronson dre cosls form a significmt proporuon of the dict of even non- and benefilsolprey seledio4 liulc atlentionhas bccn paid sp€.iaiistpredalors. as to how snak€slocate cr'?lically hidden prcy, and r!h2( Man c( al. (1982) also suggesledtbat thc unusual beluvioural and morphologicaladaptatrons lhey nray usc10 tsagmcnlalion of head shielals in \rpers. boids and o{ract pr€y itcrns Iiom their reueals. ThiE despitc&c fact Pythmodipsas catinata My promol€ th€ rvide gapc tlut many snakesare known to f€ed on prey tlMt have rcquirEdto €nguflflargc prcy. Tl.€ pPM of PrthonoApsN y" dificrcnt lemporalactivity paltcrm. BesidesS,rrorozrpsa! coihata ftngcs lrctn 0.5 lo 42.87q although re a\€r"ge caflrala, a munbcr of othcr snak€slr.rve noclumal habi6 is 28.8'1,and oron re.orclsmnSc fron l8 - 25 %. Ho$'e\cr. and le{ lced al osl cxchsivelr on diumal lizirds Tlrcsc onlv liznrd p.ey was soJficicntll inlact for RPMS lo bc include Africz \,tolt snakes (Lycophkliotr sp.; Branclr c]llflrlalcd.and ( is likcl) tllal rodentpnr) \ould compnsc 1976),and Nonll Arnericu llrc snakes(Zrn,!o/p,to.1on qr.: lnrgerRPMS. RPM lus bc€ndoql|nentcd for \,eryfcw other Temafl 1985). Menbers of tro& gencm lnve grcall) soullEm Africiin sDakes(ctarnpl.:s in Slin€ cl al. 1996a_ c. enlargedanterio. rna\illary teeth an4 like Plhohodipsa\ 199?). bur il docs not se€,n l;kely t\at Pythanodipsas ca,,rrard,also bavedonovenlnlly flattenedheads. we rus .zrtrla is exceplionalin tlns resp€cl In51ea4we proposg s.rgg€stlhat in all threc 8cner4 €nlargedanlcnor teeu (on thal drc fmgmentedhead slields of Pythonodips.L,coinata any of $e dentig€rous bones), funclion in exliactng ma!,pennit geater dislorlion of rc lEad and allow easier sleepingdiunal preyfrom fteir refeals in burrors and rock preymanipulalion ilidin the confinesofa lizrrd's retreal. crevices- Thc flattencd heads of these snakesruy also permitensier rnaniputation of.prey in r€strictedspaces. Aner Rctroduction: Duc to the lack of sexually-activefernalcs. e\lradior\ prey may bc suMued cidEr by constridion our invcs0gations resoh€ only ccnain as?ec1s of (blcophidion a\d Pythonodips8 .a,"t?ard) or by venom reprcducrionin Pythonodipsascan,ala. Aidlough few intrcducedvia grooYedposlerior t2'.,8s(Tiinorphodon atrd dclailedstudies on rcprcduclior)in sou$em Afiican snakes V'ssibly Pythonodipsascal,,/,i?ro). Currenl phyloSen€0c ha\€ bc$c.llundenakcn (Duberia lutix. Kofron 1990i h$olheses (e.g., Sch?itti& Mccadry 1987; Cadlo 1988) lsa,rroprrr sp.,Flcnmring I 994;n/arelva sp., Shine€t al. place these three genera in ditrcrent subfamilesor tnb€s 1996^:,Aspidelaps s.rral,s, Shine et al. 1996b; and

Ai. J. Herpetol. 1997,46e): a9-9j 96

Thelotomiscapeneis, Shine cr aJ. 1996c),ir is er4denrfrom Frrnalcs of vivifJarousspccies are usuaflyslr gr"vid lDegcneEl lirenturc (Broadlcy& Cock I97j; . . Branch1988; \uur |.ate-termernbryos in latesummcr {Janu?ry - March). ry90) th?t a cofiunon reprcducti\€ loaorcy slrares, for $iul_Drnnoccrmng in rc Fcbnury _ Aprit penod. No spccies in the tcrrrf,cmrcrcgions ot p.t/ronodipns Tgarous sourlEm oerarrsof rcproductiofrin tanaata c}..{j'rtin ,1y","* marura on of csss and maLjns 1l: - in €arty drc literaturc e\cept for MaEis' (1992) ctaim Out jl is ocrobefl.ess layins ,n tale oMf,arous. :!ll8 1o"c,5- +rins Wc a.reunaware of any dala to suppon rtus rsepcnber Novernbcr).willt erncrgcnceof hatclJrngs in slalcment AII southcm African muscun collccdonslack h|are$st)mmer - March). That -(Janua4/ ryUono(tipsal ruture females from the criticaj pcrio4 SeDrcmoc.lo to dris rcprcduclivc sensonatiry lo**o.?^ol* is D€cembcr.Hoi{:vcr, our dala do not connjd wjth a l}?ic:rt supponedbyi (i) tlrc fact tlut s€r€ral of lhc feinaleswe oviparous sqsonal brcedjng qcle to( plthonotltpros e\aminedshoir€d clerr evidenceof s€xualmatfity (i e.. had carinata. T\e att.F.rceof embryosin lhrec nulure fernalcs Iafgemuscular oviducts, over ltris _ spnng srlrnmcrpcnod: collectedin Januaryalso mirigatcsagairrsl ano OD u|e ocorrrence viviparjty, br{ of rc.entt)_hatchcd)oung in tjr morcdata are requir€d. March10 July pcriod.

Acknowledgcmelrts

We {\a* Ixdsc Dn*rcw (SoulhAlii@r MwM, CapeToM), EDr critin (St2teMBm, Whdhcr), wull.Heckc (Ttu$vet M@um,Prelori4, md.rcns vnduD (cslifonliaAodm!.rs"i"*., i,* r-oi*y r.,;*;;;*,, ",. for ,"raits of ,hlrdill i,, 'n€trcae we alsorb"''* Mike crifrn (NmibianNa.ue cor*ali"n wi'ar'*r .'a r'.i",v i*i"v qcoo"t.o Lr".""r n"*"r, sr,t;,1 for facilihlinsou sludies\r'rrilsr in Namibia.Tle Fox.daljorr iRB) for Re*{ch andDevelopmenr c\ dd rhcAushrun Rd?rch courcir ES) providcdtinmciat epport for rlxssrudy.

References I.Z$1.31:5349. BROADLEY, D.c. l9?1. A rwision of drc Afrialn ARNOLD, St 1993. Foraging snake drcory ano pre!._slze_ BencaliJ,4nblyodip.vsat\d X"nocatanus pr@aror-sr/eretarions in snakes.pp. tc.,tnbndae) 87_lt5 /, RA. Occ Pap.NaL Mus. RJrod Ser. 8.4,3r.62e-!y7 seigel. and J.T. Coltms (Eds.,. Sn"kes Ecotog, and BROADLEY, D.G. 1990. FilzsiNons, Snat€sof Soudrenr Behaviour Mccraw-Hill, Ncw york. (rEviseded.). Delta Book, Johann€sburg BAUER A.M. 1990. ptrytogen)and biogeogralh) -- ,Aftica ot rhe BROADLEY, D.c. t991. A rcvierv of the NamibiaJr 8ed(osot soudrcmAfrica and rheEtands of rt|e $eslem snakcs of thc gcnus /-r.ofrlJo, Indi"nOcean: A preli'niM4 anatlsrs.pp. {scrpenrcs 275-28t /, C-olubridad.wirh tt,e dcscriprionof a ncu u. peiers.and R Huflerer (E/s.), cndernrc Ven€bralcsrn rhe $cocles.Ann. Tnn-sva.1tMus. 35: 209-215. Tropics. ZmlogischesForschungsinstitut und Mu-s€urn BROADLEY,D.G., AND E.V COCK. 1975. AlexanderKoenig, BorIIl. Snakesof Zmbabwe.Lngmalln Zimbab[€, Harare. BATJE& A,]\4, W-R BRANCII AND WD. HAACKE. CADLf. J.t je88 phylogendicrctationships 1993. The herp€tofaunaof f,rnanjab arnong and adjaccnt aclvanccdsnlkes_ Univ. Cal. pubt. Zool. I I 9: + DanuratandNarrubia. Madoqua x ?7pp. t8: i I7_14) DE GRAAFF.C. 1981. Thc rcdenls BOULENCERG.A t888 of sourhcnrA,iiar On newor tiutc_kno$nSourtr Bullenvonn,Dulban. 267pp Alricarrepdtcs. Ann. Mag. Na1.Hist. (6) 2: ltO_t_ FITZSIMONS,VFM. 1962 BRANCH, W.R Tlrc Sn.*cs of So[lhcrr t976. Ttrc \lotf sn,llicaLqphi(tion Afncl. Punietl ,^.pe,* , andSons. Joinnnesburg 42lpp. ll b.ophidnn vancBatu (Rcpiha. FLEMMINC.A F s€rpcntcs l99r Reproducrj\ccyites ot rllcsndJ,cs C_olubridae,in Sourtr Africa. J. HerperolI0: ,"a,nkiofh6 p. l-ll. cru.il.r. I?iAhtoni Diaasahs and p natonidus (SetTf.ntes:Cotubridae) in the BRANCE W.R. 1988. Fietd OmngeFree Cuide lo rhe Snakesand Statc.Sourh Africa. J. Herfrrot. Odrcr Reptiles publishers, fusoc AJr 4J t9_27 of SouthemAfrica. Suuik CUNTHFR A 1868. CapeTorln. Sirr.fi accourr of new spcciesof snak€sin rl|e colle.lion of thc British BRANCI{, W.R 1994_ Mllseum. A r Herpctofaua of the Sperrgebeil Mag.Nar. His1. (4) l:413428. regionof souDcmNamibia. Herpetol Nar. Hisl. 2tt). lRIStl I 1994. l-ll The biomesofNamibia, as de(erminedby objectivc caicgorisation. Navors. BRANCT! w'R, A M BAUE& nat. Mus. AND D.A Clo()D. 1996. Bloemfontein.l0: 549-592_ A revierv of thc Nanaqua g?,*o. pachydactvlus KING,RB. 1989.Sc\uat dinorphisN ir snale(ail lengt: naalaqucns^ tRcfit|,a: Cckto idrc) frorn sourl)em S€\:!.ll sclccrior. nalural setectjon, Afrjca.wrh lhedescnplion or morpholoSiel of rirc newspccies. S.Afi €onsl,rixt?Bio: J :-in. Soc 38:133-154 Brdch, Shine,lturlow & webb 9'l

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