SYSTEMATIC REVISION OF THE GENUS PAROEDURA G~~NTHER(REPTILIA: SQUAMATA: GEKKONIDAE), WITH THE DESCRIPTION OF FIVE NEW SPECIES
RONALD A. NUSSBAUM AND CHRISTOPHER J.RAXWORTHY
MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOC~UNIVERSITYOF MICHIGAN, NO. 189
Ann Arbor, May 3 1,2000 lSSN 0076-8405 MIS(:bLLrZNbOUS PUBI.l(:ATIONS MlJSEUM OF ZOO1,OC;Y. IlNlVEKSITY OF MICHI(;AN NO. 189
The p~rblicationsol'the Mlrsrunl ol' Zoology, Tlle Uriivrl-sity olMicllig;ul, consist prirri;rt.ily of two series-tlrr Occilsiollal Papers anti tlie Miscrllatieo~rsPublic;rtiotls. Botli scr.ics wc1.c forrlitlcd by 1)n Bryant Walker, MI-.Bt-atlshaw H. Swales, and DL W. W. Newcolnl). 0ccasioll;rlly tlic Musc~uli~~rrl~lisli~s cotitril)l~tiollso~ttsitle of'tliese set-ies; begimiilig ill 1990 tlicsc arc titled Sl~cci;lll'lrblications and arc. ~ll~lril~ered.All sul~riiittcd ~riaril~scr.ipts rcccivc extct.lral rcvic\r. The Miscell;lncor~sI'ublic;ltions, which incll~dctnonogl-;~phic stttdies, papet-s on licltl i~licl11111- selun techniques, ;lntl other contt-il111tions not witllin the scolle of' the Occ.;rsio~lalPallet-s, are 11111j- lishcd soarately. It is not intendetl that they be grouped into \,olr~mes.Each ~ir~lnbrt-h;~s ;I title page and, when necrss:u-y, a tahlr of contents. The Occasional Papers, pllhlication of which was begl~nit1 1913, srl-vr as a llledirlm 101- 01-igitial studies I~asedprincipally upon the collectio~isin the M~rsclttii.Thcy arc isstred scpal.atcly. When a sirflicic~~tI~LII~I~)~I. oI'pages has I)ertl 111-iritrtl to make a volulile, :I title pagr, talIlc ol'c.o~~tc~tts,arid all ilidcx ;I[-csupplietl to libr;u-ies :uitl individuals 011 tlie mailing list k)r- the set-ies. A complete list of'p~rl)lic;rtionson Bil-cis, Fishes, Insects, M;rmm:rls, Mollusks, Reptiles a~itlArn11liil)- ians, ant1 othcr topics is available. /\dtlrcss illqrrirics to tlir 1jil.cc.tol; M~lsrlltllof~%oolo~y, 111c IJliivel-- sity of' Michigan, All11 Arbol; Micliigari 48109-1079.
B;rilcy, 1i.M. & (1. (:;r~rs. 1998. 'li\follew syrll>t-;ulcl~itlfishes, i\.lo~ioj)lc,rrr.ctr,.\~rci 11-om I'eni~lsill;lt- India 211nd M. do.sil71crih.0111 Sri I>llik;~.Occ. Pirp. 726. 18 pp., 7 figs. $2.00. N~rssl)a~rtri,1i.A. & M.E. PSI-enden 1998. Rcvisiori ol'thc Afric;w c;lcciliall gelrc~sSclti.r/o~rco/o/),(m I'arkcr. (t\nipliibia: (;ytiitrophiotia: (:irccilii(lirr). Misc. 1'ul)l. 187. :35 I)!)., 15 ligs.. 15 tal~les,2 colot- plates. $7.00. Wcl)l), S.A. and R.li. Miller. 1998. %oogo~r(~licrrs/ocl,tiln, ;I ncw goodcid fish (( ~y~~r~i~~odon~ili~r~r~ics)fi.otii tlic Atlrcca dl.;rirt;rgc ol'Mcxico, a~itl I-ccli~rgt~osisol'the getirls. Occ. 1';lp. 725. 23 1111, 5 ligs. $2.00. N~rssl~;ll~~~i,li.A., (::I. li;rxwortlly Xc 0.1'1-onk. 1998. 'l'he ghost geckos ot M;~dilgascar:a fi~t.tlil~rrc\~i- siori of tllc R/lal;rgasy l~af'to~ilgcckos (Krptilia, Sqlramat;l, (;c\kkonid;rr). Misc. I'l~bl. 1 86. 26 pp., 25 ligs., 5 tal~lcs.$9.00. Dyet; B.S. 1907. Phylogenctic Kcvision of ~\tlict.illopsi~iac('klcostci, Atllc~l.i~iol,sitlac),with (:oln- nrclits on tile Systc~rlaticsoi'thr South Atricricirll FresIr~ilt('1-Fish (;CIIIIS Bnsilic-h,/lcys<;ir.;~~.rl. Misc. PIIIII. 185. 64 pp., 46 figs. $15.00. O<:ot~tiot;R.M., R.K. (:olwcll & S. Navein. l99(i. 'The Flowel-Mites ol'Tt-inidatl 111: 'l'he (;cnirs I~/~it~o.s~irc.\ (A~.'11 :.I. A~~'~ ~tl;re).Misc. l'ubl. 184. 32 pp, 70 figs. $10.00. I(l~rgc,A.G. & 1i.A. Nussballrn. 19!)5. A Review Of Atricall-Matl;lgasca~iGckkotiid I.i/;rl.tl I'liylogelly /\11tl Riogcogl-irplly (Sq~~amat;r).Misc. Prrbl. 183. 20 pp, 11 ligs. $8.50. Raxwortlly, C.J. & K.A. Nussl)a~~tn.1!)'.)4. A review of the M;rd;lgascan Sriakc (;crlcl-a IZ~~rclo?c)lrho/)u,\, I'(~rrcrlrnn'~rcno,r,and Hp/cwliocion (Sqlrarrlata: (:ol~~l~ridar).Misc. PIIIII. 182. 37 1111, 25 tigs. $11.50. Goslilic, W.A. I!)!):). A srrrvcy o1'1rl)l)crjarv rn~~sclrlat~~rein 1iighc.t- trleoste;l~i fishes. Occ. I';III. 724. 2(i 1'11, 9 ligs. $2.20. Ducll~ii;r~i,W.E. XcJ.A. (:ampbell. 1992. Hylid ti.ogs of tlic gcntrs I'loc/rohy/cr: Systclriatics ;rtitl pliyloge- tretic relationsliil~s.Misc. P~1111. 181. 38 pp, 21 ligs. $9.10.
'I'HE KE(;ENTS 01; IHE IJNIVEKSITY Systematic Revision of the Genus Paroedura Gunther (Reptilia: Squamata: Gekkonidae) , with the Description of Five New Species
Ronald A. Nussbaum and Christopher J. Raxworthy
Division of Reptiles and Amphibians Museum of Zoology The University of Michigan Ann Arbor, Michigan 48 109-1079, USA
MlSCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOCTY, UNlVERSITY OF MICHIGAN, NO. 189
An11 Arbor, May 31, 2000 ABSTRACT
Ntr.s.slicc~rVL, R. A. cr,nul Rnxiuorll~,~~,C/. 2000. S~slotnnticrrvisio.n o/ lhr gcl'c~:u,sParocdura Giin 1h.r ( Rq+~lilicr:Sqrrcrlnc~ln: Gdlio7~itlao) , with th~ dc?vcri/)lio,~~[fivc: ntqo s/)c~cic!,s.Mist. P~cbl.iI4us. Zool. lJniv. i\/lichige~n,189: 1-26, 12,/2g\, 7 tnl~h.The gekkonine ge nus Prrrocldurc~Giin thei-, c~ldc~llicto Madagascar and the Co~rioroIslands, previously co~itainedten species divided into two subgroups. Tlic first group consisted ofalargely northern ellclave of specics defined by haviiing the nostril ill contact with the rostral scale. This gl-oup included the type species, I! snri,clzjohe~nn~i\,So~md only in the Comoros, and six Madagascan species inclrlding 1-1 gmcilis, 1-1 g~i/)vn~,I! honmlorhinc~, I! mc~soOc: I? o-oicc;l,s,and P stun~#~//i.The second group, defined by having the ilostril cxcludcd fi.01~1contact with the rostral scale by interposition oS a prenasal scale, included three sot~tliernspecies, I? antlroy~~n,si,s,I? Dn.\lartli, and P j'icte~. Exa~niiiatioilof' type inaterial and recently collected specimens yielded considerable new information about tllc distribution and taxonomic relationships within the genus. I'(~roc~kumprihvor is show11 to be a,junior synonym of I? bnslnrcli. Five ncw Matlagascan specics are described, inclt~tlingthl-ce in the .scr.nclijoknnnzs-Gro~~pand two ill tlie pictc~-Gro~~p,bl-inging the total species for the gc.1111~to 14. These new species occur in nortllcrn, west-central, and southwestern Madagasca~;li~r the nlost part in transitional and dry fi)rest habitats. A distribution map for the new species and a key to the species oI'I'(~7-o~c/~rmis prese~lted. Iky words: Reptilia, Sql~tmata,Geltkonidac, I'(rrood1~7-e1,systematics, Madagascar, I? ~~~(~iilgolierII.SP., I! limi:t/o/)hilnn.sp., I? lavjnkn n.sp., I? 71nhiny~i.sp., I? ucczimhn 11.~11. CONTENTS
PACE
ABSTRACT ...... ii INTR0I)UCTION ...... 1 METIIODS AND MATERIAIS ...... 1 SI.I\I [rs or; 1?11io1..1)1nt, I c:r/~li~~ir:DIX~N ANI) KIWI I...... 1 ~'ill FIGUIII;: PAGE Atlrllt P(I~O~(~ILT(Lmoingokn at type locality ...... Cover . . Molotypc of I'uroetl~rrc~~nc~ingokn in delcns~vepostlrre ...... 2 Jllvenilc P(~t-oedurnrrtaii~goltokn ...... 3 Gcogi-aphic distribution of new species of I'cl?-oedurn ...... 5 Holotype of I'r~r~~rd.umltnr:slophil(~ ...... (5 P(~ro~drrr.(~kar:stof,hik(~. light phase from Namoroka ...... 7 I-'nrvodurrr kcr..rslophilrr. dark phase fro111 Narnoroka ...... 10 Holotypr of' Prrrordurm lrrnjaka ...... 11 ... Holotyl~eof Paro~~durcrlanjnltcr 111 hfe ...... 12 Holotypc of' Paruedurn 71c~hiny...... 17 I-'crrord.u.rcc vclhiny in lile compared to I? c~nt1royen.si.s...... 18 Holotype of I'nroed.u.rcl 71r~:i~rn~Da...... 20 Holotypc of Pcrrordura 71nzi1nbuin life ...... 21 TABLES TABLE PAGE I Morpho~nctricand meristic data for Pal-ordura mningoka ...... 4 2 Morphomctric and meristic data tin- I'(L~UI>dlircc karslol~hilnfi-oin Narnoroka ...... 8-9 3 Morphomctric and meristic data for Parar&ura karstof,hiln from Ankal-alla...... 9 4 Morphometric and meristic data for I'aro~durn lar,jakn fi-om Bemaraha ...... 13 5 Morphometric and meristic data for Pam~durnlai7jnka from Namoroka ...... 1415 6 Morpho~netricand meristic data for I'amedum vazirnhn ...... 22 7 Statistical compariso~lof I'(~roedur(~linr:stol)hil(~ and I-lvc~:in~/)(~...... 22 INTIIODUCTION rnanus; Sdlp = number of subdigital lamellae on digits I-V of pes. Counts are listed left-right. Snout-vent and tail lengths Dixorl and ICroll (1 974) resurrected I'erro~durccGiinther, 1879, were measured with a I-uler to the nearest 1 mm; other ;I gcnns of' Matl;tgascan-C011101-oa1lgeckos, SI-om the widely dis- measl~i-cmcntswere done wit11 dial calipers and recorded to the tributed and I~cterogcneol~sgenus I-%yllorlnctylu,s,described a nelv nearest 0.1 mm. Maturity was estimated by direct exainiilatio~l species, and p~.ovidedthe most recent revie~vof the group. They of tlle gonads. Fciuales with yolked or slightly yolked ova, recognized nine species of'l'crro(~rl7trcr:I? cindroyensis (Grandidier), whether ovarian or ovid~~ctal,were considered to be mature. I? hrt.s/rrtrli (Mocql~ard), I? gr(rcili.r (Boulenger) , P guzbme Dixon Males with enlarged testes and coiled and enlarged vasa kuid ICroll, P Irot~~crlorlt~inn(Angel), I? ovic:eI).s (Boettger), I? pictn deferentia were coilsidered to be mature. Smaller individuals (Peters), I! .scr,tc/ijoh~cn,rti.sGfintllcr, and P stuv~pJ]i (Boettger) . wit11 unyolked folliclcs or l~ndcvelopedtestes ancl vasa defere~rtia One ol' tllese, I? .rccnclijolrcrn~nis,the type species of I'uroetlzcr(r by were cor~sideredto be immature. Latitudes and longitudes were nlonotypy, is cnclcmic to the Comoros. The remaining eight recol-ded to the nearest 0.001' when determined by satcllitc slxcics arc Madagascan. Nl~ssbaunland Kax-cvorthy (1994) communication and to the nearest 0.10' when estimated from described a tc~~thspecies, I! ,mcc,soOe, fro111 Zahaincna Reserve, maps. Specimens are identified by catalog numbers of' the Madagascar and hriclly summarized ecological data for thc Museum of' zoo lo^^, The University of Michigan (UMMZ), the M.,I( 1.agascan .. Sorins. Muscum National d'Histoire Natut-elle, Paris (MNIHN) , and by When Dixo~iant1 Iholl (1974) revived Pc~rord~tm(feminine) field nlrmbcrs (RAN, RAX) . 1.1-om l%,j~lloelo.c./j~lrt,s(~nascl~linc), they failed to account for the gentler cllangc ill their spclling of' specific names. Most subsequent authors (c~g:,Nussbaum and Kaxworthy, 1994; 61aw and Vencrs, 1994) also did not correct the spellings. The names Dixon and ICI-011 (1974) cstablishcd Pc~roedumgxihrnr on the af'li-ctcd arc I? "li~orncrlot-l~,inu.s,"which should be I! ho~nnlor/t,incc, basis of a scrics of' speci~llensfi-om near Betroka in solrtlrcrn and I? "Picl~r?correctetl to I! picr(1. Madagascar. They apparently believed thcsc specimens repre- Mocqt~arcl (1909) aild sltbscql~cntauthors (Angel, 1942; sented the tnainland form of I? slunzjfj, and that the latter spe- 611ilj@,1056) recognized that I'(~rorc1umcan be divided into trvo cies was othc~.\wiseconfined to its type locality on the northern groupsby the position of' tlre ~lostrilrelative to sl~rro~~ndingsatellite island named "Nosy Be." This was erroneous, however, scales. In one gro~~p,here rckrrecl to as the picla-GI-oup, thr because I? .s/un~/)/Fhad long been known SI-om the northern nostril is cxclutlctl Sroni co~~tactwith the rostra1 scale by tlre mainland adjacent to Nosy Be (Guibk, 1956). Ketroka, on tlre posiiio~rof' the ~)~.c~lasal(= sr~pranasal) scale. This group in- other hand, is 1,150 km south of the northern rainforest habitat cludes I? cr~r~lro~~~~n.si,s,I? hcr.sl(~rcli, and P picla. In the second group, of I? slumpJJli in xeric vegetation. here iratllccl tlrc ,scr?rc.lijolrcmni,s-Gr(111p,the nostril contacts the Dixon and Kroll's (1974) photograph of the holotypr and tlorsolatc~xlcorner of the I-ostral scale. This group iilcludes I? their description of l'(~ro~dur(~pribeuuled us to suspect it is based pcrcilic, I? ho.n~rrlorlt.i.n(i,I! ,rnrrsobc: I? ovicrps, !? sturnpffz, and P on specimens of I? Daslnrdi, the nlajor diff'erence being the pibeerr. 111 general, the flic.l(i-Groupis restricted to south-so~~th- position of the ~lostrilin regard to the rostra1 scale. In I! Duxlardi, western Madagascar and occurs in drier habitats, whereas the the nostril is separated fiom the rostral scale by interposition of .scr~rrcl~jolrcr~t~~t~i.s-Grc~~~pis distribr~tcd in west-ccntral and northern a small prenasal scale, whereas I? p~ihmrwas described as hav- M;rclap~scar and generally occurs in wetter, often rainforest, ing the nostril in contact with the rostral scale, as in I? .slurnp//z habitats. Othcl- than the nostril position, there is no evidence and several other northern spccics of the genus. We examined that eithcr of'thcsc two groups is monophylctic. all of'the type material of I?p~ihecr~a~rd fount1 illat in every case Our Iicld studies in Madagascar yielded live new species of the nostril is clearly excluded from the rostral by the prenasal as I'ecroc.rlrrr(railt1 additio~lalclam on the distribution, variation, and in I? ha.rlardi and two other southern species, I! nn.dmyensis and systematics ol' the genus. Wr describe tlie new species and I? picla. It is 11ncIear why Dixon and KI-011 (1974) thought the Ix"'Xt11 other new information on I'rcroc!rlur(~in this paper. nostril of' the Betrolia specinle~lscontacts the rostral, but it is clear that this error was the reason they conrpared the specimens MET1 101)s AND MATERIALS in detail only to I? stu~nI,Sfiand missed tlre otherwise obviorls siinilarities to I? hnstnr(1i. Because we could not find characters General fieltl technicll~esarc described in Raxworthy and to distingt~ishthe types of l? gwibenr fiom l? Bastnrdz, a species Nnssb;uum ( 1994). Specimens were euthanized by injection with common in tlie area of the type locality of I! guiuibme, wc consider cl~lorol~~~taiiol,lixetl in 10% buf'f'ered formalin, soaked in water I? ti), ~~llc~tctl8-10 Dcccml~cr 199.5 iit tlic t\,pc locillit\. I->\.tllc sitmc depression bet\vccn prominent canthnl ridges. Postcrolatcri~l collectors. border\ of occiput sq11;1rcd. Eiir opening vertical slit. Rods \TI-\? Other specimens. UMMZ 224237-9 (RAN .57408-10). 14 No- slightlv deprc\scd. 0.72 times snout-17ent length, ncarlv vcriil,cr 1997, coastal limestone cliffs 8 krii S Itanipolo, round in c~-o\\scction antcriorl\v, increa\inglv latcrallv 24"43.18'S, 43"59.1 O'E, l(i0 m elevation, t2mpanihv compressed posterio~.l\v,sIi;t~-pl\~ pointed tip; ventral prlgal section Fivondroni~na.Toliara (Tulear) Pro\,incc. Madagit~~;-~r.R. 1-1. with pair of' postcloaca1 wcs. Forelimb laid forward reaches to Nussl~at~~n. midwav benvccn cvc ilnd nostril, hindlimb to about 5 mni bcliind Identification. 11 mcdium-si7cd P(iro~(/rtrcl(;~Illlt six range 37- asilla. Diqits sliglitl\. cup211ldcd at tips. 71 mm SVI.); nostril cscl~~dedfrom contact with rostra1 1,v Rostra1 \tale rect;-ingula~;~videl- than tall. slightlv wider tliiin interposition of prenasal; dorsal t~tberclestrihedrxl to ~ver~klv mental. Nostril in contact with large prenasal anteriorly smaller tctl;~licd~-al;larger dorsal bod\. tubercles arranged in about 12 second supranasal dorsall\., four small postnasals, and dorsal longitudinal r-olvs, larger tubercles scpar;~tcclhv 2-3 small flat process of first supr;~lal->ial.First supralabial largest, labial\ sc;~les;~nd/or smi~ller tubercles; light tlans\.ersc 11;11id across back smooth. Snout i111d intero~.bit;\lscalesj~staposed, some slightl\, of licacl cstendi~igantcroventrall~: ending bclo~vc1.e; ncck and raised, scale row in front of orbits tuberculate, as are larger lateral 11ody with 5 transverse, dorsol;~teral,dark I~andsalternating with occipital scale5. Dorsal occipital skin co-ossified with skull. 5 light bands edged I->\. narrow dark 131-ownlines; ~vllitc\.crteh~-a1 Dorsolater-al neck ilnd bod\. wales lietcrogeneous with ;1b011t 12 stripc on ncck and body interrupted brr narro~vdark brown lines longitudinal (midbodt.) rows of cnlargc-d tetrahedral tubercles, thitt I~ordcrlight transverse bands; unregenerated tail with 8-9 enlarged tubercles ~cpi~ntcdmostl\' b\' small flat scales and fewer \vhitc b:~ndsalternating with 8-9 dark l>ro\vn I~ands;prominent small tubercles. Dors11 scales of forelinib large111flat and weaklv \rcrtic;tl or i~titt>rio~.lycur\red tail displav when disturl~cd. imbricate, escept li~rgcl.\tales of I-adio-ulnar segment, wliicli Description of holotype. See Fig. 1. MTcllprcscwcd. snlall slit are sliglitl\~tubcrc111;ttc. Dorsal scales of hincllirnb strongly postcrol;itcrall\~on venter. Hemipcrics extr~ldcd.Large ~vliite tuberculatc wit11 $mi~ll,gri~nular, interposed scales. Pre- i~ntl tc3stc's (5.8 s 3.7 nim); enlarged. ~vliitcvasa dcfkrcntia. postaxial scales of forcli~nl>not differentiated from vcnt~tlscalcs. Mciis~lrcmentcand counts in Table 1. Head wider thiln ncck. Preaxials of hindlimbs large, flat, sliglitlv imbricate; postaxii~ls ill1011t ;I*;wide as I->od\: Snout short, nci~rl\'as long as postorbital smallcr. g~-anulal: Dol-s~~lpvgal scalcs like dorsal bocl~rwale\; ~~~~~~tion of head. i~ngl~dstccpl\, do\vn~vilrdto tip, slight laterovcntral p\.g;~l\flat, sliqlitlv imbricate, wit11 a pail- of li~teri\l NUSSI~AUMAND RAXWORTI-IY 3 Fig111-c2. .J~~\.cnilrI'nro~dtrrn rnni~l,~oSoltn from tvpc localitv, in life, illustrating.j11vc1iilecolor pattcl-n (compare to Fig. 1). tr~l,c~rcl(~\.First fin~rpo\tp\'gal tail segments ei~ch1vit1i trans- subcanthill ~vhitcstripe and most of supralabials 1-6; anterior \T'I.\Crow of't~rl~c~rcl~~c1orsolilt~raIl~~; posterio~-11' tail \vitli sliglitl~. ciliaries bright white, forming a conspicuous white line in front iml~ricatc,fl;~ttcn scale.;. of eve. 1,ateral region of lo~ver.ja~vlargelvwhite with some bro~rqi Mcntal pcntilgo~;ll,l~ordc-~.cct postel-iorl\. 1x1 pilir of elongate spotson infi-i~labials;tlirec light brown streaks running l>ost~ncnt:~ls.Postmcnti~ls contact mcntal, first infralabial, one posterovcntrallv fi-om infralahials onto sublabials, first beginning <.~ili~~-g(. UMMZ 211210 211211 211212 211213 211214 211215 211216 211217 211218 211219 211220 liolo p;11-21 para pal-a pal-a pal-a pal-a pal-a pa 1-21 pal-a p“u Sex male fcmalc female rnalc Matllrity mature mature immaturc Illatlll-c SVL (i7 65 50 47 TI, 41" 50 3 9 - I3 L 23.2 21.7 18.0 18.5 HW 15.7 15.5 11.6 11.0 MD 10.2 9.8 7.4 7.7 SI . 8.1 -1.1 - (i .(i 6.9 ED 4.7 4.6 3.8 3.4 EO:,:*: 3.2x1.2 2.7x0.9 1.7x0.7 2.1x0.6 AGI> 28.2 28.0 23.0 21.4 Forclimb 22.0 21.7 16.8 16.1 I-Iintllimb 28.0 25.7 22.6 21.4 Supralab 9-10 11-10 9-9 8-8 Inl~alab 9-8 9-9 8-8 8-8 Sdlnl I 10-9 11-10 9-9 8-8 Sdl~u11 13-12 13-13 12-1 1 10-10 Sdlm I11 14-13 1415 12.1 2 12-12 Sdl1111V 12-12 13-14 12-1 1 12-1 2 Sdlm V 11-1 1 11-12 10-1 1 11-11 Sdlp 1 10-10 10-10 9-9 9-9 Sdlp I1 11-12 13-14 11-1 1 10-11 Sdlp 111 15-14 17-16 13-19 13-13 Sdlp IV 15-15 17-17 1413 1414 Sdlp V 15-15 17-17 15-15 15-15 :I:T'11 '1 1ege11crated. or broke11with pal-t lrlissiug ""llcigh~x lcugth are no other obviol~ssexual differences in coloration and by interpositioll of a large prenasal scalc (nostril contacts rostral niorpholo~y. in latter seven species). I'(~ro~dum,rrt,c~i.ngokn shares the nostril The most important variation in coloration concerns the ex- position with P bc~slnr(li,I? picla, I? uahiny n.sp., and F! andryensis. pression of light and dark bands on the necli and body. 111 some I'nroedurm mninpka differs from the latter fo11r species in having individuals, tllcsc bands are boldly expressed (Fig. 2) and in a boldly banded tail and S~lllyexpressed defensive vail display. others they are nearly completely obscured (Fig. I). Kegellcrated Pr~~-orclu~-cr.mningokn dif'el-s fi~rthel-from I? picm in having larger tails may llavc a variegated brown and white color pattern rather terininal pads that are wider than the proximal portion oS the than a banded pattern. digits (not wider in 1-1 Picto), dorsal tubercles sel~aratedby a I-Iatclllings and small juveniles are unlinown, b11t smaller lnixtr~rcof smaller cltbcrcles and sniall flat scales (separated by illdividrlals tend to be Inore brightly colored with greater con- flat scales only in I? j)ic.m), and in having vertically transverse trast between light and dark markings. This is especially notice- dark and light body and neck ba11ds (angled sllarply able in the banding of the tail and body. Bolder arid brighter anteroventrally in 1-1 picla); firrtl1er from 1-1 Dos[c~rdiinhaving large color patterns of young are also typical of I't~rordurr~hn.stnrtli and dorsal tnbercles separated by smaller scales (large dorsal I? pictn. tubercles j~~xtaposedin F! hn.slnr(li), and a white vertebral stripe Etymology. The specific name, "mcringokn" (mine goo kuh) , a and light and dark, transverse neck and body bands (missing in Mala~asyword for "scorpion", is used as an unlatinized specific adult F! baslardi); filrther from I? unndroyensisand I? uahiny in epithet. It refers to the striking behavior of individuals of this larger size (71 compared to 47 mn1 maxiinum SVL), presence species when const-onted, in which the tail is immediately curved of white vertebral stripe and body and neck bands (lacking in antcriorly over the baclz in a scorpion-like manner (Fig. 1). the latter two forms), and absence of multicarinate scales Distribution. KIIOWII only from the type locality at (present in 1-1 clndro)len~icand I? uahiny) . Tsilnalla~llpetsotsaand near Itampolo (Fig. 3). The tail display of 1'nrordu~-(Lmningoka is ~~ndoubtedlya Habitat. Illdividuals were found only at night (20:30-23:50 hr) defensive behavior, which has been reported for other lizards, on the ground among limestone bol~ldersor on limestone rocks especially geckos (Bustard, 1967; Arnold, 1988). A striking in xeric vegetation dolllillated by baobabs and baobab-like legu- parallel to the present case was reportrd for the Chinese gecko, rninous trees of short stature. 72.r~clo.rcir~c:u.srobora7usltii (Autumn and Man, 1989). The latter Remarks. Paroed,urtr mningdzn diI'Sers from 1-1 grrrcilis, 1'. authors believed the behavior is involved in Batesian inirnicry of lro,ttz(rlorhinn, 1'. knr.~lophiln11. sp., I? mmasobe, I? ovicrps, P scorpions. It is also possible, howevel; that in association with sc~~rclijohnnnis,F! .slun~j?/rj,I? lmljczkrr 11. sp., and P ucrzimhn 11. sp. the vividly banded color pattern, the tail acts as a lure to attract in having the nostril excluded from contact with the rostral scalc thc attc~ltionof predators to the most expendable part of the as holotype. UMMZ 219280-1 (RAN 55614-5), 3 July 1996, Anibinorabe-Belaka, Namoroka Reserve, 16"28.225'S, 45"19.515'E, 150-180 In elevation, Soalala Fivondronana, Ankarana (P karstophrla) Mahajanga Province, Madagascal-, J. Rafanomczantsoa, L. Rakotozafy, andJ. B. Ramanamanjato. UMMZ 219282-7 (RAN 55652-7), 8July 1996, Beritra Lakc, Nainoroka Reserve, 150-180 , _I0- - -- m elevation, Soalala Fivondronana, Mahajanga PI-ovince, Madagascar, J. Rafanomczantsoa, L. Raliotozafy, and J. B. Rainanamanjato. UMMZ 222020-37 (RAX 29-31, 91-4, 1524, Namoroka 193, 264, 271-4), 2-7 December 1996, Namor-oka Reserve, (P fanjaka) (P karstophila). 11i028.189'S,45"20.906'E, 150-180 nl elevation, Soalala Province, Mahajanga Pro\~iiice,Madagascar, J. B. Ramanarnanjato, C. J. Raxworthy, and Angelin and Angcluc Razafiininantsoa. UMMZ 2 19245-7 (RAN 55035,55037,55039),26 April 1996, Campcrncnt des Anglais, Ankarana licscrvc, Ambilobe Fivondronana, Antsiranana PI-ovince, Madagasca~; A. P. Riselinla~iana,J. 8. Ramanarnanjato, A. Razafiininantsoa. Identification. A small I'aroedurcl~~pto 55 nnn SVL,; nostril in Dabara (P vahrny) - contact with rostl-a1 scale; eye large, horizontal dianleter equal to or slightly greater- than distance between eye and ear opening; snout relatively long, 61.74% as long as head width; no vertebral CI-cstor ridge with double row of closely spaced tubercles; limbs and digits rnoderate length; apprcssed hindlimb docs not reach antcrior to axilla; fol-elimb laid forward extends to loreal region or nostril, nevcl- extending beyontl tip of snout; double row of' snbdigital larnellae; 10-14 lamellac beneath 4th digit of manus, 13-17 beneath 4th digit of pes; large dorsal tubercles of body trihctlral, sepal-ated by smallcr tubcrcles, arranged into longitu- dinal rows; four distinct, dark, dorsal trailslicrse band on neck ltampolo (Prnaingoka) a~idbody separated by lighter patches of ground color; light patch betrveen first and second dark dorsal bands extends ventrally and slightly anterior-ly on each side to antcrior fbrelimb inscrcions; no dark eye mask (Fig. 5). Figul-c 3. Ccoji~.apliictlistl.il~~~~ioi~ ol'fivc new spccics ofM;rtlagascan Description of holotype. MTell presevecl, no damage, s~liall I'rr,n~(l~on. posterolatcral slit in body wall, tail original, attached to speci- body, which is I-cadilyarrtotomizctl (Johnson and Brotlic, 1974). men. Mat~~remale, enlarged white testes, enlarged white vasa In the case of I! n~nir~,gokn,we ol~servedno obvious animals, tleferentia. scorpions or othcrwisc, at the typc locality that might scrvc 21s Mcas~~rcmcntsand counts in T'al~lc 2. Head oviform, wider baildctl ~rlodcls.I-Iowever, ~~niformlycolored scorpions of niany (9.8 mrn) than neck (6.1 mm), same width as body. Posterior specics are ab~uiclanton the lirncstone slopes and plateaux of' craninm flat, shelf'-like. Snout long, slightly longer than so~~thwcstcrnMadagascar. postorbital scgment of' skllll, sloping rnodcrately downward I'(L~P~,IC~(L~mcrin,goltn was recently found near Itampolo, 70-80 anteriorly; shallow, narrow concavity betwecii distinct canthal lim SE ofihc typc locality, in limestone hahitat and scrnb I'orest. ridges. Eyes large, 11orizoiit;~ldiameter cqnal to distance between This spccics probably occurs tliro~~gho~rtmuch of the largely cyc and ear, 0.63 ti~ncssnollt length; directed slightly anteriol-ly. ~~ns~u.vcyctl,calc;u-eo~~s Mahafaly I'latcau along thc sot~thwest- Ear vertical, oval slit; small, not visible from above. 1,imbs ern coast. Altllough rare in collections, its occurrence in a moderate length; forelimb cxtcnds Sonuard to point between resrrvc (Tsimanampctsotsa), its secretive habits, and its prcfci-- eye and nostril; hindlirnb reaches Sol-ward to forelimb insertion. cncc 101.I-orrgh terrain, wl~icliis diffic~~ltfor h-r~n~ansto spoil, Fingers and toes of moderate length; third finger longcst, only ~xovitlcsome protection lor this spccics. I? mnin,gnka is probably slightly longer than fourth; third toe longest, only slightly longer not crn.rently thl-eatened with extinction. than fourth. Tips of digits slightly expanded with small toe pad; no separate claw-bearing segment. Claws small, not visible from I'(~ro(l01,1ir~ karslo/~hil(~n . sp. above, curving down between terminal larnellac. Original tail Fig~~rcs4, 5, and 6 0.92 times SVI,, postpygal portion ronlided in cross-section proxinially, slightly laterally compl-essed distally, tapering Holotype. UMMZ 219252 ([IAN 556OO), matllrc male, gradually to a sharp tip. collcctctl 4 .Jt~ly1996 at Amboronaon~by,Na~noroka Iiescrve, Nostril contacts rostral, first supralabial, hvo snpranasals, and 1(i028.321'S, 45"20.825'E, 150-180 m elevation, Soalala lonr postnasals. Supranasals adjacent to rostra1 separated by Fivontlronai~a,Mahajanga Province, Madagascar by J. single gran~~larscale. Rostral rectangular, wider than tall, with Ralinomczantsoa, L. Iiako~ozafy,and J. B. Ramanarnanjato. pair of dorsolateral processes that contact nostl-ils. Supraoculars Paratypes. UMMZ 211-1248-5l (IIAN 55595,55597-9),same data lteeled. Snperciliaries weakly keeled on lateral edges. First NUSSRALIMAND RAXWORTHY C Figiirc .5. I'clm~rlitrclIi~tc/o/)l/rl~ from N;~~noroka:light color ~>lin\cwith fullv expressed dorsal nir~rkings.Notc that the first dorsal light marl\ (;ihovc tlic forclinlh) cstcnds down tlic \idc to thc area near the forelimb insertion. This i\ ;In important kc\. cliaractcristic that di\tinguisIicu I? Itnr~to/thrlr fronl I? r~nrr~trhcr(scc Fig. 1:'). s~~pl.;~l;~l>ialtallest. S~~pra-and infralabials sniootli, without cen- posed granular scales. \'en tral surface of digits with two ro~lsof tral raised area. Mental tri;~ng~~lar,same width anteriorly as lamellac. Pair of pilose terminal pads 011 each digit, each pad r.o\tl.al, nc;~rl\*contacting first s~lpralabialwhen mouth closed. about 0.5 nim wide.. Postnicntals piiired, longer than wide, irreq;ularl\?shaped; contact Color (after 36 months in alcoliol):Top of head tannish brown; c;~cl~other riicdiallr., mental anteriorl\r, first and second narrow dark brown line estending posteriorl!. from each nos- inf~~labialslaterall\; and enlarged chin shields posterolaterallv tril, ending above ear. Iris black with faint brassv cast. Supra- aricl postcriorl\: and infralabials whitish, heavilv dusted witli brown Cranial skin co-ossifiecl with underl\.ing hones. Tubercles of melanophores. Dorsuni of neck and body with four dark brown \nout, intcrorhital region, and anterior clorsal cranium homo- transverw bands on lighter tannish ground color. Anterior-most geneous. T~~berclesof lateral cranium and dorsolatcral neck, dorsal band just behind liead, covering most of neck; second I,od\: and tail heterogeneous with smaller tubercles interspersed over shoulders; third at midbodv; fourth above groin. Light among larger promintxnt t~lbcrcles.Prominent tubercles of hod\, ground color separating first and second dark dorsal bands on trili~clr.al,arranged into 12 longitudinal rows at midbod\: fewer neck extends vCntrall\ and sliglitlv anteriorly on each side to rows ;tntcriorlv and posteriorlv on bod]: LTppersurfaces of prosi- point near anterior edge of forelimh insertion. Tail with nial segment of forelimb \zit11 large, homogeneous, trihedral alternating light and dark bands dorsolaterall~:Vpper surfaces tubcrclcs; distiil scgnicnt hctcrogencous, with smaller tllbercles of limbs nearly ~tnifor-mlight tan. \Tc~itrals~u-faces of head, neck, a~iiongtlie I;~rgertubercles. Both proximal ant1 distal segments bodv, and limbs whitish; microscopic csariiination reveals of hindlimbs witli Iicterogeneous tubercles. Tops of hands, feet, numerous tinv, brown melanopliores on ventral surfaces. and digits co\.ered \vi th smooth, imhricate scales. Dorwlateral Underside of postp%qaltail ~vliite,hcavilv d~~~teclwith brown. s~lrfiiccof postpvgal tail with transverse rows of spin\' tubercles Variation. Morphonietric and ~neristicvariation among hpes diniinishing in si7e posteriorlv. Ventral scales regionallv is \~lnirnarii-edin Tables 2-12. There is little within population homogc'neot~s.Chin scales small, granular, juxtaposed. Posterior (Namoroka and Ankarana) variation. The si7c of the dominant t1iro:it and ventral scales of bodv larger than chin scales, sniootli, dorsal triliedr-al tubercles varies aniong indi~idt~als,as docs the flattcncd, sliglitlv imbricate. Undersurfaces of limbs and tail distinctiveness of tlie longit~~dinalrowrs formed bv the large tu- with sni:~ll,sliglitlv imbricate scales. Palms and soles ~vith.justa- bercles on the dorsuni. Analvsis of interpopulation riati ti on is Table 2. Morp1iomct1-ic;mcl meristic variation among Namol-oka types ol Pr~ro~rlul-(1Itnrclopl~~lt~. Mcaslu-cmcnts in Inm UMM% 219248 219249 219250 2192.51 2192.52 219280 219281 219282 219285 219284 219285 pan para para para holo para para para para p:11-:1 pill-a Sex female Mat~u~ty IIlatLlr(' SVI> 49 TL - ML 16.4 HW 10.4 HD 6.3 S1, (5.9 ED 4.2 EO* ' 1.5~1.O AGL 20 1701climb 20 H~ndl~nlh 28 Sup1al,~b 17-17 In11al,ib 13-14 Sdl~nI 7-7 Sdllll I1 9-1 0 Sdlm 111 12-14 Sdlm IV 13-10 Stllln V 9-10 Sdlp 1 7-7 Sdlp I1 12-10 Sdlp I11 14-1 3 Sdlp IV 16-15 Sdlp V 1415 'Missing par1 or rcgcnmtcd **height x length Table 2. (C:onti~~ucd.)Morphomctric >~ndmcristic variation in Namoroka Isnmcd~~rokurs/of)/~~lo. UMMZ 219286 219287 222020 222021 222022 222023 222024 222025 222026 222027 222028 par-a 1~~11-:I para par-a para pal-a par-a 1~ar.a pal-;I pal-;I 1x11-a - Sex Semalc Scmalc male male rnalc rnalc fctnalc fctnalc ~nalc malc Scmalc Mntl~rity ~n;~ture niatilre mature rnature mature rnature mature Iiiatilrc Inature niature rna[ure SVL 47 47 45 46 44 49 48 49 45 48 48 TL - :I(i 1(j:~ 47 5 1 :;: - 34:!: - "I;::" - - H1. l(i.0 15.7 I(i.0 16.7 15.7 16.0 16.5 16.3 15.0 16.9 16.5 HW 9.9 9.8 10.0 10.1 9.4 10.2 9.4 10.2 9.8 10.2 10.4 HD 5.9 5.7 5.4 5.7 5.7 5.8 5.9 6.2 5.1) (3. I 5.7 S1. 6.7 6.0 6.(i 6.5 6.0 7.0 6.9 6.7 6.2 6.9 6.7 E 1) 4. I 4.2 4.1 4.2 4.4 4.5 4.1 4.4 4.3 4.6 4.4 LOM: 1.4x1.0 1.5x0.9 1.5x1.0 1.7x0.8 1.8x0.7 1.8x0.9 l.7x0.8 2.0x0.8 1.5x0.1) 1.7x1.0 1.8x1.2 AGI, 20 21 18 18 19 20 22 24 20 22 21 Forelimh 19 17 18 18 18 I9 18 19 19 11) 19 Hindlilnh 25 24 2(3 25 26 27 26 26 24 28 25 Strpri~lab 17-17 17-17 18-17 17-17 17-17 I(;-16 16-17 16-I(; 16-17 16-16 18-17 lnffi~lah 11-11 11-12 14-13 12-12 12-11 12-12 11-11 11-11 12-1 1 11-10 12-1 1 Scllln I 7-7 7-7 7-7 (i-7 7-7 7-8 8-8 8-8 7-7 7-7 7- Sdlnr TI 10-10 8-9 9-9 9-8 9-9 10-10 10-10 9-9 9-10 10-9 9-9 Scllnl I11 12-13 10-11 12-11 11-11 11-11 13-13 11-11 12-11 12-12 11-11 12-12 Sdlrn IV 11-13 11-11 11-11 11-11 12-11 11-13 11-11 12-12 13-12 11-11 10- Sdl~nV 10-11 9-9 10-9 9-8 10-9 10-10 10-9 10-9 11-10 9-1 0 9- Sdlp I 8-7 7-7 7-7 7-7 8-7 7-8 8-8 7-7 8-7 7-7 7-7 Sdlp 11 11-10 10-9 10-10 10-9 10-1 1 10-1 1 10-11 11-1 1 10-1 1 9-10 9-5) Sdlp I11 15-15 13-12 14-15 12-13 14-14 1414 13-14 14-13 14- 13-14 15-13 Sdlp 1V 15-16 14-15 15-15 I413 1414 16-15 15-15 14-15 15-16 1415 15-15 Sdlp V 15-16 13-13 15-14 12-14 13-13 15-14 15-15 1413 15-15 15-15 14-15 -- "Missing part or rcgcnc~~tcd. w:llcih~ht x Icngth (Continuccl on next page.) Taljlc 2. (Continued.) Morphomctric and ~neristicval-iation in Namol-oka Pumetlu~nknrrtophiln UMMZ 222029 222030 222031 222032 222033 222034 222035 222036 222037 para para pala para para para 11a1-a pal-a para Sex male nialc ~~~alemale female female M;~turity III~~UI-c mature mature matul-c IlliltU1 C mature SVI. 45 45 4 (5 46 48 47 TI, 38:" 43 2'7" 72:': 32% 39% 1-11, 16.4 15.1 15.5 16.3 15.7 16.5 1-1 W 10.9 9.5 10.0 10.1 10.0 9.9 1-11) 6.2 5.6 (5.0 5.9 6.0 6.4 SI. (5.9 6.4 6.3 6.3 6.7 6.8 E 1) 4.1 4.5 4.4 4.1 3.4 4.5 l:.~:k:~: I .6x0.8 1.5x0.8 1.Gx0.5 1.3x0.4 1.5x0.6 1.3x0.8 A(;12 19 18 2 1 2 1 23 22 Fol-climb 19 18 20 18 20 21 I-Ii~idlimb 25 2(i 25 26 24 26 Suplxlab 18-19 18-17 18-17 17-17 1415 18-18 111l'~lI~ll~ 11-11 12-12 12-12 13-12 12-12 12-13 ,- h Stllrn I 7-7 8-7 7-7 I- 1 7-7 8-7 Stll~n11 10-9 10-10 9-8 9-9 10-10 10-10 Sdlm 111 11-10 12-12 10-12 11-11 11-11 12-12 Sdl~n1V 10-11 12-13 11-12 11- 11-13 12-12 Stlln~V 9-10 10-9 9-10 9-51 9-9 10-10 Stllp I 7-7 8-7 7-7 6-6 7-7 9-8 Sdlp I1 10-11 11-12 9-1 0 9-10 10-12 12-11 Stllp 111 12-13 1414 13-13 13-12 1413 15-15 Stllp IV 14-14 15-14 1413 13-15 15-14 1G17 Stllp V 1414 15-15 13-14 1413 13-14 1G17 "Missing part or regenel-atcd. Table 3. Mol-phomctric and meristic variation in f'umedum knrctophilo fi-orn Ankarana UMMZ Sex female male Maturity niaturc? Illalllrc SVI , 44 55 TL - 44:': 13L 15.3 18.0 HW 8.8 10.1) FIL) 5.9 6.7 SI, 6.5 7.6 ED 4.0 4.5 E():l:" l.7x0.7 2.1x0.8 AGL 17 23 Forcli~nb 1(i 18 I-Iindlimb 2 1 27 Supralah 18-16 17-1 (i Illlialab 15-14 13-14 Sdlm I 9-1 0 10-9 Sdlrn I1 12-12 12-11 Sdlrri I11 13-13 12-14 Sdlnl IV 13-13 13-14 Sdln~V 12-12 12.1 2 Stllp I 9-9 10-10 Sdlp 11 13-12 12-12 Sdlp 111 15-15 15-14 Sttlp IV 1G16 17-16 Sdlp V 1Ct17 18-16 "'Missing part 01-regenerated ""'height x length Mlsc. PURI-.Mus. ZOOL..UNIV. MICH., NO. 189 Figurc (5. J'crrn~~drtrctknrcto/)hiln from Na~noroka;dark phase ~vithfaintlv cxprcssed dorsal marking\. hampered by tlie small siirc of tlie ilnkarana sample. The larg- and Ankarana Reserves in northwestern Madagascar (Fig. 3). cst i~idividual,55 nini SW,. is a male from the Ankarana popula- Etymology. The name "knr.stoj)hiIn" refers to the apparent tion (n = 3). The largest indi~idualamong the Namoroka sam- preferred habitat of this species. ple (11= 31 ), also a male, is onl!. 50 lrini SVL. These data strongly Remarks. P(11-o~dltmIrnrrtoj~hiln is most similar to I? vozimhn 11. suggest that the northern population has a larger average size. sp., and the two species differs from other species of Poro~dtl1-0 ~llthoughthe two largest indi\idl~alsare males, several females b!. many of the same character states (see the "Remarks" section from Namoroka are nearlv as large (49 nim SVL) as the largest below for I? vrtzimhn) . male (50 mm) froni that population. The single female from All individuals were collected at night, 19:30-2330 lir, when the Ankarana population is only 4 nini shorter than the largest they were active on tlie surface. All were first observed on tsingy of the two males. It seems unlikely that significant sexual di- rock, 0.5-7.0 m above ground in open deciduous forest. morphism in size exists in this species. There is no detectable morphometric and meristic variation between the two Paroedzlra tanjaka n. sp. populations. Figures 7 and 8 Some individuals within population are more darklv colored, dorsally and \rentrally than others, arid the distinctiveness of the Holotype. UMMZ 224225 (RAN 54507), mature male, transverse, dark, dorsal bands varies froni slightlv expressed to collected 18 March 1996, Reniaraha Resenre, Ambalarano T$ing)!, prominent (Figs 5-6). The light band separating the first and 18"59.230'S, 44'45.566'E, 100-140 m elevation, Antsalova second dorsal hands, which extends ventrally along tlie sides to Fivondronana, Mahajanga Province, Madagascar by J. R. the insertions of the forelinibs, is consistentlv present and is an Ramanamanjato, A. P. Raselimanana, C. J. Raxworthv, arid important identifiing feature of the species. Angelin and Angeluc Razafirninantsoa. Hatchlings and small .juveniles are unknown. Over the size Paratypes. UMMZ 224232-3 (RAN 54468,54527), 17-19 March range of the samples, there is no evidence of ontogenetic 1996, Ambalarano (see data for holotypc). UMMZ 22422651 variation in coloration, nor is sexual dichromatism evident. (RAN 54202,54205-(5,54242,54324,54439),8-1 2 March 1996, Distribution. Known onlv from tlie karst habitats of Namoroka Remaraha Reserve, Antranopasasy River, 18'42.481 'S, fade to Fig111-c8. I'nrnrt/rrr.n tcr~ljoknliolot~l~c in life. 44O-42.SH1 '1.:. I.'ZO-200 m c.lc\.;ltio~i..\nt\;ilo\.;l Fivondronana. adult pattern during ontogen\ of ji~\.cl~ilc\;i11(1 \t~l~i~dt~lts; LJMMZ 22.42,3-1-.', (RIN .54.58.5-(i). 20 March 1 SWi, Rcmaraha unregeneratcd tails with alternating liglit and dark I>ands that Kcserve, Mi~naml~olo.l!)OOH'S, -\4O.iO'E, 100-300 m clcvation, usuall\. do not crow \.cnt~.;tlst~rfiicc: rcgcncrxtctl tail lcs\ dis- .Ints;~lo\.aFivo~id~.on;~na. Ir\4M% 2 19232-8 (Rely .?.5.',C).W, .?.jC,2S tinct11 banded, c is ti all\. with il-rc*gt~Ia~-light ant1 dark markings; '7) 4-'i.Jt~l\.l - - ~p - -- UMMZ 224225 224231 224232 224233 224234 224235 224226 224227 224228 224229 224230 holo para para para I~I-a para para II~IA para pal-a para Scx malc Irmalc male M;I( I1l;ltlll-c mature ? SVI. 102 9 0 70 '1.1, 7(y: 84"'" '37" 131, 34.9 32.3 26.4 1-1 W 24.4 23.1 16.4 111) 13.7 11.8 9.0 SI> 13.3 12.3 10.1 1'1) 9.2 7.4 5.9 p,<)*::l: 5.0x2.2 4.8x1.6 3.4x1.7 A(;I, 44 4 0 29 Forclimb 38 35 28 I-Iintllimb 50 52 36 Supfi~lal~ 12-14 1412 12-12 I ~~J'r~~lalj 11-12 10 10 10-11 Stllm 1 11-9 9-9 9-9 Stllnr I1 13-13 13-13 12-12 Stlllll I11 15-15 15-16 1414 Stllm IV 15-16 l&15 15-14 Stll~nV I 3-1 2 12-12 10-10 Stllp I 9-9 9-9 8-9 Sdlp I1 15-15 14-14 12-12 Stll[' 111 111-18 17-18 1616 Stlll' IV 20-20 18-19 18-1 7 Stllp V 21-21 19-20 18-17 :I: Missing p;trc or ~rcgcncratrd. :6:l:hcil,dl1 X lcllgth ~x~stnasals,one largcr supranasal; scales aronnd nostrils and l't~sionsand other irregularities, smaller near terminal pad, but cant11;ll bridge near nostrils flat, s~nootl~;remaining head scales ultimate subdigital enlarged, lying at base of pad; pair of terminal abovc supl-alabialsand row of small scales bordering supralabials pilosc pads large, each pad rhombic, nearly as long as wide, sides wealzly tubercnlate, those ol' cranial table especially weakly of' single pad measu~-ing2.1-2.4 mm, minimum digit width at tubercnlatr, solne Ilat; supralabials largely smooth, faintly base of pad 1.7 mm, width of both pads 4.9-5.1 mm including t~~bcrculatepostel-iorly; supraoculars Ilat; outer superciliaries space hetwccn them. large, rcctanglllar, Ilat, smooth, arranged in singlc rrnilbrm row; Color after 30 months in alcohol: cranial table and canthal inncr cilial-icssq11a1-c, Ilat, aspinous anteriorly and anterodorsally, bridge irnmacltlate light ycllowish tan, cranial table delimited b11t cach Ilearing single short spine beginning at posterodorsal behind and on sides by lighter border; sides of head light grayish corner 01' eye and behind eye; mental triang~~la~;as wide as brown with white markings; eyelids white anteriorly; iris light 1.ostra1; pair ol' clongate postrnentals in mcdial contact, cach tannish gold; dorsolateral neck and body light gray with distinct borclcrcd laterally by one elongate chin shield; three slightly light brown marbling and crossbands; lower sides grayish white cnlarged chin scales behind postmentals, central one largcst; with brown spots; upper sl~rfacesof limbs colored similarly to lirst lbrlr inhlabials much larger tllan posterior infralabials; body; tail grayish dorsolarerally with few bold light brown cross- posterior inl'l-alabials weakly t~~berculate;scalcs directly below bands and bars; uppcr digits gray; venter of head, neck, and inl'ralabials slightly larger than cllin arid throat scales; body nearly ~vhite,heavily dusted with tiny brown spots on chin dorsolateral neck and body with scattered, nloderatelyenlarged and throat, lightly dusted wit11 brown posteriorly on venter and trilledl-al tnbcrcles separated by much smaller tubercles and under limbs; palms and soles brown, subdigital scales brown; small flat scales, cnlarged tubercles not arranged in obvious ventral surface of tail uniform, light grayish brown. parallel, longirl~clinalrows; upper surfaces of limbs with some Variation. Meristic and morphomctric variation for the cnlargecl trihcdr-al tubercles; upper digits with flat, slightly paratypes from Bemaraha and Namoroka Reserves are overlapping scales; large central scale over claw; pygal region summarized in Xxbles 4 and 5, respectively. Six of 11 specimens tubcrcnlatc dorsally, bearing pair of conical tubercles, one on from Bemaraha are 90 mrn SVIJ or longer, whereas the largest each sidc, abovc lateral edges ol'cloaca; tail scalcs squarish, Ilat, of' 32 specinlens from Nan~orokais only 84 mm SVL. This srnootli,,juxtaposcd, lacking spines and tubercles (regeneraled); strongly suggests that the Bemaraha population has larger vcuiral scalcs sruooth, granular and juxtaposed on chin and average adult body size than the Namoroka population. Because thl-oat, larger and flatter posteriorly on venter; scales under limbs there is an obvious allometric increase in stoutness with size, smooth, flat, increasing in size and slightly imbricate distally; the adults of Bemaraha appear to be more robust than the larger scalcs of palms and soles smooth, small, slightly raised, Namoroka adults. With current samples sizes, there is no de- ,juxtaposctl; subdigital lanlellae ge11e1-allyin double row, but wit11 tectable geographic variation in nleristic and coloration charac- Table 5. (Continued.) Morphomctric and meristic variation in I'(~roedurntn?j7k(z: pamtypcs Srom Namoroka. UMMZ 221998 221999 222038 222039 222040 222041 222042 222043 222044 para para para para para pal-a para para 11:lr~ Sex fcmale female Mat~lrity mature mature SVL 73 82 TL - 73 I-IL 24.7 26.8 HW 16.1 17.5 H 1) 9.6 10.0 SI. 10.4 10.6 El) 5.5 6.7 1 *:Missingpart or rcgcncl-;ltcd. "'"height x Icngth tcrs. There is no evident sexual dimorphism other than that gasy word meaning "strength," and is used as an unlatinized spc- typically associated with the pygal portion of the tail in geckos. cific epithet. Scxrlal dichro~natisrnis not apparent. Remarks. P(~rocdumtnnjaltu difSers from other members of the Three of the Benlaral~aparatypes have two internasal snnctijohannis-Group as follows. It lacks thc spiny vertebral ridge (postrostral) scales between the prenasals instead of one, as in of l? grncilis and the spiny tail and white head spots and white the holotype. All but two of the 32 Na~norokaparatypes have a transverse bands of l? nzn.robe. It is more robust and has shorter single internasal scale; the two exceptional individ~~als11al;e no limbs than I-'. homtllorhinn. The slender hindlirnbs of l? internasals. hon~c~lorl~inareach forward to the back of the head, whereas they I-latchlings and ad~lltsdiffer markedly in dorsolateral colora- fall short of the axilla in l? tanjnka. I'aroedura tnnjaka has small tion of the body. The pattern of color change with age is paral- dorsal t~~berclesnot arranged into regular rows, in contrast to I? leled by other species of I'aroedura, notably Pnroedura Da.ttardi sanctijohanii,& l? .sluin/,ffi, P knrsloplziln, and l? uaziinbn, all of which and 1-1 piclrc. The change is Srom a distinctly white banded pattern have the larger dorsal tubercles disposed into longitudinal rows; in a dark, nearly black, matrix in llatclllings to one of obscured and it is much larger (up to 102 mm SVL) than all of them. banding or dorsal blotching in older (larger) individuals. A I'arord.urr~tanlakn is larger than l? ouiceps (maximum SVL = 68 hatchling paratypc (UMMZ 221997), 31 rnrn SVL, has a light mm) and differs from it in coloration. The dorsal ground color brown head, light brown limbs, and tail (upper surfaces), and a of I! tnnjnlta is light tannish brown compared to darker brown nearly black (dark p~lrplcbrown) body with three vivid, white, in l? ouicef).~.The head of l? ouice/)s does not differ markedly in even-edged transverse bands continuous with the white ventral coloration from the neck and body (brow11with irregular darker coloration. The first white band is over the shor~lder,the second markings), whereas the cranium of l? tanjaka is clearly is much wider and is positioned at midbody, and the third, which demarcated as a squarish, immaculate light brown or tannish is narrow like the shoulder band, is over the hindlimbs. The shelf: Thc snout of I? tanjnlza is also often immaculately tan- venter is grayish white. The three white bands ofjuveniles are nish. still clearly evident, but less vivid in small adults. The bands Superficially, Pnroedura laizjaka resembles 1-1 hnstardi and may have changed to light gray, are edged in dark brown, and have have been conf~~sedwith this species. darker, brown, irregular markings within tllern. The light border Pnroedurcc tanjalta has been found only at relatively low eleva- of the cranium is more distinct in larger specimens, and the tail tions (100-300 m) in dry deciduous forest habitats associated is more irregularly banded. with karst topography. All individuals were collected while they Distribution. I'aroerl,ur(~ tarlj~ltnis fo~nldonly in Bemaraha and were active at night (20:00-22:30 hr). Most were found on the Namol-oka lieserves in west-central Madagascar (Fig 3). bases of tsingy cliffs from 1.0 to 4.0 m above ground. At Etymology. Tlrc specific nanle "tmn~nkn"(tawn zaka) is a Mala Bemaraha, some individuals were collected on branches and tree UNIV.MICII.. No. I89 tr~uiks0.5 to 2.5 111 above gl-ountl. One specinle~l(UMMZ moderate, forelimb reaclrcs barely past eye to loreal region; 224231) ~vasfound dlu-ing the day, ill a cave, 30 111 fi-om the Iiindlimb laid fi)t-\vard to reach 21xilla by 5 mm; fi~lgcrsasltt entrance. Anotlicr (UMMZ 221997) was taken SI-on1the stonl- toes short, s~rbcylindrical,tips bat-ely cxpandcd, small claws ach of a snake (I,~c.otlrnsg(~irt~crrcli, UMMZ 222438). between pair of' tiny terminal, ~~ilosepads, each pad I-lrosnbic with rounded corners, abo~~t0.4 lrirrl across; tail short, 0.81 times Paroedura vahiny n. sp. snout-vent length, constrictctl at I-otrndrd base, swollen ant1 Fig~lres9 and 10 slightly latcrally compressed in midsectio~i,tapering abruptly to short, sharply pointed tenni~lalpol-tion. Holotype. UMMZ 224236 (RAN 51463), nlaturc fcmalc, Rostral scale rectangulal; bordered dorsol;~te~-allyI)y s~nootli collected 27 January 1996 at Dabara Forest, 20°22.446'S, prenasal, middorsally by threc srnall keeled tubercles bctwccn 44"50.854'E, 150 111 elevation, Mahabc Fivondronana, Toliara the prenasals; nostril bordered anteriorly by prcnasal, vcntl-ally Province, Mac1ag.ascar by A. P. Raselimanana, (:. J. Raxworthy, by two, small, flat, smooth scales, postcl.iorly by one flat, smootl~ and Angclin and Angclnc Raz;liimin:i11tso~l. postnasal, above by two, small, ~li~rlticarinatctnbcrcles; Paratypes. None. dorsolateral head scales nearly uniformly small, tnbercnlate, Other specimens. MNI-IN 1984-420 (2001/L), 23January 1969, mnlticarinate; Sew enlarged t~rberclcson side of cranium hehind "15 kin snd(?)Manja." Manja is a town lying about 130 km SSMT eyes; skin on top of craninm co-ossified with underlyi~lgbone; ol' the type localtiy of this spccies, Manja Fivondronana, Toliara supraocn1a1- and s11pe1-ciliaryscalcs 1iinl~icarit1ate;mental Province, Madagascar. This specinlen, discovered in the Paris smooth, triangulal; slightly narrower at lip than rostral, two Muscum arter the holotype was described, fits the description smooth postmentals, rougl~lyhexagonal, partially separated OSI'(~roe(llir(~ vnlri~~y in every detail. anteriorly by apex ol' nlcntal, 1x1-tiallyseparated posteriorly by Identification. A small 1'ur.oc~du.rc~(42 mtn snont-vent length) ; first small chin scale, contacting first infralabial and small, nostril excluded 1'1-om contact with rostral scale by prcnasal scale; sniooth chin shield laterally; dorsolateral scales oSnccli and body digits nearly cylindrical, slightly tlorsoventrally flattened; terminal heterogeneo~~s,with scattered, enlargctl, approxiinatcly pads small, barely wider than digits; hcad scales small, trihedral, rrinlticarinate tubcrclcs among n~uchsrrlaller t-r~berculate,~nulticarinatc, hosnogeneons, except few enlarged multicarinate tltbcrclcs of similar shape; each enlargeti dorsola- scales on side of' head behind eyes; dorsolatcral ncck and body tcral tubercle sel~aratedhy 2-6 smaller tubercles; rcsnaining scales with scattered cnlargcd, suulticaritrate tubercles separated scales, except subdigitals and those on palms and soles, by 2-6 much smaller tubercles; all remaining scalcs (except multicarinate and regionally Iromogeneous (no enlarged spines subdigitals and those of palrns and soles) srnall, regionally or t~rbcrclcsamong smaller scales), I-egionallyvatying SI-omsnlall, uniform in size, tnbcrcnlate, largely non-overlapping, and jnxtaposed to larger; flatter, and slightly overlapping posteriorly nlulticarinatc; hcad uniform brown c1orsolate1-ally,except (distally on limbs); scales of palms and solei raised, rounded, s~~pralabials,infi-alabials asid rostral, ~vhichare boldly spotted smooth; subdigital scales smooth, rounded, generally al-I-ayccl with white; dorsolateral necli, body, upper strrhces of limbs, in double row, penultimate snhdigital pait- flattened, enlarged, nearly nniform brown, except Sour very Faint M- or MI-sllapcd resting at base of srnall terminal, pilose pacts. dorsal crossbands along neck and back; tail darli brown witlr Sr~v Color, after 18 nlonths in preservative: dorsolateral snrf'accs lighter; dorsal crossbands anteriorly and more numerous, s~ilaller generally dark brown, head uniSormly so, ncck witlr very faint, white crossbands ventrally; chin, throat, and lower jaw region light dorsal patch, body 1vith three, light, M- or W-shaped dorsal consl~icuously~narked with large, oSten circular, white patches crossbands, first behind forelimbs vely failit, second in front of separated with narrow brown borders; ventral s~~rfaccsof ncck, hindlimhs gray and PI-onlinelit,third over hindlirnbs hint; lowel- body, and lisnbs dirty white (white with numerous, nearly sides of body with scattered, small, white spots usually coniined to a single scale; upper surfaces of limbs dark ~I-OTYIIwith fi.~ microscopic, brown spots). Distinguished from all otlie~. snlall white spots; dorsal surface of fingers and toes dark brown l'(iroc,durcrby presence of multicarinate scales on ventral surf'aces. with fewwhite spots; tail dark brown with two broad, liglitc~.gray Description of holotype. M'ell-preserved, tail co~npletc,original crossbands anteriorly and dorsolaterally, more f'reqnent, (Fig. 9). Small slit on left side of posterior abdomen. Left ovary narrower, ~iliitecrossbands and spots on ventr;~lsurt'ace; pupil with developingwhite ova, right side .with one white, nearly I-onnd black; antcrodorsal eyclaslr scales white; supralabials, mental, (5.8 x 4.2 mm), shelled, ovidncial egg. and inl'l-alabialswit11 white spots; chin, anterior thl-oat, and lower Measurements (inm);snout-vent length tail length 34; head jaw area conspict~ouslymarked with large, of'ten circular white length 12.7; head width 8.2; snout length 4.8; horizontal eye areas outlined with dark brown pigmentation; ventral surSaces diameter 3.2; ear opening x 0.5; axillagroin length 18.0; 1.0 oS neck, body, and limbs whitish ~vithtiny dark brown spots; forelinlb length 13.2; hindlimb length 18.5. subdigital scales, palms, and soles darli I~rown. The color in Connts: supralabials (IcSt, right) 1s-12;infralabials 10-11 ; preservative has hardly changcd from the color in life. subdigital lamellac, manus, I-V (left, right) (5-6, 9-9, 10-10, 10- Etymology. The specific name "vnhiny" (vnh he nee), a 10, 8-8; pes 6-6, 8-8, 12-11, 11-12, 16-11, Malagasy word suggesting a newcomer or strangel; refel-s to the Head ovate, mncl~wider than necli, narrower tl~anbody; "new one" among the I'c~ro~cl~imspecies. cranium flat dorsally; snout short, 1.5 times longer than Distribution. Ihown only fs-on1 the type locality and neat- Manja horizontal eye diameter, sloping sharply downward in Ss-ont oS in west-central Madagascal-. eyes; distinct canthal ridges, cantha1 bridge concave near eyes, Habitat. The holotypc was collected at 22:00, 7 cm above flat near nostrils; pupil vertical, borders crenate; ear small, ground on a plant stem in deciduous Sorest. vertical; body nlodcrate builtl, slightly depressed; lirubs Remarks. P~~roeclurcc~I(L/L~TL~ is likely to be confirsed only with I? 18 Mrsc. PURL.MUS. ZOOL.. UNIV. MICH.. NO. 189 Figure 10. Holotypc of' I'nro~rlrrrn rlnhinjl (upper) in life comparcd to living I'nm~tlrrrcl nndro?~nsi.s lo^\-cr) 11-on1 sou~licastcrnM;rtl;tgasc;~t-. Thc smoother skin of I? .~~niriny,resulting from smaller more uniform dorsal scales, is apparent. nntlro~~~~n~.ri.s,which is similar in size, shape, digital ~norphology, m elevation, Marovoay Fivondronana, Mahajanga Province, and coloration. I-Iowevel-, I? vcrhiny has a much smoother ap- Madagascar by C. J. Raxworthy and Angelin and Angeluc pearance because oThaving smaller scales on the head and body RazaGminantsoa; UMMZ 216551-8 (RAN 5 1924-6, 51957-60, (Fig. lo), smaller enlarged tnbel-clcs on the neck and body, and 54008), 17-19 Fcbrualy 1996, Antsaravy River, Bora Reserve, s~nalle~;homogeneous tail scalcs. The enlarged dorsolateral 14"51.047'S, 48"13.582'E, 100 m elevation, Antsohihy t~~bercleson the neck and body of I? crn~dro~~en,si,sare arranged Fivondronana, Mahajanga Province, Madagascar, C..J. Raxworthy rougl~lyin longit~tdinalrows, whereas those of Z? unhiny are and Angelin and Angeluc Razafirninantsoa. ~untlonilyscattered. 130th male and fenlale I? crndt-oy~n.sithave Other specimens. None. sncccssive transverse rings or enlarged spines along the tail, one Identification. A small P(zrordum (maximum snout-vent length ring at the anterior margill of each tail whorl, separated by 2-4 49 mm); rostral with dorsolateral processes that contact nostrils; (11s11ally3) rows of rnuch smaller scales. These rings of spines first supralabial in contact with nostril; sllpraocular and arc not present on regenerated tails of P ~1androy,yc..nsis.The tail of supercilary scales keeled; dorsal trihedral tubercles moderate the si~lgleknown specimen of I? 7~crhinyappears to be original, size, arranged in poorly defined rows, each tubercle separated ant1 it lacks rings of spines, which is consistent with having from others by 1-3smaller scales; body somewhat dorsoventrally s~naller,more ho~nogeneousscalcs on other parts of the body. flattened, no vertebral crest or ridge with double row of closely Tllc postmental scales of I? ucrhiny are much longer than wide spaced tubercles; limbs and digits I-elativelyshort; forelimb laid ant1 arc bordered posteriorly by the small, granular chin scales; forward extends to loreal region or nostril, never extending wl~cr~i~sin P nndroyonsis, the postmentals are nearly as wide as anterior of snout tip; double row of weak subdigital lamellae; long ;uid arc borclel-ecl posteriorly by at least one enlarged chin fewer than 12 lamellae beneath 4th digit of manus, fewer than sr;dc., often centrally placed. I'(cro~(tumvalziny has multicai-inate 15 bclow the 4th digit of pes; grayish brown dorsurn with Pour vcntl-al scalcs (smooth in H cmdroyensis) , a uniformly dark colored broad W- or Mshapcd dark transverse bands, first on neck, Sourth Ileatl dorsolatcrally (dorsum of head lighter than side of head above hindlimbs; light patch between first and second dark dorsal in I! trntlroy(~nsis), and a less boldly marked dorsun~(Fig. 10). transverse bands sq~~arish,confined to dorsum, without lateral I'crroo~/~rr(~(IIZF~~~~T)~O~PIIS~'.S and I? vnhiny are the only species of'the extensions to insertions of forelimbs (Fig. 12); no dark mask gcnlls with mu1tic;u-inate scalcs. The keeling of the scales is Inore from nostril to back of head. strongly rxpressecl in Z? vcrhircy. Generally, there are three types Description of holotype. (Fig. ll),well preserved, no dam- of' mu1tica1-inatescales in these two species. Firstly, the larger, age, a small slit posterolaterally on venter. nearly trihedral, dorsal tltbercles have a large, longitltdinal, Mcasuremen~sand connts in Table 6. Head only slightly wider ccntral ltcel on a ridge with several smaller, secondary kccls of than neck, slightly narrower than body, 1.5 times longer than variable length extending Srom the base of the scale up toward wide. Skull flat behind orbits. Snout nearly same length as the cc~ltlxllieel on a11 sides oP the central keel. Often the postorbital section of skull, sloping sharply downward anteriorly, enlarged dorsal tubercles of I? a.nctroytn.ri.r have only slightly shallo~vdepression between distinct canthal ridges in front of ror~ghcnccls~~r~~ces arountl the central keel rather than distinct orbits. Eyes directed slightly anteriorly, horizontal diameter 0.63 sccol~clarykccls, and the strength of expression of secontbry tinles snout length. Ear opening small, nearly vertical slit, not ltccls wries between individnals in this species. Secondly, the visible fi-om above. Fore- and hindlimbs relatively short, smaller, nearly granular scales lack a primary central keel, are appi-esscd forelimb barely reaches nostril, appressed hindlimb conical rather than trihcdml in shape, and l~avcseveral small, Falls short of axilla by about one toe length. Fingers and toes i1-rcg111al.keels extending up the slope of the cone on all sides. moderately long, third finger longest, only slightly longer than Thirdly, the flatter scalcs, Ihr example on the posterior abdomen second and fi)urth; third toe longest. Tips of fingers and toes antl clistal unders~~rlaccoS the limbs of I? vccl~i.ny,h usually have slightly expanded with an enlarged pair of terminal lamellae tl~rcc,11e;lrly parallel, longit~~dinalkeels. M~~lticarinatescales is below each terminal phalanx, no separate claw-bearing segment. a scc~ninglyderivetl ch;tl.actcr state and strongly suggests that i? Claws small, curved down between terminal lamellae, do not cr~rtl~q~tr.ti,sa~~dI? 71nlritcy arc sister-species. Keeling on the ventral extend beyond tips of digits. Tail complete, original, 0.87 times scales is a f~~rthertlcrivcd state found only in I? vcrhiny among sno~lt-ventlength, postpygal portion slightly baterally compressed I'cr,nc,cl~rI-cr spr cies. anteriorly (4.2 mm wide, 4.5 mm deep), round posteriorly, ta- pering abruptly then gradually to a fine point. Paroedura uazimba n. sp. Nostrils in contact with rostral, first supralabial, two Figures 11 and 12 sc~pranasals,and four postnasals. Supranasals adjacent to rostral separated by single granular scale. Rostra1 rectangular, much I-Iolotype. UMMZ 21051% (RAN 49568), adult Pe~nalc, wider than tall, with pair of small dorsolateral processes torich- collcctcd 13April 1995, at Roranical Garden R, Ampijoroa Forest ing nostrils. Supraocular and superciliary scales keeled. First Station, lGOIY'S, 4G049'E, 90 m elevation, Mat-ovoay s~~pralabialtallest. Supra- and infralabials smooth (not Fivonondroana, Mahqjanga Province, Madagascar by Angclin tuberculate) thro~~ghout.Mental triangular, narrower than and Angeluc Razafimanantsoa. rostral, cxcluded from contact with first supralabial. Postmentals Paratypes. UMMZ 21051 (3-8 (RAN 49658, 49731-2), adult fc- paired, hexagonal, longer than wide, in contact with each other malcs, collectrd 9-1 8 April 1995 at type locality by C.J. kwworthy medially, anteriorly with rnental and first infralabials, laterally ;uld Angelin and Angcl11c Raz;-lfimanantsoa; UMMZ 210519 with an elongate chin shield, posteriorly with a large central chin (IWN 49747), adult l'e~nale,collected 19 April 1995 at Ampijoroa shield, and posterolatcrally with a smaller chin shield. Forcst~.yStation, kvaran, Marovoay Kivc~;16'1 7'S, 4G054'E, 90 Skin on top of ct-anium co-ossified with underlying bones. 20 Mrsc. Ptl131. Mrls. Zoor ... UNIV.Mrcrr.. No. 189 Nr.ssn~r~~AND RAXWORTHY 2 1 Fig~ll'c' 12. I'crrordlcrrr 7rnzrrrrhn in lifc from t\lx localit\: Notc t1i;lr tlic first light colored dors;il m;irk (ahovc the forclimbs) docs not cxtcnd down to tli(* ;I~C;Iof'thc forclirnl, inwrtion. Comp;trc this color pattc1.n to t11;lt of I? kor~toj,hiIn(Fig. -5). I~orsoIi~tc~.;~Iscales ~IC~C~O~C~IICOII~;tl11)ercles on back of hc~ld, tral surfaces of head, neck, bod\; and limbs white with numer- ~ic~ck,hoclr: and uppcr s~~rfi~ccsof lim11s separated b~.1-3 smaller ous tiny brown spots especiallv dense laterallv on ventral sur- \c;11cb\; 1i11-gclvtrihctlral t~~l~crclcson hod\. in about 12 poorly faces of bodv and lateral regions of llnderside of head; ventral dcfinctl longitudinal row\. Top\ ofliatids. feet. antl digits with surface of tail banded, but less distinctlv so than dorsolateral im1,ricatc scales. lacking tuhtarclcs. Dorsolateral sc:~lcs of surfaces. The color has faded only slightlv in presenative; the post p\'~aItail ~vitli2 1 t l-;~nsvc~.\cro~vs of tubercles dccrea\ing pattern is unchangccl. 11-0111 (i to 2 tl~l~crclesper I-ow posteriorlv, rows of tubercles Variation. Morphometric and meristic variation among tlie ~<'l>;\~'i\t~-dhv ahol~t 3 rings of smaller imbricate scales, last 12 hpes is summari7ed in Table 6. Sorne parat\.pes have a slightly mni of'tail without tubcrclcs. Vc'ntral scales homogeneous. Chin darker brown dorsal ground color than tlie liolotype. In two and throat scales gr:~nular,slightlv separated; chest and bellv specimens. the darker dorsal bands of the body are divided at scalcs flat, sliglitl\r separated orjuxtaposed, larger in un~bilical midline hv a thin vertebral white line; in UMMZ 210517, these rcgion. Untlersurfacc of lilnhs and tail with small, slightlv linesjoin to form a thin, light, vertebral stripe. The apparentlv i~nhricatcscales. Palms ilnd soles wit11 granularjuxtaposed sc;xlcs. regenerated tail of ITMMZ 210519 is without bands, instead it is CTnderside of digits with double row of lamellae, pair of pilosc dark brown with light grav mottling, darker dorsallv; there are tcnninal pads on each digit, each pad about 0.5 rnm wide. no transverse rouls of enlarged tubercles as on the original tail <:olor (;tficr 48 month\ in i~lcohol):Top and sides of liead of the holotvpe. light l~ro~vnwith irrcg~~lard;xrkcr markings, especiall\.above the The eight paratvpes (IJMMZ 216551-8) from Rora Resene, ot.l>its;\llpra- and infr;ll;~bialswhite with dark brown spot\; (lor- 220 km NNE of tlic hrpe localit): are similar in si7e (28-48 mm sllni of 11cck and botl\, with fb111.broad, dark, transverse hands snout-vent length) and morphometric characteristics to the type with ir~.cgul;lrbortlcrs (M- 01.Mr-sli;~pcd), thc first just bchind series, but diffPr in coloration. The dorsolateral pattern is the the hc;~d,thc sccond.just l~cliindtlic shoulders, third at midhod\: same, but more faintlv expressed, and difficult to discern in a fourth o\.txrgroin; dorsal surfaces of limbs gralr with irrcgl~lar few indi\idr~als.~vliicli are nearlv unicolor. Generally the Rora lightcar and clarkcr markings: tail with alternating dark brown specimens have a lighter ground color, with fewer brown spots, ant1 white hands dorsolatcr-;~Il~;tlic white bands covering the especiallv ~~entrolater-;~ll\-.tljuvenile (UMMZ 2 1655,5), not listed t~.i~ns\.<.~-\~row\ of t~~bcrclcswhere tlic\. occur proxim;~ll\: 1i.n- in Table 6, measures 29 mm S\/'L. Despite its small she, it is ven. 22 Mlsc. PuI~I..Mus. ZOOL.,UNIV. MICII., No. I89 Tahlc 6. Morphomctric and ~nrriaticvariation il~~ro~lgtypes of I'rtwcirl~rrcr rlrrzi~nhcr.Mcasu~-cmcnts in mm 210516 210517 210518 210519 216551 2 16556 21 (5557 para 1x1"~ para par-a para para pal-a Sex ferrralc Scinalc I'cmalc Ielnale l'e~nale fcmalc f+rrralc Matm-i~y mature mature lnatLlrc matlu'c mature lllaturc ITI~IU~C SVI> 39 4 3 47 41) 4 7 46 4 (i TL - - - 28" - 50 4 (i 1-1 1> 14.0 14.1 15.6 1 (i. 1 15.8 1 4.9 15.2 I-IW 7.5 9.0 1) .(i 10.3 9.2 9.3 8.9 1-1 1) 5.0 5.8 5.9 5.9 6.1 5.9 5.5 SI. - 5.6 6. 1 5.7 6.2 5.1) 5.9 ED - 3.1 4.4 3.5 4.1 3.9 3.7 KO:&*: 1.1x0.2 1.Ox0.2 1.2x0.3 1.1x0.3 1.2x0.3 1.2s0.6 0.1-jx0.5 AC;L 17 20 22 22 22 2 1 21 Forclimb 14 16 I (i 16 17 I7 15 I-1indli111b 18 18 21 19 23 24 22 Sup~alab 17-16 15-16 15-15 17-16 16-17 1G17 18-19 Inf'i~lal) 11-12 11-1 1 10-10 12-10 12-11 11-10 12-1 3 m ,- Sdlm I 8-8 7-7 7-7 7-7 7-7 I- I 7-7 S~IIIII1 9-9 9-9 10-9 9-9 8-8 8-8 8-8 Stlllll Ill 11-11 10-9 10-9 11-9 11-9 10-10 10- Scll~n1V 11-10 11-11 10-1 0 10-10 10-10 1 I- 10-1 0 Sdlrn V 8-9 8-9 8-8 9-9 9-8 8-8 8-8 Sdlp 1 7-8 7-7 7-7 7-7 (i-7 (5-6 (5-7 Stllp 11 10-9 10-9 9-9 9-9 10-8 9-9 -8 Sdlp 111 13-12 12-11 12-12 12-12 11-11 11-10 11-10 Sdlp 1V 12-12 13-12 14-14 13-13 11-12 11-12 12-11 Sdlp V 13-13 13-12 I 3- 1:3 13-13 12-12 12-1 2 12-12 "Missing part or rcgcncmted "'"hciglit x Irngth sinlilar in coloration to the adults. This is in contrast to the .ocrzirnba),and i? ~/unz,b)/fialmost invariably has a light vel-tebral general ontogcnctic pattern in Pcero~rlurccin which the j~~veniles stripe running through both the dark and lighter bands 011 tllc are tilore vividly marked than adults. dorsum. I'nroecLum vc~zinrbnhas no light vertebral stripe. 13uroc:eLum Etymology. The name "unzirnha" (pronol~nced"v~th zini bull") vcczimha most closcly resembles I? Itccrstol)hilr~,hut differs as follows. is a Malagasy word referring to small, mystical, and ~nythologi- Table 7. Analysis of cow1riancc of' Sour ~norphomctriccharacters of' cal people of' the forest, and is used as an unlatinized specific I'ulwdur-n knr-ctopltilrr ant1 I! vrtzimhn. Tl~cnlcans are adjustccl to si~c cpitlict in relerencc to the small size and forcst habitat of this (covariatc = S\il>). F-tests inclici~tcho~noge~lcity of slopes between species. species Sol- all four char;ictc~-s.Scl~efSe tests for diSScl-encesin acljt~stcd Distribution. Known only Lrom ihc type locality near means bctwccn species arc all highly significant at p Checklist of P(rro~r1urnGunther, 1879, Species 7a. l>ol-sal ~uherclcstetrahedral; large, adults, up to 72 mrn SVI>..... 8 A. pir.i/r-Group 7b. Dorsal t~tbcrclestrihedral; small, adults up to 55 mln SV12...... 9 1. I'aroetlnr(i nnclroy~!n,sf.s(Grandidier) , 1867 2. I'ctroeclurcr baslardi (Mocquard), 1900 &a. Dorsal tubel-clcs prominent, arranged into regular longitudinal 3. I'(~ro~~c1ur.amairrgoka Nussbaum and Raxworthy, 2000 rows; Soul-largc, prolninent, light-colored, do~rsolateralbands with 4. Pamadurn fiictcr (W. Pctcrs) , 1854 dark edges on neck and body (the fourth over the hincllimbs); 5. P(rroedu,rrcunkiny Nussbarnrl and Raxworthy, 2000 thin, light-colored vertebral stripc usually pl.cscnt; dark cyc mask normally present, extending from nostrils across eye to 13. ~nrtrtijolmnnis-Gro~~p occiput ...... 1'1~roed~urnslu~npfii 6. I-'o,vnl~uragrticili.\ (Bol~lcngcr), 1896 8b. Dorsal t~~bcrclcsless pron~incnt,arranged into irregular longitudinal 7. P(rro~(lurolt,omnlorltir~n (Angel), 1936 rows; dorsrrm brownish with Sour faint, light-colored dorsal bands 8. I'ctmrdurcr mctsobc. Nusshaunl and 12ax~vorthy199.1 9. P(~roc!eluraovicc.l,~ (Uocltgcr), 1881 without dark cdgcs on neck and body; no light-colored vertebral 10. Purnerluva knrslopl~il~rNussbaum and Raxworthy, 2000 stripe; no dark eye ~nask...... I'arorclurtl sanctilohannh I 1. Paroedurn tnnc.lijohnn.nisGiinther, 1879 (type species) 12. l'(rroetl,um .tftr,r~k[F (Bocttger), 1879 9a. Light dorsal neck band narrow, extends ventrally and slightly 13. I'(trol!drrr.f~ la?!jakci Nussbaum and Raxworthy, 2000 anteriorly on each side to a point ncar insertion of forelimbs; eyes 14. I'aro(,elzcrci ?~(rzim,haNussbaum and Raxworthy, 2000 large, horizontal diameter equal to or nearly equal to distance bctwccn eyr and car opening ...... Parordura kontofihila 24 M~sc.PUBL. Mus. zoo^.., UNIV.MICH., NO. 189 Sb. Ligllr dorsal neck band wide, squarish or shcilcl-shaped, co~~linetl DISCUSSION dorsally, 1101 cstcndirlg vrnt~.allyto forclirnb insel-tion; eyrs stnall, I~ori~ontaldiamctc~- less tllall clislaricc bctwce~~eye and car. The nuinbcr of described spccics of I'aroed~cmhas nearly doll- ...... P~~roedzil-c~7~a~imb(~ bled (from 8 to 14 species) since 1994. As a result of' these dis- coveries, Pcrroed~~rclis becoming 1-ecogni~ctlas an important corn- 10a. Long slender li~nhs, apprcsscd liir~tlli~nhreachcs occiptlt, poncnt of the terrestrial vertebrate Fauna of' Madagascan Be- aplxcsscd l'orelinlh extends bcyol~dtip ot snout; cycs largc, callse of its newly discovered diversity and its broad distribution horizontal diameter grcatcr than distance belween posterior border oS cyc and posterior border of car opening; snout elongate and in a variety of habitats, I'uroehrcl has the potential to reveal Ilattcncd, spatula-like; 20 or more subdigital la~nellacon fourth historical distributional patterns within Madagascar. This ~oc;ground colol- ol'head, necli, and body gcncl-ally light, usually potential depends on detailed cladistic analyses witlliil the gc- wit11 poorly dclined darkcr crossbands or il-reg~~lardarkcr markings; nus using appropriate outgroups. original tails boldly lx~unclcd...... I'c~roettzcrc~homalorhi?~a The relationships within the 1'~~roedurccllavenot bccn cxplored lob. Moclcratc limb length, apprcsscd hindlimb docs not reach beyond cladistically, and the relationships of Pc~roed.urato other genela ax ill;^; i~ppressedforelimb usually extends ro loreal region or are poorly understood. Ratier's (1990) cladistic analysis of Afro- ~naxi~nallyto nostril (I! grucilbis exceptional; forelimb may cxcccd Madagascan genera of' geckos, based on morpl~ological lip of snotrt in this species);eye variable in size, lrori7ont;1l di~ur~rter characters, placed Puroedurcc in a terminal tricliotomy with two not grcater than distance bctwccn postel-ior edge of rye and antcrior other Madagascan endcrnics, IBenuzvia and lJ)-o11lutus. This tri- edgc of car opcning, usually less thall distallcc to anterior edge ol chotomy was estiiriatcd to be the sister-group to a polytomy of car opening; snout not elongate and flattened; fcwcr than 20 sttbdigital larrlellac on fourth toe (ral-cly 20 in P LnrLjaku); ground five additional gcklionid genera (Ailuronyx, Grtltok~~~i.~,Hmnof~kolis, color of head, neck, arid body generally darker ...... I I 12ygodaclylu.s,~Ji-ocotylrcLon) from Africa, Madagascar, and other islands of the western Indian Ocean. These resl~ltsled Baucr lla. Pattern of white spots on dark puvplish bl-orvn gro~uldcolor of (1990) to hypothesize that most of' the ~ucsternIndian Ocean head, neck, body, and upper surfhces 01 lirribs (absent TI-om t;~il); gelikonid farina cvolved as a singlc unit. while spots especially dcnsc on head, usually corlfirred to singlc Khlge and Nussbaum (1995) also studicd the cladistic rela largc tubcrclc; body with 3-4 light, transverse bands; original tail tionships of the geklzonid genera of Afi-ica and the wester11Iiidiail with Ihur light b;nlcls; eyes dark, ~learlyblack, directed strongly Ocean with results some~vhatat variance with those of Bauer Sorwarcl; 01-igirii~ltail laterally compressed, wit11 2-3 rings of spincs just posterior to basal constriction, usually witllirl first light-colorccl (1990). Kluge and Nussba~uri(1995) showed that wit11 the ad- mil band; cloublc row ofdorsal spines or1 original tails; three parallel dition of genera and characters, Bauer's iiifcrrcd history of the rows of'subdigiral lamcllac; large, up to 107 nlrn SVL ...... ASro-Madagascan gekkonid radiation becomes more complex ...... I'nroe(1uro mntobr wit11 the suggestion of' repeated dispersals betrueen All-ica, Mada- 11b. No pattcr~~of while spots on dark purplish gl-ountl color; eyes not gascar, and other islands of the western Indian Ocean. In addi- remarkably darli ;md dil-cctcd strongly forrvarcl;original tails without tion, Kluge and Nussba~lm(1995) found that the available data double row of dorsal spines; kwcr than three rows of subdigital are inadequate to I'ornlulatc a robust (stable) pl~ylogcnetictree la~nellae(rrsually two) ...... 12 for the genera oS geckos that occlu- in Africa, Madagascar, and islands of' the western Indian Ocean. Although I'aroc:(Lwa may 12a. Raised vertebral ridgc on body bearing double row of cnlal-gcd, prove to be part of'a inonophyletic radiation ofhtro-Madagascan alternating tubcrclcs; lorclimbs long, exterldirl~forward beyond gcckos, this remains to be demonstrated. tip ol'snout; darli brown ground color with in-cgular darker brown Tlie two informally named groups of I'(~roe(kurcl species, the rnarkir~gson cithcr side of vertebral ridge; vcrtcbral I-idgcusually liglll colored; often with two additional, irregular, paravcrtcbral scrnclijohannis- and $ict(c-Groups, may or may not prove to be light stripes on body; head coloration riot markedly dil'ferent Srorrr monophylctic. However, it is noteworthy that the newly discoverd coloration of body and neck; venter nearly as dark as dorst~ni..... species described in this paper support the biogeographic ...... P(~rorrlorrrgrac-ili~ pattern previol~slydefined by thcsc two groups. The two new 1%. No raised vertebral ridgc on body l)earing double row of enlarged, members of the pictc~-Group,I? maingoha arid P vahiny, occur in alternating tubercles; lorelimbs shorter, not reaching beyond drier southwestel-11Madagascar where the other spccies of this nostrils when appresscd; no light-colored vertebral stripe; ventral group occur; and the three new members of the sancl~jolzann,is- ground much lighter than dorsal grouncl color ...... 13 Group, P kccr.rtofihila, 1-1 lanjalza, and P vcczimbn, occlrl- in wetter parts ofwest-central and northwestern Madagascai-, closer to the 13a. 1,argc and robust, up to 102 mm SVI.; uppel- surlaces of neck, body, geographic range of the other members of this group. Tlie rec- tail, and limbs of adults light gray with irregulal; coarse, brown mark- ognition that P guibeue is synonymous wit11 P bastnrdi removes ings; tail gray, banded with darkcr brown; cranial shelf ilnmacr~latc tannisll, clearly demarcated in coloration frorn neck and body; snout what wol~ldhave been a biogeographic anomaly in regard to also often irnrnaculatc tannish; I~atchli~igsand juvcnilcs vividly this pattern. I'aroedwrrr pibmowas misidentified as a lrleinbcr of Ix~ndcd...... Pnro~~durf~tonjf~ka the northern group occurring in southern Madagascar. The five new species described in this paper arc restricted to 1%. Smallel; up to 68 rnm SVL; dorsal surfaces brown with darkcr brown markings; neck and body with Soul-,il-regulal; M- or W-shaped, dark rather specializetl, relatively dry habitats in wcsterli Madagascar. brown, dorsal hands, often broken up into brown spots; upper Three of the five species are linowil oiily fronl karst habitats, surSaccs oS limbs brow~lwith irregular darker brown markings; and none is known f'rom more than hvo localities. The restricted original tails with alternating dark gray and darli brown hands; head distributions of these spccics are, however, not rlecessarily rcasons brown clorso'~tcrally with irregular dark brown markings, not dis- for conceril about their survival. In general, the biota of'western tinctly different in coloration Srom neck and body ...... Madagascar is poorly known, and all five species may prove to ...... I'uroed~t.raouic.c,fj.r have a rnucli broader distribution than is currently lznown. l'r~lnorli~rtt(~(~~lrtjtigoktr is probably I-estricted to tlle li~nestollehabi- the Parortlum species, and undoubtedly new species will be dis- tats oS the Mahafaly Plateau of southeastern Madagascar. The covered. Understandiilg the evolution of this genus and its la~unaant1 flora oP [lie Mahafaly Plateau is I-ecordcdfrom only a biogeographic revelations await detailed cladistic analyses. f'ew sl~rvcysalong the readily accessible, coastal limestone cliffs that nlarlt the sotitl~wcstei-11boundary of this exte~lsivelimestone pl;ltcat~. The interior oP the plateau is virtually unkno~vn,but reccnt discoveries along its edge suggest that the plateau may Wc arc greatly indchtrd lo Jran Baptistc Iia~manamanjato,Achille 1ial.bor ;I locally endemic fauna ant1 flora. These discoveries Raselimanana, and Anglin and Angeluc Razali~nanantsoafor help in ii~cl~~tle1'. m/ri(~(~golicr; the gekko~iid1:'D~nnuia ,~n,ainlimrcinty the field. Brian I7ang assistccl with recording and analyzing data and (Nussb;~~~mand Raxwortliy, 1998) ; and the striped mongoose, helped to clariiy thc sti~tusof solue populations. Mark Wilkinson pro- vided data concerning the holotype of I'arocdu~a sa?7,clijohr~nni.s.This (~rrlirlic./i.\gr.ccr~,rlirlielr,\is (Wozcncralt, 1986). I-cscarch was r~iadcpossible through the cooperatioil of' the Malagasy I'rrm~clurct ktr~slol~hilt~is known only fi-om two isolated ltarst Ministere de I'Enscigncment SupCricul; the Minist?re de la Production ~nassils,Narnoroka and Ankarana. The dry tlecidl~oltsforests of Ani~naleet des Ei~uxct Fori-t, and the Ministere dc la licchei-chc Nilrnor-oka and Ankarana arc separated by the Salnbirano rain- Scientiliquc ct Teclrnologie pour le Developpcmcnt. Our rescarch was li~rcsts,wllicli currently pose a dispersal 1~1rric1-to relatively arid- supported in part by grants (DEB 90-24505, DEB 93-22600, DEB 96- irtl;~ptcclspecies lilie 1'. hrcnlo)hiln. Other arid-adapted 25873) fi-om the National Scicncc Foundation, the National Geographic liei-pctoS:~nnal species, such as I:clr(:iJ,~-o7i,\tnk~li, H(~looliot1on 11. Society (539691), and Eal-th~vatch. Logistic support was pro~.idcclby SIX (Nussbarun ;~ndIiaxworthy, in prcss), L~iohrl(~rodonwzorlestn, the Mbrltl Wide Fund foi Nat~u-c 0j)litrrc.s c.i171i/~i,and Zo~rosrcurzis1rrticccurlnlri.r arc also restricted to opposite sicks of' the Sambirano. This distributional pattern LITERATURE CITED sr~ggcststhat xeric corridors once spanncd the Sambirano ~.cgio~~.I'trroor1rcr.r~ lorv:slol,hilr~ nlay yet be fonnd in the lililestonc Angel, F. 1942. I.cs li-~ardsdr Madagascar. Mi-m. L'Acadi-mie Malgachc regions oP the largcly unknown Iklifely I'latca~r 10 the soutl~west 96, 193 pp., 22 pls. ol'N;~n~o~-olta.It scenis unlikely, however; that it will be forind Arnold, E. N. 1988. Caudal alllotonly as a dcfcnsc. Pp. 259-273. In (:. li~rtlie~.soutlr in thc Bclnaraha karsts, as Benla~.;~llais relatively Gans and R. B. I-l~lcy(ctls.) , Biology oS the licplilia, Vol. 1G, ecology well s~~rvrycd. 13. Allrr~R. I.iss, Inc., New Yo]-k,659 pp. I'rr~r~(~rho-rrlrrlcjrrlirr is cllrrcntly rccortled only from the two lime- Autumn, K. and B. Hau. 1989. Mitnicry olscorpions byjuvenile lizards, stone regions of Bemaraha and Namorolta, a clistributional pat- 'l>rc~loscinc.~~\mboro~osltii ((;ekkonidac). Chinese Hel-pctological Rcscarch 2:(i0-64. tern know11 f'or other lizards, noteably the new sciilcid species Baucl; A. M. 1990. Phylogeny and biogeography oS the gcckos oS I\/Ir107ly!~/1Ltlt7drr;/imrt (Nussbaum (:l al., 1999). The region between soutlrcln Ati-ica and the islands of the western Indian 0ce;in: a prc- Rc~narahaant1 Nanloroka is not well known, and I? tnnjnka, M. liminal-y analysis. Pp. 27.5-284. In G. Pctcrs and R. Huttcl-cr (cds.), I~ttrrlr~f/tn~,and othc~.spccics with similar disjunct distribr~tional Vel-tcbl-arcsin the Ti-opics. Museurn Alexander Kocnig, Bonn. ~x~ttcrnsmay eventually he found in this gap. The calcareous Bustar-cl,R. H. 1967. Defensive tlisplay behavior of the Austl-alian gccko, I