A REVIEW OF AFRICAN-MADAGASCAN GEKKONID LIZARD PHYLOGENY AND BIOGEOGRAPHY (SQUAMATA) ARNOLD G. KLUGE AND RONALD A. NUSSBAUM MISCELLANEOUS PUP'.ICAT'3NS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 183 MIS(XLLANE0lTS PUBI,I<:ATIONS MUSElJM OF %OOI,OGl: LrNIVEKSI'lY OF MICHIGAN NO. 18?1 .I . lnc ~x~l)lic;~lionol'tl~e Muselur~ of Zoology, The Univrl-sity of Michigan, co~~sistprim;trily of' two series-tllc Occasional I'aprl-s and the Mi>.cclla~icorrsPublic;~tions. Both series welt foundetl by 1)n Rt-yant Walker, Mr. Bt.adshaw H. Swales, ancl Ilr. W.W. Ncwcon~h.Occasionally the Muscun~put)lisIncs contl-il~~~tionsor~tsidc ol these sel-irs; heginning in I990 thcsc at-c titled Special Publications and arc nr~mbercd.All submitted rnan~rscriptsreceive cxtet-nal review. The bliscellancous Publications, which incluclc nlonographic stltdies, papers on firltl and mu- seum techniqries, iintl othcl- contributions not within [he scopr of the Occasional Papers, are pub- lished separately. It is not inlendctl that they be gl-ouped into vol~~n~cs.Each number has a title page ;und, when ileccssrry, ;I table of contcnts. The C)cc;lsional Papel-s, publication of w11ic.h was begun in 19 13, serve as a ~nediu~nfor original studies based principally upon the collections in the Muserrm. They are isstled separ;rtely. Wl~cna slrfficirnt nrrn~her-of pages has bc.t:~i pri~~tetlto makc a volr~~rte,a title page, table ofcontet~ts,and ;III indvx arc s~rpplictlto lil~rariesand indivitl~r;tlson the rnailing list for tl~eseries. Publicatio~~of. the Occasional Papers has heen strspentled. A conrplete list of p~~hlic;itionson Rirds, I:isl~es, I~lsects,M;ur~n~als, Moll~rsks, Reptiles alrd A~lnpl~il)- ims, ;uncl otl~crtopirs is ;rvailable. Atltlt-rss inq~~iriesto tl~c Ilirccto~; Museum of Loology, The Univer- sity of' Michigan, An11 ArI)ot; Miclnig;~~~48 109-1 079. Kol~ll',F.1. K. F.1,. 13ookstei11(ctls.). 1990. I'rocccdi~~gsof' the Michigan Mo~.~~l~o~r~ct~-icsM~I-ksl~op. Spec. I'l~bl.2. :480 pr). With softw;i~-c,$25.00. Witho~rtsoftware, $17.50. i\lc=~~~tlcl;R.D. 1990. kIow tlitl hurn21ns e\,olvc? licflcctions on the rrniq~rrlyIIII~~II~ species. Spec. P11hl. I. :?A pp. $4.00. liaxwol-thy, (:,j. & 1i.A. Nlrsst)aum. A rrvirw of' tl~cxMadaptscan SII;I~~(;cnc~-;r I'cc~r~clo~t.lro/)r~c, I'(~i.ccrhor/i,rtcrc.cc, i~ndff~/~roliodoir (Stl~~aniirtir: (:olrrb~-idae). Misc. Puhl. 182. 37 pp, 25 figs. $1 1.50. (-Tosl~nc, . W.A. 1993. A survey ol'upl~rrjaw IIIIIS(.II~;~~III-~ill highel. telcostcai~fishes. Occ. Pap. 724. 2ti pp. 9 Jigs. $2.20. I)t~ell~nan,W.E. &,!.A. (:i~mpbcll. 1992. Hylicl 11.ogsof'tl~c gc1111s I'/f~c./tiiltyln: Systrnnatics ;i~~tlpltyloge- lietic- rcl;rtionsl~ips.Misc. I'uhl. 181. 38 pb), 21 figs. $9.10. Mclcitl-ick, b1.C. I!)!)I. Phylogcncric analysis of ;IV~:III I~indlirnbrn~rsc~~la~urc. Misc. 1'11hl. 179. 8') PI), 3 figs. $9.60. I<lugc, I\.(;. 1991. Boinc s~~akcphylogeny ;III~I ~.csc;i~-clicyclrs. Misc. Pl~bl.178. (52 pp, 14 figs. $6.00. I'rl~m,R.O. 1990. A test of'the tnonopl~ylyoftl~c Mat~akins (I'ipridac) ;lntl ol'the (:otingas ((:oti~~gitl:rc) I,i~scdon n~o~.plnologyOcc. I'ap. 723. 44 pi), ti tigs. $3.70. S~~ttkr~s,R.D. and K.M. Bailey. 1990. (:haraCtcrs. I-cliitiorlsl~ips,distl-ihr~tio~~, ;III~ I)iolo~y of No/ro/~i.\ ttt~/~rr~o.t~ott~ri.~,a ~-c-~enily ~l;ln~ctl cyprinid fish tl-0111 so~1t11raste1.nUnitctl St;~trs.0c.c. Pap. 722. 15 pp, 3 tigs. $2.20. Mycl-s, l', ,].I.. k';ltlo~t, i111cI M.1:. Srr~ill~.1990. '4 rc\lic\v of' tl~e/)~/IT)~(,II.YL,\ gl-01111 01 ./\/{o(/oti (MIII.~~:I(~: Sig~norlo~~tinac),,with cmplr;rsis 011 PC~II;~ntl 15olivi;i. 108 PI), 21) figs. $13.20. Railcy, R.M. & 11.A. Ktnicr. 1989. (:om~nents011 111csuhgencra of'dartc~-s(Pel-cidac) with descl-iptions of tbvo new species of lil/~ro.s/orncr((lloc.m/rzr) 1'1-o~nsor~thc;istcr~~ Ur~it~tl States. Misc. Plrl)l. 175. 52 pp, I color 111;1tt., 2 figs. $8.60. I3ca11cBaker, AIIIIArbor I'ar~l W. Kro\v~~,Mackinac lsl;~nd Sl~il-IcyM. McFee, Kiittle (:leek Neil1 D. Niclscn, Brighton l'hilip H. I'owc~; All11 Arl)o~- Vcronic;~1,;itta Sn~itl~.(;rosse IIc Nellie M. Va1.ne1; Ilctl-oit ,];~nles I.. Waters, Muskegon 1;rrnesJ. I)~~dc~.sradt,c.?i oflit io A Review of African-Madagascan Gekkonid Lizard Phylogeny and Biogeography (Squamata) Arnold G. Kluge and Ronald A. Nussbaum Museum of Zoology and Department of Biology The University of Michigan Ann Arbor, Michigan 48109-1079 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 183 Ann Arbor, October 23,1995 ABSTRACT Kluge, A.G., and K.A. N~~ssDaun~.1995. A Hminu OJAJi-ican-Mccdagascan Geltkonid Liznrd Phylogen? And Biog(:ography (Squamala). Aksc. Pu61. Mus. Zool. Univ. Micl~igan,183:l-20, 11 figs. The phylogenetic and biogeographic history of African-Madagascain gekkonid lizards is assessed with 34 morphological charac- ters. The cladistic analysis employs 22 i~lgroupand seven outgroup genera as terminal taxa, and the best- fitting hypothesis leads to the following corlclusions: (1) the ingroup forms a clade; (2) the generally rccog~lizedPachydactylus assemblage of genera forms a clade, as do the northern and southern African subgroups, (C:rYc;lton%a,Tarentola) and (Chondnxiactylus, (;olopus, Pachydactylus, I'almatogecko, lUloptrol,us), re- spectively; (3) the Madagascan-Seychelles sa~mpleforms a historical entity, except for the ambiguously related Gedtokpis; (4) the sample oS Madagascan-Seychelles endemics is derived from African gekkos, and some of the African fauna has a Madagascan ancestry; (5) a Lygodactylus-Phrlsuma clade (including Rhoptrof~ella)is delimited; (6)Blaesodaclyl~~s and Homopholis are not sister taxa. Adding other "padded-toed" gekkos, Gehj~ra,Geklio, and EZemidar.lylus, to the ingroup leads to a substantially different set of phylogenetic and biogeographic conclusions, and reveals the tenuous nature of most of' these conclusions. The close phylogenetic relationship between Ge/~,yraand Paragehym implied by Angel (1929) is not confirmed. It is clear that synapomorphies are more import;ant than geographic proximity in recovering the histoly of' gekkos. These results are largely at odds with Joger's (1985) and Rauer's (1990a) recent findings. ICcy words: I<eFtilici, Squ,crmatcl, Gokkonidac: .4frica, Madagccscai; cladzstics, Oiogeography, 2Y,phylogeny. CONTENTS PAGE Introduction ....................................................................................................................................................... 1 Recent Research on the Phylogeny ol' Gekkonids from the Ethiopian Region .............................................. 1 Methods. Ingroup Taxa and Characters ........................................................................................................... 3 Outgroup Taxa ................................................................................................................................................... 4 Character Descriptions ...................................................................................................................................... 6 Other Potential Evidence ................................................................................................................................ 12 Phylogenetic Results ........................................................................................................................................ 13 Biogeographic Results ...................................................................................................................................... 15 Status ol' Pmagef~ym............................................................................................................................... 15 Conclusions ...................................................................................................................................................... 18 Acknowledg~nents............................................................................................................................................ 18 Rel'erences ........................................................................................................................................................ 18 11.1. USTKATIONS FIGURE PAGE Joger's (1985.fig. 3) hypothesis of phylogenetic relationships .............................................................. 2 1Sallcr's (1990a.fig. 1) hypothesis ol'phylogenetic relationships ...........................................................3 Phylogenetic hypothesis reslilting fro111 a reanalysis of Bar~er's(1990a: appendix 1) data .....................................................................................................................................4 1Sony palate oS gekkos ............................................................................................................................10 Pectoral girdle of gekkos ....................................................................................................................... 11 Phylogenetic hypothesis ol'gekkos restricted to the Ethiopian Region ..................................... ... 12 l'hylogenetic hypothesis of gekkos restricted to the Etliiopian liegion. and i~lcludillgthe cxtralimital Gehyrc~.C.eliltko. and Hemidnctylus ......................................... 13 Area cladogram Lor the gekkos analyzcd hyJoger (1985.fig. 3) ..........................................................16 Area cladogram for the gekkos analyzed by Bauer (19tlOa.fig. 1) ....................................................16 Area c:ladograrn Sor the gekkos restricted to the Ethiopian Kegion .................................................