MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 173 Cladistic Relationships in the Gekkonoidea (Squamata, Sauria) Arnold G. Kluge Museum of Zoology and Department of Biology University of Michigan Ann Arbor, Michigan 48109-1079 Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN April 3, 1987 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 173 The publications of the Museum of Zoology, The University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, initiated in 1913, serve as a medium for original studies based principally upon the collections in the Museum. They are issued separately. When a sufficient number of pages has been printed to make a volume, the Museum will supply a title page, table of contents, and an index to libraries and individuals on the mailing list for the series. The Miscellaneous Publications, which include papers on field and museum techniques, monographic studies, and other contributions not within the scope of the Occasional Papers, were established in 1916 and are published separately. It is not intended that they begrouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, Ann Arbor, Michigan 48109- 1079. MISCELLANEOUS PIJBLICATIONS MlJSEUM OF ZOOLOGY, LJNIVERSITY OF MICHIGAN NO. 173 Cladistic Relationships in the Gekkonoidea (Squamata, Sauria) t)~ Arnold G. Kluge Museum of Zoology and Department of Biology University of Michigan Ann Arbor, Michigan 48109-1079 Ann Arbor MUSEUM OF ZOOLOGY, IJNIVERSITY OF MICHIGAN April 3, 1987 ABSTRACT Kluge, Arnold C;. 1987. Cladistcc Relationships in the Gtjkkonozden (.Squamnta, Sauria). Misc. PIL~I~.MUS. 2001. liniu. hfzrhzgnn, 173:I-54, fzgs. 1-12.-Thr phylogcrlctic relationships among gekko and pygopod lizards arr investigated. The most parsimonious hypothesis contains srvel-a1 well-corroborated tladcs. 1-wcnty-seven synapomorphies rhararterizr the group gckkos + pygopods. Eublr- pharines rrprcsent the most primitive linragc, and its sister-group is diagnosecl by four shared derived features. Thus, gckkos arr no longer consiclrred a monophy- lctic assemblage. Tllc sister-group relationship between diplodactylines ar~d pygopods is based on ;I single apomorphy, :I rnratal closure muscle that encircles or nearly crrcircles the rxterr~al auciitol-y rnratus. The majority of grkkos- gekkonirles and sphaerodactylines-form an assemblage diagnosrd by tour ape- morphies, with thc genus Tert~torcincusbeing the sis~er-lineageto all others. Sphaelodactylines and the gcnus Pristurus corrstitu~e;t group which is set apart by six synapomorplries. Prlslurus and Gontctodes are sistcr-taxa, in that ortier, to the remaining sph;lerodactylincs. The absence of the second cer;~tobr.anchial arch di;ignoses a large group of gckkonines, mostly Afl-ican in distribution. A strictly monophylctic classititation is proposed, and thr phylogmctic- hypothesis on which it is founded is cotlsistrtlt with sever;ll major rverlts in the brr;tkup of P;ul~ge;i. Key words: Bzogrography, cladisties, Eublepharldae, (;ekkonidae, gekkos, phy- logmy, Pygopodzdnr, pygopods. CONTENTS PAGE INTRODIJC1710N ................................................................ 1 MET'IODS AND MATEKIA1.S ...................................................6 ANCIENT GEKKOTANS ........................................................ 10 CHARACTER ANALYSI .: S ....................................................... 11 Informative Charartcrs ....................................................... 11 Qucstionablc Evidence ....................................................... 33 CONCLIJSIONS ................................................................ 36 Most P;~rsimonious(I:latlo~qam ............................................... .36 Cllarat ter F.volution. ......................................................... 36 Evitlcncc lor Sistrr-Group Relationships ...................................... 38 (:l;~ssification ................................................................ 39 Biogrog~.:cphy ................................................................ 40 AClKNOWLEDGMENTS ......................................................... 42 1.I.I'ERATIIRE CITED .......................................................... 47 APPEN1)IX I: Crntrurn Shapc Ot Adult Trunk Vertcbrac .......................... 54 AI'PI<NDIX 11: Diplodactylines l<x;~minrdFor Vrrtrbral Processes ................... 54 ILL. IJSTRATIONS FIGITRE 1'AC;I . I Recently proposrtl pliylogcnctic hypothcscs of tllc major groups ol gekkonoids ............................................................... 3 2 Grou~~tl-pl;rntlivc~-gcncc diagl-a~n ........................................... 1 :I Cl.rtlog~.am illustrating the conccpt ol syn;rpornorpliy ........................ 7 1 C:l;itlogr;rrn illr~st~;iting( 11;~r;rctrr t ongruencc anti 11ornopl;isy ................. 7 5 C;ri~plr 01 condyle d~v~l~pnicritin sphir~~otl:ic~ylinrs ........................ 15 (i Ch;rr;tt ter-st;rtc tree of nic~;~t;~lrrrusc-lc tr;rnsfo~-mationsc.ric.s .................. 20 7 P;iirctl prcm;rxill;re ol day-old l~ygopods................................... 22 8 Vcnu;~lview5 of p;il;rtc ................................................... 26 9 (;r;rph of variation in ~iurnhrrsol sclrr;rl ossiclcs ........................... 27 10 Skull and cntlolynipliatic sacs 01 Phyllodnc.tylus ruropnc. us .................. 52 11 C.l.rtlogr.am ol gckkotioitls (based on evitlcrrcc sumrn;~ri/cdill Tal)le 3) ........ 37 12 (:l;ltlo~r;lm 01 ~c~l;~tio~isl~il)s;i~nong CCI-rain "ptyod;~ctylincs" ................. 11 TABLES TABLE 1'11 C, I. I Nurnl)~~ol sc lrl;rl osaic lcs pcr c.yc.l).~ll...................................... 13 2 X-ray tlillrac-tion ;~rialysisof gc.kkor~oitlegg sliell .rntl "(;~lcium~nilli" ........ 45 3 <;ckkonoid data mahis ................................................... I6 INTRODUCTION More than 25% of all living genera and species of lizards are placed in the Gekkonoidea,' and much has been written in the last 30 years about their phylogenetic relationships. Underwood's seminal papers on gekkos were based largely on eye charactcrs (1951, 1954, 1955). Hecht ( 1976) and Russell (1979) inferred three principal lines of evolution from Underwood's Linnean classification (1954): Eublepharidae, Sphaerodactylidae and Gekkonidae. The latter was subdivided inlo Gekkoninae and Diplodactylinae on the basis of differences in pupil shape. In 1957, Underwood documcnted a sister- group relationship of the cosmopolitan gekkos with the snake-like pygo- pods of Australia and New Guinea (Fig. 1J; see also McDowell and Bogert [1954]). Werner (1961) accepted Underwood's (1954) major terminal taxa of gekkos, but proposed an alternative phylogenetir interprcration (Fig. IF; Ychudah Werner, pers. comm.). I concluded that the pupil-shape evidence emphasized by Underwood was more variable than he recorded (Kluge, 1967a). Further, I examined his hypothesis of gekko relationships for congruence with a different data set of 18 characters, mostly taken from the skeletal system (Kluge, 1967a).Wagner's (1961) ground-plan divergence method of phylogenetic inference was used to analyze those data. My major conclusions (Figs. lA, 2) were: (1) The five eublepharine genera were relatively more primitive than all other living gekkos. However, I was unable to discover evidence that the five shared a more recent common ancestor than the one that gave rise to thc remaining gekkos. (2) The Diplotlactylinae + Gekkoninae + Sphaerodactylinae group shared a common ancestor; those gekkos had certain derived states in common which were not shared with the eublcpharines. (3)The Gekkoninae t Sphaerodactylinae was also delimited by shared derived features, and, presumably, formed a natural assemblage. (4) Likewise, there was evidence that the five sphaerodactyline genera constituted a historical entity. (5) No sharcd derived features were discovercd that provided unambiguous evidence lor either of the two largest groups of gekkos, Diplodactylinae and Gek- koninae. Furthermore, I altered much of the generic composition of Under- wood's Diplodactylinae and Gekkoninae in my analyses (contrary to the claims of Hecht [1976], and Hecht and Edwards [1977]), the former sub- family being restricted to the 13 genera endemic to the Australian Region (Table 1). 'popularly referred to as gckkos ;~ndpyxopods (flap-footed lizards), they account for over 90 genera and 850 species (Table 1). In all sections of this paper, except Classification and Biogeography (pp. 39-42), I use current nomenclature, unless stated otherwise. Thus, the present ch;lractcr analyses c;ln br morc efficiently summ;rrised and readily related to the data and interpretations of previous ;~uthors.Gekkota refers to bo~hgckkonoids and Jurassic- bavarisaurs. The taxonomic composition of the other hisher taxa follows Kluge (1967a; 1974). While I enclorsc Wilcy's (1981a) proposal that undiagnosed taxa be placcd in shutter quotcs, I have not employed that convention when citing previous research in order to simplify the presentation. My Linnean classification was not totally consistent with, nor fully informative of, the phylogenetic hypothesis on which it was based