Norntatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

NorntatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3139, 23 pp., 11 figures, 3 tables June 29, 1995

Cladistic Relationships of Sphaerodactyl Lizards

ARNOLD G. KLUGE'

CONTENTS Abstract ...... 2 Introduction ...... 2 Acknowledgments ...... 2 Materials and Methods ...... 3 Ingroup Terminal Taxa ...... 3 Outgroups ...... 7 Sources of Specimens ...... 7 Sources of Data ...... 8 Hypothesis Formulation ...... 8 Characters ...... 9 Other Observations ...... 13 Cladistic Results ...... 13 Homonomy-Heteronomy, Individual Variants, and the Evolution of the Ungual Sheath ...... 14 A Gekko in Amber ...... 18 References ...... 20 Appendix. Cleared and Stained Specimens Examined ...... 22

' Curator and Professor, Museum of Zoology and Department of Biology, University of Michigan, Ann Arbor, Michigan 48109. Copyright © American Museum of Natural History 1995 ISSN 0003-0082 / Price $2.50 2 AMERICAN MUSEUM NOVITATES NO. 3139

ABSTRACT A cladistic analysis of 25 internal and external tylus, Coleodactylus, Pseudogonatodes) clade is morphological synapomorphies results in the fol- judged in terms of the homonomy and heteron- lowing most parsimonious hypothesis of sphae- omy relations ofthe individual scales ofthe digits. rodactyl lizard relationships: (Gonatodes (Lepi- An Oligo-Miocene gekko in amber from the Do- doblepharis (Sphaerodactylus (Coleodactylus, minican Republic is almost certainly a Sphaero- Pseudogonatodes)))). The value ofthe ungual sheath dactylus. in diagnosing the (Lepidoblepharis, Sphaerodac-

INTRODUCTION The historical relationships of the major on, 1970; Russell, 1972; Hoogmoed, 1985), clades of gekkonoid lizards continue to be the interrelationships of these taxa remain reexamined and reformulated with the dis- poorly understood, as indicated by compet- covery of new synapomorphies and the re- ing hypotheses of affinities. The following interpretation ofcharacters employed in ear- propositions of sister-group relationships, in lier studies (Kluge, 1967, 1987, 1994; Bauer, parenthetic notation, illustrate some ofthose 1990; Kluge and Nussbaum, 1995). The tax- different propositions: (Gonatodes (Lepido- onomy of these family-group categories has blepharis (Pseudogonatodes, Sphaerodacty- also been altered with the application of a lus))) after Noble (1921: fig. 8); (Gonatodes strictly monophyletic nomenclature (Kluge, (Lepidoblepharis, Pseudogonatodes (Coleo- 1987). However, some major groups of gek- dactylus, Sphaerodactylus))) after Parker kos have received relatively little review in (1926)3; (Gonatodes (Lepidoblepharis (Pseu- recent years and are ripe for further study. dogonatodes (Coleodactylus, Sphaerodacty- For example, there is the New World Sphae- lus)))) after Vanzolini (1968b); (Gonatodes rodactylidae of Underwood (1954), which ((Lepidoblepharis, Pseudogonatodes) (Coleo- now consists of approximately 132 species dactylus, Sphaerodactylus))) after Russell (table 1) in five genera, Coleodactylus, Gon- (1972). The present paper begins with a re- atodes, Lepidoblepharis, Pseudogonatodes, examination of the cladistic affinities among and Sphaerodactylus (Kluge, 1993). Sphae- the five genera, including the hypothesis that rodactyls, as I will informally refer to this the New World radiation is monophyletic. group of New World endemics, account for Such a historical accounting is then used to approximately 14% of all gekkonoid species assess the evolution ofcertain details ofdigit recognized currently (including pygopods), scalation in the sphaerodactyl clade, and to with Sphaerodactylus being the most speciose classify a gekko preserved in Oligo-Miocene genus-group ofall gekkos (Kluge, 1993).2 Al- amber from the Dominican Republic. though the monophyly ofeach sphaerodactyl genus appears to be well corroborated (Par- ACKNOWLEDGMENTS ker, 1926; Vanzolini, 1968a; Huey and Dix- I wish to thank Darrel Frost for his counsel on the correct identification of Sphaerodac- tylus dommeli. David A. Grimaldi generous- 2 My use of sphaerodactyl is not to be confused with ly gave me the opportunity to report on the Russell's (1972) Sphaerodactylus Group, which excluded American Museum of Natural History Gonatodes. Currently, 90 species ofSphaerodactylus are recognized. Other species-rich gekkonoid genera include Sphaerodactylus in Dominican amber, and it Cyrtodactylus (sensu stricto; 62 species), Hemidactylus was his preliminary description of the spec- (75), Lygodactylus (51), and Phyllodactylus (56). The to- imen which provided the basis for my further tal number ofgenera and species ofgekkonoids listed by investigation. Wolfgang Bohme and Dieter Kluge (1993) is 102 and 962, respectively. These numbers represent approximately 12 and 29% increases in the recognized genus and species group taxa in the past 3 Actually, Parker (1926: 301) implied that Lepido- 30 years, since Wermuth (1965). blepharis and Coleodactylus are paraphyletic. 1 995 KLUGE: SPHAERODACTYL LIZARDS

TABLE 1 Cleared and Stained Sphaerodactyls Examined Genera Species Species Specimens recognized recognized sampled examined Coleodactylus 5 3 5 Gonatodes 17 14 35 Lepidoblepharis 16 4 11 Pseudogonatodes 5 3 9 Sphaerodactylus 90 47 111 Totals: 133 71 171

Schlee provided information on the original material ofS. dommeli. Carlos Rivero-Blan- co generously provided information from his doctoral thesis on Gonatodes systematics. The late Albert Schwartz was especially helpful in obtaining material of sphaerodactyls for Fig. 1. Geographic range of Coleodactylus. clearing and staining. Others contributing material included Richard Zweifel, American Museum of Natural History; Jens Vindum, species, respectively, constitute the ingroup California Academy ofSciences; Jay M. Sav- terminal taxa employed in this study. Junior age, private collection; Pedro Ruiz, Instituto synonyms are placed in parentheses (cur- de Ciencias Naturales Renovables y del Am- rently recognized subspecies are indicated biente, Bogota; William E. Duellman, Mu- with an asterisk). The accompanying i.e. and seum ofNatural History, University ofKan- n.n. notations specify an invalid emendation sas; John Wright, Los Angeles County Mu- (e.g., lapsus calami or substitute name) and seum of Natural History; Doug Rossman, nomen nudum, respectively. Complete cita- Louisiana State University, Museum of Zo- tions to taxonomic authorship can be found ology; Carlos Rivero-Blanco, Museo de Cien- in Kluge (1993). cias Naturales, Caracas; Ernest E. Williams, Museum of Comparative Zoology, Harvard Coleodactylus Parker (1926a) University; David B. Wake, Museum ofVer- Species content (and synonymy): 1. ama- tebrate Zoology, University of California, zonicus Andersson, 1918 (zernyi Wettstein, Berkeley; Jose Ayarzaguena, Museo de His- 1928); 2. brachystoma Amaral, 1935 (pfri- toria Natural La Salle, Caracas; Richard Tho- meri Miranda-Ribeiro, 1937); 3. guimaraesi mas, private collection; James R. Dixon, Vanzolini, 1957; 4. meridionalis Boulenger, Texas A&M University; Walter Auffenburg, 1888b; 5. septentrionalis Vanzolini, 1980. University ofFlorida, Florida State Museum; Geographic distribution (fig. 1): Most ofthe Tom Uzzell, University of Illinois, Natural species are now known from several geo- History Museum; George Zug, United States graphically close localities (Vanzolini, 1980; National Museum. L. Lee Grismer and Greg Hoogmoed, 1985). The remaining disjunct K. Pregill reviewed the pentultimate draft of species distributions may not be due to lack this paper, and the manuscript was improved of considerably as the result of their efforts. collecting. Gonatodes Fitzinger (1843) MATERIALS AND METHODS Species content (and synonymy): 1. albogularis A. M. C. Dumeril and Bibron, 1836 INGROUP TERMrNAL TAxA (albigularis [i.e.] Fitzinger, 1 843;fuscus* Hal- The following five sphaerodactyl genera, lowell, 1855; varius A. H. A. Dumeril, 1856; which contain 5, 17, 16, 5, and 90 recognized notatus* Reinhardt and Liitken, 1862; ma- 4 AMERICAN MUSEUM NOVITATES NO. 3139

Fig. 2. Geographic range of Gonatodes. Fig. 3. Geographic range of Lepidoblepharis. culatus Steindachner, 1867; braconnieri represents recent human introductions. Spec- O'Shaughnessy, 1875; bodinii* Rivero-Blan- imens from the Cayman Islands, Jamaica, co, 1964); 2. annularis Boulenger, 1887a (an- and Haiti are considered representative ofan nularis Boulenger, 1887b; boonii Lidth de endemic subspecies (notatus) of the wide- Jeude, 1904; beebei Noble, 1923a); 3. antil- spread G. a. albogularis. The coastal Cuban lensis Lidth de Jeude, 1887; 4. atricucullaris records of G. a. fuscus may represent human Noble, 1921a; 5. caudiscutatus Gunther, introductions; however, that interpretation 1859b (collaris Garman, 1892); 6. ceciliae does not seem to apply to interior localities Donoso-Barros, 1965a; 7. concinnatus (Schwartz and Henderson, 1988). The Ga- O'Shaughnessy, 1881 (buckleyi O'Shaugh- lapagos material is believed to represent a nessy, 1881; ligiae Donoso-Barros, 1967); 8. mainland South American species, G. cau- eladioi Nascimento, Avila-Pires and Cunha, discutatus, which probably represents one or 1987; 9. falconensis Shreve, 1947; 10. hase- more accidental introductions by humans. mani Griffin, 1917 (spinulosus Amarfil, 1932); The two disjunct South America records are 1 1. humeralis Guichenot, 1855 (ferrugineus referable to the widespread and otherwise Cope, 1864 [1863]; incertus W. Peters, 1872a continuously distributed G. humeralis to the [1871]; sulcatus O'Shaughnessy, 1875); 12. north (Vanzolini, 1968a, 1968b; Rivero- ocellatus Gray, 1831a (condifentatis [n.n.] Blanco, 1979). Marcuzzi, 1950); 13. petersi Donoso-Barros, 1967; 14. seigliei Donoso-Barros, 1965a; 15. taniae Roze, 1963; 16. tapajonicus Ro- Lepidoblepharis Peracca (1897b) drigues, 1980; 17. vittatus Wiegmann, 1856 Genus synonymy. Lathrogecko Ruthven, (gillii Cope, 1864 [1863]; roquensis* Roze, 1916. 1956). Species content (and synonymy): 1. buch- Geographic distribution (fig. 2): Venezuela waldi Werner, 191 Ob; 2. colombianus Mech- and the Dutch West Indies represent the area ler, 1968; 3. duolepis Ayala and Castro, 1983; of greatest species diversity (Rivero-Blanco, 4. festae Peracca, 1897b; 5. grandis Miyata, 1979). Although Gonatodes is present on 1985; 6. heyerorum Vanzolini, 1978a; 7. in- Trinidad and Tobago, representatives are termedius Boulenger, 1914a; 8. miyatai La- otherwise absent from the Lesser Antilles, and mar, 1985; 9. microlepis Noble, 1923b; 10. much of the Greater Antilles. The Florida montecanoensis Markezich and Taphorn, distribution (mainland and Keys) probably 1994; 11. oxycephalus Werner, 1894; 12. per- 1995 KLUGE: SPHAERODACTYL LIZARDS

tratus* Shreve, 1968; enriquilloensis* Shreve, 1968; lucioi* R. Thomas and Schwartz, 1983b); 2. argivus Garman, 1888 (bartschi* Cochran, 1934; lewisi* Grant, 1941 [1940]); 3. argus Gosse, 1850 (henriquesi Grant, 1940; andresensis* Dunn and Saxe, 1950); 4. ar- masi Schwartz and Garrido, 1974; 5. armstrongi Noble and Hassler, 1933 (hypsi- nephes* R. Thomas and Schwartz, 1 983a); 6. asterulus Schwartz and Graham, 1980; 7. beattyi Grant, 1937 (seamani* R. Thomas and Schwartz, 1966a); 8. becki K. P. Schmidt, 1919b; 9. bromeliarum G. Peters and Schwartz, 1972; 10. caicosensis Cochran, 1934; 11. callocricus Schwartz, 1976; 12. cel- icara Garrido and Schwartz, 1982; 13. cinereus Wagler, 1830 (steinegeri* Cochran, 1931); 14. clenchi Shreve, 1968 (apocoptus* Fig. 4. Geographic range of Pseudogonatodes. Schwartz, 1983); 15. cochranae Ruibal, 1946; 16. copei Steindachner, 1867 (anthracinus Cope, 1862a [1861]; picturatus* Garman, accae Boulenger, 1908; 13. ruthveni Parker, 1887; asper Garman, 1888; cataplexis* 1926a; 14. sanctaemartae Ruthven, 1916 (fu- Schwartz and R. Thomas, 1965 [1964]; en- gax Ruthven, 1928); 15. williamsi Ayala and ochrus* Schwartz and R. Thomas, 1965 Serna, 1986; 16. xanthostigma Noble, 1916. [1964]; polyommatus* R. Thomas, 1968; as- Geographic distribution (fig. 3): The dis- treptus* Schwartz, 1975; pelates* Schwartz, junct part of the range at the mouth of the 1975; websteri* Schwartz, 1975; deuterus* Amazon River represents L. heyerorum, and Schwartz, 1975); 17. corticola Garman, 1888 that which occurs in Venezuela is L. mon- (aporrox* Schwartz, 1968a; campter* tecanoensis. All species of Lepidoblepharis, Schwartz, 1968a; soteir Schwartz, 1968a); 18. other than L. festae and L. heyerorum, are cricoderus R. Thomas, Hedges and Garrido, found outside the Amazon Basin, the major- 1992; 19. cryphius R. Thomas and Schwartz, ity of which are from Colombia (Mechler, 1977; 20. darlingtoni Shreve, 1968 (noblei* 1968). Shreve, 1968; bobilini* R. Thomas and Schwartz, 1 983a; mekistus* R. Thomas and Pseudogonatodes Ruthven (1915) Schwartz, 1983a); 21. difficilis Barbour, 1914a (lycauges* Schwartz, 1983; euopter* Schwartz, Species content (and synonymy): 1. bar- 1983; typhlopous* Schwartz, 1983; peratus* bouri Noble, 192 1 a; 2.furvus Ruthven, 1915; Schwartz, 1983; diolenius* Schwartz, 1983; 3. guianensis Parker, 1935a (amazonicus* anthracomus* Schwartz, 1983); 22. dunni K. Vanzolini, 1967); 4. lunulatus Roux, 1927; P. Schmidt, 1936; 23. elasmorhynchus R. 5. peruvianus Huey and Dixon, 1970. Thomas, 1966; 24. elegans MacLeay, 1834 Geographic distribution (fig. 4): The species (punctatissimus* A. M. C. Dumeril and Bi- are allopatric (Huey and Dixon, 1970), and bron, 1836; alopex Cope, 1862a [1861]); 25. lack of collecting probably does not explain elegantulus Barbour, 1917; 26. epiurus R. entirely the highly fragmented distribution. Thomas and Hedges, 1993; 27. epizemius Garrido and Jaume, 1984 (docimus Schwartz Sphaerodactylus Wagler (1830) and Garrido, 1985); 28. fantasticus Cuvier, 1836 (ligniservulus* W. King, 1962b; anidro- Genus synonymy Sphaeriodactylus Gray, tus* R. Thomas, 1964; fuga* R. Thomas, 1831a. 1964; hippomanes* R. Thomas, 1964; ka- Species content (and synonymy): 1. alta- rukera* R. Thomas, 1964; orescius* R. Tho- velensis Noble and Hassler, 1933 (breviros- mas, 1964; phyzacinus* R. Thomas, 1964; 6 AMERICAN MUSEUM NOVITATES NO. 3139 tartaropylorus* R. Thomas, 1964); 29. gai- Gosse, 1850 (dacnicolor* Barbour, 1910); 59. geae Grant, 1932; 30. gilvitorques Cope, pacificus Stejneger, 1903; 60. parkeri Grant, 1862a (1861); 31. glaucus Cope, 1866 (1865) 1939; 61. parthenopion R. Thomas, 1965; 62. (inornatus W. Peters, 1873; torquatus Strauch, perissodactylius R. Thomas and Hedges, 1887); 32. goniorhynchus Cope, 1895 (1894); 1988; 63. plummeri R. Thomas and Hedges, 33. graptolaemus Harris and Kluge, 1984; 34. 1992; 64. ramsdeni Ruibal, 1959; 65. randi heliconiae Harris, 1982; 35. homolepis Cope, Shreve, 1968 (methorius* Schwartz, 1977a; 1886 (carinatus Andersson, 1916; imbricatus strahmi* Schwartz, 1977a); 66. rhabdotus Andersson, 1916; mertensi Wermuth, 1965); Schwartz, 1970; 67. richardi Hedges and Gar- 36. inaguae Noble and Klingel, 1932; 37. in- rido, 1993; 68. richardsonii Gray, 1845a (gos- termedius Barbour and Ramsden, 1919 (dra- sei* Grant, 1939); 69. roosevelti Grant, 193 la; petiscus Schwartz, 1958); 38. klauberi Grant, 70. rosaurae Parker, 1940; 71. ruibali Grant, 1931a; 39. ladae R. Thomas and Hedges, 1959b; 72. sabanus Cochran, 1938; 73. sa- 1988; 40. lazelli Shreve, 1968; 41. leucaster manensis Cochran, 1932; 74. savagei Shreve, Schwartz, 1973; 42. levinsi Heatwole, 1968; 1968 (juanilloensis* Shreve, 1968); 75. scaber 43. lineolatus Lichtenstein and von Martens, Barbour and Ramsden, 1919; 76. scapularis 1856 (casicolus Cope, 1862a [1861]); 44. ma- Boulenger, 1902; 77. schwartzi R. Thomas, crolepis Gunther, 1859a (imbricatus Fischer, Hedges and Garrido, 1992; 78. semasiops R. 1881; grandisquamis* Stejneger, 1904 [1903]; Thomas, 1975; 79. shrevei Lazell, 1961; 80. danforthi Grant, 1931a; parvus* W. King, sommeri Graham, 1981; 81. sputator Sparr- 1962b; ateles* R. Thomas and Schwartz, man, 1784 (pictus Garman, 1887); 82. sto- 1966a; guarionex* R. Thomas and Schwartz, reyae Grant, 1944; 83. streptophorus R. Tho- 1966a; inigoi* R. Thomas and Schwartz, mas and Schwartz, 1977 (sphenophanes* R. 1966a; mimetes* R. Thomas and Schwartz, Thomas and Schwartz, 1983a); 84. thomp- 1966a; phoberus* R. Thomas and Schwartz, soni Schwartz and Franz, 1976; 85. torrei 1966a; spanius* R. Thomas and Schwartz, Barbour, 1914a (spielmani* Grant, 1958); 86. 1966a; stibarus* R. Thomas and Schwartz, townsendi Grant, 1931a; 87. underwoodi 1966a); 45. mariguanae Cochran, 1934; 46. Schwartz, 1968a; 88. vincenti Boulenger, 1891 microlepis Reinhardt and Liitken, 1862 (me- (festus* Barbour, 1915; monififer* Barbour, lanospilos A. H. A. Dumeril, 1873; thomasi* 1921; adamas* Schwartz, 1964; diamesus* Schwartz, 1965); 47. micropithecus Schwartz, Schwartz, 1964;josephinae* Schwartz, 1964; 1977b; 48. millepunctatus Hallowell, 1861 pheristus* Schwartz, 1964; psammius* (1860) (continentalis Werner, 1896d); 49. Schwartz, 1964; ronaldi* Schwartz, 1964); 89. molei Boettger, 1894b (buergeri Werner, williamsi R. Thomas and Schwartz, 1983b; 1900b; venezuelanus Roux, 1927; boettgeri 90. zygaena Schwartz and R. Thomas, 1977. [n.n.] Donoso Barros, 1968); 50. monensis Geographic distribution (fig. 5): Only one Meerwarth, 1901; 51. nicholsi Grant, 1931 a; species, S. molei, occurs on Trinidad and To- 52. nigropunctatus Gray, 1845a (decoratus* bago, and that taxon is also widely distrib- Garman, 1888; flavicaudus* Barbour, 1904; uted on mainland South America (Harris, gibbus* Barbour, 1921; alayoi* Grant, 1959a; 1982: fig. 5). No Sphaerodactylus occurs in atessares R. Thomas and Schwartz, 1966b; the Dutch West Indies, or the Grenadines granti* R. Thomas and Schwartz, 1966b; lis- and Grenada, which is similar to the geo- sodesmus* R. Thomas and Schwartz, 1966b; graphical distribution of Gonatodes. Two ra- ocujal* R. Thomas and Schwartz, 1966b; diations ofSphaerodactylus, West Indian and strategus* R. Thomas and Schwartz, 1966b; mainland groups of species, have been men- porrasi* Schwartz, 1972); 53. notatus Baird, tioned in recent research (Hass, 1991); how- 1859 (1858) (exsut* Barbour, 1914a; amau- ever, synapomorphies have yet to be discov- rus* Schwartz, 1966 [1965]; atactus* ered which diagnose the latter group of en- Schwartz, 1966 [1965]; peltastes* Schwartz, demics (Harris and Kluge, 1984). Although 1966 [1965]); 54. nycteropus R. Thomas and S. pacificus, which is endemic to Cocos Is- Schwartz, 1977; 55. ocoae Schwartz and R. land, is quite likely to be part ofthe mainland Thomas, 1977; 56. oliveri Grant, 1944; 57. radiation, other species geographically close omoglaux R. Thomas, 1982; 58. oxyrhinus to the mainland do not appear to be part of 1 995 KLUGE: SPHAERODACTYL LIZARDS

southwest Asian Pristurus was delimited by six synapomorphies, including absence ofthe , splenial bone, and the individuality of that assemblage was formally recognized by plac- ~~~~~~~~~~Sphaerodasctylus ing all six taxa in Sphaerodactylini (Kluge, 1987). A recent review of African-Madagas- can gekko relationships (Kluge and Nuss- baum, 1995) suggested that Pristurus belongs to a clade of African gekkos, which includes km Narudasia, Quedenfeldtia, and Saurodacty- lus. The absence of cloacal sacs and bones was found to be diagnostic of that group, which would also include sphaerodactyls (Kluge and Nussbaum, 1995; their characters 11 and 24, respectively). Also, the absence of -0 1000 preanal and/or femoral pores is characteristic of that assemblage (Parker, 1926; see also Kluge and Nussbaum, 1995; their character Fig. 5. Geographic range of Sphaerodactylus. 22). Further, Russell (1972) suggested that Gonatodes is similar, ifnot closely related, to Cnemaspis, particularly the Indian-Oriental that history. For example, S. argus is thought species-group, and therefore I have also in- to be part of the Antillean radiation (Hass, cluded the latter taxon as an outgroup. Thus, 1991), but that species is present on Corn I have expanded the outgroups in this study, Island, Honduras, and Isla San Andres, Co- to include Cnemaspis, Narudasia, Pristurus, lombia. The former location probably rep- Quedenfeldtia, and Saurodactylus, in order to resents a human introduction, whereas the render the ingroup character polarity hy- later may not (Schwartz and Henderson, potheses more globally parsimonious, and to 1988). The Swan Islands records for S. no- more generally test the individuality of the tatus also indicate a species that is considered ingroup (Clark and Curran, 1986; Nixon and part of the West Indian radiation, and has Carpenter, 1993; Kluge, 1994). The fact that probably dispersed toward the mainland. Kluge and Nussbaum's (1995: figs. 6,7) anal- Sphaerodactylus and Anolis (sensu lato) ex- yses led to different sister-group hypotheses hibit certain geographic similarities. The for Narudasia, Pristurus, Quedenfeldtia, and numbers of species of each taxon found on Saurodactylus, which also differed from those West Indian islands are highly correlated4. In delimited herein, strongly suggests the need addition, the three most widespread Anolis for further research on those outgroup taxa. species (A. carolinensis, A. distichus, and A. sagrei) have geographic ranges very similar to the three most widespread Sphaerodacty- lus species (S. argus, S. copei, and S. notatus). SOURCES OF SPECIMENS The skeletons ofmore than halfofthe cur- OUTGROUPS rently recognized species of sphaerodactyls were examined (table 1; Appendix). The re- The sister-group relationships ofthe sphae- pository abbreviations for that material are rodactyl lineage to the northeast African- as follows: AMNH, American Museum of Natural History; AS and ASFS, Albert Schwartz private collection (now KU); CAS, 4r = 0.846, df = 390; the data were taken from Ma- California Academy of Sciences; CR, Costa clean et al. (1977). Only those islands that had at least Rica Collection, Jay M. Savage private col- one species ofeither genus were scored. The islands south lection, currently housed at the University of of St. Vincent, including those in close proximity to the Miami; ICN, Instituto de Ciencias Naturales mainland of South America, were not sampled. Renovables y del Ambiente, Bogota; KU, 8 AMERICAN MUSEUM NOVITATES NO. 3139

TABLE 2a SOURCES OF DATA Data Matrix: Character by Taxon The vast majority of the characters used

A B C D E F G H I have been mentioned in other research on sphaerodactyl relationships (Noble, 1921; 1. 0 0 0 0 1 1 0 0 0 0 Parker, 1926; Underwood, 1954; Kluge, 1967, 2. 0 1 0 0 0 0 1982, 1983, 1987; Vanzolini, 1968a, 1968b; 3. 0 1 1 0 0 0 0 4. 0 1 1 0 1 1 0 0 Russell, 1972), or in studies on other gekkos 5. 0 0 0 0 1 0 0 (e.g., Bauer, 1990; Kluge and Nussbaum, 6. 0/1 1 1 0 1 0/1 0/1 1995). Some ofthe evidence cited most often 7. 0 0 0/1 0 0 1 0 0 comes from the scalation and share of the 8. 1 0 0 1 0 0 digits, particularly the ungual sheath. Unfor- 9. 0 1 0 0 0 0 tunately, an inconsistent terminology and 10. 0 l 1 1 1 0 1 0/1 several arbitrary systems of letter and num- 11. 0 2 0 0 0 0 2 0 0 ber labels for those scales have been em- 12. 0 2 0 0 0 0 0 0 ployed over the years (Noble, 1921; Parker, 13. 0 1 1 0 0 0 0 0/1 1l 1926, Mechler, 1968; Vanzolini, 1968a, 14. 0 1 0 1 1 1 0 0/1 1 15. 0 0 1 0 0 0 0 1968b; Russell, 1972; Hoogmoed, 1985). I 01 16. 0 0 1 1 0 1 0 0 believe our understanding ofthe evolution of 17. 0 0 1 0 1 1 sphaerodactyl digits will improve substan- l 1 1 18. 0 0 1 ? 1 1 1 tially by adopting a standardized nomencla- 19. 0 1 0 0 0 ture. Further, a simple system of names and 1 0 20. 0 0/1 0 0 l 0 1 labels based strictly on anatomical location, 21. 0 0 0 0 0 1 1 1 rather than presumed homology, is to be pre- 22. 0 1 0 0 0 0 1 ferred because topological similarity is im- 23. 0 0 0 0 0 0 portant in determining which variants are 24. 0 0 0 0 0 scored as shared I 25. 1 1 0 1 1 apomorphies. Thus, apply the following terms and labels in my discus- a The taxa, by columns, are A = Cnemaspis, B = Co- sions of the sphaerodactyl ungual sheath: leodactylus, C = Gonatodes, D = Lepidoblepharis, E = (1) Dorsals (labeled d in the illustrations Narudasia, F = Pristurus, G = Pseudogonatodes, H = Quedenfeldtia, I = Saurodactylus, and J = Sphaerodac- employed herein). These scales form one or tylus. Data are analyzed as missing when they appear in two (rarely) proximal- to distal-oriented rows the matrix as ? (unknown) and 0/1 (variable). on the dorsal midline ofthe digit. Dorsals are usually larger than laterals. (2) Ventrals (labeled v). Often termed subdigital or inferior lamellae in the literature Museum of Natural History, University of on squamates. These scales form one or two Kansas; LACM, Los Angeles County Muse- (rarely) proximal- to distal-oriented rows on um of Natural History; LSUMZ, Louisiana the ventral midline of the digit. They are al- State University, Museum of Zoology; most always wider than the adjacent scales. MCNC, Museo de Ciencias Naturales, Ca- (3) Laterals (labeled 1). These scales form racas; MCZ, Museum of Comparative Zo- two or more proximal- to distal-oriented rows ology, Harvard University; MVZ, Museum between the dorsals and ventrals. of Vertebrate Zoology, University of Cali- The scale (or series) adjacent to the dorsals fornia, Berkeley; MHNLS, Museo de Histo- is referred to as superolateral(s), whereas in- ria Natural La Salle, Caracas; RT, Richard ferolateral(s) applies to that scale (or series) Thomas private collection; TCWC, Texas adjacent to the ventrals. The terms pre- and A&M University; UF, University ofFlorida, postaxial, or lateral and median, refer to the Florida State Museum; UINHM, University side of the digit on which the lateral scale ofIllinois, Natural History Museum; UMMZ, occurs. University ofMichigan, Museum ofZoology; USNM, United States National Museum. The HYPoTHEsIs FORMULATION skeletal material for the outgroups is listed The best-fitting hypothesis of sphaerodac- in Kluge and Nussbaum (1995). tyl relationships was determined with the ie 1995 KLUGE: SPHAERODACTYL LIZARDS

2 mm 1 mm Fig. 6. Ventral view of the palate (reproduced from Kluge and Nussbaum, 1995). A. The plesiomorphic condition in gekkos. B. The apomorphic state, which is typical of Narudasia and Pristurus. m = maxilla; p = premaxilla; pa = palatine; v = vomer.

(implicit enumeration) command in cording to Kluge and Nussbaum (1995; their Hennig86 (Farris, 1988). Fit to data is as- character 29), the apomorphic state is re- sessed in terms of consistency and retention stricted to Narudasia and Pristurus (fig. 6). indices. Character consistency, c, is defined The plesiomorphic condition was illustrated as m/s, where s is the minimum number of originally by Kluge (1987: fig. 8). steps a character can exhibit on a given tree 2. Maxillary process of palatine long (0) hypothesis, and m is the minimum number or short or absent (1). A long maxillary pro- of steps that character can show on any tree cess, state 0, is similar in length to the vo- hypothesis (Kluge and Farris, 1969). Char- merine process of the palatine. acter retention, r, is defined as (g-s)/(g-m), 3. The choanal canal on the ventral surface where g is the greatest number ofsteps a char- of the palatine is short and shallow (0) or long acter can have on any tree (Farris, 1989). The and deep (1). Russell (1972) referred to the ensemble consistency, C, and ensemble re- similar palates ofsphaerodactyls, and he contention, R, indices used to choose among al- sidered Gonatodes to be a part of that New ternative phylogenetic hypotheses are simply World radiation largely because of that sim- the quantities for a single character, m, g and ilarity. Unfortunately, Russell did not specify s, summed over all characters in the matrix, the exact nature of the palatal similarity; thereby providing corresponding totals, M, however, the size and shape of the choanal G and S. Thus, C = M/S, and R = (G-S)/(G- canal seem to have been a part of his assess- M). ment. Polarity was realized with the outgroup op- tion in Hennig86 (Farris, 1988); Cnemaspis, 4. Splenial present (0) or absent (1). Ac- Narudasia, Pristurus, Quedenfeldtia, and cording to Kluge (1967, 1987), the splenial Saurodactylus were designated as outgroups is present in most lizards, including all gek- to the sphaerodactyl ingroup. The two mul- konids, except Pristurus, Ptyodactylus, and tistate characters, 1 1, 12, were treated as ad- sphaerodactyls. This is character 33 in Kluge ditive. Table 2 summarizes which character and Nussbaum (1995). Hecht and Edwards states have been attributed to the terminal (1976) used the absence of the splenial as taxa. evidence for sphaerodactyl common ancestry; however, they overlooked that same state CHARACTERS in Pristurus and Ptyodactylus, and the py- gopod Aprasia (Kluge, 1976). The two gek- 1. Maxillae separated or in narrow (0) or konids do not seem to share any derived states broad (1) contact posterior to premaxilla. Ac- (Kluge and Nussbaum, 1995), besides the ab- 10 AMERICAN MUSEUM NOVITATES NO. 3139 sence of the splenial, that could be used as 9. Vertebrae amphicoelous (0) or procoe- evidence oftheir common ancestry, and there lous (1). The apomorphic state is actually de- can be little doubt that Aprasia is part of an fined as well developed procoely (see Kluge, independent line ofevolution. The loss ofthe 1987: fig. 5). Noble (1921: fig. 1) briefly dis- splenial in sphaerodactyls and Pristurus may cussed the procoelous nature of some sphae- not be convergent; however, that interpre- rodactyls, and Underwood (1954: 476) im- tation will be judged later in this study, in plied that Gonatodes was strictly procoelous. the context of the analysis of all of the rele- 10. Number of attached sternal and xiph- vant available evidence and character con- isternal ribs more than four (0) or less than gruence. five (1). According to Kluge and Nussbaum 5. Coronoid tall (0) or short (1). The size (1995; their character 31), Narudasia and of the coronoid is assessed relative to the Quedenfeldtia exhibit state 1, Pristurus state general contour ofthe mandible, when viewed 0, and Saurodactylus is variable (0/1). All laterally. The tall state much exceeds that sphaerodactyls are recorded as state 1 (see contour, whereas the short condition is hard- also Noble, 1921: figs. 4,5); however, the fol- ly elevated above that outline. lowing individual variation was observed: one 6. Neural arch of atlas paired (0) or fused of the three Gonatodes albogularis fuscus (1). Kluge and Nussbaum (1995) observed specimens examined exhibited both states (their character 8) that the fused condition (UMMZ 127793); the one G. caudiscutatus occurs in all sphaerodactyls, Pristurus, some examined had both states; one oftwo G. tan- Quedenfeldtia (variable in Q. moerens), and iae examined possessed state 0 (UMMZ some Saurodactylus (single in S. fasciatus, 124303); one offour G. v. vittatus specimens paired in S. mauritanicus). The arch is single had state 0 (CR 453); the one Sphaerodac- in almost all Cnemaspis (C. affinus, C. afri- tylus altavelensis enriquilloensis studied ex- cana, C. indica, C. kandiana, C. nigridia, C. hibited both states; two of seven S. a. argus ornata, C. quatturoseriata, C. siamensis, and investigated possessed state 0 (UMMZ C. wynadensis). The paired state was ob- 127809, USNM 40510); the one specimen of served only in C. boulengerii, and possibly S. becki and S. corticola soter had both states; one individual of C. kendallii. Thus, I have one ofthree S. klauberi exhibited both states scored Cnemaspis as variable. The arch is (RT 3902); one oftwo S. lineolatus possessed paired in Narudasia. state 0 (UMMZ 63739); one of five S. mil- 7. Lateral projection of hyoid cornu pres- lepunctatus had both states (UMMZ 152733); ent (0) or absent (1). The projection is well one of eight S. molei exhibited state 0 (RT developed in Cnemaspis, Coleodactylus, 1191); one of three S. oxyrhinus possessed Gonatodes (absent in one of three specimens both states (UMMZ 85911); one of two S. of G. humeralis examined), Narudasia, Pris- roosevelti had state 0 (RT 4107); both S. ro- turus, Quedenfeldtia, and Saurodactylus. It is saurae exhibited state 0. variable in Lepidoblepharis-absent in one of 11. Fourth phalangeal element in fourth three L. microlepis (UMMZ 131865) and all finger long (0), short (1) or absent (2). I treat three L. s. sanctaemartae examined (the pro- this transformation series as additive (see fig. jection is well developed in both L. s. fugax 7). The apomorphic condition in the forefoot available). It is recorded as present in Pseu- ofPseudogonatodes appears to have been re- dogonatodes; however, it is not always well ported first by Gasc (1976: 22). developed. It is uniformly absent in Sphae- 12. Fourth phalangeal element in fourth rodactylus. toe long (0), short (1), or absent (2). I also 8. Second ceratobranchial arch present (0) treat this transformation series as additive or absent (1). According to Kluge and Nuss- (see fig. 7). baum (1995; their character 7), the arch is 13. One or more large fenestrae present (0) typical ofall ofthe taxa included in this study, or absent (1) in clavicle. Noble (1921: figs. except Cnemaspis and Pristurus. Noble (1921) 4,5; see also Parker, 1926) appears to have appears to have been the first to call attention been the first to draw attention to this vari- to the presence ofthe second ceratobranchial ation in sphaerodactyls. All Coleodactylus and arch in Sphaerodactylus, as well as its absence Pseudogonatodes studied have no conspicu- in such taxa as Cnemaspis. ous fenestrae (tiny openings may be present). 1995 KLUGE: SPHAERODACTYL LIZARDS

Length* of Fourth Element in the Fourth Finger Fourth Toe Gonatodes N = 35 Lepidoblepharis N = 12£ Sphaerodactylus N = 109 _

Coleodactylus absent

Pseudogonatodes absent absent

5 10 15 20 25 10 15 20 25 *% of total length of elements 2-5

Fig. 7. The length of the fourth phalangeal element in the fourth digit of the fore- and hindfoot of sphaerodactyls. The horizontal line is the observed range of variation; the vertical line is the mean; the dark rectangle is one standard deviation. N = sample size.

All of the representatives of Gonatodes and and the sternum (Noble, 1921: figs. 4,5). Long Lepidoblepharis examined have no discern- and short (or none) "arms" describe the two ible fenestrae, except for two of five G. an- states, 0 and 1, respectively. Some gekkos tillensis (UMMZ 127795a,b) and the single have nearly straight interclavicles, which ex- L. buchwaldi and L. intermedius examined, pand considerably at or posterior to their in- which have one or more tiny openings. While sertion into the sternum (e.g., Pristurus). the vast majority of Sphaerodactylus ob- These examples are recorded as state 1. Al- served have state 0, like the outgroups, there though I have scored Coleodactylus, Lepi- are several exceptions which cause me to rec- doblepharis, and Pseudogonatodes as state 1, ord that taxon as variable (0/1). With the the lateral margins ofthe interclavicle exhibit exception of the single S. corticola soter and a noticeable projection. Like the previous one ofthe 13 S. n. notatus examined (UMMZ character, there is obvious variation in 95583), almost all of this variation occurs Sphaerodactylus, which forced me to record among mainland endemics (Harris and Kluge, that taxon as variable (0/1). Moreover, al- 1984). For example, although all S. glaucus most all of the exceptional taxa are again (8 specimens) and S. heliconiae (1) have state mainland endemics. For example, all S. dun- 0, S. dunni (1), S. homolepis (2 of 2), S. mil- ni and S. molei clearly exhibit state 1, and lepunctatus (5 of 6; UMMZ 152733 is the the interclavicle arms in S. homolepis are exception), and S. molei (5 of 8; AMNH nearly equally short. All other mainland en- 15617-8 and RT 1191 are the exceptions) are demics, including S. millepunctatus, have usually state 1. Also, although the size of the much more cruciform-shaped interclavicles. fenestrae in both representatives of S. lineo- 15. Hypoischium long (0) or short or ab- latus and S. pacificus studied is not tiny or sent (1). Noble (1921: fig. 6) appears to have absent, the opening is much smaller than is been the first to draw attention to the much typical of most Sphaerodactylus species. reduced hypoischium in sphaerodactyls. 14. Interclavicle shape cruciform (0) or 16. Meatal closure muscle L, (0) or 0- (1) more daggerlike (1). The focus is on the lat- shaped. The muscle borders the posterior and eral margin of the interclavicle, which lies ventral margins of the external auditory me- approximately halfway between the clavicles atus in state 0, whereas the muscle encircles 12 AMERICAN MUSEUM NOVITATES NO. 3139

A B C D E

a .)

..s. -. -s b ..; .:-

,., .% C b

.5 mm Fig. 8. Lateral (a), dorsal (b), and ventral (c) views ofthe fourth toe ofrepresentative sphaerodactyls. A. Gonatodes. B. Lepidoblepharis. C. Pseudogonatodes. D. Coleodactylus. E. Sphaerodactylus. entirely, or nearly completely, that opening conclusion that similarly modified scales are in state 1. According to Kluge (1987: fig. 6; present in Quedenfeldtia. The type of gen- after Weaver, 1973, 1974, 1978), state 1 ap- eration gland has yet to be determined for plies to Pristurus and sphaerodactyls. Pristurus. If that gland is of the B type (see 17. Clutch size two or more (0) or one (1). Arnold, 1976: pl. 2) then there is additional Even though the modal values are obviously evidence for relating Pristurus to the sphae- two and one for states 0 and 1, respectively, rodactyl group. there is some variation (see Kluge, 1987). 19. Beta generation glands present (0) or 18. Escutcheon generation glands absent absent (1). This character was reviewed by (0) or present (1). Escutcheon glands are found Kluge (1983; see also Kluge, 1987; his char- in most sphaerodactyls (Maderson, 1972; acter 32). Menchel and Maderson, 1971), and similar 20. Supraciliary spine absent (0) or pres- holocrine epidermal modifications in the pre- ent (1). Parker (1926: 298; see also Russell, anal region have been observed in some (Co- 1972: 241) apparently was the first to em- leonyx brevis, C. variegatus, and Hemithe- phasize the possible phylogenetic informa- conyx taylorO), but not all, eublepharine gek- tiveness ofthis character when he stated that kos. If Maderson's (1972) conclusion is cor- the absence of a "supraciliary spine" distin- rect, that the escutcheon scale types in guished Coleodactylus from Sphaerodactylus. sphaerodactyls and eublepharine gekkos are Indeed, the spine is extremely large in almost not homologous, then this epidermal modi- all Sphaerodactylus (e.g., Harris and Kluge, fication provides additional evidence that 1984: fig. 3); however, smaller projections are sphaerodactyls form a natural group (Kluge, generally present in Gonatodes and some 1983, 1987; his character 33). However, the Lepidoblepharis. Spines appear to be absent weight of the generation gland evidence for in both Coleodactylus and Pseudogonatodes. monophyly also depends on how the absence Single spines also seem to be absent in all of of an escutcheon (state 0; Vanzolini, 1968b) the outgroups, except Saurodactylus (Bons in Coleodactylus and Pseudogonatodes is in- and Pasteur, 1957a: fig. 2). Although the en- terpreted, as plesiomorphic or an evolution- tire margin of the eyelid is spinose in Que- ary reversal. I have recorded state 1 for Gon- denfeldtia, I have recorded that taxon as state atodes, Lepidoblepharis, Pristurus, and 0, because no single projection is particularly Sphaerodactylus (Harris and Kluge, 1984: fig. long, as it is in sphaerodactyls. 15). I tentatively accept Russell's (1972: 182) 21. Supraciliary plate(s) absent (0) or pres- 1995 KLUGE: SPHAERODACTYL LIZARDS

ent (1). Vanzolini (1968b: 86) described some tests of homology (Rieppel, 1980a; see also sphaerodactyls as having "the anterior gran- Patterson, 1982). The omitted characters in- ules ... [of the eyelid] ... enlarged into flat clude several of those mentioned by Noble scales." Although such plates occur in most (1921: fig. 6), Underwood (1954: 470, 476, sphaerodactyls, there is considerable varia- 480, 488; 1970; see also Vanzolini, 1968b), tion among species of Gonatodes and Sphae- and Kluge (1967). These characters are: (1) rodactylus, and they are never particularly true eyelids absent, spectacle covering eye; large in any case. However, those plates are (2) parietals paired; (3) absence of vocaliza- especially noticeable in all other sphaerodac- tion; (4) presence ofsexual dichromatism; (5) tyls (see figures in Mechler, 1968), and I have temporal fovea present; (6) presence of oil recorded only these species as having state 1. droplets in visual cells; (7) premaxilla devel- 22. Ungual sheath of digit absent (0) or ops from single center of ossification, paired present (1). Russell (1972) stated that the condition absent from adults; (8) calcified en- sheath is present only in Coleodactylus, Lep- dolymphatic sacs present in postcranial re- idoblepharis, Pseudogonatodes, and Sphae- gion; (9) supratemporal absent; (10) scleral rodactylus, and Noble (1921) and Parker ossicles approximately 14 per eyeball; (11) (1926) also commented on the importance of angular absent; (12) frontal single; (13) nasals the ungual sheath in diagnosing that sub- paired; (14) squamosal rarely absent; (15) group ofsphaerodactyls (fig. 8). Noble (1921) pectineal process ofthe pelvic girdle well de- claimed the sheath merely represents an en- veloped. largement ofthe distalmost scales ofthe digit, such that they nearly completely surround the claw. 23. Distalmost superolateral scales in digit CLADISTIC RESULTS separated by dorsal scale (0) or in contact (1). There are three equally most parsimoni- This character applies generally to the fourth ous, best-fitting, hypotheses of sister-group toe (fig. 8). relationships for the characters summarized 24. Tip of fourth toe straight (0) or bent in table 2 (S = 36; C = 0.75; R = 0.83), with laterally (1). State 1 is also obvious in terms Cnemaspis, Narudasia, Pristurus, Queden- of the asymmetrical size and distribution of feldtia, and Saurodactylus included as out- the scales which make up the ungual sheath groups. All ofthe variable relationships occur (fig. 8). among three outgroup taxa, Cnemaspis, Na- 25. Pupil narrowly elliptical and conspic- rudasia, and Saurodactylus, and a strict con- uously emarginate (0) or more nearly circular sensus topology of those results is shown in and straight (1). Underwood (1954; see also figure 9. A posteriori iterative weighting did Parker, 1926) referred to the former state as not improve the resolution. the gekko-type. State 1 does include an oval Each resolved clade is corroborated by two pupil with slightly "puckered" margins (e.g., or more characters assumed to be indepen- S. parkeri, Underwood, 1954: 487; Sauro- dent pieces of evidence (fig. 9). Further, the dactylus, Bons and Pasteur, 1957a, 1957b). individual character consistency and reten- Other observations on pupil shape come from tion indices summarized in table 3 indicate Underwood (1970: 20), Werner (1977), Fran- that most of the diagnostic variables have kenberg (1979), and Thomas (1982). had unique and unreversed histories on the best-fitting hypothesis of sphaerodactyl re- OTHER lationships. The following diagnosis sum- OBSERVATIONS marizes the evidence: (Pristurus, sphaero- Several other characteristics have been used dactyls): splenial absent, meatal closure mus- in previous discussions of sphaerodactyl cle 0-shaped; (sphaerodactyls): maxillary monophyly or intergeneric relationships; process of palatine short or absent, choanal however, I have not employed them in this canal on ventral surface of palatine long and study because they are (1) plesiomorphic, (2) deep, fenestra absent in clavicle, hypois- too variable, (3) unknown for many taxa, chium short or absent, beta generation glands and/or (4) have not passed the preliminary absent; (Lepidoblepharis, Sphaerodactylus, 14 AMERICAN MUSEUM NOVITATES NO. 3139

0P I Op Sp

11(2),18(0)*, 23

Fig. 9. A strict consensus ofthe three equally most parsimonious cladograms for the data summarized in table 2 (S = 36, C = 0.75, R = 0.83). All ofthe variation in relationships occurs among the outgroups. The numbers refer to the diagnostic characters discussed in the text; state 1 can be assumed unless indicated otherwise in parentheses. The asterisk refers to a homoplastic state.

Coleodactylus, Pseudogonatodes): procoelous employed as evidence of intergeneric rela- vertebrae, ungual sheath present; (Sphaero- tionships. For example, based on toe pad sca- dactylus, Coleodactylus, Pseudogonatodes): lation, Noble (1921) hypothesized a single coronoid short, fourth phalangeal element in line of progression from Gonatodes to Lepi- fourth finger relatively short, fourth phalan- doblepharis to Pseudogonatodes to Sphaero- geal element in fourth toe relatively short; dactylus (the two Coleodactylus species rec- (Coleodactylus, Pseudogonatodes): fourth ognized at that time, C. amazonicus and C. phalangeal element in fourth finger absent, meridionalis, were included in Sphaerodac- escutcheon generation glands absent, distal- tylus), whereas Parker (1926) derived Pseu- most superolateral digit scales in contact. dogonatodes from Lepidoblepharis and, in- dependently, Sphaerodactylus from Lepido- HOMONOMY-HETERONOMY, blepharis by way ofa common ancestor with INDIVIDUAL VARIANTS, Coleodactylus (see also Vanzolini, 1968a). AND THE My reinterpretation ofthe evolution ofthe EVOLUTION OF THE individual scales ofthe ungual sheath, which UNGUAL SHEATH is to follow, assumes that the scalation The presence ofthe ungual sheath has long throughout much of the length of the free been recognized as diagnostic ofa large group digit, including those epidermal modifica- of sphaerodactyls, Coleodactylus, Lepidob- tions covering the ultimate phalangeal bone lepharis, Pseudogonatodes, and Sphaerodac- and the claw, form a homonomous series. My tylus (fig. 8). The sheath is believed to be references to homonomy are simply obser- absent only in Gonatodes. Further, numerous vational and have nothing to do with spec- authors have used the general shape of the ulations about possible underlying develop- digits and the terminal toe pad scalation to mental mechanisms for the iterated parts delimit each sphaerodactyl genus. These (Riedel, 1978; Roth, 1984, 1988; Wagner, characteristics of the digits have also been 1989a, 1989b). I use the similarity of a re- 1995 KLUGE: SPHAERODACTYL LIZARDS

TABLE 3a Individual Character Performances According to Figure 9 1 2 3 4 5 6 7 8 9 10 11 s 2 1 1 1 1 1 1 2 1 2 2 c .50 1.0 1.0 1.0 1.0 1.0 1.0 .50 1.0 .50 1.0 r .00 1.0 1.0 1.0 1.0 1.0 1.0 .00 1.0 .00 1.0 12 13 14 15 16 17 18 19 20 21 22 s 2 1 2 1 1 1 2 1 3 2 1 c 1.0 1.0 .50 1.0 1.0 1.0 .50 1.0 .33 .50 1.0 r 1.0 1.0 .66 1.0 1.0 1.0 .55 1.0 .00 .50 1.0 23 24 25 s 1 2 1 c 1.0 .50 1.0 r 1.0 .00 1.0 a s = number of steps; c = consistency index; r = retention index. peated series of elements, proximal to distal trageneric and individual variation. Lastly, oriented rows of digit scales in this study, as my reinterpretations conform to the pattern part of my preliminary test of similarity of of sister-group relationships provided by the particular scales in the ungual sheath (Riep- best-fitting hypothesis (fig. 10). The following pel, 1980a; Patterson, 1982). My use of ho- remarks pertain to the fourth toe, unless stat- monomy might be considered a special case ed otherwise; the terminology was outlined of two of Remane's (1952) similarity crite- under Material And Methods. ria-similarity in relative position, and sim- Gonatodes (Figs. 8, 10) exhibits little in- ilarity oftransitional form. The basic idea in terspecific variation in toe shape and scala- my application is that the similarity ofhighly tion. The digits are long, angulate, and com- differentiated repeated parts among organ- pressed throughout their length. The claw is isms can be judged in terms of the less- or well developed and exposed-the ungual undifferentiated parts of the comparable se- sheath is said to be absent (Russell, 1972). ries. It is the homonomy (the identity of re- There are single well defined rows of dorsals peated parts within and among organisms) and ventrals, between which are located two that serves as the basis in similarity for saying or three rows of smaller and more variably that heteronomous conditions or organisms shaped laterals. Of the 10 species surveyed, are part of the "same" synapomorphy rela- only G. ceciliae and G. taniae are character- tion. Thus, similarity of highly modified un- ized by three pairs of laterals (see Rivero- gual sheath scales in different sphaerodactyls Blanco, 1979, for additional details). The claw is not only assessed in terms ofthe similarity in Gonatodes is almost always surrounded by ofthe particular scales, but the series ofwhich two scales. I hypothesize the upper scale to they are assumed to be a part (fig. 8). I believe be a dorsal, at least in part, on the basis of this basis for assessing similarity will be even similar anatomical position and shape ofthe more important once the developmental ba- scute, relative to the homonomous dorsals sis for digital scale patterning is understood immediately proximate to it. The relatively (Hinchliffe and Johnson, 1980; Raynaud, larger size ofthe terminal dorsal may indicate 1985; Rosenberg et al., 1992). that it includes part of each distalmost su- My reinterpretation ofthe evolution ofthe perolateral; however, I have no other obser- individual scales of the ungual sheath also vations to corroborate that proposition. The takes advantage ofpreviously overlooked in- ventrally located scale surrounding the claw 16 AMERICAN MUSEUM NOVITATES NO. 3139

d d n~~~~~~~ v

l v+l.J dd d Pseudoonato3 n~~~~~~~e v dVv+ Pseudogonatodes

d d a I I

I I I

Lepidobl

I V+l Lepidoblepharis b nn II d

c n~~~~ Gonatodes

Fig. 10. Sphaerodactyl sister-group relationships (see fig. 8), with dorsal, lateral, and ventral views of the scalation of the fourth toe (see fig. 8). d = dorsal; 1 = lateral; n = claw (nail); v = ventral. is much taller than the homonomous ventrals and L. xanthostigma, have less angulate digits. immediately proximal to it. The additional Much of the claw in Lepidoblepharis is hid- height on each side is approximately that of den between two pairs of enlarged scales, an inferolateral. Rarely observed adult var- which I assume to be laterals (at least in part), iants with partial or complete lateroventral and a row oftwo dorsals, the distalmost being division of the most distal ventral supports tiny. The ungual sheath of Lepidoblepharis the hypothesis that the more ventrally locat- does not include a separate ventral. The in- ed scale surrounding the claw is a composite ferolaterals contact each other distal and (ventral + inferolaterals). Also, the distal- proximal to the claw, and their extreme depth most ventral is frequently notched anteriorly, may indicate that they have had ventrals fused and occasionally that ventral is completely to them during ontogeny. The individual divided into two symmetrical scales. These variation observed in Gonatodes (see discus- individual adult variants may also suggest sion immediately above) also supports my that the more ventrally located scale sur- contention that the inferolateral ofthe ungual rounding the claw includes more than one sheath is a composite. The fact that two dor- ventral (see discussion below). sals in a row form the ungual sheath may Lepidoblepharis (figs. 8, 10) species have indicate that the claw is covered laterally by undilated digits like Gonatodes; however, they two fused supero- and inferolaterals per side. tend to be much more round in cross section. The single alcoholic specimen of L. inter- Short-toed congeners, like L. sanctaemartae medius examined (UMMZ 127205) did not 1995 KLUGE: SPHAERODACTYL LIZARDS 17 have a single well defined row ofdorsals prox- tylus is more like that in Sphaerodactylus. imate to the ungual sheath. Mechler (1968: Given the same arguments that I used for fig. 17) illustrated L. colombianus as having Lepidoblepharis and Pseudogonatodes, I hy- two small rows of dorsals. pothesize that the inferolaterals ofthe sheath The remaining sphaerodactyl genus with in Coleodactylus are a composite, including undilated digits, Pseudogonatodes (figs. 8, 10), one or two ventrals. The enlarged, pilose scale is much more similar to Lepidoblepharis than in the sheath of Coleodactylus is almost cer- it is to Gonatodes. The digits of Pseudogon- tainly the postaxial ventral + inferolateral, atodes tend to be even shorter, stockier, and and the asymmetry ofthe digit appears to be less angulate than those of Lepidoblepharis. a simple function of the larger size of that The claw ofPseudogonatodes may be entirely member of the pair. The number of speci- hidden. The ungual sheath consists of a pair mens is not sufficient to evaluate Vanzolini's of large supero- and inferolaterals (probably (1957) scenario of digit evolution in the ge- a composite ofventral + inferolateral) and a nus, beyond concluding that the presence of single tiny dorsal distal to the claw. Unlike four scales in the ungual sheath of some Co- the ungual sheath of Lepidoblepharis and leodactylus species (Vanzolini, 1968a: fig. 4) Sphaerodactylus, the more posteriorly locat- is apomorphic. ed dorsal is absent in Pseudogonatodes. There Most Sphaerodactylus species have a great- is no evidence that the lack of a dorsal is a ernumber ofscales ofvarious sizes and shapes result offusion with the superolaterals. There forming the ungual sheath than do other is considerable variation in Pseudogonatodes sphaerodactyls. In addition, the size of the in number of laterals proximal to the ungual sheath in most Sphaerodactylus is much larg- sheath: one or two pairs were observed in P. er and more markedly asymmetrical. These barbouri, two in P. lunulatus, and three in P. species have made it particularly difficult to furvus. Moreover, the latter species did not identify the homonomy and heteronomy re- have a single well defined row ofdorsals. Huey lations to the ungual sheath scales, which in and Dixon (1970) apparently overlooked turn makes individual scale comparisons be- these interspecific differences. tween representatives of different sphaero- Coleodactylus and Sphaerodactylus (figs. 8, dactyl genera even more problematical. In 10) resemble each other in terms of the di- order to minimize the effects of extreme dif- lation and asymmetry oftheir ungual sheaths. ferentiation, I have relied mostly on species, However, the best-fitting hypothesis ofsister- like S. klauberi, which have relatively straight group sphaerodactyl relationships suggests and undilated digits. I assume that these taxa (optimization ofthe characters in question is are plesiomorphic relative to those with ambiguous) that those two features of size markedly bent and terminally wide digits. and shape might have evolved independent- Usually, the ungual sheath ofSphaerodac- ly. A detailed comparison of the individual tylus consists ofan extremely large pad, a row scales ofColeodactylus, Pseudogonatodes, and oftwo dorsals (Noble, 1921, claimed the more Sphaerodactylus adds support to the homo- proximal dorsal was a lateral; however see plasy hypothesis. Parker, 1926: 300), and two or three rows of The ungual sheath of at least some species laterals. The superolateral scale in the sheath ofColeodactylus (C. meridionalis and C. sep- is much shorter than the inferolateral, which tentrionalis; Vanzolini, 1968a: fig. 3, and I assume is a developmental composite (in- Hoogmoed, 1985: fig. 2, respectively) concluding a ventral; see previous arguments for sists of five scales, which nearly cover the Lepidoblepharis and Pseudogonatodes). The entire claw. I interpret these to be a single preaxially located composite is nearly verti- dorsal and two pairs of laterals. This pattern cal, with its median counterpart almost hor- is the same as that observed in Pseudogon- izontal. At least in some species of Sphae- atodes-the more proximate dorsal of the rodactylus, such as S. klauberi, the horizontal sheath is assumed to be absent, while the plate appears to cover another equally large more distal dorsal is present. However, the scale completely; this second horizontal scute small size of the superolateral in Coleodac- is pilose and forms the conspicuous pad of 18 AMERICAN MUSEUM NOVITATES NO. 3139 the ungual sheath. This interpretation applies 14.0 mm, tail length about 13.6 mm. The most obviously to species like S. klauberi specimen is mostly cleared, and is entirely where there is relatively little asymmetry. surrounded by a reddish "halo," which is typ- Moreover, in species like S. klauberi, there ical of some Dominican amber, particularly is a small scale without pilosity located at the those with pyritized inclusions. The lizard postaxial, proximal border of the pad, which itselfis not pyritized. The piece had originally I interpret to be the counterpart to the larger been cracked and appears to have been re- ventral. Thus, I am not only hypothesizing paired with glue. The fracture extends through that the pilose pad of Sphaerodactylus is a the specimen, fromjust behind the left foreleg ventral, but that the pad is the preaxial ven- to the base of the right foreleg. tral. This scenario of the asymmetrical evo- The right side of the head is occluded by lution ofthe pilose pad in Sphaerodactylus is some fractures and by black and white gran- consistent with the other less proximate ular debris, perhaps a product of decompo- homonomous ventrals being either notched sition. The left side ofthe head is most visible or completely divided. Such a subdivision of (fig. 1 1). A small part ofthe tongue protrudes the ventral has also been observed in Gon- from the base of the left side of the mouth, atodes, Lepidoblepharis, and Pseudogona- with a small drop "hanging" from the tip. A todes. I hypothesize that all other small scales small bubble protrudes from the tip of the at the proximal pre- and postaxial margins mouth. The eyes are very large, partly sunk- of the pilose ventral in Sphaerodactylus are en, and point forward. The face and nose are inferolaterals. Variation in the rows of scales short; the cranial portion of the head just proximal to the ungual sheath (4-6, mode of behind the eyes is high and dome-shaped. 5; see also Parker, 1926) and the absence of Scalation on the throat is observable, but the a well defined row of dorsals at the base of whitish granular substance (presumably de- the ungual sheath make this proposition cayed tissue) on the remainder of the head highly speculative. The study of ontogenetic obscures the scalation. series may provide the evidence necessary to The trunk, abdomen, legs, tail, and feet are test this hypothesis because Sphaerodactylus mostly cleared, some small portions with tis- is known to undergo considerable transfor- sue debris in them. One can see scalation over mation of digital shape during ontogeny most ofthe body when properly reflected and (Werner, 1971). In summary, I propose that transmitted fiber-optic light is employed. The the pilose pad is a modified preaxial ventral dorsal body scales are small and only slightly in Sphaerodactylus, whereas the pilose part imbricate. Curiously, there is no evidence of of the sheath is assumed to be the postaxial bone in the parts that are most transparent ventral + inferolateral in Coleodactylus. If (e.g., feet, legs, tail). Perhaps this is a juvenile this interpretation is correct, then there is specimen, with little ossification, and the soft additional evidence for the independent evo- bones decayed with the other tissues. The lution of the dilation and asymmetry of the right foreleg and foot are bent back against digit in Coleodactylus and Sphaerodactylus the trunk; the left foreleg is forward, with the (fig. 10). foot flat and the toes splayed. Both hind legs stick out from the body. The toes are clearly A GEKKO IN AMBER visible. The trunk and tail are filled with a pocket of air, but the posterior third of the A dark, clear yellow piece of amber was tail is not so obscured. The tail is complete, forwarded to David A. Grimaldi, American and covered below with enlarged ventrals. Museum of Natural History, by Ms. Susan Air pockets occur at the tips of some toes; Hendrickson, who acquired it in the Domin- however, details offoot scalation are evident ican Republic. It is a slightly asymmetrical (fig. 1 1). There is an ungual sheath and the tear-shaped piece, 53 mm long by 11 mm tip ofeach digit is obviously asymmetrical in wide at the broadest point, with some deep shape (figs. 8, 10). There are approximately longitudinal scratches. The piece contains an 13 ventrals covering the fourth toe, two of entire specimen of a tiny lizard, SVL about which are paired at the origin of digit. 1 995 KLUGE: SPHAERODACTYL LIZARDS 19

.5 mm Fig. 1 1. Illustrations of AMNH lizard preserved in Oligo-Miocene Dominican amber (provided by D. A. Grimaldi). A. Left lateral view of the head. B. Dorsal view of right forefoot. C. Ventral view of left forefoot. D. Ventral view of left hindfoot.

Given the presence of a conspicuously doblepharis and Sphaerodactylus), 27 and 37 asymmetrical ungual sheath (fig. 11), there mybp, respectively, are too low. This dis- can be little doubt that the AMNH gekko in crepancy may be due to the evolutionary sat- amber is a Sphaerodactylus. The small size uration ofthe genes responsible for the clock. of the dorsal body scales, and their slightly Judging from the description of Sphaero- imbricate nature, are conditions similar to dactylus dommeli (B6hme, 1984), also known those of other Hispaniolan Sphaerodactylus, only from Dominican amber, the AMNH such as S. callocricus, S. cinereus, S. elegans, specimen is certainly representative of a dif- and S. samanensis (Henderson and Schwartz, 1984). Unfortunately, I cannot be certain that ferent species. In fact, the general habitus the fossil exhibits any autapomorphies, and (proportions of limbs, shape of head) of S. therefore I cannot determine whether it rep- dommeli suggests that it is probably an an- resents a new species. That it is a highly de- oline lizard (Darrel Frost, personal com- rived form is suggested by its similarity to mun.). The better photograph of S. dommeli species in the cinereus species-group (Hass, (of the paratype specimen in the Staatlisches 1991). That such a derived Sphaerodactylus Museum in Stuttgart, Schlee, 1980; 1990) also fossil is believed to be of Oligo-Miocene age corroborates this tentative placement. The (23-30 mybp; Grimaldi, 1995) seems to in- type material of"Sphaerodactylus" dommeli dicate that Hass' (1991: 547; see also Hedges must be reexamined in order to correctly re- et al., 1992) albumin clock estimates for times classify the species; comparison with Riep- of origin for sister lineages Lepidoblepharis pel's (1980b) green anole from Dominican and Sphaerodactylus, and Gonatodes (Lepi- amber is essential. 20 AMERICAN MUSEUM NOVITATES NO. 3139

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APPENDIX Cleared and Stained Specimens Examined Coleodactylus amazonicus (UMMZ 127803a,b); Pseudogonatodes barbouri (UMMZ 127807,8); C. brachystoma (UMMZ 144467); C. meridionalis P. guianensis guianensis (KU 121951, TCWC (UMMZ 10305 la,b). 41231,2); P. lunulatus (MCZ 48894, UMMZ Gonatodes albogularis albogularis (CR 538, 124312); P. peruvianus (LSUMZ 27359, MVZ UMMZ 127790a,b); G. a. fuscus (UMMZ 82137). 127792,3,148120); G. a. notatus (UMMZ 127794); Sphaerodactylus altavelensis brevirostratus G. annularis (UMMZ 53894); G. antillensis (ASFS 44073); S. a. enriquilloensis (RT 771); S. (MHNLS 2231, UMMZ 57325,6, 127795a,b); G. argivus argivus (UMMZ 143257); S. argus argus atricucullaris (TCWC 28333, UMMZ 127797a,b, (KU 157180, UMMZ 127809a-c, USNM 40510, 127798); G. caudiscutatus (MCZ 74175); G. ce- 192526); S. armstrongi (AS 30444); S. beattyi ciliae (MCZ 79808); G. concinnatus (CR 1949, beattyi (UMMZ 143256); S. becki (AS 6277); S. MCZ 77390); G. hasemani (MCZ 119418); G. hu- caicosensis (UMMZ 143263); S. callocricus (UF meralis (CR 1096, UMMZ 127800, 128141); G. 21559, USNM 41252); S. cinereus cinereus ocellatus (UMMZ 127801); G. seigliei (CR 1001); (AMNH 49566, RT 4108, UMMZ 95582a-j, G. taniae (CR 900, UMMZ 124303); G. vittatus 127810,11); S. clenchi (AS 34334); S. copei ca- vittatus (CR 453, UMMZ 54687, 54893, taplexis (UMMZ 143260); S. corticola soter 127802a,b); G. sp. nov. (MCN 3740). (UMMZ 143259); S. darlingtoni noblei(AS 28627); Lepidoblepharis buchwaldi (MCZ 145149); L. S. difficilis (RT 1418); S. dunni (LACM 47302); festae(LACM 44678); L. intermedius(CAS 13260); S. elegantulus (UMMZ 143255); S. fantasticus L. microlepis (UMMZ 127804,5, 131865,6); L. fantasticus (AS X5385); S. gaigeae (UMMZ 73606, sanctaemartae (UMMZ 125043, 125045, 143254); S. glaucus (UMMZ 70448, 143261, 127806a,b). 151524, 151529-32, USNM 113126); S. gonior- 1995 KLUGE: SPHAERODACTYL LIZARDS 23 hynchus (UMMZ 127812, USNM 42054); S. hel- oi (UINHM 44223); S. n. gibbus (UMMZ 127823, iconiae (ICN 3217, 3226, UMMZ 171649) S. 117018); S. notatus notatus (UMMZ 95583a-j, homolepis (UF 31305, 37188, UMMZ 127818); 148121); S. n. exsul((USNM 23361-3, 23358); S. S. inaguae (UMMZ 127815); S. klauberi (RT 3902, oxyrhinus oxyrhinus (ASFS 20050, UMMZ 85911, UMMZ 73594, 143252); S. lineolatus (UMMZ 143258); S. pacificus (UMMZ 127819a,b); S. par- 63738,9); S. macrolepis macrolepis (UMMZ keri (UMMZ 127820,1); S. randi randi (ASF 127816a,b); S. mariguanae (UMMZ 127817, V39837); S. roosevelti (ASFS 4290, RT 4107); S. 148123); S. microlepis microlepis (UMMZ rosaurae (LSUMZ 22385, UF 28557); S. sabanus 143250); S. millepunctatus (KU 157110, LACM (ASFS 19776); S. savagei savagei (ASFS 35019, 47784, UMMZ 117873,143262,151525,151527, USNM 40973,4); S. sputator (ASFS 19956); S. 152733); S. molei (AMNH 15616-21, RT 1191, streptophorus streptophorus (ASFS 42503); S. tor- UMMZ 65168); S. monensis (UMMZ 143253); S. rei torrei (AS V15725); S. underwoodi (ASFS nicholsi (UMMZ 143251); S. nigropunctatus alay- 10934); S. vincenti vincenti (ASFS 18139). Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates, Bulletin, and Anthropological Papers published during the last five years are available free of charge. Address orders to: American Museum ofNatural History Library, Department D, Central Park West at 79th St., New York, N.Y. 10024. TEL: (212) 769-5545. FAX: (212) 769- 5009. E-MAIL: [email protected]

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