GECKO CLASSIFICATION 



117 



morphology (Bauer and Russell 1995). In addition, Cosymbotus, 

 Briba and Teraiolepis are also very similar and are almost certainly 

 share a common ancestry with Hemidactylus, or are derived from 

 within it. 



Bauer (1990b) found some evidence for the recognition of a 

 Madagascan radiation including several genera of leaf and fan-toed 

 geckos including Uroplatus, Ebenavia and Paroedura. Kluge and 

 Nussbaum (1995) did not retrieve identical patterns of relationship, 

 but these genera nonetheless grouped closely when only Afro- 

 Malagasy geckos were included in the analysis. An expanded Indian 

 Ocean lineage, including these taxa plus Ailuronyx, Blaesodactylus, 

 Homopholis, and Geckolepis was retrieved by Bauer (1990b), al- 

 though not by Kluge and Nussbaum (1995). 



Another putatively monophyletic group is the Lygodactylus com- 

 plex (Pasteur 1964), which includes two additional genera, at least 

 one of which, Millotisaurus, is probably derived from within 

 Lygodactylus (Pasteur, 1995; Kriiger, 2001). Lygodactylus itself 

 clustered with Phelsuma in analyses constrained to include only 

 Afro-Malagasy genera (Bauer 1990b; Kluge and Nussbaum 1995). 

 Kriiger (2001 ) also clustered Lygodactylus and Phelsuma together. 



Although some genera have been revised at the alpha level, and 

 numerous new taxa have been erected, most revisions have merely 

 proposed species groups, without providing explicit hypotheses of 

 relationship (e.g. Pasteur 1964; Brown and Parker 1977; Nussbaum 

 and Raxworthy 2000). These, like many of the other groups, share 

 digital similarities and geographic cohesiveness. Among those gen- 

 era for which some idea of relationships exist, there are several for 

 which species level phylogenies have been proposed, including 

 Uroplatus (Bauer and Russell 1989) and Gchyra (selected species 

 only; King 1979, 1983). 



Rhoptropella has been associated with several different genera by 

 different authors. Russell (1977b) used digital morphology to argue 

 that it was in fact a Phelsuma, with no direct affinities to Rhoptropus, 

 with which it had previously been associated (e.g. Boulenger 1 885). 

 Russell and Bauer (1990) found additional support for this from 

 histological investigations and Good and Bauer (1995) presented 

 allozyme evidence for Rhoptropella' s links to Phelsuma. Both 

 Bauer (1990b) and Kluge and Nussbaum (1995) found the two 

 genera to be sister taxa when a generic analysis was conducted. 

 Rosier (2001 ), discussing pholidosis, also concluded that Phelsuma 

 and Rhoptropella are sister taxa. Roll (1999), however, using oph- 

 thalmological and digital surface data, interpreted it as displaying 

 features of both Rhoptropus and Phelsuma, which, if true, could 

 suggest affinities between the chiefly African Pachydactylus group 

 and the putatively monophyletic Indian Ocean complex. A variety 

 of character types also suggest that Bogertia and Thecadactylus may 

 be allied (Russell and Bauer 1988; Abdala and Moro 1996). 



Cnemaspis, Narudasia, Quedenfeldtia. Saurodactylus and 

 Pristurus have been proposed as gekkonine taxa basal to the 

 sphaerodactyl lineage (Arnold 1993; Kluge 1995), demonstrating 

 the paraphyly of the Gekkoninae. Although Kluge (1995) did not 

 claim any specific relationships among these taxa, his analysis did 

 yield patterns in which Pristurus was the sister group of the 

 sphaerodactyls, and Narudasia, Saurodactylus and Cnemaspis 

 formed a clade. Arnold (1993) advocated the pattern ({((Pristurus, 

 Quedenfeldtia) sphaerodactyls) Saurodactylus) Narudasia). Behav- 

 ioural apomorphies unique to this cluster were documented by 

 Rosier and Wranik (2001), who noted reproductive morphological 

 apomorphies shared by Quedenfeldtia and the sphaerodactyls to the 

 exclusion of Pristurus. Arnold (1993) provided a species level 

 phylogeny for Pristurus. The African members of this group were 

 also clustered together in an anlysis of Afro-Malagasy taxa by Kluge 

 and Nussbaum (1995). Roll and Schwemer (1999) identified a 



unique crystallin ligand common to several of these taxa (plus 

 Lygodactylus), that they interpreted as synapomorphic. This was 

 subsequently found in Cnemaspis (Roll, in press), but whether this 

 indicates affinity or convergence among secondarily diurnal forms 

 remains to be determined. 



The naked-toed geckos have proved especially difficult to deal 

 with. Szczerbak and Golubev (1984, 1986) provided evidence of 

 relationship among some Palearctic forms, such as Tenuidactylus, 

 Mediodacrylus, Asiocolotes, and Cyrtopodion. Macey et al. (2000) 

 found evidence for the monphyly of Cyrtopodion and Mediodacrylus 

 and hypothesized relationships among a small number of species in 

 each group. The generic allocation of certain Himalayan members 

 of the group has proved especially problematic (Khan 1993; Khan 

 and Rosier 1999). 



Another group of naked-toed geckos including Agamura, Bunopus, 

 Alsophyla.x, Crossobamon, Microgecko, and Tropiocolotes has been 

 even less well investigated (Leviton and Anderson 1972; Szczerbak 

 and Golubev 1977; Golubev 1984; Golubev and Szczerbak 1985). 

 The New World naked toed forms, Gymnodactylus and Homonota, 

 have been included in analyses by Abdala (1996) and Abdala and 

 Moro ( 1 996) but these investigations included only South American 

 gekkonines. Abdala ( 1988) also provided a species level phylogeny 

 for Homonota (see also Vanzolini 1968). 



While some degree of resolution for the gekkonine taxa outlined 

 above has been reached, certain other gekkonines remain enigmatic 

 and without any sound indication of affinities. Teratoscincus is 

 highly unusual in its morphology, and appears to be the sister group 

 of all remaining gekkonines ( Kluge 1 987). A species level phylogeny 

 for this group has been generated (Macey et al. 1 999). Stenodactylus 

 has sometimes been considered to be allied to Teratoscincus (Kluge 

 1 967a; Kluge and Nussbaum 1 995 ). but its position remains equivo- 

 cal (Arnold 1980). 



Another perplexing padless genus is Ptenopus, a southern African 

 endemic. Both Bauer (1990b) and Kluge and Nussbaum (1995) 

 found little evidence for particular affinities, and constrained or 

 retrieved a basal placement among African gekkonines. Ptenopus 

 possesses a large number of autapomorphic traits (Haacke 1975; 

 Rittenhouse et al. 1998; Russell et al. 2000). This mirrors the 

 situation that plagued analyses of Uroplatus in that many features 

 segregate these geckos from other taxa, but those traits that are 

 shared are chiefly primitive ones. 



Four pad-bearing genera, which appear unrelated to one another 

 and have no obvious affinities to previously discussed groups, are 

 also problematic. These are Afroedura, Aristelliger, Calodactylodes, 

 and Paragehyra. Paragehyra was long known from a single speci- 

 men of a single species, but a second species was recently discovered 

 (Nussbaum and Raxworthy 1994). The availability of additional 

 material allowed the relationships of the genus to be investigated in 

 more detail, but this has not yielded any definitive statements about 

 its position within the Gekkoninae (Kluge and Nussbaum 1995), 

 although Nussbaum and Raxworthy (1994) noted the similarity of 

 the digits of this form to those of another enigmatic taxon, the West 

 Indian Aristelliger. 



Russell (1972) grouped Afroedura and Calodactylodes in the 

 same digitally defined cluster. Loveridge (1944) had initially segre- 

 gated Afroedura from the Australian Oedura, and this was reflected 

 in Underwood's (1954) placement of the genera in different sub- 

 families. Some question as to the distinctiveness of these taxa 

 remained, however, until Cogger (1964) conducted detailed osteo- 

 logical comparisons. Despite some similarities in digital design, 

 Russell and Bauer (1989) concluded that Calodactylodes and 

 Afroedura were more likely convergent than related. Bauer and Das 

 (2000) noted some superficial similarity and geographic proximity 



