Accepted by S. Carranza: 20 Nov. 2012; published: 8 Jan. 2013
ISSN 1175-5326 (print edition)
1175-5334 (online edition)
Copyright © 2013 Magnolia Press
Zootaxa 3599 (4): 301–324
A preliminary phylogeny of the Palearctic naked-toed geckos (Reptilia:
Squamata: Gekkonidae) with taxonomic implications
AARON M. BAUER
, RAFAQAT MASROOR
, JAMES TITUS-MCQUILLAN
, MATTHEW P. HEINICKE
JUAN D. DAZA
& TODD R. JACKMAN
Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA
Zoological Sciences Division, Pakistan Museum of Natural History, Garden Avenue, Shakarparian, Islamabad 44000, Pakistan
current address: Department of Natural Sciences, University of Michigan-Dearborn,125 Science Building, 4901 Evergreen Road,
Dearborn, Michigan 48128, USA
Palearctic naked-toed geckos are a group of gekkonid geckos that range from North Africa to northern India and western China,
with their greatest diversity in Iran and Pakistan. Relationships among the constituent genera remain incompletely resolved and
the monophyly of key genera remains unverified. Further, competing classifications are in current use and many species have
been allocated to different genera by different authors. We used both mitochondrial (ND2) and nuclear genes (RAG1, PDC) to
explore relationships among representatives of all but one genus in the group (Rhinogecko), including four genera not
previously included in phylogenetic analyses (Asiocolotes, Altigekko, Indogekko, and Siwaligekko). Siwaligekko (and
presumably other Tibeto-Himalayan species often referred to Cyrtopodion) are more closely related to tropical Asian
Cyrtodactylus than to Palearctic naked-toed geckos. Sampled species of Asiocolotes and Altigekko are sister taxa, but both
genera are here considered junior subjective synonyms of Altiphylax. Cyrtopodion sensu lato is non-monophyletic;
Mediodactylus and Tenuidactylus, which have variably been considered as subgenera or synonyms of Cyrtopodion are both
valid genera. Indogekko is embedded within Cyrtopodion and is here treated as a subgenus. Bunopus and Crossobamon are
closely related to one-another, and with Agamura are interdigitated among taxa previously assigned to Cyrtopodion. Our data
confirm the previous identification of a Saharo-Arabian Stenodactylus/Tropiocolotes/Pseudoceramodactylus clade and verify
that Microgecko and Alsophylax are not members of the main clade of Palearctic naked-toed geckos. Osteological differences
between Tropiocolotes and Microgecko, formerly treated as congeneric, are discussed and illustrated. The divergence between
Cyrtodactylus and the Palearctic naked-toed clade predates the initial collision of the Indian and Eurasian plates, but deeper
divergences within both groups are consistent with mountain building in the Himalayas and adjacent ranges as promoting
cladogenic events. Miocene divergences within Tenuidactylus are consistent with vicariant speciation caused by uplift events in
the Iranian and Transcaspian regions. Taxonomic implications of our phylogenetic results are discussed and a preliminary
allocation of all species of padless Palearctic gekkonids to genus is provided.
Key words: Cyrtopodion, Tenuidactylus, Mediodactylus, Indogekko, Altigekko, Altiphylax, Siwaligekko, Asiocolotes, Bunopus,
Crossobamon, Tropiocolotes, Stenodactylus, Pseudoceramodactylus, Cyrtodactylus, Microgecko, Alsophylax,
biogeography, Himalayas, timetree
“Palearctic naked-toed geckos” including the so-called “angular-toed geckos,” are a large group (101 species) of
gekkonid geckos distributed from North Africa across southwestern and Central Asia to northern India, western
China, and southern Mongolia (Fig. 1), united by their shared lack of adhesive subdigital pads. This group includes
taxa that have variously been assigned to the genera Agamura Blanford, 1874, Alsophylax Fitzinger, 1843, Altigekko
Khan, 2003c, Altiphylax Jerem
enko & Szczerbak, 1984, Asiocolotes Golubev, 1984, Bunopus Blanford, 1874,
Carinatogecko Golubev & Szczerbak, 1981, Ceramodactylus Blanford, 1874, Crossobamon Boettger, 1888,
Cyrtodactylus Gray, 1827, Cyrtopodion Fitzinger, 1843, Garzoniella Perret, 1976, Gonyodactylus Kuhl & van
Hasselt, 1822, Gymnodactylus Spix, 1825, Indogekko Khan, 2003c, Mediodactylus Szczerbak & Golubev, 1977,
Mesodactylus Szczerbak & Golubev, 1984, Microgecko Nikolsky, 1907, Pseudoceramodactylus Haas, 1957,
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302 · Zootaxa 3599 (4) © 2013 Magnolia Press
Rhinogecko de Witte, 1973, Siwaligekko Khan, 2003c, Stenodactylus Fitzinger, 1826, Tenuidactylus Szczerbak &
Golubev, 1984, Trachydactylus Haas & Battersby, 1959, Trigonodactylus Haas, 1957, and Tropiocolotes Peters, 1880.
The taxonomy of these geckos, despite decades of assiduous research by herpetologists around the world,
remains poorly resolved and unstable (Auffenberg et al. 2010; Ahmadzadeh et al. 2011), partly because many
genera are delimited by qualitative characters, including degree of tuberculation and angle of toes. These characters
do distinguish several morphological groupings of genera: straight- to slightly bent-toed, weakly tuberculate
geckos with long, thin limbs and tail (Agamura, Rhinogecko); straight-toed, atuberculate geckos (Asiocolotes,
Microgecko, Tropiocolotes); straight- to slightly bent-toed, variably tuberculate geckos with enlarged nasal scales
(Alsophylax, Bunopus); straight- to slightly bent-toed, weakly tuberculate geckos with scales forming lateral
fringes on toes (Crossobamon, Pseudoceramodactylus, Stenodactylus); distinctly bent-toed, variably (often
heavily) tuberculate geckos (Carinatogecko, Cyrtodactylus, Cyrtopodion, Indogekko, Mediodactylus, Siwaligekko,
Tenuidactylus). In many cases, generic assignment of species within these rough groupings has been problematic,
and some Palearctic naked-toed geckos are intermediate between two or more of these groups. For example,
Siwaligekko is intermediate in morphology between Cyrtodactylus and Cyrtopodion, species typically assigned to
Altigekko or Alitphylax are basically intermediate in morphology between Alsophylax and Cyrtopodion, and
Asiocolotes has been alternately considered congeneric with Tropiocolotes and Altiphylax. Further, based on both
results of previous phylogenetic analyses (e.g., Macey et al. 2000;
ervenka et al. 2008) and osteological
differences, most or all of these rough morphological groupings are not monophyletic. Much of the convoluted and
anastomosing taxonomic and nomenclatural history of these geckos has been reviewed by Szczerbak & Golubev
(1986, 1996), Anderson (1999), and Krysko et al. (2007). While we do not wish to repeat what these authors have
already painstakingly detailed, a certain minimal review of selected issues in the group is necessary to place the
taxonomic chaos of these geckos into perspective.
FIGURE 1. Composite distribution of the Palearctic naked-toed geckos. The genera Alsophylax and Microgecko, which fall
outside this clade (Gamble et al. 2012), are not included, but have largely overlapping distributions. Shading of the base map
depicts elevation, with high-elevation areas darker. Key geographic and physiographic regions, including countries, mountains,
plateaus, deserts, and tectonic plate boundaries, are labeled.
Microgecko and Asiocolotes have variously been regarded as subgenera within Tropiocolotes (Minton et al.
1970; Golubev 1984; Szczerbak & Golubev 1986, 1996) or as full genera (Kluge 1983, 1991, 1993, 2001); recently
Asiocolotes has also been treated as a junior synonym of Altiphylax (Sindaco & Jerem
enko 2008). Both
Tropiocolotes and Bunopus have had species reassigned to Carinatogecko (Golubev & Szczerbak 1981), which has
itself been synonymized with Mediodactylus (
ervenka et al. 2010). Bunopus and Alsophylax have been at various
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PALEARCTIC NAKED-TOED GECKO PHYLOGENY
times confused with one another or considered as synonyms (see Leviton & Anderson 1963; Mertens 1965; Clark
et al. 1969; Szczerbak & Golubev 1977, 1986, 1996; Anderson 1999) and both have subsumed species previously
assigned to Crossobamon or Stenodactylus. Altiphylax was erected as a subgenus of Alsophylax by Jerem
Szczerbak (1984), with Alsophylax tokobaejvi Jerem
enko & Szczerbak as its type species. However, a
subsequently allocated species, A. (Altiphylax) boehmei Szczerbak, has since been synonymized with Cyrtopodion
stoliczkai (Steindachner) (Auffenberg et al. 2004), implying that at least part of Altiphylax is inappropriately
included within Alsophylax.
Stenodactylus, which is relatively easily characterized by its digital morphology (digits with reduced
phalangeal complement and with lateral fringes), has remained otherwise relatively stable following the
synonymization of Ceramodactylus, Trigonodactylus, and Pseudoceramodactylus by Kluge (1967), with the
exception of the recent resurrection of the last of these genera for S. khobarensis Haas (Fujita & Papenfuss 2011).
Agamura and Rhinogecko have been considered as distinct from one another (Anderson 1999; Krysko et al. 2007;
Khan 2006; Sindaco & Jerem
enko 2008) or as synonymous (Szczerbak & Golubev 1986, 1996; Kluge 1991,
1993, 2001) and Gymnodactylus gastropholis Werner has been allocated either to Agamura (Minton 1966) or to
Cyrtopodion (Szczerbak & Golubev 1986; Anderson 1999; Krysko et al. 2007).
The greatest confusion concerns a core group of bent-toed geckos with their distribution chiefly in Central
Asia, northern Pakistan, and adjacent India. Until the mid-20
century most padless geckos that lacked obvious
digital apomorphies such as lateral scaly fringes or tuberculate lamellae were lumped together in the large genus
Gymnodactylus. Annandale (1913) initially recognized distinctions among the Asian geckos then assigned to
Gymnodactylus, allocating them to five different informal groups. Following the subdivision of the genus by
Underwood (1954), both Palearctic and tropical Asian “Gymnodactylus” were assigned to the genus Cyrtodactylus
with Gymnodactylus restricted to some New World geckos now allocated to the Phyllodactylidae.
Szczerbak & Golubev (1984) removed from Cyrtodactylus 14 Palearctic species that differed in a suite of
characters, including overall body form, tail segmentation, and degree of tuberculation, from the chiefly tropical
Asian forms, erecting for them the genus Tenuidactylus. Within this genus they recognized three subgenera, the
nominotypical Tenuidactylus with Gymnodactylus caspius Eichwald as its type species, Mediodactylus, with
Gymnodactylus kotschyi Steindachner as its type, and Mesodactylus, with Gymnodactylus kachhensis Stoliczka as
its type. Böhme (1985) and Kluge (1985) subsequently noted that an older name applicable to this group,
Cyrtopodion (type species Stenodactylus scaber Heyden) had been overlooked. Szczerbak & Golubev (1986)
consequently presented a new species allocation, in which Mesodactylus was regarded as a junior subjective
synonym of Cyrtopodion, although they continued to treat Cyrtopodion as a subgenus of Tenuidactylus. Not
assigned to subgenus by Szczerbak & Golubev (1984, 1986) were members of the “Tibeto-Himalayan” group of
naked-toed geckos, a heterogeneous group of taxa that were geographically peripheral to these authors’ main area
Subsequent work by Khan (1988, 1991, 1993a, 1993b; Khan & Tasnim 1990) and others (Zhao & Li 1987;
Baig 1998) resulted in the discovery of many new bent-toed gecko species from northern Pakistan and adjacent
regions. Those described by 1991 were placed, along with Szczerbak & Golubev’s (1986) Tibeto-Himalayan
group, in the genus Gonyodactylus by Kluge (1991), who had earlier (1985) recognized this as a senior objective
synonym of Cyrtodactylus. As argued by Mees (1987) Gonyodactylus is a nomen nudum, and Kluge (1993, 2001)
consequently allocated these and subsequently described taxa to either Cyrtodactylus or Tenuidactylus.
Anderson (1999), considering chiefly Iranian taxa, retained all of Szczerbak & Golubev’s subgenera within
Cyrtopodion and recognized four species groups: the scabrum group (= Cyrtopodion sensu stricto), the
agamuroides group, the caspium group (= Tenuidactylus), and the kotschyi group (= Mediodactylus), providing
diagnoses for each.
Khan (1988) recognized within Pakistan two distinct lineages of Cyrtodactylus, a Palearctic stock, and a
Tibeto-Himalayan stock. Following the work of Szczerbak & Golubev (1986), he subsequently (Khan 1989, 1991,
1993b, 1993c, 1994; Khan & Tasnim 1990; Khan & Baig 1992) employed the name Tenuidactylus for the majority
of the Palearctic naked-toed geckos of Pakistan and adjacent regions (allocating C. kachhensis [sic] (Stoliczka),
scaber [sic], and C. watsoni (Murray)) to Cyrtopodion (Khan 1993c, 1997) and retained within Cyrtodactylus the
Tibeto-Himalayan taxa, which he considered to include C. stoliczkai, C. chitralensis (Smith), C. dattanensis
(Khan), C. mintoni (Golubev & Szczerbak), and C. himalayanus (Duda & Sahi). Khan (1993a) later added C.
battalensis Khan, C. tibetinus [sic] (Boulenger), C. laderanus [sic] (Stoliczka), and C. fasciolatus (Blyth) to the
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304 · Zootaxa 3599 (4) © 2013 Magnolia Press
latter group. Subsequently he transferred the species Tenuidactylus kohsulaimanai Khan and T. montiumsalsorum
(Annandale) from Tenuidactylus to Cyrtopodion (Khan 1997) and described C. potoharensis in this genus (Khan
2001). Khan & Rösler (1999) subdivided their Circum-Himalayan (= Tibeto-Himalayan) group of Cyrtodactylus
into stoliczkai, tibetinus [sic], and walli subgroups, including in the last of these C. kirmanensis (Nikolsky). Khan
(2000, 2001, 2003a) placed all members of the stoliczkai and walli subgroups in Mesodactylus but retained the
tibetinus [sic] subgroup members in Cyrtodactylus. Within the Palearctic stock he (Khan 2001) segregated the
sandstone geckos, in Tenuidactylus, from all other taxa, which he retained in Cyrtopodion (Khan 2003b).
Khan (2003c) synonymized Tenuidactylus with Cyrtopodion and considered Mediodactylus first as a subgenus
(Khan 2003c) and later (Khan 2004, 2006) as a full genus. However, Khan (2003c) also described three new
genera: Altigekko (type species Tenuidactylus baturensis Khan & Baig), Indogekko (type species Cyrtodactylus
indusoani Khan), and Siwaligekko (type species Cyrtodactylus battalensis) and allocated to these genera species
occurring chiefly in northern Pakistan and portions of Indian Kashmir and western Nepal, some of which had
previously been considered as members of the Tibeto-Himalayan group (Szczerbak & Golubev 1986). Some
members of these putative genera have been especially peripatetic, being variously assigned to one of Khan’s
genera, or to Cyrtopodion, Cyrtodactylus, Mediodactylus, or Altiphylax. Recognition of Khan’s genera has been
sporadic and Krysko et al. (2007) concluded that his classification, although based on morphological similarity,
was not reflective of phylogenetic patterns. These same authors concluded that the addition of new information,
including genetic data would undoubtedly result in the generic reassignment of many of the Pakistani members of
this group and did not themselves propose a comprehensive generic system for them. Auffenberg et al. (2010) and
Shi & Zhao (2011) echoed this conservative approach and considered that placement of new or existing Palearctic
naked-toed taxa in any genus other than Cyrtopodion sensu lato (Cyrtopodion, Tenuidactylus, Mediodactylus,
Altigekko, Indogekko, and Siwaligekko) was premature.
Most recently, Sindaco & Jerem enko (2008) accepted that there were natural subgroups within Cyrtopodion,
but given the incomplete knowledge of many of the constituent taxa, they retained these within Cyrtopodion, using
the subgeneric allocations applied by previous authors. With respect to Khan’s (2003c) groupings, they treated
Altigekko as a junior synonym of Altiphylax and considered Khan’s (2003b) Indogekko to belong to Cyrtopodion
(subgenus Tenuidactylus), although they considered that the lack of femoral pores in males of C. indusoani (Khan),
the type species of Indogekko, might preclude it from close relationship to other members of the group. They
interpreted Siwaligekko as polyphyletic, with S. mintoni being assigned to Altiphylax, and most other species
belonging not to the Palearctic naked-toed radiation, but to the chiefly tropical Asian clade to which typical
Thus far, the Palearctic naked-toed geckos have been included to a very limited degree in only a few
phylogenetic analyses. Szczerbak & Golubev (1986, 1996) provided tree diagrams of proposed phylogenetic
relationships, although these were not explicitly character-based, nor were they fully consistent with monophyletic
genera. Likewise, Khan (2009) presented a view of intergeneric relationships, but this is primarily a phenetic
interpretation. Macey et al. (2000), using allozyme data, demonstrated that Cyrtopodion (represented in their
sample by C. caspium, C. fedtschenkoi (Strauch), C. longipes (Nikolsky), and C. elongatum (Blanford)) and
Mediodactylus (represented by M. russowii (Strauch) and M. spinicauda (Strauch)) were each monophyletic but
that they were not sister taxa. Additional phylogenetic analyses by
ervenka et al. (2008, 2010) using
mitochondrial DNA sequences (approximately 700 bp) have further demonstrated the paraphyly of Cyrtopodion
sensu lato as well as of the subgenus Cyrtopodion as recognized by Szczerbak & Golubev (1986, 1996), and have
shown that Carinatogecko is embedded within Mediodactylus. Gamble et al. (2011, 2012) have clarified that the
primarily tropical bent-toed geckos, Cyrtodactylus, are sister to Hemidactylus and that most Palearctic naked-toed
geckos are part of a monophyletic group that is sister to these two together. However, Alsophylax and Microgecko
are sister taxa and basal within the entire gekkonid clade (Gamble et al. 2012). Within the clade of remaining
naked-toed geckos the pattern (((Tenuidactylus (Crossobamon, Bunopus)) (Cyrtopodion, Agamura))
((Stenodactylus, Tropiocolotes) Mediodactylus)) is well-supported, confirming the non-monophyly of Cyrtopodion
sensu Szczerbak & Golubev (1986), and establishing a backbone phylogeny for the group. Wood et al. (2012) have
further identified that at least one member of Tibeto-Himalayan group, Cyrtodactylus tibetanus, is more closely
allied to Cyrtodactylus than to Palearctic naked-toed species.
Despite these phylogenetic advances, patterns of relationships among naked-toed geckos remain largely
incomplete. Although they sampled 107 genera of gekkotans, Gamble et al. (2012) lacked material of Altiphylax
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PALEARCTIC NAKED-TOED GECKO PHYLOGENY
(including Altigekko and Asiocolotes), Indogekko, Pseudoceramodactylus, Rhinogecko, and Siwaligekko. Further,
the assignment of particular species to genus remains contentious (Anderson 1999; Krysko et al. 2007) and existing
taxon sampling in molecular phylogenies has been sparse. The poor sampling is largely the result of the distribution
of this group across a wide area of the western Palearctic, encompassing both remote and difficult to access areas
and zones of intense human conflict. We here present phylogenetic data based on a limited sampling of Palearctic
naked-toed geckos that includes representatives of a number of genera that have not previously been included in
molecular phylogenetic studies: Altiphylax (including taxa also sometimes assigned to both Asiocolotes and
Altigekko), Indogekko, and Siwaligekko, as well as several previously unstudied species of naked-toed geckos. In
combination with previously collected data (
ervenka et al. 2008, 2010; Fujita & Papenfuss 2011) this permits us
to take one step closer to the establishment of a more representative phylogeny for the group and the stabilization
of generic assignments within Palearctic naked-toed geckos. Specifically, with this study we aim to 1) explicitly
test the monophyly of Cyrtopodion sensu lato, 2) determine whether sampled “Tibeto-Himalayan” taxa are
associated with either the tropical Cyrtodactylus or Palearctic Cyrtopodion radiations, 3) determine the
phylogenetic placement of other problematic species and previously unsampled genera, and 4) identify rough
historical biogeographic patterns and examine the plausibility of previous biogeographic hypotheses and
interpretations (e.g., Khan 2009) in relation to the tectonic history of Central and South Asia.
Materials and methods
Thirty-six individuals representing 28 species of Palearctic naked-toed geckos were sampled from all of the
currently recognized genera in the region except Rhinogecko (Table 1). In addition, representative taxa from the
genera Hemidactylus (six species) and Cyrtodactylus (seven species), known to be sister to one another and
collectively sister to the main Palearctic naked-toed gecko clade (Gamble et al. 2012) were sampled as were an
additional 16 gekkotan and two non-gekkotan outgroups, including representatives of all major gekkotan clades.
When referring to sampled taxa as well as other taxa mentioned in the text, authors of taxonomic names are given
at first usage. Spelling of author names follows that used in the original publications. In some cases, this results in
the same author name having multiple spellings, depending upon the transliteration used in the original
publications (for instance, the same name is alternately transliterated from the Cyrillic alphabet as Jerem
Yeriomtschenko, or Jerjomtschenko).
Identification of samples to species was, whenever possible, based on direct examination of fluid-preserved
specimens by one of the authors and comparison of the relevant specimens to published descriptions and recorded
collection localities of named species. All specimens originating from Pakistan and Afghanistan were examined
directly, as were some, but not all, specimens originating from other geographic regions. These remaining species
identifications are therefore based on identifications of the original collectors. As species boundaries are not always
clearly defined and new species of Palearctic gecko continue to be discovered, identifications of a few specimens
may require subsequent revision, but all specimens can be considered accurate to genus and species group. We treat
this as a preliminary study that may help to inform ongoing taxonomic revisions, and caution that careful revision
of some species groups is still needed, especially those occurring from Iran eastwards.
Genomic DNA was isolated from tail or liver tissue samples preserved in 95–100% ethanol with the Qiagen
DNeasy tissue kit (Valencia, CA, USA). We used double-stranded PCR to amplify 2,817 aligned bases of one
mitochondrial (ND2 and five adjacent tRNAs – 1,345 bases) and two nuclear (1,077 bases of RAG1 and 395 bases
of PDC) loci using the primers listed in Table 2. Amplification of 25
l PCR reactions was executed on an
Eppendorf Mastercycler gradient thermocycler.
Amplification of genomic DNA began with an initial denaturation for 2 minutes at 95 °C followed by 95 °C for
35 s, annealing at 50 °C for 35 s, and extension at 72 °C for 150 s with 4 s added to the extension per cycle for 32
cycles for mitochondrial DNA and 34 cycles for nuclear DNA. When needed, annealing temperatures were
adjusted to increase or decrease specificity on a case by case basis. Products were visualized with 1.5% agarose gel
electrophoresis. Target products were purified with AMPure magnetic bead solution (Agencourt Bioscience,
Beverly, MA, USA) and sequenced with either the BigDyeR Terminator v3.1 Cycle Sequencing Kit (Applied
Biosystems, Foster City, CA, USA) or the DYEnamic™ ET Dye Terminator Kit (GE Healthcare, Piscataway, NJ,
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306 · Zootaxa 3599 (4) © 2013 Magnolia Press
TABLE 1. List of samples used in this study giving sample locality, museum voucher specimen or collector’s field number, and
GenBank accession numbers for each gene. Non-gekkotans and non-gekkonid gekkotans were used as outgroups and/or in
establishing calibrations for the timetree analysis and are not shown in Figure 2. Collection abbreviations: AdS = Anslem de
Silva field series, AMS = Australian Museum Sydney, BPBM = Bernice P. Bishop Museum, CAS = California Academy of
Sciences, CES = Centre for Ecological Sciences, Indian Institute of Science, Bangalore, FMNH = Field Museum of Natural
History, ID = Indraneil Das field series, JB = John Boone private collection, JEM = Jane E. Melville field series, JFBM = James
Ford Bell Museum of Natural History, LSUHC = La Sierra University Herpetological Collection, LSUMZ = Louisiana State
University Museum of Natural Sciences, MAM = Mohammed Al-Mutairi field series, MCZ = Museum of Comparative
Zoology, MVZ = Museum of Vertebrate Zoology, PMNH = Pakistan Museum of Natural History, RAH = Rod A. Hitchmough
field series, ROM = Royal Ontario Museum, TG = Tony Gamble private collection, YPM = Yale Peabody Museum of Natural
History, ZSM = Zoologische Staatssammlung München.
Species Musuem No. Locality GenBank Accession Numbers
ND2 RAG1 PDC
Agamura persica FMNH
Pakistan, Balochistan, Makran district,
JX440515 JX440675 JX440625
Altiphylax levitoni 1 PMNH 2431 Afghanistan, Logar Province, Aynak
KC151974 KC152022 KC151997
Altiphylax levitoni 2 PMNH 2432 Afghanistan, Logar Province, Aynak
KC151964 KC152021 KC151996
Alitphylax stoliczkai 1 PMNH 2323 Pakistan, Gilgit-Baltistan, Skardu,
KC151971 KC152018 KC151993
Alitphylax stoliczkai 2 PMNH 2326 Pakistan, Gilgit-Baltistan, Skardu,
KC151972 KC152019 KC151994
Alsophylax pipiens CAS 238804 Mongolia, Khovd, 1 km N of Bulgan KC151973 KC152020 KC151995
Anolis carolinensis n/a n/a EU747728 AAWZ
Bunopus tuberculatus CAS 228737 United Arab Emirates, Sharjah, 1 km S
Sharjah Desert Park exit on Sharjah-Al
Bunopus tuberculatus MVZ
Iran, Qeshm Island HQ443540
Namibia, Gai-as spring JN393945
CAS 193884 Namibia, 30 km N Swakopmund EU293645 EU293712
Crossabamon orientalis ID 7607 India, Rajasthan, Sam KC151975 KC152023 KC151998
Cyrtodactylus angularis FMNH
Thailand, Sa Kaeo, Muang Sa Kaeo JX440523 JQ945301 JX440632
CAS 216459 Myanmar, Rakhine State, Than Dawe
Dist., Gwa Township
JX440526 JX440685 JX440634
Cyrtodactylus fasciolatus CES 091196 India, Uttarkhand, Mussoorie HM622351 HM622366
Philippines, Romblon Island JX440550 JQ945304 JX440660
Solomon Islands, New Georgia I., Mt
Javi, 5km N Tatutiva Village, Marovoa
JX440556 JX440715 JX440665
Cyrtodactylus tibetanus MVZ
China, Lhasa Municipality, Lhasa, at
base of mountains, ca. 3 km (by air)
WNW of the Potala Palace
PMNH 2301 Pakistan, NWFP, Battagram City,
Chaphar Gram Bridge
KC151983 KC152035 KC152007
PMNH 2303 Pakistan, NWFP, Battagram City, Kass
Bridge on Ooghi Road
KC151984 KC152036 KC152008
AdS 35 Sri Lanka, Yakkunehela JX440522 JX440682 JX440631
continued on next page
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PALEARCTIC NAKED-TOED GECKO PHYLOGENY
TABLE 1. (continued)
Species Musuem No. Locality GenBank Accession Numbers
ND2 RAG1 PDC
Pakistan, Dera Ghazi Khan, Khar
Cyrtopodion scabrum 1MVZ
Pakistan, Chagai District, Batto Village
15km W Nushki, Sayap Chashma
KC151977 KC152025 KC151999
Cyrtopodion scabrum 2 ID 7614 India, Rajasthan, vic. Sam KC151976 KC152024
Cyrtopodion sp. FMNH
Pakistani Border KC151978 KC152026 KC152000
PMNH 2392 Pakistan, Dera Ghazi Khan, Khar
KC151980 KC152028 KC152002
PMNH 2391 Pakistan, Dera Ghazi Khan, Khar
KC151979 KC152027 KC152001
Gehyra sp. AMS
Australia, Western Australia,
JN393929 JN393973 JN394007
Gekko gecko MVZ
Thailand: Phuket Island AF114249
Gekko gecko CAS 204952 Myanmar, Ayeyarwady, Mwe Hauk
Hemidactylus angulatus MVZ
Ghana, Volta region, Togo Hills EU268367 EU268306 EU268336
Hemidactylus fasciatus CAS 207777 Equatorial Guinea, Bioko Sur Prov., ca
3.6 km N of Luba
EU268371 EU268310 EU268340
Hemidactylus frenatus CAS 229633 Myanmar, Tanintharyi Division, Kaw
HM559629 HM559695 HM559662
Hemidactylus garnotii CAS 223286 Myanmar, Rakhine State, Taung Gok
EU268363 EU268302 EU268332
Hemidactylus mabouia YPM 14798 USA, Florida, Monroe County, Little
HM559639 HM559705 HM559672
Hemidactylus turcicus LSUMZ H-
USA, Louisiana, Baton Rouge EU268360 EU268299 EU268329
Lialis burtonis JFBM 8 Australia (captive) GU459540 GU459742
Lialis jicari n/a Australia AY369025
Lygodactylus mirabilis ZSM 388/
Madagascar, Ankaratra, above
JX041382 HQ426300 HQ426211
PMNH 2165 Pakistan, NWFP, Battagram City,
Chaphar Gram Bridge
KC151981 KC152029 KC152003
Mediodactylus kotschyi 1MCZA
Turkey, Mugla, Cicekli koyu-ula KC151967 KC152031
Mediodactylus kotschyi 2MCZA
Turkey, Mersin, Alanya-Anamur arasi
Mellec Anamur'a 26 km kala
Mediodactylus russowii JEM 863 Kazakhstan, Ili River, 44°03’05”N,
JX440517 JX440678 JX440627
Mediodactylus spinicauda CAS 228709 Iran, Khorasan Prov., ca 4 km S (by rd)
of Jaanbaazaan Square (Birjand),
Bande-dare Spring (dam)
KC151968 KC152032 KC152004
JB 5 captive KC151982 KC152034 KC152006
JB 27 captive KC151969 KC152033 KC152005
Nactus vankampeni BPBM
Papua New Guinea, East Sepik
EU054295 EU054279 EU054263
Oedura marmorata AMS
Australia, Queensland GU459951 EF534779 EF534819
Phelsuma inexpectata JB 56 Réunion (captive) JN393939 JN393983 JN394016
continued on next page
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308 · Zootaxa 3599 (4) © 2013 Magnolia Press
USA). Sequencing reactions were purified with CleanSeq magnetic bead solution (Agencourt Bioscience, Beverly,
MA, USA) and analyzed with an ABI 3730 XL automated sequencer. The accuracy of sequences was ensured by
incorporating negative controls and sequencing complementary strands. Sequences were aligned by eye in Text
Wrangler 3.1, and protein-coding genes were translated to amino acids with MacClade (Maddison & Maddison
2005) to confirm conservation of the amino acid reading frame and check for premature stop codons.
Phylogenetic relationships among the samples were assessed with parsimony, likelihood, and Bayesian
optimality criteria using the combined dataset. The maximum parsimony (MP) analysis was conducted in
PAUP*4.0b10 (Swofford 2002). A heuristic search algorithm was used with tree bisection-reconnection (TBR)
branch swapping, zero-length branches collapsed to yield polytomies, and gaps treated as missing data. Each base
position was treated as an unordered character with four alternate states. We used nonparametric bootstraps (1,000
pseudoreplicates) to assess node support with TBR branch swapping and five random addition replicates per
For model-based ML and BI methods, a ten-partition scheme was used, with each gene partitioned and the
protein-coding genes further partitioned by codon position. We used the Akaike Information Criterion (AIC) in
jModelTest 0.1.1 (Guindon & Gascuel 2003; Posada 2008) to find the model of evolution that best fit the data for
each partition, which were identified as GTR + Γ or GTR + I + Γ for each partition. The ML analysis was
Species Musuem No. Locality GenBank Accession Numbers
ND2 RAG1 PDC
Phelsuma rosagularis n/a Mauritius EU423292
Phelsuma rosagularis JB 109 Mauritius (captive) HQ426306 HQ426217
Phyllodactylus xanti ROM 38490 Mexico: Baja California Sur JN393940 EF534807 EF534849
United Arab Emirates, Dubai, Jebel Ali HQ443551
Ptyodactylus guttatus TG 00072 Egypt (captive) JX041426 EU293636 EU293703
Pygopus nigriceps MVZ
Australia, Northern Territory JX440518 EF534783 EF534823
Python molurus n/a n/a AEQU
Python regius n/a n/a AB177878
CAS 198428 USA, Puerto Rico JN393943 EF534785 EF534825
Sphaerodactylus torrei JB 34 Cuba JX440519 EF534829 EF534788
Stenodactylus doriae JB 20 captive KC151985 KC152037 KC152009
Stenodactylus slevini MAM 3066 Saudi Arabia, Ibex Reserve KC151986 KC152010
Mauritania, Nouadhibou, Cansado, 10
KC151987 KC152038 KC152011
Tenuidactylus caspius CAS 228602 Iran, Semnan Prov., Touran Protected
Area, Delbar Field Station
KC151988 KC152039 KC152012
Tenuidactylus elongatus JB 127 China, Gobi JX440516 JX440677 JX440626
JEM 346 Uzbekistan, 5km from Nurata, Aktau
Mtns., 40°33’18.7”N, 65°40’04.7”
KC151989 KC152040 KC152013
Tenuidactylus longipes CAS 228830 Iran, Yazd Prov., 23 km N Tabas,
KC151990 KC152041 KC152014
Teratoscincus roborowskii CAS 171203 China, Xinjiang AF114252
Teratoscincus roborowskii TG 00070 China (captive) EF534799 EF534841
Tropiocolotes nubicus 1 JB 123 Egypt KC151991 KC152042 KC152015
Tropiocolotes nubicus 2 JB 130 Egypt KC151970 KC152043 KC152016
Tropiocolotes steudneri JB 28 Captive JX440520 JX440680 JX440629
Tropiocolotes tripolitanus MVZ
Niger, Agadez, Tafokin, 13 km NNE
KC151992 KC152044 KC152017
Woodworthia maculata RAH 292 New Zealand, Titahi Bay GU459852 GU459449 GU459651
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PALEARCTIC NAKED-TOED GECKO PHYLOGENY
performed using RAxML HPC v7.2.3 (Stamatakis 2006). As RAxML can only implement a single GTR model, the
GTR + I + Γ model was used. Support was assessed with 1,000 non-parametric bootstraps.
The Bayesian analysis was conducted using MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003). Each partition
was given a mixed substitution model with invariant sites and a Γ parameter, with all parameters unlinked across
all partitions. Analyses were initiated with random starting trees and run for 20,000,000 generations; Markov
chains were sampled every 1,000 generations; the first 5000 trees, representing 25% of all trees, were discarded as
burn in. We used Are We There Yet (AWTY) (Nylander et al. 2008) to verify that each chain ran for a duration that
adequately sampled the posterior distributions, as well as referring to estimated sample sizes and standard
deviations of split frequencies.
In addition to the combined analyses, single-locus ML analyses were also performed to ensure that individual
loci exhibited no strongly conflicting phylogenetic signal. Parameters of the single-locus analyses were identical to
those of the combined analysis, except the data were divided into three (for RAG1 and PDC) or four (for
We performed a Shimodaira-Hasegawa (SH) test (Shimodaira & Hasegawa, 1999) in RAxML to test the non-
monophyly of Cyrtopodion sensu lato, as recovered by
ervenka et al. (2008, 2010). Our best-scoring likelihood
tree was tested against the best-scoring likelihood tree in which Cyrtopodion sensu lato is monophyletic. This
constrained tree was identified using the ConstraintTree option in RAxML, in concert with a likelihood search
where all other run and model parameters were identical to those used in the combined unconstrained ML analysis.
Our constraint tree enforced monophyly of a clade including all sampled Cyrtopodion, Indogekko, Mediodactylus,
and Tenuidactylus, corresponding to Cyrtopodion sensu Sindaco & Jerem
BEAST 1.7.2 (Drummond & Rambaut 2007) was used to estimate times of divergence among Palearctic
naked-toed geckos. The same partition scheme used in the ML and BI phylogenetic analyses was employed in the
timing analysis. The analysis used an ultrametric starting tree which was estimated using PathD8 (Britton et al.
2007) and employed Yule tree priors and a relaxed uncorrelated lognormal clock. Five previously-used calibration
points (Heinicke et al. 2011) were used to date the tree: divergence of cinereus-series Sphaerodactylus from other
Sphaerodactylus (exponential prior, mean=3, offset=15); divergence of New Zealand from Australian
diplodactylids (exponential prior, mean=17, offset=16); diversification of crown-group pygopods (exponential
prior, mean=10, offset=20); diversification of the Mauritius-Réunion Phelsuma clade (uniform prior, 0–8 Ma); and
divergence of Gekkonidae from other gecko families (lognormal prior, mean=3, S.D.=1, offset=97). The analysis
was run for 50 million generations, sampling every 5,000 generations, with the first million generations discarded
as burn-in. Effective sample sizes were consulted in Tracer 1.5 (Rambaut & Drummond 2007) to ensure adequate
TABLE 2. Primers used in this study.
We recovered a strongly supported Hemidactylus/Cyrtodactylus/Palearctic naked-toed gecko clade that included all
of the naked-toed gecko genera except Alsophylax and Microgecko (Fig. 2), broadly agreeing with the pattern
observed by Gamble et al. (2012). All remaining Palearctic naked-toed genera clustered in a single clade except
Primer Gene Reference Sequence
PHOF2 PDC Bauer et al. (2007) 5'-AGATGAGCATGCAGGAGTATGA-3'
PHOR1 PDC Bauer et al. (2007) 5'-TCCACATCCACAGCAAAAAACTCCT-3'
L4437B ND2 Macey et al. (1997) 5'-AAGCAGTTGGGCCCATACC-3'
L5002 ND2 Macey et al. (1997) 5'-AACCAAACCCAACTACGAAAAAT-3'
Trpr3a ND2 Greenbaum et al. (2007) 5'-TTTAGGGCTTTGAAGGC-3'
H5934a ND2 Arévalo et al. (1994) 5'-AGRGTGCCAATGTCTTTGTGRTT-3'
R13 squamate RAG1 Groth & Barrowclough (1999) 5'-TCGAATGGAAATTCAAGCTGTT-3'
R18 squamate RAG1 Groth & Barrowclough (1999) 5'-GATGCTGCCTTCGGCCACCTTT-3'
RAG1 F700 RAG1 Bauer et al. (2007) 5'-GGAGACATGGACACAATCCATCCTAC-3'
RAG1 R700 RAG1 Bauer et al. (2007) 5'-TTTGTACTGAGATGGATCTTTTTGCA-3'
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FIGURE 2. Combined gene (ND2, RAG1, PDC) phylogeny of Palearctic naked-toed geckos. Non-gekkotans and non-
gekkonid gecko outgroups are not shown. Shaded boxes indicate taxa previously allocated to Cyrtopodion sensu lato. Stars
indicate Palearctic naked-toed geckos that are not members of the main clade (see also Gamble et al. 2012). Lettered circles
indicate dated nodes and stems corresponding to Table 3. Support values at nodes are in the format BI posterior probability/ML
bootstrap/MP bootstrap support. Taxon names correspond to generic allocations as proposed in this paper. See Table 1 for
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PALEARCTIC NAKED-TOED GECKO PHYLOGENY
FIGURE 3. Dorsal views of (A) Tropicocolotes tripolitanus (CAS 123467) and (B) Microgecko helenae (CAS 120795)
illustrating numerous osteological differences between the two genera. Abbreviations: j = jugal; flp = anterolateral process of
frontal; n = nasal; pma = premaxilla-maxilla aperture. Scale bar equals 10 mm.
Siwaligekko which, along with Cyrtodactylus tibetanus (placed in the Tibeto-Himalayan group of
Tenuidactylus by Szczerbak & Golubev 1986), are basal members of the Cyrtodactylus clade. The main Palearctic
clade, which receives strong support, is divided into four monophyletic groups. Basal relationships among these
four groups are not resolved, and this portion of the tree is characterized by exceptionally short internodes and
varying branching order depending on the analysis used. The first of these groups includes the genera
Pseudoceramodactylus, Stenodactylus, and Tropiocolotes. This is sister, albeit with weak support, to a second
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312 · Zootaxa 3599 (4) © 2013 Magnolia Press
strongly-supported group, Mediodactylus (including its type species, kotschyi). The two sampled species of
Altiphylax form a strongly supported third group, confirming the association of species often assigned to
Asiocolotes (in part) and Altigekko. The fourth major group of naked-toed geckos that receives strong support
comprises a monophyletic Tenuidactylus, including its type species, caspius, with strong support under Bayesian
inference, as sister to a more weakly supported clade including all remaining genera sampled. Within this group
Bunopus and Crossobamon are strongly supported as close relatives, with Agamura obtaining moderate support as
sister to these two. This clade, in turn, is sister to a Cyrtopodion clade also incorporating Indogekko. Within this
clade there is strong support for Indogekko plus C. scabrum and an unidentified congener, but only weak support
uniting these with C. kohsulaimani. Constraining the ML analysis to recover a monophyletic Cyrtopodion/
Mediodactylus/Tenuidactylus/Indogekko clade results in a phylogeny with a much lower likelihood score (lnL
–58948 constrained vs. –58817 unconstrained). Based on the SH test results, monophyly of a Cyrtopodion/
Mediodactylus/Tenuidactylus/Indogekko clade is rejected (p < 0.01).
Inspection of the BEAST output in Tracer suggested adequate mixing of the chain: traces of all parameters
revealed no evident patterns, and ESS values for all but one parameter were much greater than 200 (ESS for the
standard deviation of the uncorrelated lognormal distribution for ND2 was 160). The branching pattern is similar to
that recovered in the ML and BI phylogenetic analyses, though interrelationships among the four basal groups
within the main Palearctic clade differ from those obtained in some of the other analyses. Divergence time
estimates (Table 3) suggest that the split between Cyrtodactylus + Hemidactylus and the Palearctic clade occurred
in the late Cretaceous, 70 (79–61) million years ago (Ma). Divergences within the Palearctic clade itself are
restricted to the Cenozoic, with the earliest divergence estimated to have occurred 56 (64–48) Ma. Divergences
among Cyrtodactylus species are extremely similar to those estimated by Wood et al. (2012), and suggest that the
Tibeto-Himalayan species C. tibetanus and Siwaligekko battalensis diverged from tropical Cyrtodactylus about 50
Ma (composite 95% HPD 71–40 Ma), roughly contemporaneous with the collision of the Indian and Eurasian
Plates (Rowley 1996). The divergence of the montane western Himalayan genus Altiphylax from other Palearctic
genera also occurred in this general time frame, 52 (61–45) Ma. Other divergences of Himalayan taxa (e.g.,
Indogekko) from other Palearctic taxa are more recent.
TABLE 3. Estimated divergence dates among selected Palearctic geckos, with 95% HPD values given in parentheses. Node
labels refer to Figure 1. mrca = most recent common ancestor.
Our broader results are largely consistent with those of Gamble et al. (2012), who sampled the majority of all
gecko genera. Like them we found Alsophylax and Microgecko to be outside of the main radiation of Palearctic
naked-toed geckos. Gamble et al. (2012) found strong support for these two taxa as sister to one another and
together as sister to all other gekkonid geckos. Their placement in our phylogeny was different, but we attribute
Node/Stem Description Divergence Time (Ma)
A Palearctic naked-toed geckos vs. Cyrtodactylus + Hemidactylus 70 (79–61)
B Cyrtodactylus tibetanus vs. other Cyrtodactylus 54 (71–43)
C Siwaligekko vs. other Cyrtodactylus 49 (59–40)
D mrca Palearctic naked-toed geckos 56 (64–48)
E stem age of Altiphylax 52 (61–45)
Fmrca Agamura/Bunopus/Crossobamon/Cyrtopodion/Tenuidactylus 35 (42–28)
Gmrca Agamura/Bunopus/Crossobamon 29 (36–22)
H Indogekko vs. core Cyrtopodion 23 (30–16)
I Tenuidactylus longipes vs. other Tenuidactylus 16 (21–11)
J Tenuidactylus caspius vs. Tenuidactylus fedtschenkoi 12 (17–7)
Kmrca Mediodactylus 38 (47–29)
Lmrca Pseudoceramodactylus/Stenodactylus/Tropiocolotes 46 (56–36)
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this to our poor taxon sampling within the Gekkonidae, which resulted in a lack of support for most relationships
outside of the Hemidactylus/Cyrtodactylus/Palearctic naked-toed gecko clade. This latter clade, which receives
strong support here, has had universal support from those previous molecular phylogenetic analyses that included
each of these taxa (Han et al. 2004; Feng et al. 2007; Gamble et al. 2011, 2012).
Our data support the non-monophyly of Cyrtopodion, as first suggested by Macey et al. (2000) and
subsequently confirmed by
ervenka et al. (2008, 2010) and Gamble et al. (2012). Mediodactylus is particularly
distinct with respect to the other groups originally proposed as subgenera of Cyrtopodion (Szczerbak & Golubev
1986). Our sampling included the type species, M. kotschyi, as well as M. russowii, M. spinicauda (the three
species explicitly included in this group by Macey et al. 2000), and M. brachykolon (Krysko, Rehman &
Auffenberg). Based on
ervenka et al. (2010), this clade should also include M. heterocercus (Blanford), M.
sagittifer (Nikolsky), and Carinatogecko spp. Torki (2011) has intimated that the Carinatogecko included by
ervenka et al. (2010), C. cf. heteropholis, might be either M. heterocercus or a similar form and on this basis and
differences in scalation maintains the validity of the genus Carinatogecko. Unfortunately, our lack of material of
any members of this group precludes further comment. Cyrtopodion brachykolon is unambiguously a member of
the Mediodactylus clade. Krysko et al. (2007) believed that this species could not be unambiguously assigned to
any particular subgenus and took a conservative approach in allocating their new species to the genus Cyrtopodion
sensu lato, although they concluded their paper by predicting that future generic reassignment of this and other
Pakistani naked-toed geckos was likely. Khan (2008b) reevaluated the generic allocation of C. brachykolon and
recommended its transfer to Altigekko. We found no support for this, at least in the context of the sole Altigekko we
sampled, A. stoliczkai.
Remaining Cyrtopodion sensu lato (i.e., Cyrtopodion sensu stricto plus Tenuidactylus) are not monophyletic.
Together they belong to a clade that also includes Bunopus, Agamura, Crossobamon, and Indogekko. This is
consistent with earlier findings (
ervenka et al. 2008, 2010; Gamble et al. 2012). The sister group relationship of
Bunopus and Crossobamon was also found by the last of these authors and had been proposed by Anderson (1999).
We examined only a single species of Bunopus, but
ervenka et al. (2008, 2010) sampled B. crassicaudus and
what would appear to be several cryptic taxa masquerading under B. tuberculatus. In our RAG1 tree the Bunopus +
Crossobamon clade was in turn sister to Cyrtopodion kohsulaimanai, but in the ND2 and combined trees the latter
species was sister to the Cyrtopodion scabrum/Indogekko clade.
The position of Agamura has varied in different analyses.
ervenka et al. (2008) recovered it as sister to
Cyrtopodion sensu lato exclusive of Mediodactylus. However,
ervenka et al. (2010) placed it as sister to
Cyrtopodion sensu stricto, and Gamble et al. (2012) found support for it as the sister to Bunopus + Crossobamon +
Tenuidactylus + Cyrtopodion. In our tree Agamura’s position is different again, although without support.
Unfortunately, we did not have samples of the two species variously assigned to Agamura (e.g., Szczerbak &
Golubev 1986, 1996; Kluge 1991, 1993, 2001) or to Rhinogecko (Anderson 1999; Krysko et al. 2007; Khan 2006;
Sindaco & Jerem
Our results support the recognition of Cyrtopodion and Tenuidactylus as separate genera. Tenuidactylus, with
its type, T. caspius, was recovered in a clade also including T. elongatus, T. fedtschenkoi, and T. longipes, the same
four taxa supported as a clade by the allozyme data of Macey et al. (2000).
ervenka et al. (2008, 2010) likewise
hypothesized a T. caspius/T.longipes clade, either as sister to Cyrtopodion sensu stricto (2008) or to Agamura plus
Cyrtopodion sensu stricto as its sister group. Sindaco & Jerem
enko (2008) hypothesized that Cyrtopodion
elongatum should be allied to Rhinogecko.
Unfortunately, our sampling of remaining Cyrtopodion sensu lato was poor. In our analyses the type species of
Cyrtopodion, C. scabrum, clusters with an unidentified species from Pakistan and with Indogekko rohtasfortai
(Khan & Tasnim). In the combined and ND2 trees Cyrtopodion kohsulaimanai also clusters weakly with this clade.
This is consistent with the results of Golubev et al. (1995) who suggested that C. kohsulaimanai was allied to those
species later placed in Indogekko by Khan (2003).
ervenka et al. (2008, 2010) recovered C. sistanense Nazarov &
Rajabizadeh and an unidentified Iranian species as the closest relatives of C. scabrum, with C. agamuroides
(Nikolsky) and C. gastrophole, neither monophyletic, outside of these.
As Fujita & Papenfuss (2011) have recently provided a phylogeny for Stenodactylus, we did not sample deeply
in this genus, although we added one species, S. slevini Haas, not included by them. Arnold (1980; repeated by
Sindaco & Jerem
enko 2008) suggested that S. slevini was sister to S. leptocosymbotes Leviton & Anderson + S.
doriae (Blanford). These authors also hypothesized affinities of other members of the genus, but these have been
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314 · Zootaxa 3599 (4) © 2013 Magnolia Press
superseded by the work of Fujita & Papenfuss (2011). We likewise did not sample deeply in Tropiocolotes sensu
stricto. We note, however, that our samples of T. nubicus Baha El Din are minimally distinct from a single sample
of T. steudneri (Peters), from which it was differentiated on morphological grounds. Tropiocolotes was not
monophyletic in our phylogeny, with the type species, T. tripolitanus Peters, as the immediate sister to
Stenodactylus, rather than to other Tropiocolotes, although support for this relationship was weak.
Our phylogenetic analysis is far from taxon complete, but it includes enough critical new samples to allow some
preliminary evaluation of four taxonomies of Palearctic naked-toed geckos that currently co-exist rather
uncomfortably: those of Szczerbak & Golubev (1986, 1996; here we consider that the varying earlier taxonomies
of these authors have been superceded by these works and also adequately reviewed by them, as well as by
Anderson 1999, Krysko et al. 2007, and Sindaco & Jerem
enko 2008); Anderson (1999); Khan (2003c; variation in
Khan’s earlier generic allocations of taxa have been summarized by Krysko et al. 2007 and Sindaco & Jerem
2008); and Sindaco & Jerem
enko (2008). It should be noted that each of these works is geographically conscribed,
with Szczerbak & Golubev (1986, 1996) concentrating on the region of the former Soviet Union, Anderson (1999)
on Iran, Khan (2003c) on Pakistan, and Sindaco & Jerem
enko (2008) on the whole of the western Palearctic, but
excluding eastern Pakistan, India, Nepal, Mongolia and China. Although naked toed-geckos from these last regions
have been treated peripherally, these have never been adequately reviewed.
Sczerbak & Golubev’s (1986, 1996) inclusion of Microgecko and Asiocolotes as subgenera within
Tropiocolotes has not been supported. Asiocolotes and Tropiocolotes are members of the same large clade of naked
toed geckos, but do not form an exclusive monophyletic group. On the other hand, Microgecko is, along with
Alsophylax, one of the most basal members of the family Gekkonidae and not related to the bulk of the Palearctic
naked-toed clade (Gamble et al. 2012). Continued recognition of Microgecko as part of Tropiocolotes (e.g., Torki
2008; Torki et al. 2008; Rajabizadeh et al. 2010) is no longer warranted given the overwhelming evidence that the
two are only distantly related. The morphological features that have been used to distinguish species of Microgecko
from those of Tropiocolotes (see keys in Minton et al. 1970; Leviton & Anderson 1972) are well established.
Further, representative taxa from each genus, M. helenae and T. tripolitanus, exhibit fundamental osteological
differences that are typically invariant within a single gekkotan genus. In this instance, the nasal bones, which are
typically paired in gekkonids (Kluge 1967, 1987; Daza 2008) are fused in Microgecko (Fig. 3). Other differences
include a much enlarged, shoehorn-shaped jugal bone in Microgecko, presence of a second ceratobranchial arch in
Microgecko (Kluge 1983), and the concealed anterolateral process of the frontal and prominent premaxilla-maxilla
aperture in Tropiocolotes, all of which are typically consistent within a gecko genus. Although we did not retrieve a
monophyletic Tropiocolotes sensu stricto, poor support for the alternative relationship (Fig. 2) does not warrant
taxonomic action at this time.
Szczerbak & Golubev’s (1986, 1996) subdivision of Cyrtopodion (their Tenuidactylus) into four subgroups —
Tenuidactylus, Mediodactylus, Cyrtopodion, and the Tibeto-Himalayan group — appears to have been
fundamentally sound, but they were mistaken in accepting the monophyly of these groups collectively. Although
not all of the taxa they evaluated have been included in molecular phylogenetic analyses, it is evident that most of
the species they assigned to Mediodactylus, Tenuidactylus, and Cyrtopodion sensu stricto are correctly allocated,
with the exception of Tenuidactylus elongatus, which they had assigned to Cyrtopodion. Their Tibeto-Himalayan
group, however, is a phylogenetic hodge-podge including taxa here assigned to both Cyrtodactylus sensu lato and
As with Szczerbak & Golubev (1986, 1996), Anderson’s (1999) identification of four monophyletic groups of
Cyrtopodion appears to be largely vindicated, although their collective monophyly is falsified. The agamuroides
group, which has not been formally recognized taxonomically, is morphologically well-characterized and to the
extent that it has been sampled,
ervenka et al. (2010) have recently demonstrated its monophyly. ervenka et al.
(2008, 2010) placed this group in two different positions relative to the Cyrtopodion scabrum group (Cyrtopodion
sensu stricto), Tenuidactylus, and Agamura. First, they reported a monophyletic Cyrtopodion (exclusive of
Mediodactylus), with the agamuroides group sister to Tenuidactylus, but with low support. Subsequently they
found this group to be sister to the C. scabrum group, with no information about clade support provided.
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Ahmadzadeh et al. (2011) thus considered that status of this clade to be unstable. With the addition of new taxa
described by Nazarov et al. (2009) and Ahmadzadeh et al. (2011), the agamuroides group contains five species
which we here retain in Cyrtopodion.
Of the genera described by Khan (2003c), we can only address implications in light of the samples we have
sequenced and if Khan’s generic groupings are not monophyletic our interpretations would certainly be different.
We conclude that Siwaligekko is a member of the Cyrtodactylus clade. Cyrtodactylus is an unwieldy genus of
approximately 170 species and it is tempting to maintain Siwaligekko as its sister genus. It does not, however, form
a monophyletic group with the single other Tibeto-Himalayan taxon sampled, C. tibetanus. Macey et al. (2000)
first placed tibetanus with the Cyrtodactylus clade based on allozyme data. Wood et al. (2012) have used a multi-
gene approach to demonstrate that this species is sister to all other members of the genus sampled, which have a
more tropical distribution. Remaining taxa assigned to Siwaligekko by Khan, as well as other Tibeto-Himalayan
taxa not considered by him (C. malcolmsmithi Constable, C. medogense Zhao & Li, C. zhaoermii Shi & Zhao)
share not only their Himalayan distribution, but also a similar morphology with one another, but it is uncertain
which of these may group with S. battalensis and which with C. tibetanus. Alternatively, better taxon sampling
may reveal that all are indeed monophyletic and sister to tropical Asian Cyrtodactylus. Because a comprehensive
phylogenetic analysis and corresponding taxonomic revision of these geckos is lacking we tentatively synonymize
Siwaligekko with Cyrtodactylus, however Siwaligekko may be conveniently used as a subgenus for S. battalensis
and its close relatives. It should be noted that Khan (2003c, 2005) also included within Siwaligekko a number of
peninsular Indian and Sri Lankan taxa that are typically assigned to Geckoella Gray. Wood et al. (2012) have
demonstrated that Geckoella is embedded within Cyrtodactylus and shares no special relationship to the Tibeto-
Himalayan naked-toed geckos (see also Fig. 2).
We here synonymize Khan’s (2003c, 2008a) Indogekko with Cyrtopodion sensu stricto. However, one of his
constituent taxa, I. longipes (and presumably its subspecies) clearly falls in the Tenuidactylus clade in our
phylogeny, whereas I. rohtasfortai, presumably a more typical
‘Indogekko’ is a member of Cyrtopodion sensu
stricto. The similar habitus of unsampled Indogekko is certainly consistent with their forming a monophyletic
group and deeper sampling within Cyrtopodion may well result in a revised position for C. kohsulaimanai, which
could reveal Indogekko as the sister to Cyrtopodion sensu stricto. With the present data we regard synonymization
as the most conservative course of action; however, we propose the recognition of Indogekko as a subgenus of
Cyrtopodion, highlighting the need to consider the taxonomic status of the sandstone geckos in any future revision
Khan’s (2003c, 2004) Altigekko, type species Tenuidactylus baturensis, also included A. stoliczkai (as well as
A. boehmei, subsequently synonymized with A. stoliczkai by Auffenberg et al. 2004) and A. yarkandensis
(Anderson). In our study Altigekko stoliczkai is certainly a genetically distinctive lineage and its closest relative
among sampled taxa is Asiocolotes levitoni (Golubev & Szczerbak). Altigekko is clearly distinctive and represents
a lineage separate from other named groups. However, the name Altiphylax has been considered as its subjective
senior synonym based on the allocation of several taxa to both names (e.g., stoliczkai; see below).
Although they did not conduct an explicit phylogenetic analysis or taxonomic revision, Sindaco & Jerem
(2008) made specific decisions in their chosen taxonomy and expressed their opinions regarding phylogenetic
relationships. Interestingly, they regarded Alsophylax and Microgecko as sister taxa, as subsequently demonstrated
by Gamble et al. (2012). However, they considered these to have a close relationship to both Altiphylax and
Tropiocolotes. They considered Altiphylax as largely congruent with the Tibeto-Himalayan group of Szczerbak &
Golubev (1986, 1996) and included in it the type species, A. tokobajevi, and A. levitoni, the type species of
Asiocolotes Golubev, 1984, A. batturensis, the type species of Altigekko Khan, 2003 as well as A. mintoni (included
in Siwaligekko by Khan 2003c), A. stoliczkai, A. walli (Ingoldby), and A. yarkandensis. They allocated the second
species of Asiocolotes, A. depressus (Minton & Anderson), to Microgecko, although without explicit justification.
Our sparse sampling, as well as our review of the available literature largely supports their interpretations in this
regard and our proposed taxonomy is congruent with theirs for these taxa, as well as for the taxa assigned by Khan
(2003c) to Siwaligekko. However, Sindaco & Jerem
enko’s (2008) placement of Indogekko spp. in Cyrtopodion
(subgenus Tenuidactylus) is at odds with our findings.
Many more questions remain in the systematics of Palearctic naked-toed geckos. The status of Rhinogecko
relative to Agamura cannot be evaluated given our data, nor can the strict synonymy of Altiphylax and Altigekko,
given that we have not compared material from the type species of either genus. Likewise, the composition of
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Cyrtopodion, Tenuidactylus and Mediodactylus are not finalized (see, for example, Masroor 2009 and ervenka &
Kratochvíl 2010) and a question still remains as to the synonymy of Carinatogekko with Mediodactylus.
The subdivision of the species into Palearctic naked-toed forms or Cyrtodactylus allies appears to be resolved and
serves to demonstrate the clear geographic ties of these two large clades. As demonstrated here and by Wood et al.
(2012) there is a west to east progression of the clades of the Cyrtodactylus group, with Tibeto-Himalayan groups
most basal. These extend as far west as northern Punjab and the North West Frontier Province in Pakistan. On the
other hand, the Palearctic naked-toed group on the southern flanks of the Himalayas penetrates only as far east as
Jammu and Kashmir and Himachal Pradesh in northern India. North of the Tibetan Plateau, however, at least
Tenuidactylus elongatus reaches Xinjiang, China and southern Mongolia. The overall distribution of these two
groups corresponds extremely well to the approximate boundaries of the Palearctic and Indo-Malayan (Oriental)
regions as outlined by Sindaco & Jerem
enko (2008). Likewise the Palearctic Naked-toed clade as a whole is
largely restricted to the western Palearctic as they define it. The two naked-toed gecko genera that are not part of
the main Palearctic clade, Alsophylax and Microgecko, have largely complementary distributions. The former is
mostly Turanian, extending eastwards into the Eastern Turkestanian region and the latter further south, mostly
Iranian and extending westwards into the Western Asian Mountain transition zone (sensu Sindaco & Jerem
2008). Among the groups in the main Palearctic clade, Stenodactylus + Tropiocolotes + Pseudoceramodactylus is
clearly a Saharan/Arabian group, whereas the Bunopus + Agamura + Crossobamon clade is Arabian/Iranian.
Remaining Palearctic naked-toed geckos (Altiphylax, Cyrtopodion, Indogekko, Mediodactylus, Tenuidactylus) for
the most part have distributions to the north of these clades, from southeastern Europe across the former Soviet
Central Asia to the western Himalayas and Gobi Desert, though there is broad overlap, for example in Iran.
The high diversity of Palearctic naked-toed geckos in Central Asia suggests that tectonic uplift, precipitated by
the continuing collisions of the Indian and Arabian Plates with the Eurasian Plate, could have played a role in
speciation and diversification of naked-toed geckos, both by promoting adaptive evolution of taxa to newly formed
high-elevation habitats and by creating impassible high-elevation barriers that promote vicariant speciation of
formerly continuously-distributed taxa. These collisions produced the exceptional topographic relief that now
characterizes Central Asia. Himalayan uplift commenced first, upon the initial collision of the Indian and Eurasian
Plates in the early Eocene (about 50 Ma), while mountain ranges to the north and west, more distant from this
collision zone, experienced uplift later, beginning about 20 Ma in the Tien Shan, for example (Rowley 1996;
Macey et al. 1999; Huang et al. 2006). The collision of the Arabian Plate, which began in the late Oligocene (about
25 Ma), has caused further uplift, most notably of the Iranian Plateau (Hearn & Ni 1994).
A scenario of uplift-associated divergence in Central Asian geckos has already been supported in analyses of
the frog-eyed geckos, genus Teratoscincus (Macey et al. 1999, 2005). Both phylogenetic relationships and times of
divergence among Teratoscincus species are consistent with successive uplift events causing vicariant
fragmentation. Inasmuch as estimated divergence times among Palearctic naked-toed geckos are contemporaneous
with or post-date the initial Indian-Eurasian plate collisions about 50 Ma, a role for mountain building in promoting
their isolation and speciation is quite plausible. As our taxon sampling is relatively incomplete at the species level,
we are generally not able to comment on which particular uplift events may have precipitated specific divergences
(although some Himalayan taxa, including Altigekko, Siwaligekko, and Cyrtodactylus tibetanus, diverged from
non-Himalayan relatives around 50 Ma). However, we can comment on the compatability of some previously
suggested biogeographic scenarios for Palearctic naked-toed geckos with our estimates of phylogenetic
relationships and divergence times.
Leviton & Anderson (1984) argued that divergences among several Tenuidactylus species (their Cyrtodactylus
caspius group), including the species T. caspius, T. fedtschenkoi, and T. longipes, were due to vicariant speciation
caused by uplift events in the Iranian and Transcaspian regions starting in the early Miocene (23–16 Ma). Our
estimates of evolutionary relationships and divergence times—divergence of T. longipes 16 (21–11 Ma), split of T.
caspius and T. fedtschenkoi 12 (17–7) Ma—are entirely compatible with this scenario. Thus, for Tenuidactylus, at
least, a Teratoscincus-type vicariant history is plausible.
In multipe papers, Khan (2003c, 2009), while discussing overall Palearctic gecko evolutionary history
(including distantly related geckos of the family Eublepharidae) suggested that Central Asian naked-toed geckos
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PALEARCTIC NAKED-TOED GECKO PHYLOGENY
represent two groups. One group, including Siwaligekko and Altiphylax (Khan’s Altigekko), was derived from a
tropical Cyrtodactylus-type ancestor that colonized the Himalayas from India upon the India-Eurasia tectonic plate
collision. The other group included Cyrtopodion, Indogekko, Mediodactylus and Tenuidactylus, and was suggested
to have colonized Asia from northeastern Africa starting in the Miocene, subsequently speciating through the
Pliocene and Pleistocene. Some aspects of Khan’s scenario are supported by our analyses. In particular, Central
Asian naked-toed geckos are divided into two groups, one of which corresponds to the main Palearctic clade and
the other of which is closely related to tropical Cyrtodactylus (though Altigekko belongs to the Palearctic group, not
the Cyrtodactylus/Siwaligekko group). Also, tectonic plate collision and resulting Himalayan uplift are plausible
agents for the divergences of Altigekko and Siwaligekko from their closest relatives. However, several aspects of
Khan’s scenario do not receive support. Most notably, based on the geographic distributions and phylogenetic
relationships of sampled species in this study and in Wood et al. (2012) both the main Palearctic naked-toed gecko
clade and the Cyrtodactylus clade probably occurred ancestrally in central Asia, rather than migrating from Africa
or South/Southeast Asia, respectively. Also, our molecular clock analysis demonstrates that the main Palearctic
naked-toed gecko clade is much older than Miocene in age, having a most recent common ancestor 56 (64–48) Ma.
Proposed Generic Allocation of Species of Palearctic Naked-Toed Geckos
Agamura (1 species)
Composition: A. persica Duméril
Distribution: Iran, Afghanistan, Pakistan
Alsophylax (6 species)
Composition: A. laevis Nikolsky, A. loricatus Strauch, A. pipiens (Pallas), A. przewalskii Strauch, A.
szczerbaki Golubev & Sattaroc, A. tadjikiensis Golubev
Distribution: Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Kyrgyzstan, northwestern China, southern
Altigekko [see Altiphylax]
Comments: Junior subjective synonym of Altiphylax fide Sindaco & Jerem enko (2008).
Altiphylax (5 species)
Composition: A. baturensis (Khan & Baig), A. levitoni (Golubev & Szczerbak), A. mintoni (Golubev &
Szczerbak), A. stoliczkai (Steindachner), A. tokobajevi (Yeriomtschenko & Szczerbak), A. yarkandensis
Distribution: Kyrgyzstan, Kashmir (India), northeastern Afghanistan, northern Pakistan
Comments: We follow Auffenberg et al. (2004) in synonymizing Altiphylax boehmei with A. stoliczkai. The
specific distinctness of A. yarkandensis remains uncertain (Khan 1994; Auffenberg et al. 2004). We follow Das et
al. (1998) and Auffenberg et al. (2004) in regarding the origin of the types of this species as Ladakh, Jammu and
Kashmir rather than Yarkand , Xinijang, China. We accept Sindaco & Jerem
enko’s (2008) proposed movement of
A. levitoni, the type species of Asiocolotes, to Altiphylax and the transfer of Asiocolotes depressus to Microgecko.
We retain, however, Khan’s (2009) allocation of walli, assigned to Altiphylax by Sindaco & Jerem
enko (2008), to
Asiocolotes [see Altiphylax and Microgecko]
Comments: Junior subjective synonym of Altiphylax fide Sindaco & Jerem enko (2008). Asiocolotes
depressus, however, is allocated to Microgecko fide Sindaco & Jerem
Bunopus (4 species)
Composition: B. blanfordii Strauch, B. crassicauda Nikolsky, B. spatalurus Anderson (B. s. hajarensis
Arnold), B. tuberculatus Blanford
Distribution: Jordan, Syria, Arabian Peninsula, Iraq, Iran, southeastern Turkmenistan, Afghanistan and Pakistan
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Comments: Bunopus blanfordii is not recognized by certain authors (e.g., Szczerbak & Golubev 1986, 1996;
Sindaco & Jerem
enko 2008) but is here considered as valid pending further investigation.
Carinatogecko [see Mediodactylus]
Comments: We follow ervenka et al. (2010) in synonymizing Carinatogecko with Mediodactylus.
Crossobamon (2 species)
Composition: C. eversmanni (Wiegmann), (C. e. lumsdeni (Boulenger)), C. orientalis (Blanford)
Distribution: northwest India, Pakistan, eastern Iran, northern Afghanistan, Turkmenistan, Uzbekistan,
Tajikistan, southern Kazakhstan
Comments: The status of Stenodactylus lumsdeni Boulenger is unstable. Anderson (1999) considered this as a
synonym of Bunopus tuberculatus, but we follow Szczerbak & Golubev (1986, 1996) in regarding it as related to
Crossobamon eversmanni. These authors regarded it as a valid subspecies of C. eversmanni and treated
Stenodactylus maynardi Smith as its junior synonym. Having not examined specimens, we withhold judgement on
its status and simply follow Szczerbak & Golubev (1986, 1996) pending further investigation.
Cyrtodactylus (170+ species, 12 of which — listed here — have been variously considered as allied to
Palearctic naked-toed geckos )
Composition: C. battalensis (Khan), C. dattanensis (Khan), C. fasciolatus (Blyth), C. himalayanus (Duda &
Sahi), C. lawderanus (Stoliczka), C. malcolmsmithi (Constable), C. markuscombaii (Darevsky, Helfenberger,
Orlov & Shah), C. martinstolli (Darevsky, Helfenberger, Orlov & Shah), C. medogensis (Zhao & Li), C. nepalensis
(Schleich & Kästle), C. tibetanus (Boulenger), C. zhaoermii Shi & Zhao
Distribution: southern Tibet, southern flanks of Himalayas from northern India and Nepal eastwards to
Southeast Asia, Sri Lanka, Indonesia, Philippines, New Guinea, Solomon Islands, northern Australia
Comments: Although Khan & Rösler (1999) included lawderanus in their stoliczkai group of “Circum-
Himalayan” Cyrtodactylus, Khan subsequently moved it to the “tibetinus [sic] group” which he later (Khan 2003c)
named Siwaligekko. All Siwaligekko are here considered to be Cyrtodactylus, although the name may be retained at
the subgeneric level for members of this basal grade of Tibeto-Himalayan geckos (see text). Cyrtopodion
medogensis is tentatively included in this genus. Originally described as a Tenuidactylus, it has recently been
moved to Cyrtodactylus by Li et al. (2010). Although body proportions and its distribution in eastern Tibet are
consistent with Cyrtodactylus, body and especially tail tuberculation are uncharacteristically pronounced for this
Cyrtopodion (22 species)
Composition: C. agamuroides* (Nikolsky), C. aravallense (Gill), C. baigii Masroor, C. belaense Nazarov,
Ananjeva, & Papenfuss, C. brevipes (Blanford), C. fortmunroi
(Khan), C. gastrophole* (Werner), C. golubevi*
Nazarov, Ananjeva, & Rajabizadeh, C. indusoani
(Khan), C. kachhense (Stoliczka) (C. k. ingoldbyi (Procter)), C.
kiabii* Ahmadzadeh, Flecks, Torki & Böhme, C. kirmanense (Nikolsky), C. kohsulaimanai (Khan), C.
mansarulum (Duda & Sahi), C. montiumsalsorum (Annandale), C. persepolense* Nazarov, Ananjeva, &
Rajabizadeh, 2010, C. potoharense Khan, C. rhodocauda
(Baig), C. rohtasfortai
(Khan & Tasnim), C. scabrum
(Heyden), C. sistanense Nazarov & Rajabizadeh, C. watsoni (Murray)
Distribution: northeastern Egypt, Sinai, eastern Turkey, Israel, Jordan, Syria, Azerbaijan, Arabian Peninsula,
Iraq, Iran, Afghanistan, Central Asia, Pakistan, northern India, introduced on Red Sea coast of Africa
Comments: We include the apparently monophyletic agamuroides clade sensu Anderson (1999) (taxa marked
by asterisks) within Cyrtopodion based on the phylogenetic results of
ervenka et al. (2010). All members of
Indogekko sensu Khan (2003c) except longipes and voraginosus are also subsumed within Cyrtopodion (see text).
We suggest the use of Indogekko as a subgeneric name for most of those species assigned to it by Khan (2003c)
(taxa marked by hash mark above).
Indogekko [see Cyrtopodion]
Comments: We synonymize Indogekko (exclusive of the species elongatus and voraginosus) with Cyrtopodion
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PALEARCTIC NAKED-TOED GECKO PHYLOGENY
but suggest its use as a subgeneric name for the taxa assigned to it by Khan (2003c) except longipes and
Mediodactylus (14 species)
Distribution: southern Italy, Balkan Peninsula, Crimea (Ukraine), Turkey, Cyprus, Lebanon, Israel, Syria, Iraq,
Iran, Pakistan, Afghanistan, Kyrgyzstan, Tajikistan, Turkmenistan, Uzbekistan, southern Khazakhstan, southern
Russia, northern Xinjiang (China)
Composition: M. amictopholis (Hoofien), M. aspratilis (Anderson), M. brachykolon (Krysko, Rehman, &
Auffenburg), M. dehakroensis (Masroor), M. heterocercus (Blanford) (M. h. mardinensis (Mertens)), M.
heteropholis (Minton, Anderson & Anderson, 1970), M. ilamensis (Fathinia, Karamiani, Darvishia, Heidari &
Rastegar-Pouyani), M. kotschyi (Steindachner), (M. k. adelphiensis (Beutler & Gruber), M. k.bartoni (Št
M. k. beutleri (Baran & Gruber), M. k. bibroni (Beutler & Gruber), M. k. bolkarensis Rösler, M. k. buchholzi
(Beutler & Gruber), M. k. ciliciensis (Baran & Gruber), M. k. colchicus (Nikolsky), M. k. concolor (Bedriaga), M.
k. danilewskii (Strauch), M. k. fitzingeri (Št
pánek), M. k. fuchsia (Beutler & Gruber), M. k. kalypsae (Št pánek),
M. k. karabagi (Baran & Gruber), M. k. lycaonicus (Mertens), M. k. maculates (Bedriaga), M. k. oertzeni
(Boettger), M. k. orientalis (Št
pánek), M. k. ponticus (Baran & Gruber), M. k. rumelicus (Müller), M. k. saronicus
(Werner), M. k. schultzewestrumi (Beutler & Gruber), M. k. skopjensis (Karaman), M. k. solerii (Wettstein), M. k.
pánek), M. k. stepaneki (Wettstein), M. k. syriacus (Št pánek), M. k. tinensis (Beutler & Frör), M.
k. unicolor (Wettstein), M. k. wettsteini (Št
pánek)), M. narynensis Jerjomtschenko, (Zarinenko & Panfilow), M.
russowii (Strauch) (M. r. zarudnyi (Nikolsky)), M. sagittifer (Nikolsky), M. spinicauda (Strauch), M.
stevenandersoni (Torki), M. walli (Ingoldby)
Comments: We here include the four species included in the analyses of Macey et al. (2000) and this paper, as
well as the three species assigned to Carinatogecko by Sindaco & Jerem
enko (2008) and Torki (2011). We follow
ervenka & Kratochvíl (2010) in assigning Cyrtopodion dehakroense to Mediodactylus, and Khan (2003c, 2006,
2009) in assigning walli to this genus. The inclusion of Cyrtopodion brachykolon to Mediodactylus is based on the
results presented in this paper.
Microgecko (4 species)
Composition: M. depressus (Minton & Anderson), M. helenae Nikolsky (M. h. fasciatus (Schmidtler &
Schmidtler)), M. latifi (Leviton & Anderson), M. persicus (Nikolsky) (M. p. bakhtiari (Minton, Anderson &
Anderson), M. p. euphorbiacola (Minton, Anderson & Anderson))
Distribution: Iran, Pakistan, Rajasthan (India)
Comments: We follow Sindaco & Jerem
enko (2008) in assigning Asiocolotes depressus to Microgecko. The
extension of M. p. euphorbiacola into Rajasthan was documented by Agarwal (2009).
Pseudoceramodactylus (1 species)
Composition: P. khobarensis Haas
Distribution: Eastern Arabian Peninsula (Saudi Arabia, United Arab Emirates), isolated records from southern
Oman and Qeshm Island, Iran.
Comments: Pseudoceramodactylus is revalidated based on the phylogeny of Fujita & Papenfuss (2011), which
identified the sole species as the sister to Stenodactylus plus Tropiopcolotes.
Rhinogecko (2 species)
Composition: R. femoralis (Smith), R. misonnei de Witte
Distribution: Iran and adjacent Pakistan
Comment: In the absence of phylogenetic data to the contrary, we tentatively accept the validity of this genus,
following the arguments of Anderson (1999).
Siwaligekko [see Cyrtodactylus]
Comments: We synonymize Siwaligekko with Cyrtodactylus based on our phylogenetic results and those of
Wood et al. (2012). Siwaligekko is applicable at a subgeneric level to those species most closely related to C.
battalensis; however, given the uncertainty of relationships among basal Cyrtodactylus (recovered as paraphyletic
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320 · Zootaxa 3599 (4) © 2013 Magnolia Press
in this paper), we suggest that Siwaligekko be retained as a subgeneric name for all members of the genus occurring
in the western Himalayan region. Khan’s (2003) inclusion of Geckoella in Siwaligekko is not supported, but this
genus is also embedded within Cyrtodactylus sensu lato (Wood et al. 2012).
Stenodactylus (12 species)
Composition: S. affinis (Murray), S. arabicus (Haas), S. doriae (Blanford), S. grandiceps (Hass), S.
leptocosymbotus Leviton & Anderson, S. mauritanicus (Duméril & Bibron), S. petrii Anderson, S. pulcher
Anderson , S. slevini Haas, S. stenurus Werner, S. sthenodactylus (Lichtenstein) (S. s. mauritanicus Guichenot), S.
Distribution: North Africa, Israel, Jordan, Lebanon, Syria, southern Turkey, Iraq, southern Iran, Arabian
Peninsula, isolated populations in northwest Kenya
Comments: Stenodactylus khobarensis was recently removed to the revalidated Pseudoceramodactylus by
Fujita & Papenfuss (2011).
Tenuidactylus (7 species)
Composition: T. caspius (Eichwald), T. dadunensis (Shi & Zhao), T. elongatus (Blanford), T. fedtschenkoi
(Strauch), T. longipes (Nikolsky) (T. l. microlepis (Lantz)), C. turcmenicus Darevsky, T. voraginosus (Leviton &
Distribution: Azerbaijan, southwestern Russia, southern Kazakhstan, Turkmenistan, Uzbekistan, Tajikistan,
Iran, Afghanistan, northern Pakistan, northwestern China, southern Mongolia, introduced in Armenia, Kyrgyzstan.
Comments: Cyrtopodion elongatum and its former subspecies, C. voraginosum were assigned to Indogekko by
Khan (2003c), but the position of the former in our phylogeny is consistent with these taxa being members of
Tenuidactylus. We have retained Szczerbak & Golubev’s (1986, 1996) subspecific use of T. l. microlepis (see also
Anderson 1999), but we regard T. voraginosus as a distinct species. Based on the similarities between T. elongatus
and the recently described Cyrtopodion daduense we tentatively place the latter species in Tenuidactylus as well, as
Tropiocolotes (9 species)
Composition: T. algericus Loveridge, T. bisharicus Baha el Din, T. nattereri Steindachner, 1901, T. nubicus
Baha el Din, T. scortecci Cherchi & Spano, T. somalicus Parker, T. steudneri (Peters), T. tripolitanus Peters (T. t.
occidentalis Parker, T. t. apoklomax Papenfuss), T. wolfgangboehmei Wilms, Shobrak & Wagner
Distribution: North Africa from Mauritania to northern Somalia to the Mediterranean, Arabian Peninsula,
Comments: Species from Iran through Rajasthan previously assigned to this genus are now recognized as
members of the distantly related Microgecko (Gamble et al. 2012; this study). Both Asiocolotes (here not regarded
as valid) and Microgecko were previously considered subgenera of Tropiocolotes.
We thank Mohammed Al-Mutairi, Jon Boone, Indraneil Das, Anslem de Silva, Tony Gamble, L. Lee Grismer, Rod
Hitchmough, Fred Kraus, Jim McGuire, Jane Melville, Bob Murphy, Alan Resetar, Ross Sadlier, Carol Spencer,
Miguel Vences, Jens Vindum, and Greg Watkins-Colwell for providing some of the tissue samples used in this
study. Sayantan Biswas and Perry Wood, Jr. provided assistance in the laboratory. High resolution x-ray CT scans
were prepared at the University of Texas at Austin, Department of Geological Sciences. This research was was
supported by grants DEB 0844523 and DEB1019443 from the National Science Foundation of the United States
and by the Gerald M. Lemole, M.D. Endowed Chair funds.
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