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INTRODUCTION
The ‘Hemidactylus group’ of geckos (Hemidac-
tylus, Briba, Cosymbotus, Dravidogecko, Tera-
tolepis) was identied by Russell (1972, 1976,
1979) on the basis of internal digital anatomy.
Bauer and Russell (1995) formally synonymized
the monotypic peninsular Indian genus Dravi-
dogecko with Hemidactylus based on a more de-
tailed consideration of pedal anatomy. A recent
species level phylogeny of Hemidactylus based
on the mitochondrial genes cytochrome b and
12S (Carranza and Arnold, 2006) revealed that
two other members of the Hemidactylus group-
Cosymbotus (two species, tropical Asia) and
Briba (monotypic, Brazil)- were in fact embed-
ded deeply within it, rendering Hemidactylus
paraphyletic and consequently requiring taxo-
nomic action. Although Carranza and Arnold
(2006) did not take this action themselves, Zug
et al. (2007) formally included the species pre-
viously allocated to Cosymbotus in Hemidacty-
lus and similar action is required in the case of
Briba. Until now the phylogenetic position of
the remaining small genus in the group, Tera-
tolepis (two species, India and Pakistan), has
not been formally assessed.
The taxonomic history of Teratolepis is
disproportionately convoluted and complex,
Hamadryad Vol. 33, No. 1, pp. 13 – 27, 2008.
Copyright 2008 Centre for Herpetology, Madras Crocodile Bank Trust.
ON THE SYSTEMATICS OF THE GEKKONID GENUS TERATOLEPIS
GÜNTHER, 1869: ANOTHER ONE BITES THE DUST
Aaron M. Bauer1, Varad B. Giri2, Eli Greenbaum1, Todd R. Jackman1,
Mahesh S. Dharne3 and Yogesh S. Shouche3
1Department of Biology, Villanova University, 800 Lancaster Avenue,
Villanova, Pennsylvania 19085, U.S.A.
Email: aaron.bauer@villanova.edu, eli.greenbaum@villanova.edu,
todd.jackman@villanova.edu
2Herpetology Section, Collection Department, Bombay Natural History Society,
Hornbill House, S. B. Singh Road, Mumbai 400 023, Maharashtra, India.
Email: varadgiri@gmail.com
3 Molecular Biology Unit, National Centre for Cell Science, Ganeshkhind,
Pune 411007, Maharashtra, India.
Email: dmahesh@nccs.res.in, yogesh@nccs.res.in
(with two text-gures)
ABSTRACT.– A molecular phylogenetic analysis (cyt b, ND4, RAG-1 and PDC genes) of the
two recognized species (fasciata and albofasciata) of Teratolepis and representatives of all
major clades of Hemidactylus reveals that Teratolepis is embedded within the Tropical
Asian clade of the latter genus. Its closest relatives are the other small terrestrial
South Asian species, H. reticulatus and H. gracilis. The monophyly of Tropical Asian
Hemidactylus as a whole is not supported, but the terrestrial clade including Teratolepis
is well supported as the sister-group to H. brookii among the taxa sampled. Hemidactylus
anamallensis and H. scabriceps are likely members of this clade as well. Synonymization of
Teratolepis with Hemidactylus follows earlier actions sinking other small (Cosymbotus) or
monotypic (Dravidogecko, Briba) genera that rendered Hemidactylus paraphyletic. This
action necessitates a new specic epithet for Hemidactylus fasciatus (Blyth, 1853) which
is a junior secondary homonym of Hemidactylus fasciatus Gray 1842, a widespread and
common West African gecko. We therefore here erect the replacement name Hemidactylus
imbricatus nomen novum.
KEY WORDS. – Gekkonidae, Hemidactylus, Teratolepis, molecular phylogeny, taxonomy.
14 Hamadryad [Vol. 33, No. 1
given that only two species, T. fasciata (Blyth,
1853) and T. albofasciata (Grandison and So-
man, 1963) are presently recognized (Rösler,
2000; Kluge, 2001). Although the latter is poor-
ly known, the former is fairly common in the
pet trade (Mudrack, 1977, 1986; Girard, 1993;
Klarsfeld, 2001; Poulíček, 2002; Henkel and
Schmidt, 2003).
Teratolepis fasciata (Blyth, 1853) was origi-
nally described as Homonota fasciata from an
unstated locality in British India. Blyth’s (1853)
allocation to Homonota, however, was incon-
sistent with Gray’s (1845) generic diagnosis of
that genus, and Günther (1869) subsequently
erected the new genus Teratolepis to accommo-
date the species, emphasizing its bizarre scala-
tion (relatively large, at, weakly imbricate
polygonal scales on the dorsum and very large
imbricate scales on tail, which is typically swol-
len basally).
Kluge (1964), in revising the genus Homono-
ta, now restricted to South America, moved
Gymnodactylus fasciatus Duméril and Bibron,
1836 into Homonota, creating the new combi-
nation H. fasciata — a junior secondary homo-
nym of Blyth’s (1853) name. Kluge (1964) was
aware of the earlier name and indicated “[non]
Homonota fasciata: Jerdon [sic], 1853, p. 468”
in his account of this species. Wermuth (1965)
subsequently provided a replacement name for
G. fasciatus, G. pasteuri, in order to deal with
another instance of secondary homonymy, that
between G. fasciatus Duméril and Bibron, 1836
and Uromastix fasciatus Ménétriès, 1832, a
junior subjective synonym of Gymnodactylus
caspius Eichwald, 1831 (now Tenuidactylus
caspius). In an addendum to the same work,
Wermuth (1965) incorporated Kluge’s (1964)
generic reallocations and transferred G. fas-
ciata Duméril and Bibron, 1836 to Homonota,
as Homonota pasteuri (nomen novum). Al-
though neither name was cited extensively in
the following decades, Vanzolini (1968) and
Cei (1978) used Wermuth’s replacement name,
whereas Kluge (1991, 1993) continued to use
H. fasciata. Abdala and Lavilla (1993) provided
further evidence to support Kluge’s usage and
subsequently, most authors (e.g., Abdala, 1993,
1998; Dirksen and de la Riva, 1999; Rösler,
2000; Kluge, 2001) have employed this name.
Abdala and Lavilla (1993) based their argu-
ment on the fact that the removal of H. fasciata
to Teratolepis by Günther (1869) obviated the
need for a replacement name.
Annandale (1906) described a second species,
Teratolepis scabriceps from Rámanád [= Ram-
nad], Madura [= Madurai] District, Tamil Nadu
but this species, which has subsequently been
collected in Sri Lanka as well (Mariccukatti [=
Marichchukkaddi], Northern Province), was re-
moved to a new genus, Lophopholis, by Smith
and Deraniyagala (1934) on the basis that it
possessed the imbricate scales, but not the undi-
vided scansors of T. fasciata. Smith (1935) also
reported the locality Adiyar [= Adayar] near Ma-
dras [= Chennai] for this species. Deraniyagala
(1953) recognized the afnity of Lophopholis to
Hemidactlyus by including both genera (along
with the dissimilar Calodactylodes) in his sub-
family Hemidactylinae and most subsequent au-
thors have allocated L. scabriceps to the genus
Hemidactylus (e.g., Loveridge, 1947; Kluge,
1991, 1993, 2001; Das and Andrews, 1997; de
Silva, 1996, 1998; Rösler, 2000; Das and de
Silva, 2005) although it has been retained as a
separate genus by some workers (e.g., Murthy,
1990; Tikader and Sharma, 1992).
Grandison and Soman (1963) described the
small terrestrial gecko Hemidactylus albofas-
ciatus from the villages of Dorle, Dabhil and
Gavkhadi in the Ratnagiri District of Mahar-
ashtra. This species has only partly divided sub-
digital scansors and imbricate scales on the tail.
Grandison and Soman (1963) suggested that
the afnities of H. albofasciatus were with H.
reticulatus, another small terrestrial Hemidac-
tylus with undivided proximal subdigital lamel-
lae. More recently Murthy (1990) suggested a
close relationship to another Indian endemic, H.
prashadi, although the basis for this is unclear
as the two taxa exhibit size, colour, scansor, and
body and tail scalation features that are greatly
dissimilar to one another. Kluge (1967) rst
transferred H. albofasciatus to Teratolepis based
on a personal communication from Jerry A. An-
derson. Subsequent workers have either retained
albofasciatus in Hemidactylus (Murthy, 1990;
Tikader and Sharma, 1992; Sharma, 2002), or
removed it to Teratolepis (e.g., Das et al., 1998;
Das, 2001; Rösler, 2000; Kluge, 2001), although
none have presented explicit justications for
their allocations.
October, 2008] Status of Teratolepis 15
The recent taxonomic history of both T. al-
bofasciata and H. scabriceps thus suggests that
Teratolepis and Hemidactylus are closely allied.
Indeed, as early as 1876 Theobald noted the
similarity of Teratolepis to Hemidactylus spp.,
especially with respect to head scalation. The
close relationship of these two genera was re-
cently conrmed by Han et al. (2004) and Feng
et al. (2007), who found that Hemidactylus and
Teratolepis shared, along with Agamura, Cros-
sobamon, Cyrtodactylus and Geckoella, a 9 bp
insertion and a 21 bp deletion in the nuclear gene
c-mos. Unfortunately, however, this analysis
used only generic exemplars and an even more
recent species-level phylogeny of Hemidactylus
based on mitochondrial genes (Carranza and
Arnold, 2006) did not include any specimens of
Teratolepis spp., nor any other endemic South
Asian taxa.
The recent rediscovery of Teratolepis albo-
fasciata by one of us (VG) at Dorle village pro-
vided genetic material to assess the relationships
of this taxon and precipitated a reinvestigation
of the status of Teratolepis and its afnities with
respect to Hemidactylus.
Materials and methods
Based on preliminary results from a broad scale
phylogenetic analysis of all gekkotan lizards
(Bauer, Jackman and Greenbaum, unpublished)
the afnities of Teratolepis with Hemidactylus
(Han et al., 2004; Feng et al., 2007) were con-
rmed. For this study, we thus included in our
ingroup the two species currently assigned to
Teratolepis, as well as representatives of Carran-
za and Arnold’s (2006) ve major clades, as well
as the West African species H. fasciatus, which
they sampled but did not include in any of their
named clades. As geographic proximity sug-
gested that afnities of Teratolepis would most
likely be with Carranza and Arnold’s “Tropical
Asian Clade”, we included representatives of all
of the constituent taxa reported on by Carranza
and Arnold (2006), as well as the Indian endem-
ics H. reticulatus and H. gracilis. These two spe-
cies have been proposed to be closely allied to
one another (Bauer et al., 2005) and the former
species had previously been predicted to be al-
lied to Teratolepis albofasciata (Grandison and
Soman, 1963). We used three representatives
of the chiey southeast Asian/Indo-Australian
genus Cyrtodactylus as outgroup taxa based on
the results of Han et al. (2004) and Feng et al.
(2007). Specimens sampled and their associated
clade membership based on Carranza and Ar-
nold (2006) are listed in Table 1.
Genomic DNA was isolated from 95–100%
ethanol-preserved tail or liver samples with the
Qiagen DNeasy tissue kit (Valencia, CA, USA).
We used double-stranded PCR to amplify 3733
aligned bases of mitochondrial (ND2, ND4, cyt
b) and nuclear (RAG1, PDC) gene sequence
data with ve different pairs of published prim-
ers (Table 2). For some key taxa, not all ve
genes could be sequenced: ND2, ND4 and
RAG1 lacking for Hemidactlyus reticulatus; cyt
b, ND4, RAG-1 and PDC lacking for H. gracilis
and Teratolepis albofasciata).
Amplication of 25 μl PCR reactions was
executed on an Eppendorf Mastercycler gradi-
ent thermocycler. Amplication of genomic
DNA occurred with an initial denaturation step
of 95°C for 2 min, followed by denaturation at
95°C for 35 s, annealing at 50°C for 35 s, and
extension at 72°C for 95 s with 4 seconds added
to the extension per cycle for 32 cycles for mito-
chondrial DNA and 34 cycles for nuclear DNA.
When necessary, annealing temperatures were
adjusted to increase or decrease specicity on a
case by case basis, and products were visualized
with 1.5% agarose gel electrophoresis. Target
products were puried with AMPure magnetic
bead solution (Agencourt Bioscience) and se-
quenced with either the BigDye® Terminator
v3.1 Cycle Sequencing Kit (Applied Biosys-
tems) or the DYEnamic™ ET Dye Terminator
Kit (GE Healthcare). Sequencing reactions were
puried with CleanSeq magnetic bead solution
(Agencourt Bioscience) and analyzed with an
ABI 3700 automated sequencer. The accuracy
of sequences was ensured by incorporating neg-
ative controls and sequencing complementary
strands. Sequences were aligned by eye in the
computer program SeqMan, and protein-cod-
ing genes were translated to amino acids with
MacClade (Maddison and Maddison, 1992) to
conrm conservation of the amino acid reading
frame and check for premature stop codons.
Phylogenetic relationships among the sam-
ples were assessed with maximum parsimony
and Bayesian optimality criteria. Based on the
missing data indicated above, three datasets
16 Hamadryad [Vol. 33, No. 1
were analyzed: RAG1, PDC, ND2, ND4 and cyt
b (all taxa except T. albofasciata, H. reticulatus
and H. gracilis), PDC and cyt b (all taxa except
T. albofasciata and H. gracilis), and ND2 alone
(all taxa except H. reticulatus).
Maximum parsimony (MP) analyses were
conducted in PAUP*4.0b10 (Swofford, 2002).
The heuristic search algorithm was used with
the following conditions: 25 random-addition
replicates, accelerated character transforma-
tion (ACCTRAN), 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 (1000
pseudoreplicates) to assess node support in re-
sulting topologies.
The Akaike Information Criterion (AIC) in
ModelTest 3.06 (Posada and Crandall, 1998)
was used to nd the model of evolution that best
t the data for subsequent Bayesian inference
(BI) analyses. The GTR + γ + I model was used
with the most parsimonious tree to estimate the
parameters, and the same conditions as the par-
simony search were used to nd the Bayesian
trees with the best likelihood scores.
Partitioned Bayesian analyses were conducted
with MrBayes 3.1 (Ronquist and Huelsenbeck,
2003) with default priors. Separate models for
each gene and codon position of protein-coding
genes were estimated (Brandley et al., 2005).
A total of 10 partitions were made: RAG1, 3
codons; PDC, 3 codons, ND2, ND4 and cyt b,
3 codon positions; and mitochondrial tRNAs.
Analyses were initiated with random starting
trees and run for 2,000,000 generations; Mark-
ov chains were sampled every 100 generations.
Convergence was checked by plotting likelihood
scores against generation, and 125 trees were
discarded as “burn in.” Two separate analyses
with two independent chains were executed to
check for convergence of log-likelihoods in
stationarity (Huelsenbeck and Ronquist, 2001).
Both analyses ended with the standard deviation
of split frequencies less than 0.01.
RESULTS
The dataset based on all genes yielded a Baye-
sian tree (Fig. 1) in which each of Carranza and
Arnold’s (2006) clades except the ‘Tropical
Asian Clade’ were monophyletic with posterior
probabilities of 1.0. The H. angulatus clade was
sister to the remaining members of the genus,
which were themselves divided into two weakly
supported clades (pp < 0.90): one consisting of
H. fasciatus as the sister group of a subset of the
Tropical Asian clade comprising H. garnotii, H.
karenorum, H. bowringii and H. platyurus (we
here follow Zug et al., 2007 in formally allo-
cating the members of the genus Cosymbotus
to Hemidactylus), and another with the remain-
ing Tropical Asian forms as the sister group to
the Arid clade plus the African-Atlantic and H.
mabouia clades. Support for interclade relation-
ships was generally weak, with only the union
of H. mabouia with the African-Atlantic clade
receiving strong support (pp = 1.0). Within the
Tropical Asian species, all relationships in both
clades, except the sister group relationship of
H. garnotii and H. bowringii were supported by
posterior probabilities of 1.0. In this analysis,
Teratolepis fasciata was strongly supported as
the sister species of H. brookii, with H. frenatus
and H. aviviridis as sequentially more distant
relatives.
In the Bayesian analysis of ND2 alone, all
of Carranza and Arnold’s (2006) clades were
monophyletic, although the Tropical Asian clade
had no signicant support (pp = 0.62). All other
interclade relationships were well-supported, but
the tree topology differed greatly from that of
the previous analysis: Arid clade ((H. mabouia,
African-Atlantic clade) ((H. fasciatus, H. angu-
latus clade) Tropical Asian clade)). Within the
Tropical Asian clade, the same two groupings
revealed by the larger data set were recovered,
but in this case H. karenorum was weakly sup-
ported as the sister of H. garnotii (pp = 0.60). In
the H. brookii subclade, relationships among the
species dealt with in the previous analysis were
identical. The added taxa, Teratolepis albofas-
ciata and H. gracilis were strongly supported
as sequential sister taxa to Teratolepis fasciata,
with all three taxa sister to H. brookii (Fig. 2A).
In the PDC and cyt b analyses, all higher
order relationships, except the monophyly of
Hemidactylus sensu lato and the union of H.
mabouia with the African-Atlantic clade (pp =
1.0), received weak support and were differ-
ent from both previous analyses: H. angulatus
clade (Arid clade (H. fasciatus (H. mabouia,
October, 2008] Status of Teratolepis 17
African-Atlantic clade) Tropical Asian clade))).
Relationships also differed within the Tropical
Asian clade, with H. aviviridis more closely
related to the H. bowringii group than to the H.
brookii group. However, the remaining relation-
ships within the H. brookii group were identi-
cal to both other analyses with respect to shared
taxa (Fig. 2B). The additional taxon for which
only PDC and cyt b were available, H. reticu-
latus, was strongly supported as the sister to T.
fasciata.
Maximum parsimony analyses of the same
datasets yielded less well-resolved trees that
were, however, fully consistent with the Baye-
sian analyses. Parsimony bootstrap support was
high (> 90%) for all clades that also had high
posterior probabilities (Fig. 1). In the ND2 and
PDC + cyt b analyses, H. gracilis and Tera-
tolepis albofasciata, and H. reticulatus, respec-
tively, were likewise strongly supported as con-
stituting a monophyletic group with T. fasciata
(Fig. 2).
DISCUSSION
Phylogeny.– With respect to the monophyly of
each of the clades identied by Carranza and
Arnold (2006), our results are consistent with
these authors’ own ndings of moderate to high
support values. On the other hand, our different
analyses yielded different patterns of relation-
ship among these clades, an unsurprising result,
given that Carranza and Arnold (2006) reported
no signicant support values for inter-clade
relationships in their analyses. However, our
data did yield consistent support for the sister
group relationship of H. mabouia and the Af-
rican-Atlantic clade. The position of H. fascia-
tus remains problematic, grouping weakly with
the bowringii group of the Tropical Asian clade
(all genes) or the H. mabouia/African-Atlantic
clade + all Tropical Asian taxa (PDC and cyt b),
or strongly with the H. angulatus clade (ND2).
Patterns within the Tropical Asian clade are
more consistent. The two clades reected in
Carranza and Arnold’s (2006) results, the H.
bowringii and H. brookii groups, are always
retrieved, although in the PDC/cyt b tree, H.
aviviridis clustered weakly with the H. bow-
ringii group, rather than with the H. brookii
group. Patterns within the H. bowringii group
varied slightly between analyses and from
those reported by Carranza and Arnold (2006).
However, patterns within the H. brookii group
were consistent and in the ve-gene analysis,
Teratolepis fasciata was the sister group to H.
brookii. Based on the ND2 data, the two species
of Teratolepis are each other’s sister species and
are nested deep within the Hemidactlyus tree,
closer to H. gracilis than to H. brookii. Although
we lack ND2 data for H. reticulatus, this spe-
cies is strongly supported as the sister group to
Teratolepis by cyt b data (lacking for H. gracilis
and T. albofasciata). With the available data, it
is not possible to resolve the relationships of H.
gracilis, H. reticulatus and Teratolepis to one
another, but it is clear that all four are members
of a single, well-supported clade.
Teratolepis is just one of several small gen-
era of geckos that have long been recognized
as allied to Hemidactylus, but which have been
segregated because of their possession of one or
more recognizable features, which although di-
agnostic, are probably best regarded as autapo-
morphic and not indicative of higher relation-
ships. Parker (1942), who studied the speciose
and morphologically diverse Hemidactylus of
the Horn of Africa, considered generic arrange-
ments that partitioned Hemidactylus-like gen-
era into different groups to be largely arbitrary.
However, prevailing systematic views of the pe-
riod favored the recognition of morphologically
distinctive groups, even if this rendered other
groups paraphyletic.
The strongly supported inclusion of Tera-
tolepis within Hemidactylus is not surprising.
Earlier authors (Grandison and Soman, 1963;
Murthy, 1990) hypothesized close relation-
ships between T. albofasciata and South Asian
Hemidactylus, and Anderson (1964) noted simi-
larities in the vocalizations of T. fasciata in the
Indus Delta plain to those of sympatric Hemi-
dactylus. Russell (1972, 1976, 1979), who ar-
gued that internal digital anatomy was a more
reliable indicator of homology and afnity than
external form, dened a ‘Hemidactylus group’
of geckos (Hemidactylus, Briba, Cosymbotus,
Dravidogecko, Teratolepis) based on a series
of apparently derived internal digital structures:
dorsal interossei muscles robust — with eshy
bellies extending as far as the digital inection,
tendinous insertion of dorsal interossei muscles
onto distal margin of each scansor, and antepe-
18 Hamadryad [Vol. 33, No. 1
Table 1. List of samples used in this study and their membership in the clades of Hemidactylus identied by Carranza and Arnold (2006). Collection abbreviations: AMB =
Aaron M. Bauer eld series, BNHS = Bombay Natural History Society, CAS = California Academy of Sciences, FK = Fred Kraus eld series, FMNH = Field Museum of
Natural History, ID = Indraneil Das eld series, JFBM = James Ford Bell Museum, University of Minnesota, St. Paul, JS = Jay Sommers (Kansas City), KU = University
of Kansas Natural History Museum, LLG = L. Lee Grismer eld series, LSUMZ = Louisiana State University Museum of Natural Sciences, MCZ = Museum of Com-
parative Zoology, Harvard University, MVZ = Museum of Vertebrate Zoology, University of California, Berkeley, MZUSP = Museu de Zoologia da Universidade de São
Paulo, WRB = William R. Branch eld series.
Sample
Clade Member-
ship (Carranza &
Arnold, 2006)
Museum No. Locality
GenBank Accession Numbers
cyt b ND2 ND4 RAG-1 PDC
Cyrtodactylus
ayeyawardyensis —/outgroup CAS 216446 Myanmar, Rakhine State, vic.
Kanthaya Beach EU268380 EU268348 EU268411 EU268287 EU268317
Cyrtodactylus consobrinus —/outgroup LLG 4062 Malaysia, Sarawak, Niah Cave EU268381 EU268349 EU268412 EU268288 EU268318
Cyrtodactylus loriae —/outgroup FK 7709
Papua New Guinea, Milne Bay
Province, Bunisi, N slope of Mt.
Simpson
EU268382 EU268350 EU268413 EU268289 EU268319
Hemidactylus cf. angulatus H. angulatus MVZ 245438 Nigeria, Togo Hills, Nkwanta EU268399 EU268367 EU268430 EU268306 EU268336
Hemidactylus bowringii 1 Tropical Asian CAS 206649 Myanmar, Sagaing Division,
Alaungdau Kathapa Natl. Park EU268405 EU268373 EU268436 EU268312 EU268342
Hemidactylus bowringii 2 Tropical Asian CAS 228109 China, Yunnan Province, Nujang
District, Liuku EU268406 EU268374 EU268437 EU268313 EU268343
Hemidactylus brasilianus African-Atlantic MZUSP 92493 Brazil, Piauí, Parque Nacional
Serra das Confusões EU268383 EU268351 EU268414 EU268290 EU268320
Hemidactylus brookii 1 Tropical Asian LLG 6755 Malaysia, Pulau Pinang, Empan-
gon Air Hitam EU268398 EU268366 EU268429 EU268305 EU268335
Hemidactylus brookii 2 Tropical Asian LLG 6754 Malaysia, Pulau Pinang, Empan-
gon Air Hitam EU268397 EU268365 EU268428 EU268304 EU268334
Hemidactylus brookii 3 Tropical Asian CAS 206638 Myanmar, Mandalay Division EU268407 EU268375 EU268438 EU268314 EU268344
Hemidactylus fasciatus 1 not placed in clade WRB no number Gabon, Rabi EU268402 EU268370 EU268433 EU268309 EU268339
Hemidactylus fasciatus 2 not placed in clade CAS 207777 Equatorial Guinea, Bioko Island,
3.6 km N of Luba EU268403 EU268371 EU268434 EU268310 EU268340
Hemidactylus aviviridis 1 Tropical Asian FMNH 245515 Pakistan, Punjab Province EU268387 EU268355 EU268418 EU268294 EU268324
Hemidactylus aviviridis 2 Tropical Asian ID 7626 India, Rajasthan, Kuldhara EU268388 EU268356 EU268419 EU268295 EU268325
October, 2008] Status of Teratolepis 19
Hemidactylus frenatus 1 Tropical Asian LLG 6745 Malaysia, Pulau Pinang, Empan-
gon Air Hitam EU268390 EU268358 EU268421 EU268297 EU268327
Hemidactylus frenatus 2 Tropical Asian AMB 7411 Sri Lanka, Pidipitiya EU268389 EU268357 EU268420 EU268296 EU268326
Hemidactylus frenatus 3 Tropical Asian AMB 7420 Sri Lanka, Rathegala EU268391 EU268359 EU268422 EU268298 EU268328
Hemidactylus garnotii 1 Tropical Asian CAS 223286
Myanmar, Rakhine State, Taung
Gok Township, Ma Ei Ywa Ma
Village
EU268395 EU268363 EU268426 EU268302 EU268332
Hemidactylus garnotii 2 Tropical Asian CAS 222276
Myanmar, Mon State, Kyaihto
Township, Kyait Hti Yo Wildlife
Sactuary
EU268396 EU268364 EU268427 EU268303 EU268333
Hemidactylus gracilis not included BNHS 1592 India, Maharashtra, Pune —EU268379 ———
Hemidactylus greefi African-Atlantic CAS 219044 São Tome and Principe, São Tome
Island, Praia da Mutamba EU268401 EU268369 EU268432 EU268308 EU268338
Hemidactylus haitianus H. angulatus CAS 198442 Dominican Republic, Nacional
Dist., near Santo Domingo EU268404 EU268372 EU268435 EU268311 EU268341
Hemidactylus karenorum Tropical Asian CAS 210670
Myanmar, Mandalay Division,
Kyaukpadaung Township, Popa
Mt. Park
EU268394 EU268362 EU268425 EU268301 EU268331
Hemidactylus mabouia H. mabouia MCZ R-184446 South Africa, Limpopo Province EU268393 EU268361 EU268424 EU268300 EU268330
Hemidactylus palaichthus African-Atlantic LSUMZ H-
12421 Brazil, Roraima State EU268400 EU268368 EU268431 EU268307 EU268337
Hemidactylus persicus Arid CAS 227612 Oman, Wilayat Nazwa, 4.5 km N.
of Tanuf, Wadi Tanuf EU268409 EU268377 EU268440 EU268316 EU268346
Hemidactylus platyurus Tropical Asian KU 304111 Philippines, Lubang EU268384 EU268352 EU268415 EU268291 EU268321
Hemidactylus reticulatus not included AMB 5730 India, Tamil Nadu, Vellore EU268410 ———EU268347
Hemidactylus robustus Arid MVZ 248437 Pakistan, Thatta District, 40 km S
of Mipur Sakro EU268408 EU268376 EU268439 EU268315 EU268345
Hemidactylus turcicus Arid LSUMZ H-
1981 USA, Louisiana, Baton Rouge EU268392 EU268360 EU268423 EU268299 EU268329
Teratolepis albofasciata not included BNHS 1579 India, Maharashtra, Ratnagiri
District, Dorle Village —EU268378 ———
Teratolepis fasciata not included JS 11 Pakistan (captive specimen) EU268385 EU268353 EU268416 EU268292 EU268322
Teratolepis fasciata not included JFBM 2 Pakistan (captive specimen) EU268386 EU268354 EU268417 EU268293 EU268323
20 Hamadryad [Vol. 33, No. 1
nultimate phalanx on digits III and IV of manus
and III-V of pes short and erect. In addition, all
members of this group that have paraphalanges
possess the Hemidactylus type, lying within
the lateral digital tendons (Russell and Bauer,
1988).
Subsequent phylogenetic work has borne out
the evolutionary reality of this cluster of genera;
Carranza and Arnold (2006) demonstrated that
both Cosymbotus and Briba were embedded
within major clades of Hemidactylus — a fact
corroborated here. Both of these groups have
typical Hemidactylus-type divided subdigital
scansors and are arboreal.
The two Hemidactylus that are particularly
closely related to Teratolepis, H. gracilis and H.
reticulatus are poorly known, but both are pri-
marily terrestrial (Sanyal et al., 1993; Murthy,
1990; Bauer et al., 2005) as are the Teratolepis
species (Grandison and Soman, 1963; Anderson,
1964; Minton, 1966). As early as 1912 Annan-
dale suggested that H. gracilis (as H. platyceps)
was most closely related to H. reticulatus, and
in 1972, Russell had identied a group within
Hemidactylus that shared a distinctive pattern of
digital anatomy — this comprised H. albofas-
ciatus, H. gracilis and H. reticulatus, as well as
the Socotran species H. pumilo, which is also
terrestrial (Rösler and Wranik, 2000, 2003).
The remaining member of Russell’s Hemidac-
tylus group, Dravidogecko anamallensis, is also
chiey terrestrial or rupicolous (Gvoždik and
Veselý, 1998; Henkel and Schmidt, 2003). All
of these taxa share a number of morphological
features in common with each other that are
associated with their terrestrial habitus. Bauer
and Russell (1995) demonstrated that undivided
subdigital scansors were shared by T. fasciata,
H. reticulatus and Dravidogecko anamallensis
and that an intermediate pattern of divided dis-
talmost scansors occurred in T. albofasciata, as
well as a few Hemidactylus sensu stricto, such
as H. bouvieri and H. somalicus. Based on the
continuum between divided and undivided
scansors, the fact that at least one Hemidacty-
lus has completely undivided scansors, and their
shared internal digital anatomy, Bauer and Rus-
sell (1995) synonymized Dravidogecko with
Hemidactylus. Although they did not explic-
itly address the taxonomic status of Teratolepis,
identical arguments could be made for its syno-
nymization with Hemidactylus. Unfortunately,
we lacked tissue samples of Hemidactylus
(formerly Dravidogecko) anamallensis and are
unable to assess whether it is also a member of
the Teratolepis clade or if its undivided scansors
represent a convergent morphology within the
Tropical Asian clade of Hemidactylus, although
on biogeographic grounds, we suspect the
former interpretation. Bauer and Russell (1995)
concluded that H. anamallensis was a relatively
primitive Hemidactylus, but if we are correct
in our conjecture, it represents part of a highly
derived lineage that has undergone secondary
loss of some scansorial features. Hemidactylus
scabriceps may well also be a member of this
radiation as it shares imbricate scalation with
Teratolepis spp. This taxon is particularly poor-
Table 2. Primers used in this study.
Primer Gene Reference Sequence
ND4f11 ND4 Jackman et al. (2008) 5’-GCAAATACAAACTAYGAACG-3‘
Leur1 Leu tRNA Arevalo et al. (1994) 5’-CATTACTTTTTACTTGGATTTGCACCA-3‘
PHOF2 PDC Bauer et al. (2007) 5’-AGATGAGCATGCAGGAGTATGA-3’
PHOR1 PDC Bauer et al. (2007) 5’-TCCACATCCACAGCAAAAAACTCCT-3’
L4437b Met tRNA Macey et al. (1997) 5’-AAGCAGTTGGGCCCATACC-3’
L5002 ND2 Macey et al. (1997) 5’-AACCAAACCCAACTACGAAAAAT-3’
ND2f101 ND2 Greenbaum et al. (2007) 5’-CAAACACAAACCCGRAAAAT-3’
ND2r102 ND2 Greenbaum et al. (2007) 5’-CAGCCTAGGTGGGCGATTG-3’
Trpr3a Trp tRNA Greenbaum et al. (2007) 5’- TTTAGGGCTTTGAAGGC-3’
H5934a COI Macey et al. (1997) 5’- AGRGTGCCAATGTCTTTGTGRTT-3’
R13 RAG1 Groth and Barrowclough (1999) 5’- TCTGAATGGAAATTCAAGCTGTT-3’
R18 RAG1 Groth and Barrowclough (1999) 5’-GATGCTGCCTCGGTCGGCCACCTTT-3’
RAG1 F700 RAG1 Bauer et al. (2007) 5’-GGAGACATGGACACAATCCATCCTAC-3’
RAG1 R700 RAG1 Bauer et al. (2007) 5’-TTTGTACTGAGATGGATCTTTTTGCA-3’
October, 2008] Status of Teratolepis 21
Figure 1. Phylogenetic relationships among Teratolepis fasciata and representative species of Hemidactylus
based on the combined analysis of the mitochondrial genes ND2, ND4 and cyt b, and the nuclear genes RAG1
and PDC. Bayesian inference tree with branch lengths corresponding to those of tree with best likelihood score.
Bayesian posterior probabilities indicated above the branches and maximum parsimony bootstraps indicated
below. Bars at right indicate membership in the main clades of Hemidactylus identied by Carranza and Arnold
(2006). Note the lack of support for a monophyletic Tropical Asian Clade and the strong support for the inclu-
sion of Teratolepis fasciata within the brookii group.
22 Hamadryad [Vol. 33, No. 1
Figure 2. Phylogenetic relationships among members of the Hemidactylus brookii group based on ND2 alone
(A) and cyt b and PDC alone (B). The analyses of these particular partitions permitted the inclusion of key
taxa for which complete data were lacking: H. gracilis and Teratolepis albofasciata (A) and H. reticulatus (B).
Results of the phylogenetic analyses of all taxa sampled are presented in the text. Topologies depicted are from
the Bayesian tree and do not show likelihood branch lengths. Bayesian posterior probabilities indicated above
the branches and maximum parsimony bootstraps indicated below. The added taxa are strongly supported as
members of the brookii group and as close relatives of Teratolepis fasciata in particular.
October, 2008] Status of Teratolepis 23
ly known, with no new specimens having been
recorded since Smith (1935). Indeed, the valid-
ity of the Sri Lankan record, dating from 1933,
is questionable and relatively intense search
efforts by several groups of researchers (e.g.,
Manamendra-Arachchi, 1997) have yielded no
additional specimens. It has been considered
extirpated by some authors (Somaweera and
Wickramasinghe, 2006).
Taxonomic Implications.– Regardless of
the precise pattern of relationships among the
members of the Teratolepis group, it is clear
that maintenance of monophyletic groupings re-
quires their collective inclusion into Hemidac-
tylus. While this poses no problem in the case
of T. albofasciata, which reverts to the genus
within which it was initially described, it has
more far-reaching taxonomic implications for
Teratolepis fasciata which becomes Hemidac-
tylus fasciatus (Blyth, 1853), a secondary jun-
ior homonym of Hemidactylus fasciatus Gray,
1842, a common and widespread species of
West African gecko. There are no junior syno-
nyms available for T. fasciata (see synonymies
in Smith 1935; Wermuth 1965; Rösler 2000;
Kluge 2001) so a replacement name is required.
We, therefore, propose Hemidactylus imbrica-
tus Bauer, Giri, Greenbaum, Jackman, Dharne
& Shouche as a nomen novum for Hemidactylus
fasciatus (Blyth, 1853) in order to prevent sec-
ondary homonymy. The specic epithet refers to
the imbricate scales of the dorsum, and particu-
larly the tail of this species.
Comments on original description and dis-
tribution of Hemidactylus imbricatus, nom.
nov.– Blyth’s description of Homonota fasciata
has been cited as Blyth or Blyth in Jerdon “1854
(1853)” by several recent authors (e.g., Das et al.,
1998; Kluge, 2001) and many modern authors
(e.g., Hoge and Romano Hoge, 1981; Golay et
al., 1993; McDiarmid et al.,1999; David and In-
eich, 1999) have given the date of 1854 to spe-
cies described by Jerdon in the same paper. The
1854 date is almost certainly based on that on
the title page of the entire volume in which the
paper appeared. However, the original wrappers
on the specic part of the journal containing this
paper (Volume 22[VI] = issue CCXXXVII) are
dated 1853 and in the absence of information to
the contrary, we follow Bauer (2003) and accept
this date as correct.
Although Blyth (1853) did not specify the ori-
gin of the types of Homonota fasciata, Theobald
(1876) subsequently stated that the type locality
was “Jaulnah,” Hyderabad Province [= Jalna,
Maharashtra]. It is unclear from the description
if there was more than one type specimen, but
ZSI 5981, now in terrible condition, has been
regarded as the holotype. Das et al. (1998),
however, noted that BMNH 69.8.28.32, pre-
sented by Dr. A. H. Leith from Sind, might be a
syntype. This assertion was based on a penciled
note in the BMNH loose-leaf catalogue, but is
not substantiated by any other data (C. J. Mc-
Carthy in litt. April 2007). Indeed, it is clear that
this specimen is that examined and discussed by
Günther (1869).
Boulenger (1890) and Annandale (1905) con-
sidered T. fasciata to be distributed in Sind and
the Deccan — the latter based solely on Theo-
bald’s (1876) Jalna record. The populations in
Pakistan are well documented. Smith (1935),
Anderson (1964), Minton (1962, 1966), Mertens
(1969), and Khan (2002, 2004) reported T. fas-
ciata from the Indus Delta of Sind and Min-
ton (1966) considered it restricted to the Tatta
and Hyderabad districts of the province. Khan
(1999) characterized its habitat as tropical thorn
forest and sand dunes in semi-desert areas of the
Thar Desert and later (Khan, 2006) as Salsola
and grass-dominated areas of desert scrub veg-
etation on silt. Although some of the reported
localities to the east of the Indus River approach
the Indian border quite closely, there have been
no records from adjacent Rajasthan.
Aside from the Jalna record, the only locality
in the current territory of the Republic of India
is that reported by Smith (1935) from Shillong
in the Khasi Hills of north-eastern India. Both
Indian localities were considered erroneous by
Minton (1966) and Khan and Mirza (1977), and
Das (2001) considered only Pakistani records
to be valid. Nonetheless, Teratolepis fasciata
has continued to be listed as part of the Indian
fauna by many subsequent authors (e.g., Daniel,
1983, 2002; Murthy, 1990; Tikader and Sharma,
1992; Mathew, 1995; Sharma, 2002). Tikader
and Sharma (1992) added an unspecied record
from Tamil Nadu and also incorrectly included
Sri Lanka in the range of the species. All these
records are almost certainly incorrect. However,
one of the authors (VBG) has recently discov-
24 Hamadryad [Vol. 33, No. 1
ered a new species of Hemidactylus clearly
allied to H. imbricatus from Maharashtra (to
be described elsewhere) and it is possible that
this species (or H. albofasciatus or H. scabri-
ceps) might have been responsible for at least
the doubtful peninsular localities, particularly if
identication was based on the imbricating cau-
dal scalation shared by all of these taxa.
ACKNOWLEDGEMENTS
We thank Thomas Kennedy for assistance in the
laboratory and Indraneil Das for help and com-
panionship in the eld. Tissue samples were
kindly provided by the California Academy of
Sciences (Jens Vindum), the John Ford Bell
Museum, University of Minnesota (Tony Gam-
ble), The Museum of Vertebrate Zoology (Carla
Cicero), the Louisiana State University of Natu-
ral Science (Donna Dittmann, Fred Sheldon),
the Museum of Comparative Zoology (José
Rosado and James Hanken), the Field Museum
of Natural History (Alan Resetar), the Univer-
sity of Kansas Natural History Museum (Rafe
Brown), Museu de Zoologia da Universidade de
São Paulo (Hussam Zaher), Indraneil Das (Uni-
versiti Malaysia Sarawak), Lee Grismer (LaSi-
erra University), Fred Kraus (Bernice P. Bishop
Museum), William R. Branch (Bayworld, Port
Elizabeth) and Jay Sommers (Kansas City, Mis-
souri). Channakesava Murthy (Zoological Sur-
vey of India) and Colin McCarthy (The Natural
History Museum, London) kindly provided us
with photographs of specimens in their care.
MSD and YSS would like to acknowledge the
Department of Biotechnology, New Delhi, In-
dia for providing funding and a platform to con-
duct this study. Funding for this research was
provided by grant DEB 0515909 from the Na-
tional Science Foundation of the United States
(to AMB and TJ) and Villanova University. The
manuscript beneted from the comments of
George R. Zug and Stephen Mahony.
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Received: 10 October 2007.
Accepted: 1 November 2007.
