22 Accepted by S. Carranza: 30 May 2011; published: 20 Jun. 2011
ISSN 1175-5326 (print edition)
1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2924: 22–32 (2011)
A new species of angular-toed gecko, genus Cyrtopodion (Squamata: Gekkoni-
dae), from southern Iran
, MORRIS FLECKS
, FARHANG TORKI
& WOLFGANG BÖHME
Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute, Shahid Beheshti University, G.
Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany
FTEHCR (Farhang Torki Ecology and Herpetology Center for Research), 68319-16589, P. O. Box 68315-139, Nourabad City,
Lorestan Province, Iran
Corresponding author. E-mail: email@example.com
We describe a new species of the genus Cyrtopodion from the coastal area of Bushehr Province in southern Iran based on
morphometric and pholidotic data. Cyrtopodion kiabii sp. nov. belongs to the agamuroides-group and the key characters
to distinguish this species from all other members of the group are the lower number of ventrals and the extremely slender
habitus with long and delicate legs. Resulting from pholidosis, the observed presence of sexual dimorphism, and distribu-
tion we suggest a close relationship with C. gastropholis, which is also reviewed herein.
Key words: Cyrtopodion agamuroides-group, sexual dimorphism, taxonomy
The taxonomy of the species-rich genus Cyrtopodion Fitzinger, 1843 still is very unstable. Several groups have
been defined for the Iranian species (Anderson 1999) and are largely concordant to the described subgenera Cyr-
topodion, Mediodactylus and Tenuidactylus (Szczerbak and Golubev 1984, Böhme 1985, Kluge 1985). The most
important exception is the placement of C. agamuroides and C. gastropholis into an own group (named agamuroi-
des-group). These two species were even thought to be members of the genus Agamura (e.g. Minton 1966, Kluge
1991), which share the slender, long-legged habitus. Correspondingly, subsequent authors did not assign them to
any of the subgenera of Cyrtopodion, but listed them as incertae sedis (e.g. Sindaco and Jeremcenko 2008). Molec-
ular phylogenetics showed that Cyrtopodion is paraphyletic with respect to Agamura and Bunopus, and conse-
quently, Mediodactylus was elevated to full generic status (
ervenka et al. 2008) as already proposed by other
authors (Kluge 1991, Macey et al. 2000). Though the remaining taxa of Cyrtopodion are monophyletic, the rela-
tionships between the genus Agamura and the subgenera Cyrtopodion and Tenuidactylus, as well as the agamuroi-
des-group, are not yet resolved satisfactorily. Each of them represents a distinct clade within Cyrtopodion sensu
ervenka et al. 2008), which supports the isolated position of the agamuroides-group as realized by Anderson
(1999). Two recently described species—C. golubevi and C. persepolense—were assigned to the agamuroides-
group and it was assumed that this group is a large complex composed of eight to ten cryptic species not described
yet (Nazarov et al. 2009).
Here we describe a new Cyrtopodion species from a coastal area in southern Iran. This species clearly belongs
to the agamuroides-group as currently defined (Anderson 1999, Nazarov et al. 2009), adding a little more knowl-
edge to the presumably underestimated diversity of the Iranian gecko fauna in general and this group in particular.
Zootaxa 2924 © 2011 Magnolia Press · 23
NEW SPECIES OF ANGULAR-TOED GECKO FROM IRAN
Material and methods
We examined a set of 13 morphometric, twelve meristic and three categorical characters and compared these char-
acters with specimens of C. gastropholis, including the type series (see appendix), and took the data given for C.
agamuroides, C. golubevi and C. persepolense by Nazarov et al. (2009) into account for our analyses. Specimens
were deposited in the herpetological collection of the Zoologisches Forschungsmuseum Alexander Koenig in
Bonn, Germany (ZFMK) and in the private collection of F. Ahmadzadeh (FA).
The following characters were used for morphological analysis, abbreviations and measurement details are
given in parentheses: snout-vent length (SvL; from tip of the snout to cloaca), tail length (TaL; from cloaca to tip of
the tail, only measured when not partially or completely missing), head length (HeL; from tip of the snout to rear of
the skull), head width (HeW; at widest point), neck width (NeW; at narrowest point), snout length (SnL; from tip of
the snout to anterior margin of the orbit), orbit diameter (OrD; from anterior to posterior margin of orbit), ear length
(EaL; at widest point of the ear opening), orbit-ear distance (OeD; from posterior margin of the orbit to anterior
margin of the ear opening), length of upper forelimb (UfL; from shoulder to elbow), length of lower forelimb (LfL;
from elbow to carpus), length of upper hindlimb (UhL; from pelvis to knee), length of lower hindlimb (LhL; from
knee to heel). All measurements were taken with digital callipers to the nearest 0.1 mm by the second author.
For better comparison of the species, several ratios were calculated. These are head ratio (HeR; head width
divided by head length), neck ratio (NeR; neck width divided by head width), orbit ratio (OrR; orbit diameter
divided by snout length), forelimb ratio (FlR, sum of upper and lower forelimb length divided by snout-vent
length), and hindlimb ratio (HlR; analogous to FlR).
Meristic and categorical characters include number of transverse ventral scales (TVe; across midbody), number
of longitudinal ventral scales (LVe; between mental and cloaca), number of subdigital lamellae of fourth finger
(SLm), number of subdigital lamellae of fourth toe (SLp), number of active preanal pores (PPo; in males only),
number of postcloacal tubercles (CTu, both sides summarized), number of supralabials (SLa), number of infralabi-
als (ILa), number of internasals (INa; all scales between the nasals that border the rostral), number of postmentals
(PMe); number of transverse dorsal tubercles (DTu; across midbody), number of longitudinal rows of transversely
enlarged subcaudals (SCa), anterior pair of postmentals in contact (PMC), presence of enlarged femoral scales
(EFS), nasals swollen (NAS).
Cyrtopodion kiabii sp. nov.
(Figs. 1 to 5)
Holotype. ZFMK 91834 (field no. 138), adult male, collected 4.5 km SW of Nayband village at a distance of
approx. 300 m to coast of Persian Gulf, 27°21’9.5’’ N, 52°37’56.5’’ E, 108 m above sea level, Bushehr Province,
southern Iran, on 14 July 2007 by F. Ahmadzadeh.
Paratypes. ZFMK 91835 (field no. 190), adult female, same locality and collection data as the holotype;
ZFMK 91836, adult male, same locality as the holotype, collected on 30 June 2010 by F. Ahmadzadeh.
Other specimens. FA 1001, adult male, same locality as the holotype, collected on 30 June 2010 by F.
Diagnosis. A small and slender Cyrtopodion species of the agamuroides-group with angular-shaped toes and a
maximal snout-vent length of 46.9 mm. Limbs are very long and thin, so that knee and elbow—at least in males—
meet when pressed alongside the straightened body. The head width is less than two thirds of the head length and
the neck is about half as wide as the head, underlining the overall appearance, which is even more delicate than in
the other species of the agamuroides-group. The eyes are large in relation to the short snout, which is barely longer
than the diameter of the orbit (fig. 2). Due to the swollen nasals, the nostrils seem to be mounted. Postmentals are
distinctly larger than the posterior gular scales, whereas Agamura has small, indistinguishable postmentals. The
anterior pair of postmentals may be in contact or separated from each other by a diminutive median scale (fig. 2).
Ventrals are large, with ten to twelve scales transversely across midbody (fig. 3), which is less in number than in the
related species (14 to 18 in C. gastropholis, 22 to 28 in C. golubevi, 28 to 32 in C. persepolense, and 28 to 38 in C.
agamuroides). Longitudinally, there are 98 to 104 scales between the mental and the vent, which is at the upper end
of the variation observed within C. gastropholis and less than in the remaining species. Subdigital lamellae are not
AHMADZADEH ET AL.
24 · Zootaxa 2924 © 2011 Magnolia Press
tuberculous as in Bunopus. Numbers under the fourth finger and fourth toe range from 22 to 24 and 25 to 28,
respectively. Males possess four active preanal pores (fig. 4), separating the species from C. agamuroides which
has only two. The subfemoral scales are larger than the surrounding scales, but no femoral pores or femoral tuber-
cles are developed, distinguishing it from all members of the subgenus Tenuidactylus. Dorsal tubercles are roundish
and equal to or larger than interspaces (versus trihedral and smaller than interspaces in the subgenus Cyrtopodion)
and arranged in nine rows along the back. The tail is whorled and distinct caudal tubercles are present, both charac-
ters further discriminating this species from Agamura. The tubercles form the terminal row of each whorl opposed
to being located in the middle of each whorl as in Mediodactylus. The subcaudal scalation is homogenous, a char-
acter practicable to distinguish all members of the agamuroides-group from the otherwise similar C. kirmanense
which has a heterogeneous scalation under the tail. Subcaudals are arranged in a single row of transversely
enlarged scales, as in C. gastropholis and C. golubevi, whereas, in C. persepolense the enlarged subcaudals are
bifurcated and form a double row. In contrast, C. agamuroides does not have distinctly enlarged subcaudals. A
comparison of the diagnostic characters of the agamuroides-group is shown in table 1.
TABLE 1. Comparison of diagnostic characters within the Cyrtopodion agamuroides-group. Character abbreviations as
explained in the text. Values given (except for the categorical characters) are mean value, standard deviation and range. Values
for C. golubevi, C. persepolense and C. agamuroides were calculated using the data presented by Nazarov et al. (2009).
kiabii gastropholis golubevi persepolense agamuroides
n4 10 9 11 3
PMC yes/no yes/no yes no yes/no
EFS yes yes yes no no
NAS yes yes yes no yes
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NEW SPECIES OF ANGULAR-TOED GECKO FROM IRAN
FIGURE 1. Cyrtopodion kiabii sp. nov. in life.
FIGURE 2. Dorsal (a), lateral (b) and ventral (c) views of the head of the holotype of Cyrtopodion kiabii sp. nov. Scale bars
represent 2 mm.
Description of holotype. Snout-vent length 46.9 mm, tail length 46.3 mm with the distal 26 mm regenerated.
Forelimbs (measured from shoulder to carpus) are approximately half as long as the distance between snout and
vent and hindlimbs (measured from pelvic hinge to carpus) are two thirds of the snout-vent length. The head is 1.6
times longer than wide and twice as wide as the neck. The orbit is nearly as large as the snout length. The nostril is
AHMADZADEH ET AL.
26 · Zootaxa 2924 © 2011 Magnolia Press
pierced just above the suture of the rostral and first labial, contacting both and further surrounded by three smaller
nasals. These scales around the nostril appear to be swollen. The rostral is large and has a long median groove from
its posterior end to nearly the tip of the snout. Three internasals are bordering posteriorly with the median one
being slightly embedded into the rostral groove. Numbers of supralabials total eleven and twelve on left and right
side, respectively; ten infralabials on each side. The mental shield is large and triangular to pentagonal, followed
posteriorly by three pairs of large postmentals, with the median pair divided from each other by a diminutive scale.
There are twelve ventrals transversely across midbody and 104 ventrals in a longitudinal median row between
mental and cloaca. 22 subdigital lamellae under the fourth finger and 26 under the fourth toe. The obviously adult
male specimen has four active preanal pores within very large scales, no femoral pores are present, but subfemoral
scales are enlarged. The scales directly behind the preanal pores are also larger than other surrounding scales. Two
postcloacal tubercles are located on each side of the base of the tail. Nine rows of dorsal tubercles run across the
middle of the back. These tubercles are roundish and often slightly keeled. The distance between tubercles is equal
to or larger than the size of the tubercles. The surface of the femur is also marked with round tubercles. Original tail
with one row of plate-like, transversely enlarged subcaudal scales, two of these per whorl. The dorsal and lateral
tail scalation is heterogeneous, small granular scales are mixed with larger, trihedral tubercles, the latter arranged in
rows consisting of six tubercles around the tail. The tubercles located in the terminal row of each whorl are the
FIGURE 3. Dorsal (a) and ventral (b) scalation of the holotype of Cyrtopodion kiabii sp. nov. and comparison of the ventral
scalation with the holotype of C. gastropholis (c). Scale bars represent 2 mm.
FIGURE 4. Anal region of the female paratype (a) and the male holotype (b) of Cyrtopodion kiabii sp. nov. Scale bars repre-
sent 2 mm.
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NEW SPECIES OF ANGULAR-TOED GECKO FROM IRAN
FIGURE 5. Dorsal views of the female paratype (left) male holotype (right) of Cyrtopodion kiabii sp. nov., illustrating mor-
phometric differences between sexes. Scale bar represents 1 cm.
The dorsal colouration consists of a light, sandy brown ground with darker blotches on the back, which are
arranged in seven bands between neck and groin; limbs and original part of the tail with brown crossbars, which are
much more distinct in the latter. The ventral side is uniformly whitish coloured.
Variation. All paratypes and the additional specimen are generally similar to the holotype concerning morpho-
metrics, pholidosis and colouration. Notable variation of scale characters includes the internasals, which are only
two in number in all of these specimens in contrast to three in the holotype. The arrangement of the internasals is
not symmetrical, but rather one of the two scales is slightly enlarged and projecting into the rostral as does the
third, median internasal of the holotype. In ZFMK 91836 and FA 1001, the median pair of postmentals is bordering
each other and not divided by a diminutive scale. Ventrals are even less in the paratypes than in the holotype, rang-
ing from ten to eleven when counted transversely and from 98 to 102 when counted longitudinally. The female
paratype ZFMK 91835 differs in some morphometric characters: the head is relatively shorter and the neck rela-
tively wider than in the male types. Furthermore, the legs are slightly shorter (fig. 5). Complete measurements of
all specimens are presented in table 2.
AHMADZADEH ET AL.
28 · Zootaxa 2924 © 2011 Magnolia Press
TABLE 2. Measurements for Cyrtopodion kiabii sp. nov. Character abbreviations as explained in the text. Values for metric
characters are given in mm. Asterisk indicates regenerated tail.
Distribution. So far, the species is only known from the type locality.
Etymology. We name this new species in honour of Dr. Bahram Hassanzadeh Kiabi (Shahid Beheshti Univer-
sity of Tehran, Iran) to appreciate his indefatigable work as an ecologist who contributed greatly to the knowledge
of Persian wildlife.
Ecological notes. The collected specimens were found in two abandoned buildings, situated in a cliffy, but not
mountainous area approximately 100 m from the coast of the Persian Gulf (fig. 6). Activity was only observed dur-
ing night and we assume this species to be strictly nocturnal as Cyrtopodion species are in general. Sympatric liz-
ard and snake species comprise Hemidactylus flaviviridis, Hemidactylus persicus, Pristurus rupestris, Asaccus sp.,
Laudakia nupta, and Echis carinatus.
ZFMK 91834 ZFMK 91835 ZFMK 91836 FA 1001
holotype paratype paratype
sex male female male male
HeW 8.1 7.9 7.5 7.3
NeW 4.2 4.5 3.8 4.0
SnL 4.7 4.4 4.2 4.1
OrD 4.5 4.2 4.1 3.9
EaL 1.4 1.3 1.3 1.2
OeD 2.8 2.7 2.6 2.6
UfL 10.5 9.7 9.9 9.3
TVe 12 11 10 11
LVe 104 102 102 98
ILa 10 9 9
PMC no no yes yes
EFS yes yes yes yes
NAS yes yes yes yes
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NEW SPECIES OF ANGULAR-TOED GECKO FROM IRAN
FIGURE 6. Habitat of Cyrtopodion kiabii sp. nov.
FIGURE 7. Distribution of Cyrtopodion kiabii sp. nov. (star) and C. gastropholis (circles). The blank circle marks the dubious
locality of the latter species discussed in the text. Names of single localities are as follows: Basht (1), Chahak (2), Bandar-e-
Gonaveh (3), Kazerun (4), Dalaki (5), Bushgan (6), Fasa (7), Khos Hangan (8), Nayband (9).
AHMADZADEH ET AL.
30 · Zootaxa 2924 © 2011 Magnolia Press
We observed a sexual dimorphism in C. kiabii: males have a more slender body, a relatively longer head, a nar-
rower neck and slightly longer legs than the female. This is based on a single female available for examination.
Nevertheless, we consider it plausible, especially as the same dimorphism was also detected in a larger series of C.
gastropholis. At least for these two species, this does not affect the diagnostic quality of the characters, as differ-
ences between sexes are minor compared to differences between species. Apparently, this sexual dimorphism is not
present in C. golubevi and C. persepolense, but females of these two species seem to grow larger than males (com-
pare morphometric data given by Nazarov et al. 2009). However, some morphometric characters can be problem-
atic for the identification of juveniles, e.g. the overlap of the variation in the neck ratio of C. kiabii and C.
gastropholis (see table 2) is solely due to a juvenile specimen of C. gastropholis. There are no obvious differences
between sexes in scalation, except for the preanal pores, which are also present in females, but are less developed
and not active as in males (fig. 4). Probably owing to that circumstance, Szczerbak & Golubev (1996) erroneously
considered the holotype of C. gastropholis to be a male.
Bearing the findings on the phylogeny of Cyrtopodion inferred from molecular data (
ervenka et al. 2008) in
mind, we do not assign C. kiabii to any of the described subgenera of Cyrtopodion, but place it into the informal
agamuroides-group. The only autapomorphy of this group is the darkly pigmented peritoneum (Anderson 1999).
Nazarov et al. (2009) further characterize it by several morphometric traits and the relatively low number of prea-
nal pores. A key character to identify C. kiabii is the low number of large ventrals across midbody, which are
twelve or less. Otherwise, it can be distinguished from the remaining species of the agamuroides-group by having
more subdigital lamellae under the fourth finger and toe, and by a more slender habitus and relatively longer legs
(see table 1). Identification of the currently described members of this group is reliable even if one only compares a
set of two characters, namely the number of ventral scales across midbody and subcaudal scalation. Within the aga-
muroides-group, we suggest a close relationship of C. kiabii and C. gastropholis. This is based on distributional
proximity, the probably exclusive presence of sexual dimorphism, and – most notably – on pholidosis. Especially
the low numbers of both transversely and longitudinally counted ventral scales shared by these two species could
be considered as an autapomorphy.
As this character is crucial for identification, it is necessary to comment on a problem associated with C. gas-
tropholis. In the original description Werner (1917) counted ten ventral rows in the holotype and ten to 14 in the six
paratypes. As already noted by other authors (Wettstein 1951, Anderson 1999), this is below the actual numbers,
because Werner may have only considered the very large median ventrals. Unaware of the problematics associated
with the type series of C. agamuroides (Szczerbak and Golubev 1996, Nazarov et al. 2009), Wettstein (1951) syn-
onymized C. gastropholis with C. agamuroides solely based on the number of ventrals, which he by mistake
thought to be equal in both species. However, C. agamuroides possesses 28 or more ventrals and differs in further
characters (e.g. subcaudal scalation and number of preanal pores) from C. gastropholis and all other members of
the agamuroides-group (Nazarov et al. 2009). A re-examination of the type series of C. gastropholis showed that
typical C. gastropholis have 16 to 18 ventrals. We examined further material from Bushgan, Bushehr Province,
which could be identified as C. gastropholis and features a range of 14 to 17 transverse ventrals. Nazarov et al.
(2009) examined specimens of C. gastropholis from Basht, Kazerun and Fasa (all in Fars Province) with 16 to 19
Although not mentioned in the original description, Werner’s specimens originate from “between Kazerun-Shi-
raz-Persepolis” in Fars Province according to their label. Hence, the distributional range of C. gastropholis proba-
bly includes that of C. persepolense, but both species are clearly distinct in scalation. The most prominent
differences are the single row of subcaudals in C. gastropholis (versus two rows in C. persepolense), the lower
number of ventrals and the presence of enlarged subfemoral scales and swollen nasals in C. gastropholis.
The documented occurrence in Fars Province disproves the suggestion that the latter species might be restricted
to Bushehr Province (Leviton et al. 1992). In contrast, we would not be surprised, if the localities published for C.
gastropholis from the coastal plains of Bushehr Province (Anderson 1999,
ervenka et al. 2008, Frynta et al. 1997)
would actually refer to C. kiabii, as earlier authors might have been misguided by Werner’s original diagnosis. If
so, C. gastropholis would be restricted to the Zagros Mountains. The single record of C. gastropholis from Hor-
mozgan Province (
ervenka et al. 2008) is far to the east of the species’ known range. Most certainly, this repre-
sents another species and our assumption is supported by the phylogeny produced by the same study. Whether this
Zootaxa 2924 © 2011 Magnolia Press · 31
NEW SPECIES OF ANGULAR-TOED GECKO FROM IRAN
species is C. kiabii or not remains unclear until further research is conducted. For now, we consider C. gastropholis
to occur at all the localities in question, as C. kiabii is only known with certainty from its type locality (fig. 7). Sim-
ilar cases might apply for the records of C. agamuroides from Fars Province, which should either turn out to be C.
persepolense or yet another new species.
Besides the puzzling taxonomy, another common error associated with C. gastropholis is the spelling “gastrop-
hole,” but since “gastropholis” (ancient greek, from gastron and pholis) obviously is a noun in apposition it must
not be conjugated.
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gekkonov (Reptilia, Gekkonidae, Tenuidactylus gen. n.). Vestnik Zoologii, 1984(2), 50–56. [in Russian with English
Szczerbak, N.N. & Golubev, M.L. (1996) Gecko Fauna of the USSR and Contiguous Regions. SSAR, 233 pp.
Sindaco, R. & Jeremcenko, V.K. (2008) The reptiles of the Western Palearctic. Edizioni Belvedere, 579 pp.
Werner, F. (1917) Reptilien aus Persien (Provinz Fars). Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen
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32 · Zootaxa 2924 © 2011 Magnolia Press
Besides the specimens of C. kiabii already mentioned in detail in the text, we examined material of C. gastropholis for compar-
ison (see table 3). This includes the type series, formerly in the collection of the Museum Göttingen, of which three para-
types are lost. Two specimens of C. gastropholis in the collection of the California Academy of Sciences (CAS 100472-
100473) have the same locality and collector’s data as the types and may originate from that series. Unfortunately, we were
not able to examine these two specimens in the course of this work.
[Appendix] TABLE 3. Measurements for Cyrtopodion gastropholis. Character abbreviations as explained in the text. Values
for metric characters are given in mm.
holotype paratype paratype paratype
between Kazerun-Shiraz-Persepolis Bushgan, Bushehr Province
sex female female female male female male female male juvenile female
SvL 49.2 – 47.4 42.6 47.8 44.7 50.0 49.5 31.8 46.6
HeL 13.0 13.2 12.6 12.8 12.5 12.0 12.8 13.1 9.3 11.8
HeW 9.1 9.1 8.9 8.3 9.0 8.5 9.2 9.4 6.3 8.4
NeW 5.8 6.0 5.7 5.2 5.8 4.9 6.8 6.0 3.4 6.0
SnL 5.0 5.2 5.1 5.1 4.6 4.4 5.0 4.9 3.4 4.4
OrD 4.1 4.2 4.0 4.3 4.0 3.8 4.1 4.2 3.1 3.5
EaL 1.3 1.6 1.2 1.5 1.4 1.4 1.3 1.5 0.9 1.3
OeD 3.4 3.4 3.4 3.2 3.2 3.0 3.5 3.5 2.1 3.1
UfL 10.3 10.4 9.9 10.5 9.6 8.5 10.3 9.8 6.3 8.5
LfL 8.9 9.2 9.3 10.2 9.2 7.9 9.6 9.8 5.7 8.6
UhL 12.5 13.1 11.5 12.3 11.8 11.8 12.7 12.4 8.1 11.7
LhL 12.6 11.3 12.2 11.6 11.6 11.5 11.9 12.6 8.2 11.4
LVe 93 – – 101 96 94 94 99 100 84
SLm 20 18 18 – 22 21 20 22 21 20
PPo– – – 4 –4–4––
CTu4 4 4 4 444444
ILa 9 9
DTu 10 – 9 9 10 10 10 12 9 11
SCa1 – – 1 111111
PMC yes yes yes yes yes yes no yes yes yes
EFS yes yes yes yes yes yes yes yes yes yes
NAS yes yes yes yes yes yes yes yes yes yes