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Description of four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey

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One new Asaccus from the Anatolian plateau and three new Asaccus from the Iranian plateau are described as follows. (1) Asaccus barani sp. nov.: diagnosed by strongly heterogeneous dorsal tubercles; (2) Asaccus iranicus sp. nov.: diagnosed by small body size and digits (forelimbs) parallel joint to palm; (3) Asaccus tangestanensis sp. nov.: by having enlarged trihedral tubercles all over the dorsal body; (4) Asaccus zagrosicus sp. nov.: secondary postmentals are not in contact with lowerlabials (100% of specimens). Other important data on the new Asaccus are given in detail in the text.
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Amphibia-Reptilia 32 (2011): 185-202
Description of four new Asaccus Dixon and Anderson, 1973
(Reptilia: Phyllodactylidae) from Iran and Turkey
Farhang Torki1, Faraham Ahmadzadeh2,3, Çetin Ilgaz4, Aziz Avcı5,*, Yusuf Kumluta¸s6
Abstract. One new Asaccus from the Anatolian plateau and three new Asaccus from the Iranian plateau are described as
follows. (1) Asaccus barani sp. nov.: diagnosed by strongly heterogeneous dorsal tubercles; (2) Asaccus iranicus sp. nov.:
diagnosed by small body size and digits (forelimbs) parallel joint to palm; (3) Asaccus tangestanensis sp. nov.: by having
enlarged trihedral tubercles all over the dorsal body; (4) Asaccus zagrosicus sp. nov.: secondary postmentals are not in contact
with lowerlabials (100% of specimens). Other important data on the new Asaccus are given in detail in the text.
Keywords:Asaccus barani sp. nov., Asaccus iranicus sp. nov., Asaccus tangestanensis sp. nov., Asaccus zagrosicus sp. nov.,
Iran, Phyllodactylidae, Turkey.
Introduction
The genus Asaccus comprises at least 12
species in southwestern Asia (Arnold, 1972;
Dixon and Anderson, 1973; Martens and Kock,
1991; Leviton et al., 1992; Arnold and Gard-
ner, 1994; Rastegar-Pouyani, 1996; Anderson,
1999; Rastegar-Pouyani, 2006; Torki and Shar-
ifi, 2007; Torki et al., 2008, in press; Afrasiab
and Mohamad, 2009; Torki, 2009, 2010, in
press), as follows. Four species are distrib-
uted in the Oman-Emirate Mountains [A. mon-
tanus Gardner, 1994 (from the mountainous
regions of United Arab Emirates); A. gal-
lagheri Arnold, 1972 (from Oman, United Arab
Emirates); A. platyrhynchus Arnold and Gard-
ner, 1994 (from Tanuf, Oman) and A. cau-
divolvulus Arnold and Gardner, 1994 (from
Jebel Ras, United Arab Emirates)], and eight
1 - Farhang Torki Ecology and Herpetology Center for Re-
search (FTEHCR), 68319-16589, P. O. Box 68315-139,
Nourabad City, Lorestan Province, Iran
2 - Department of Biodiversity and Ecosystem Man-
agement, Environmental Sciences Research Institute,
Shahid Beheshti University, Iran
3 - Zoologisches Forschungsmuseum Alexander Koenig
(ZFMK), Adenauerallee 160, D-53113 Bonn, Germany
4 - Fauna and Flora Research and Application Center,
Dokuz Eylül University, 35160 Buca, ˙
Izmir, Turkey
5 - Department of Biology, Faculty of Science and Arts,
Adnan Menderes University, 09010 Aydın, Turkey
6 - Department of Biology, Faculty of Education, Dokuz
Eylül University, 35160 Buca, ˙
Izmir, Turkey
*Corresponding author; e-mail: aavci09@yahoo.com
species are distributed in the Zagros Moun-
tains (Iran-Iraq) [A. kurdistanensis Rastegar-
Pouyani, Nilson and Faizi, 2006 (from Kurdis-
tan Province, western Iran); A. kermanshahensis
Rastegar-Pouyani, 1996 (from 45 km north of
Kermanshah city, Kermanshah Province, west-
ern Iran); A. nasrullahi Werner, 2006 (from
Lorestan Province, southwestern Iran); A. gran-
ularis Torki, in press (from Lorestan Province,
southwestern Iran); A. andersoni Torki et al., in
press (from Illam Province, southwestern Iran);
A. elisae (F. Werner, 1895) (from Iran, Iraq,
Turkey and Syria); A. griseonotus Dixon and
Anderson, 1973 (around Islamabad, Kerman-
shah Province, western Iran), and A. saffinae
Afrasiab and Mohamad, 2009 (from Shera Swar
Cave in the Saffine Mountain near Irbil, north-
ern Iraq)]. Previously A. elisae has been re-
ported from Turkey (Birecik, ¸Sanlıurfa-Harran,
¸Sanlıurfa-Nusaybin, Mardin; see Baran and
Gruber, 1982; Tok, Atatür and Tskavak, 1997;
Franzen, Schmidtler and Bischoff, 2002; Tok,
Atatür and Cihan, 2002; Baran et al., 2003). In
this study, Harran samples were reevaluated and
described as a new Asaccus. In total we describe
four new Asaccus from the Anatolian and Za-
gros Mountains.
Materials and methods
During field trips carried out in the Iranian and Anatolian
plateaus we collected four new Asaccus from these regions.
(a) Harran, ¸Sanlıurfa, Turkey, (b) near the coastal Persian
Gulf, Bushehr province, Iran, (c) Tangestan region, Bushehr
©Koninklijke Brill NV, Leiden, 2011. DOI:10.1163/017353711X556998
186 F. Tor k i et al.
Figure 1. Type localities of four new Asaccus in the Anatolia and Zagros mountains.
province, Iran and (d) Tang-e-Haft, Lorestan province, Iran
(fig. 1).
For comparison, we examined seven species of Asaccus
inhabited in Iran as follows: A. elisae complex, A. kurdista-
nensis,A. kermanshahensis,A. griseonotus,A. nasrullahi,
A. granularis and A. andersoni. Harran specimens were
marked with ZDEU (Zoology Department Ege University)
number, and kept in the Zoology Lab of the Department
of Biology at Buca Education Faculty, Dokuz Eylül Uni-
versity, ˙
Izmir, Turkey. Holotypes of the three new Iranian
Asaccus species were marked with ZFMK (Zoologisches
Forschungsinstitut und Museum Alexander Koenig) num-
ber and kept in the ZFMK. Paratypes of the three Iranian
species are deposited in Farhang Torki Ecology and Her-
petology Center for Research (FTHM).
Characters were selected to optimize comparisons with
data reported by Dixon and Anderson (1973), Arnold
and Gardner (1994), Rastegar-Pouyani (1996), Anderson
(1999), Rastegar-Pouyani, Nilson and Faizi (2006), Werner
(2006), Torki (in press), Torki et al. (in press). Measure-
ments were taking using a dial caliper with 0.01 mm pre-
cision. Additionally in this study we studied on other char-
acters that are important for taxonomy of Asaccus such as
tubercle size, shape (fig. 2) and distribution (Torki, in press;
Torki, Fatinia and Rostami, in press). Characters that were
used to describe the four new Asaccus are as follows: SVL:
snout-vent length (in mm); TL: tail length (in mm); HL:
head length (taken axially to the rear border of the ear, using
the Goren and Werner (1993) calipers – in mm); HW: head
width (in mm); HD: head depth (in mm); HI: head index
(100*length/width); HF: head flatness (100*length/depth);
SW: snout width at 3rd (in mm); EHD: eye (horizontal) dia-
meter (in mm); EVD: ear (vertical) diameter (in mm); NY:
nostril-eye distance (in mm); EE: eye-ear distance (in mm);
NE: nostril-ear distance (in mm); FLL: forelimb (includ-
ing fingers) (in mm); HLL: hind limb (including toes) (in
mm); AG: axilla-groin distance; LT: number of lamellas on
4th toe; DTR: number of dorsal tubercles rows (widest re-
gion); PM: number of postmentals; SPM: number of scales
surrounded postmentals; SL: number of scales between 3rd
upperlabials; IS: interorbits scales; NES: number of scales
from nostril to eye.
Figure 2. Five type tubercles in Asaccus. (up-left) simple;
(up-right) pointed; (mid-left) keeled tubercle type a; (mid-
right) keeled tubercle type b; (down) keeled tubercle type c.
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 187
Asaccus iranicus sp.nov.(g.3)
Holotype. ZFMK 91933, female, collected at
coastal Persian Gulf, Assaloye City, Bushehr
Province, Southern Iran, on 1 May 2008,
(2718N-5242E, altitude 50-221 m a.s.l.), by
F. Torki and Faraham Ahmadzadeh.
Paratype. FTHM 002691. Adult male, other
data as for holotype.
Diagnosis. Small Asaccus; forelimb fingers
parallel joint to palm; dorsal body covered all
over by tubercles; small dorsal tubercles keeled
and trihedral type a and b; dorsolateral tuber-
cles keeled, pointed or simple, much smaller
than the middle; tubercles on the middle of the
neck keeled, laterals pointed; occipital tuber-
cles pointed or simple; upper head tubercles
very small and simple; simple tubercles extend
between interorbit and upper eyelash; several
coarse tubercles (sharp or simple) in front of the
eyes; simple tubercles cover the arm and fore-
arm, tubercles not extend into palm and digits;
thigh covered with keeled tubercles, laterals co-
vered with simple or pointed tubercles, smaller
than the middle; foreleg tubercles extend into
palm and upper part of the 1st and 2nd digits.
Additional characters are shown in table 1.
Description of holotype. Depressed body,
small body size (32.22 mm), tail completely
flattened; fingers (forelimbs) parallel joint to
palm (fig. 4); large eye; vertical pupil; verti-
cal ear (higher than wide). The dorsal body
covered all over by tubercles; small dorsal tu-
bercles, keeled and trihedral (mostly type b
and partly type c), 14-15 irregular rows, 2-6
scales between large tubercles; dorsolateral tu-
bercles are keeled, pointed or simple signifi-
cantly smaller than the middle; tubercles on the
middle of the neck are keeled type b, pointed
laterals; occipital tubercles pointed or simple;
upper head tubercles are very small and sim-
ple; simple tubercles extend between interor-
bit and upper eyelash; several coarse tubercles
(sharp or simple) in front of the eyes; no tuber-
cles between ear-neck and ear-eye; simple tu-
Figure 3. Holotype of Asaccus iranicus sp. nov. (a) dorsal
view; (b) ventral view; (c) ventral view of palm of right
forelimb.
188 F. Tor k i et al.
Tab l e 1 . Morphological characters of Asaccus iranicus sp. nov. *: characters
in mm; other characters are numerical.
Characters Holotype Paratype Characters Holotype Paratype
SVL* 32.22 41.83 NE* 8.59 11.14
TL* 40.75 FLL* 15.85 19.17
HW* 6.42 7.80 HLL* 20.29 27.56
HL* 10.99 13.56 AG* 14.6 17.75
HD* 3.87 5.60 LT 10 9
HI 171.18 173.84 DTR 11 10
HF 283.98 242.14 PM 2 2
SW* 3.22 3.95 SPM 16 19
EHD* 2.63 3.38 SL 16 18
EVD* 1.05 1.52 IS 26 27
NY* 4.41 5.46 NES 12 13
EE* 2.26 3.23
bercles cover the arm and forearm, tubercles not
extended into palm and digits; thigh is covered
by keeled tubercles, laterals are covered by sim-
ple or pointed tubercles, smaller than the mid-
dle; foreleg tubercles extend into palm and up-
per part of 1st and 2nd digits; dorsal trihedral
tubercles extend into tail; tail tubercles whorl
not extended into ventral side, 6 tubercles form
1st to 3rd whorls, tubercles in each rows re-
duced from 6 (proximal tail) to 1 (distal tail) or
0 (end of tail); 5 scales between whorls; dor-
sal scales not overlapping; scales on upper head
and especially between the eye are partly glob-
ular; five scales around the nostril (1 nasal, 1st
supralabials, 2 scales on posterior, and rostral);
internasal scale in one point are in contact, two
large scales on posterior of internasals and one
medium scale between internasals and posteri-
ors; 10 rows of scales (coarse) between nostril
and eyes; 23 supralabials (12 large and the oth-
ers very small; 11 to mid-eye); 18 (8 large, and
10 very small) scales in lower labials; two post-
mentals, first postmental is larger than the sec-
ond, 1st postmentals are in contact with each
other, and in contact with the first lowerlabi-
als; secondary postmentals are distinguished by
5 scales and in contact with the 2nd lower-
labial; 18.25/7; chin scales are small and partly
overlapped; neck scale is elevated and partly
overlapped; ventral scale rounded or hexago-
nal or tetragonal, not overlapped, more than
2.5 time larger than dorsal scale; ventral scale
on arm is smaller than forearm, arm and fore-
arm scale is rotund or tetragonal and not over-
lapping; forelimbs palm scale partly globular
shape and are similar to lamella; thigh and fore-
leg scales are cobble-stone shaped, and tetrag-
onal or partly rounded (in some part); a few
of the foreleg scales are larger than the thigh
scales; the palms of hindlimbs are partly glob-
ular like, and have similar ornamentation to
lamellas; there are large scales between inter
hindlimbs; the ventral tail scale is large and
most of the ventral side is covered; mental pen-
tagonal (length/width: 1.79 mm/1.88 mm); ros-
tral width (length/width: 0.64 mm/1.68 mm)
and not divided in up; 6 lamellas underneath
the 4th finger; 10 lamellas underneath the 4th
toe; claves in front of scansors. The background
color of the body is whitish; there are several
dark crossbars on the tail, some of them ex-
tend onto the ventral side; spots cover the dor-
sal body; the color of ventral site is whitish; the
color of limbs and head are white without any
pattern; the upper eyes are dark.
Variation. Variations in most external mor-
phological characters are given in table 1. Num-
ber of lamellas on 4th toe 9; number of dor-
sal tubercles rows 10; number of postmentals
2; number of scales surrounded postmentals 19;
number of scales between 3rd upperlabials 18;
interorbits scales 27; NES: number of scales
from nostril to eye 13. The snout-vent length
41.83 mm; tail regenerated. For other morpho-
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 189
logical characters see table 1. The color and
color-pattern feature of paratype was similar to
the holotype.
Habitat and ecology. Specimens were col-
lected at midnight from two destroyed homes
in the coastal Persian Gulf. Collection site is
approximately 100 m from the sea. There are
no mountains or hills in this region. The geo-
morphology of this habitat is different for all of
the new species that are described in this pa-
per as well as the other Asaccus inhabitants of
Iran. Specimens belonging to this new species
of Asaccus appear to be nocturnal.
Distribution. Asaccus iranicus sp. nov. is at
present known only from the type locality.
Sympatric lizards and snake. The sympatric
lizard and snake species comprised Hemidacty-
lus flaviviridis,Hemidactylus persicus,Pristu-
rus rupestris,Cyrtopodion sp., Laudakia nupta
and Echis carinatus.
Etymology. The species is named for Iran in
which the type locality (Assaloye City, Bushehr
Province) is found.
Comparison. Asaccus iranicus sp. nov. is eas-
ily distinguishable from other Asaccus as fol-
lows: The direction of the fingers to forelimbs
palm of the A. iranicus sp.nov.isdifferent
with other species of Asaccus (fig. 4). Scansors
do not extend beyond claws and tubercles are
present on the arms for A. iranicus sp. nov.; this
is in contrast to A. gallagheri,A. platyrhynchus,
A. caudivolvulus (Khasab population), A. kur-
distanensis,A. kermanshahensis,A. nasrullahi,
A. granularis,A. andersoni,A. griseonotus and
A. saffinae (e.g., Arnold and Gardner, 1994;
Torki and Sharifi, 2007; Torki et al., 2008;
Afrasiab and Mohamad, 2009; Torki, 2009, in
press; Torki, Fatinia and Rostami, in press). A.
Figure 4. Forelimb (right part) of four new species.
(a) Asaccus iranicus sp. nov.: Forelimb parallel joint to
palm; (b) Asaccus tangestanensis sp. nov.; (c) Asaccus za-
grosicus sp. nov.; (d) Asaccus barani sp. nov. Forelimb par-
allel joint to palm and in other different shape for Asaccus
iranicus sp. nov.
190 F. Tor k i et al.
iranicus sp. nov. shows difference from A. cau-
divolvulus (Jebele population) by having claws
beyond the scansors. A. iranicus sp. nov. differs
from A. montanus and A. elisae by having small
size tubercles on dorsal body (e.g., Arnold and
Gardner, 1994; Gardner, 1994). A. iranicus sp.
nov. is significantly smaller than A. barani sp.
nov., A. tangestanensis and Asaccus zagrosicus
sp. nov.
Asaccus tangestanensis sp.nov.(g.5)
Holotype. ZFMK 91934, adult male, collected
in the end of southern Zagros Mountains,
Khaiiz, Tangestan City, Bushehr Province,
Southern Iran, on 4 May 2008, (2843N-
5131E, altitude 525 m a.s.l.), by F. Torki and
Faraham Ahmadzadeh.
Paratypes. Five adult specimens (3-2):
FTHM 002671 (), 002672 (), 002673 (),
002674 (), 002675 (). Same data as for holo-
type.
Diagnosis. Medium size gecko; large size
Asaccus (49-57 mm); thin body; thin, elongated
limbs; tail is longer than body; trihedral tu-
bercles (type b) all over the dorsal body in-
cluding the back, limbs, head; special tubercles
(strongly keeled, elevated and trihedral) in front
of eye and between the eyes and ears; claws in
front of scansore. Two large postmentals, first
in contact; rostral is not divided in upper re-
gion; internasal scales elongated and crescent
shaped, distinct with one scale or in contact;
several small scales on posterior of postnasal
scale; a special pattern includes six crossbands
on dorsal side (from occipital region to prox-
imal of tail); some specimens have one cross-
band on neck and one crossbands on occipital,
and in general crossbands are only present from
Figure 5. Holotype of Asaccus tangestanensis sp. nov. (a) dorsal view; (b) ventral view; (c) enlarged thrihederal tuberculs
type c; (d) paratype under captive condition – egg on abdomen.
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 191
the occipital into the tail and not extended into
other region; three crossbands on the first half
of the original tail; two or three rings on sec-
ondary part of the original tail; other regions
of the body including limbs, head, and ventral
sides are without any pattern; in some speci-
mens regenerated tails have black rings, and in
others there is no pattern (dorsal and ventral
side) and have whitish color (similar to ventral
of body). The color is very translucent in live
specimens (fig. 5), because we could see ab-
domen components such as digestive and vas-
cular system clearly visible; in live females an
egg in the abdomen was visible; dorsal cross-
bands or spots are colorless (pallid brownish);
pattern on limbs is pallid with typically no pat-
tern on limbs in most of the specimens; upper
eyes are completely emerald or green; center of
scansor is reddish.
Description of holotype. An adult male; large
Asaccus (57.52 mm), thin and flat body; thin
and elongated limbs; the length of head
(17.32 mm) is 1.7 times its width (10.0 mm);
large eyes; vertical pupil; vertical ears; the
height of ear (2.32 mm) is 1.6 times its width
(1.44 mm); the dorsal body is completely co-
vered by keeled tubercles; large, trihedral and
keeled type c dorsal tubercles, 12-13 irregular
rows, 1-4 scales between large tubercles; neck
tubercles are strongly keeled, large and mostly
type c, not extend to the laterals side; occipital
tubercles type b, upper head tubercles sharply
and weakly keeled; several sharp tubercles be-
tween the interorbits, interorbit scales are glob-
ular; several large, strongly keeled, and elevated
tubercles in front of the eyes, similar to type a,
and b (but with more elevation); tubercles near
the ear and between the ear and eye are type b;
limbs are covered by type c tubercles, high tu-
bercle density on the arm and especially on the
forearm; dorsal tubercles extend into the tail;
tail tubercles whorls do not extend onto ventral
side, 7 tubercles form each whorls, 4-5 scales
between whorls; dorsal scales not overlapped;
scales on upper head and especially between
the eye are globular shape; five scales around
the nostril (1 nasal, 1st supralabials, 2 scale
on posterior, and rostral); internasal scale elon-
gated and distinct by one large scale, two large
scale on posterior of internasals; 10 rows of
large scales between the nostril and the eyes; 16
supralabials (11 to mid-eye); 11 in lower labi-
als. Two large postmentals, first postmentals are
in contact, and in contact with first lowerlabials;
secondary distinct postmentals with 4 scales and
in contact with 2nd lowerlabial; chin scales are
small and overlapped; neck scales are large and
significantly overlapped; ventral scales are me-
dium size and not completely distinct, 3 times
larger than the dorsal scale; ventral scales are
smaller than forearm, forearm scales are ro-
tund like; forelimbs palm scales are globular
shape and are similar to lamella; thigh scales
are cobble-stone shaped, in contrast foreleg
scales are overlapped; foreleg scales are larger
than thigh scales; palm of hindlimbs are glob-
ular like, elevated and are similar ornamenta-
tion to lamellas; large scales in between in-
ter hindlimbs; ventral tail scales are large and
cover most of the ventral side; mental trihedral
(length/width: 3.23 mm/2.95 mm); rostral width
(length/width: 0.83 mm/2.42 mm) and not di-
vided up; nasal scales are elongated, crescent
shape and distinct with one pentagonal scale;
postnasal scales are large; several small scales
on posterior of postnasal scales; 6 lamellas un-
derneath the 4th finger; 11 lamellas underneath
the 4th toe; claves in front of scansors.
Upper head between the eye to the rostral is
white and without any pattern; occipital has one
irregular crossbar; limbs and fingers are without
any pattern; five crossbands on dorsal body; 3
crossbands on the first half of tail, and not ex-
tend to the ventral side (color of these cross-
bands is similar to dorsal crossbands); 3 width
rings on the secondary half of the tail (com-
pletely extended into the ventral side of the tail),
color of rings is black and are different than
the crossbands; rings are wider than crossbands;
the ventrals of the body including the head, ab-
domen, limbs and tail (except three rings) are
whitish; the scansors are cream.
192 F. Tor k i et al.
Variation. Body size is between 50-57.52
mm; tail size: 60-65 mm; in one paratype few
tubercles (type b, smaller than large dorsal tu-
bercles) intermixed with dorsal large trihedral
type C tubercles; in one paratype tubercles dis-
tributed in 3rd first part of digits (hindlimbs),
and in most paratype distributed on 1st and
2nd first part of digits (hindlimbs); secondary
postmentals in all specimens were smaller than
other postmentals, size of some secondary post-
mentals in some specimens is smaller than in
others; tail length/body size in specimens with
regenerated tail (1.2-1.25) is more than in those
with original tail (1.18). Additional characters
are given in table 2.
In most specimens there is one spot on the
lower eyes; two crossbands on the head region,
one crossbands on occipital and one in upper
head, upper head crossband combined from sev-
eral spots; a large spot occurs inside of each
dorsal crossbands, color of spots is darker than
backgrounds (crossbands). Full crossbands in
some preserved specimens strongly pallid and
in other are visible; color of regenerated part of
tail has different color than original segment.
Habitat and ecology. The Tangestan region is
at the end of the southern part of the Zagros
Mountains. This region has palm trees. Asac-
cus tangestanensis sp. nov. is distributed in a
mountainous area. The structure of this moun-
tain is sedimentary. Shelter site of the new Asac-
cus is limited clefts and caves in this moun-
tain. A lot of specimens in one cave in this lo-
cality were observed along with Hemidactylus
persicus. Conditions inside the cave are moist,
in contrast to the conditions outside the cave.
The new Asaccus in this cave have nocturnal
and diurnal activity. The egg-laying site for the
new Asaccus occurs in the roof of the cave. The
specimens of new Asaccus were kept and bred
at FTEHCR. After three months two specimens
laid eggs. Our observation in both natural and
laboratory conditions show the new Asaccus lay
one large and spherical egg, hatching occurring
after 42-50 days. Juvenile specimens have sim-
ilar color patterns to parents.
Distribution. Asaccus tangestanensis sp. nov.
is distributed in two localities as follows: (1)
at the type locality in Khaiiz, Tangestan City,
Bushehr Province, Southern Iran; and (2) Jam
region, elevation 643 m a.s.l., the coordinates
are 5217E-2747N, Jam to Kangan road,
Kangan City, Bushehr Province, Southern Iran.
Both localities are situated at the end of the
southern Zagros Mountains. Distance of both
localities to the Persian Gulf is approximately
150 km.
Sympatric lizards and snake. In the type lo-
cality we could see several lizard and snake
species, including: Lizards: Hemidactylus per-
sicus,Laudakia nupta,Trapelus agilis,Tropio-
colotes persicus,Coluber sp., Macrovipera le-
betina and Echis carinatus. In Jam region, Tro-
Tab l e 2 . Morphological characters of Asaccus tangestanensis sp. nov. (N: number of specimens; Min.: minimum value; Max.:
maximum value; SD: standard deviation; SE: standard error of the mean). *: characters in mm; other characters are numerical.
Characters N Mean Min. Max. SD SE Characters N Mean Min. Max. SD SE
SVL* 6 52.79 49.07 57.95 3.13 1.28 NE* 6 12.94 11.56 14.06 0.96 0.39
TL* 5 64.32 59.31 69.46 4.65 2.08 FLL* 6 23.76 21.89 25.83 0.54 0.63
HW* 6 9.48 8.25 10.02 0.65 0.265 HLL* 6 31.95 29.12 35.02 0.25 0.92
HL* 6 16.12 14.90 17.33 1.00 0.41 AG* 6 23.31 21.64 25.25 0.26 0.51
HD* 6 6.57 6.02 6.97 0.34 0.14 LT 6 10.33 9.00 11.00 0.82 0.33
HI 6 170.45 156.55 182.44 11.01 4.49 DTR 6 12.83 12.00 14.00 0.75 0.31
HF 6 245.33 225.52 266.67 14.02 5.72 PM 6 2.00 2.00 2.00 0.00 0.00
SW* 6 4.64 4.41 4.91 0.16 0.06 SPM 6 16.17 13.00 18.00 1.72 0.70
EHD* 6 4.35 3.99 4.93 0.4 0.16 SL 6 19.33 17.00 23.00 2.25 0.92
EVD* 6 2.14 1.79 2.92 0.18 0.07 IS 6 25.33 24.00 27.00 1.21 0.49
NY* 6 6.25 5.48 7.51 0.78 0.32 NES 6 12.33 10.00 14.00 1.37 0.56
EE* 6 3.35 2.86 3.79 0.36 0.15
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 193
piocolotes persicus,Pristurus rupestris,Cyr-
topodion sp., Laudakia nupta,Trapelus agilis
and Trachylepis aurata were observed.
Etymology. The species is named for Tanges-
tan in which the type locality (Khaiiz, Tangestan
City, Bushehr Province) is found.
Comparison. Asaccus tangestanensis sp. nov.
is easily distinguishable from other Asaccus as
follows: Scansors in the Asaccus tangestanensis
sp. nov. do not extend beyond claws and tuber-
cles are present on the arms; this is in contrast to
A. gallagheri,A. platyrhynchus,A. caudivolvu-
lus (Khasab population), A. kurdistanensis,A.
kermanshahensis,A. nasrullahi,A. granularis,
A. andersoni,A. griseonotus and A. saffinae
(e.g., Arnold and Gardner, 1994; Torki and
Sharifi, 2007; Torki et al., 2008; Afrasiab and
Mohamad, 2009; Torki, 2009, in press; Torki,
Fatinia and Rostami, in press). A. tangestanen-
sis sp. nov. is different from A. caudivolvu-
lus (Jebel population) by having claws beyond
scansors (Arnold and Gardner, 1994). A. tanges-
tanensis sp. nov. differs from A. elisae (Arnold
and Gardner, 1994; Torki et al., 2008; Torki, in
press), and other new Asaccus (A. barani,A.
iranicus, and A. zagrosicus) by having large, tri-
hedral keeled tubercles type c on most part of
dorsal body (including dorsal, neck, and limbs)
and thin (plus long) limbs. A. tangestanensis sp.
nov. (up to 50 mm) is larger than A. montanus
(less than 40 mm), and also has thinner limbs
and body than A. montanus (Arnold and Gard-
ner, 1994; Gardner, 1994).
Asaccus zagrosicus sp.nov.(g.6)
Holotype. ZFMK 91935, adult male, collected
in the western slope of central Zagros Moun-
tains, Tang-e-Haft region, Khorramabad City,
Figure 6. Holotype of Asaccus zagrosicus sp. nov. (a) dorsal view; (b) ventral view; (c) postmentals, 2nd postmentals distinct
from lowerlabials; (d) one of Paratype of A. zagrosicus sp. nov. before preservation.
194 F. Tor k i et al.
Lorestan Province, western Iran on 24 August,
2008 (3302N-4839E, altitude 647 m a.s.l.)
by Farhang Torki.
Paratypes. FTHM 002645-002650, 002652-
2669, 002765-002770. 31 adult specimens,
same data as for holotype.
Diagnosis. Medium size Asaccus (44-55 mm);
tubercles are distributed in dorsal body, neck,
occipital, limbs, and tail; dorsal tubercles are
small, keeled, trihedral type b intermixed with
pointed tubercles; 16-18 irregular rows at mid-
trunk; neck tubercles are weakly keeled or
pointed; occipital and upper head tubercles are
pointed or simple; few simple tubercles be-
tween interorbit; few large tubercles (simple or
pointed) are in front of eyes; arm and fore-
arm tubercles are pointed; thigh tubercles are
pointed or keeled; arm tubercles have low den-
sity; several simple or weakly keeled tubercles
on the forearm; 8 tubercles form each whorls
of tail; scales on upper head are cobble-stone
shaped; scale weakly globular shape between
the eyes; two postmentals, secondary postmen-
tals not in contact with lowerlabial (separated
from lower labials with 1-3 rows scales); small
and weakly overlapped chin scales; between in-
ter hindlimbs scales have same shape and size
to ventrals; mental trihedral; claves in front of
scansors.
Description of holotype. An adult male
(ZFMK 91935); medium size (52.26 mm) Asac-
cus; flat body and head; tubercles are distributed
in dorsal body, neck, occipital, limbs, and tail;
small, keeled, trihedral type b intermixed with
pointed dorsal tubercles; 16-18 irregular rows,
1-4 scales between large tubercles; neck tuber-
cles are small, weakly keeled or pointed, not
extend to laterals side; occipital tubercles are
pointed or simple; upper head tubercles pointed
or simple; few simple tubercles between the in-
terorbit; few large tubercles (simple or pointed)
in front of the eyes; tubercles do not extend
between ear-neck or ear-eyes; arm and fore-
arm tubercles are pointed; elbow without tuber-
cles; thigh tubercles are pointed or keeled; the
density of arm tubercles is low; several sim-
ple or weakly keeled tubercles on the forearm;
dorsal tubercles extend into tail; tail tubercles
whorl do not extend onto ventral side, 8 tu-
bercles form each whorls, 5 scales between the
whorls; dorsal scales not overlapped (but is con-
densed); scale on the upper head is cobble-
stone shaped; between the eye scales weakly
globular shape; five scales around the nostril
(1 nasal, 1st supralabials, 2 scale on poste-
rior, and rostral); internasal scales are in con-
tact; two scale on posterior of internasals (one
small scale is between internasals and posteri-
ors); 10 rows of large scale between the nostril
and the eyes; 11 supralabials (9 to mid-eye); 9 in
lower labials; two postmentals, first postmentals
are larger than second, first postmentals are in
contact each other, and in contact with first low-
erlabials; secondary distinct postmentals with
6 scales, not in contact with lowerlabial (sepa-
rated from lower labials with one row of scales);
chin scales are small and weakly overlapped;
ventral neck scales are large and overlapped;
ventral scales are medium size and partly over-
lapped, 2.5 times larger than the dorsal scales;
ventral scales of arm are smaller (1/4 times)
than the forearm; arm scales are rounded and
forearm scale are tetragon; ventral scales of both
limbs are cobble-stone shaped (not globular or
rotund); scales on palm of both forelimbs are
globular shape; ventral scales of thigh and fore-
leg are partly overlapped; rounded foreleg scale
shape are larger than the hexagonal thigh scales;
ornamentation of palm (both limbs especially
hindlimbs) scales are partially similar to lamel-
las; between inter hindlimbs scales have same
shape and size to ventrals; ventral tail scales are
large and cover most ventral side; mental trihe-
dral (length/width: 2.65 mm/2.64 mm); rostral
width (length/width: 1.03 mm/2.19 mm) and not
divided up; 7 lamellas underneath the 4th fin-
ger (forelimb); 9 lamellas underneath the 4th toe
(hindlimb); claves in front of scansors.
Dorsal background including dorsum, head;
limbs and tail in live specimen is darken; spots
covers dorsum, limbs, and head (few observable
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 195
in preservation); upper labials are dark; digits
are covered by small dark spots; ventral body is
dirty white; 5 dark bands on tail do not extend
onto ventral side, 4 black rings (partly extended
onto ventral side) of secondary half of tail; black
ring is weakly different to bands in preserved
specimen.
Variation. 3-9 scales between secondary post-
mentals; secondary postmentals are smaller
than first postmentals; secondary postmentals
are in some specimens smaller than others;
12-15 rows of tubercles on dorsal body; tail
length/body length in specimens with original
tail (1.11-1.43) is more than in those with re-
generated tail (0.85-1.0); length of original tail
is more than body length; in contrast in regen-
erated tail, tail is smaller than body (maximum
regenerated size is similar size to body length).
Additional characters are given in table 3.
Dorsal body is covered by full crossbands,
spots (large or small), in some specimens no any
pattern on dorsal side was observed; dorsolat-
erals of head is covered by small spots; limbs
are covered by crossbands or spots; tail is co-
vered by dark bands not extend into ventral side;
some specimens have black ring completely ex-
tend into ventral side on secondary half of tail;
the color of ventral is dirty white; lateral of chin
is dark in some specimens. All over the dor-
sal body including dorsum, head, limbs and tail
have darkened background in live specimens;
color and pattern in live specimens is easily vis-
ible; a variety of patterns were observed (Torki
and Sharifi, 2007) in this new Asaccus including
(a) spotty, (b) full crossbandsand (c) intermixed
pattern.
Sexual dimorphism. Males and females were
compared according to morphological charac-
ters (see Material and methods) in order to de-
termine sexual dimorphism. In general, males
(mean ±SE =51.23 ±0.72 mm) are longer
than the females (mean ±SE =50.49 ±
1.0 mm). According to statistical analyses re-
sults, only one characters (SPM – number of
scales surrounded postmentals) showed statis-
tically differences between males and females
(P<0.01) and females had highest SPM
value (mean ±SE =20.89 ±0.45 mm) than
the males (mean ±SE =18 ±0.7 mm).
Habitat and ecology. Specimens belonging to
A. zagrosicus sp. nov. were collected from only
several tunnels in Tang-e-Haft Region, South-
ern Lorestan. Tang-e-Haft Region has warm cli-
matic condition and is located between central
Zagros Mountains and Khuzestan Plain. There
is no tree near the type locality but oak (Quer-
cus sp.) forest is distributed in mountains of
this region. A. zagrosicus sp. nov. has egg-sites,
which is unusual within Asaccus. Shelter site
and egg-site are strongly distinct in most part of
the habitat. Biological activity of A. zagrosicus
sp. nov. was recorded from mid-winter to mid-
autumn. Most of the specimens have noctur-
Tab l e 3 . Morphological characters of Asaccus zagrosicus sp. nov. (only adult specimens; for abbreviation see table 2).
*: characters in mm; other characters are numerical.
Characters N Mean Min. Max. SD SE Characters N Mean Min. Max. SD SE
SVL* 15 50.72 45.90 54.10 2.48 0.64 NE* 15 11.98 10.92 13.04 0.64 0.16
TL* 11 55.75 45.72 66.86 7.4 2.23 FLL* 15 20.26 18.2 22.8 1.3 0.33
HW* 15 9.59 8.29 11.11 0.68 0.18 HLL* 15 27.5 24.1 31.25 2.14 0.55
HL* 15 15.39 13.43 17.15 1.03 0.27 AG* 15 22.87 20.85 27.8 1.82 0.47
HD* 15 5.63 4.69 6.3 0.47 0.12 LT 15 9.07 9.00 10.00 0.26 0.067
HI 15 160.69 143.48 170.68 8.81 2.27 DTR 15 12.60 11.00 14.00 0.91 0.24
HF 15 268.75 235.58 332.19 23.65 6.11 PM 15 2.00 2.00 2.00 0.00 0.00
SW* 15 4.74 3.8 5.56 0.59 0.15 SPM 15 20.13 16.00 24.00 2.20 0.57
EHD* 15 3.38 2.67 4.8 0.48 0.12 SL 15 18.20 16.00 21.00 1.52 0.39
EVD* 15 1.57 1.26 2.12 0.24 0.06 IS 15 20.80 18.00 25.00 2.15 0.55
NY* 15 5.4 4.7 6.0 0.39 0.1 NES 15 10.93 9.00 13.00 1.10 0.28
EE* 15 4.03 3.3 5.3 0.44 0.11
196 F. Tor k i et al.
nal and diurnal activity because of the darkened
tunnels during day. During spring and summer
egg could be seen in abdomen of females. This
new Asaccus feeds on small insect and insect
larva distributed all around of the habitat.
Distribution. Currently the new species is
only recorded from the tunnels situated at the
type locality at Tang-e-Haft region, Khorram-
abad, Lorestan, Iran. In spite of several field
trips to adjacent areas to the type locality, it was
not recorded elsewhere.
Sympatric lizards and snake. From type local-
ity the following reptile species were recorded:
Hemidactylus sp., Cyrtopodion scabrum,Pseu-
docerastes persicus fieldi. In mountains around
type locality, the other gekkonid lizard Asaccus
griseonotus was also recorded.
Etymology. The species is named for the
Zagros Mountains in which the type local-
ity (Tang-e-Haft region, Khorramabad City,
Lorestan Province) is found.
Comparison. A. zagrosicus sp. nov. differs
from other Asaccus as follows. Secondary post-
mentals are not in contact with lowerlabials in
all specimens (100% of specimens); this is in
contrast to other Asaccus (Arnold and Gardner,
1994; Gardner, 1994; Anderson, 1999; Torki et
al., 2008; Afrasiab and Mohamad, 2009; Torki,
2009, in press). Scansors in the A. zagrosicus sp.
nov. do not extend beyond claws and tubercles
are present on the arm; this is in contrast to A.
gallagheri,A. platyrhynchus,A. caudivolvulus
(Khasab population), A. kurdistanensis,A. ker-
manshahensis,A. nasrullahi,A. granularis,A.
andersoni,A. griseonotus and A. saffinae (e.g.,
Arnold and Gardner, 1994; Torki and Sharifi,
2007; Torki et al., 2008; Afrasiab and Mo-
hamad, 2009; Torki, 2009, in press; Torki, Fa-
tinia and Rostami, in press). A. zagrosicus sp.
nov. shows difference with A. caudivolvulus
(Jebel population) by having claws beyond the
scansors (Arnold and Gardner, 1994). A. za-
grosicus sp. nov. (up to 40 mm) has larger body
size than the A. montanus and A. gallagheri
(less 40 mm) (Arnold and Gardner, 1994; Gard-
ner, 1994).
Asaccus barani sp.nov.(g.7)
Asaccus elisae Dixon and Anderson (1973)
Asacus [sic] elisae Baran and Gruber (1982)
Asaccus elisae Arnold and Gardner (1994)
Asaccus elisae Tok, Atatür and Ta¸skavak (1997)
Figure 7. Holotype of Asaccus barani sp. nov. (a) dorsal
view; (b) ventral view; (c) high variation dorsal tubercles,
simple tubercles.
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 197
Asaccus elisae Franzen, Schmidtler and Bis-
choff (2002)
Asaccus elisae Tok, Atatür and Cihan (2002)
Asaccus elisae Baran et al. (2003)
Asaccus elisae Torki et al. (2008)
Holotype. ZDEU 70/2001-1, an adult male,
collected from Harran, ¸Sanlıurfa, Southeastern
Anatolia, Turkey on 31 April 2001 (3651N-
3900E, 380-390 m a.s.l.) by ˙
I. Baran, Y. Kum-
luta¸s, Ç. Ilgaz and A. Avcı.
Paratypes. ZDEU. 70/2001.2-17. 16 adult
specimens, same data as for holotype.
Diagnosis. Medium size Asaccus (34-56 mm);
the dorsal body is covered all over by tuber-
cles; keeled trihedral dorsal tubercles are be-
tween type a and b, simple and pointed tuber-
cles are intermixed with keeled dorsal tubercles;
dorsolateral tubercles are pointed or simple; tu-
bercles present on occipital and especially neck
show considerable variation in size and type;
several pointed or keeled tubercles are in front
of eyes; two small tubercles (one simple and one
keeled) are between ear-eye; several keeled or
simple tubercles are on the arm, some of them
are similar to scales; elbow and knee scales are
generally tubercular like (simple); forearm tu-
bercles extend onto elbow in most the speci-
mens; forearm tubercles also extend onto knee
in some specimens; several keeled tubercles are
on palm of hindlimbs, 1st, 2nd and 3rd digits;
two large postmentals; interhindlimbs scales are
same size to ventrals but are mostly same shape
to ventral of thigh; mental pentagonal; claves in
front of the scansors.
Description of holotype. An adult male; large
Asaccus (54.89 mm); depressed body; the
dorsal body is covered all over by tubercles;
moderate, trihedral between type a and b dorsal
tubercles, 12-13 irregular rows, 1-4 scales are
between large tubercles; several simple tuber-
cles are on dorsal body; dorsolateral tubercles
are pointed or simple; neck tubercles are keeled,
pointed, weakly keeled or simple (high vari-
ation); occipital tubercles are simple, pointed
or few tubercles keeled; upper head tubercles
are simple and small; upper ear tubercles are
pointed or weakly keeled; simple tubercles are
extend into interorbits; several pointed or keeled
tubercles in front of eyes; without any tubercles
between ear-neck; two small tubercles (one sim-
ple and one keeled) between ear-eyes; several
tubercles (keeled or simple) on the arm, some
of them are similar to scales; elbow scales are
tubercular like (simple); forearm tubercles are
pointed or weakly keeled; without any tubercles
on palm of forelimbs or digits; thigh and fore-
leg tubercles are keeled; several keeled tuber-
cles are on palm of hindlimbs, 1st, 2nd and 3rd
digits; dorsal tubercles are extend into tail; tail
tubercles whorls do not extend onto ventral side,
6 tubercles form each whorl, 4 scales between
whorls; dorsal and neck scales are simple; up-
per head and interorbits scales are globular like;
five scales around the nostril (1 nasal, 1st supral-
abials, 2 scale on posterior, and rostral); inter-
nasal scales extend and in contact each other;
two large scale on posterior of the internasals
(one small scale between internasals and poste-
riors); 9 rows of large scale between the nos-
tril and the eyes; 11 supralabials (8 large and
3 small, 8 to mid-eye); 10 in lower labials (9th
and 10th are small); two large postmentals, first
postmentals are larger than secondary, 1st post-
mentals are in contact each other, and in con-
tact with 1st and 2nd lowerlabial; secondary
postmentals are distinct with 5 scales and in
contact with 2nd lowerlabial; chin scales are
small and not overlapped; strongly overlapped
neck scales are larger than chin scales; rounded
ventral scales are medium size and not over-
lapped, 2 times larger than the dorsal scales;
ventral scales on arm and forearm are rounded
and partly overlapped; arm scales are smaller
than forearm scales; forelimbs palm scales are
globular shape and similar to lamella (ornamen-
tation); thigh scales are partly tetragonal and
not overlapped; foreleg scales are rounded and
partly overlapped; palm of hindlimbs are partly
globular like, and are similar ornamentation to
lamellas; inter-hindlimbs scales are same size
198 F. Tor k i et al.
to ventrals but are mostly same shape to ven-
tral of thigh; ventral tail scales are large and
covered most of the ventral side; mental pen-
tagonal (length/width: 3.19 mm/2.90 mm); ros-
tral wide (length/width: 1.08 mm/2.55 mm) and
not divided; 8 lamellas beneath 4th finger; 10
lamellas underneath the 4th toe; claves in front
of scansors.
The dorsum is covered all over by spots or
crossbands as follows: Dorsal body is covered
by five crossbands and several large spots; sev-
eral large spots (combined together) are on the
upper head; snout is covered by six straight
bands (from snout to eyes); two straight bands
are present between the eyes and the ears (four
bands on both side); tail is covered by three dark
bands having same color to dorsal bands and
three black rings; limbs and digits are covered
by spots (partly band-like); ventral body (except
lover labials) is white without any pattern; color
of ventral side of limbs and tail are cream (dif-
ferent color from ventral).
Variation. Dorsal body is covered by several
type tubercles such as simple, pointed, keeled
(type a and b) tubercles; simple tubercles are
on the dorsal side of the body could be seen
in all specimens but their density is change-
able; neck tubercles are simple (large) or weakly
keeled type a; upper head tubercles are simple
or partly pointed in all specimens; upper arm
tubercles are simple and elevated in most of
the specimens; tubercles in some specimens is
found on elbow and in other is not extended;
mental pentagonal; first postmental in all spec-
imens is larger than secondary postmentals; 4-
7 scales between secondary postmentals. Addi-
tional characters are given in table 4.
Dorsal body, neck, upper head and limbs are
covered by spots or bands; five crossbands on
dorsal body, dorsal crossbands are full and are
regular or irregular, most specimens have spots
on dorsolaterals; limbs are covered by spots
(band like); original tail has two or three black
rings (mostly three); regenerated tail is without
any pattern; original tail have two or three bands
the same as dorsal that do band not extend onto
ventral side; 4-6 bands extend from the nostril
to the eyes; labials (lower and especially upper)
are strongly pigmented; tip of tail is strongly
flattened in original and regenerated ones.
Habitat and ecology. The Harran plains is one
of the driest areas of SE Anatolia, with annual
rainfall ranging approximately from 300 to 600
mm, largely depending on altitude. Terra typica
is characterized by the ruins of ancient castle,
in an area covered by hardened grey sand with
intermixed blocks of sedimentary fossiliferous
rocks. The vegetation is a steppe, with scattered
shrubs mostly less than 50 cm high. The domi-
nant vegetation sample is Peganum harmala L.
Distribution. Specimens belonging to new ta-
xa were currently only recorded from the type
locality (Harran). However, Birecik and Nusay-
Tab l e 4 . Morphological characters of Asaccus barani sp. nov. (only adult specimens; for abbreviation see table 2).
*: characters in mm; other characters are numerical.
Characters N Mean Min. Max. SD SE Characters N Mean Min. Max. SD SE
SVL* 10 45.78 33.34 56.49 8.55 2.70 NE* 10 11.19 8.82 13.38 1.77 0.56
TL* 9 54.98 41.34 75.00 10.58 3.53 FLL* 10 19.11 15.07 25.25 3.41 1.08
HW* 10 8.90 6.65 10.86 1.57 0.50 HLL* 10 26.56 21.53 32.33 4.31 1.36
HL* 10 14.30 10.85 16.96 2.40 0.76 AG* 10 19.04 14.39 24.19 3.55 1.12
HD* 10 5.86 4.23 6.82 0.87 0.27 LT 10 10.10 9.00 11.00 0.74 0.23
HI 10 160.99 151.68 168.35 5.26 1.66 DTR 10 11.70 10.00 13.00 0.95 0.30
HF 10 255.78 232.83 270.69 12.76 4.03 PM 10 2.00 2.00 2.00 0.00 0.00
SW* 10 3.76 2.82 4.70 0.63 0.20 SPM 10 16.90 15.00 21.00 1.73 0.55
EHD* 10 3.32 2.65 4.25 0.50 0.15 SL 10 17.50 16.00 19.00 1.08 0.34
EVD* 10 1.74 1.25 2.40 0.36 0.11 IS 10 20.00 16.00 24.00 2.36 0.75
NY* 10 5.14 4.05 6.20 0.78 0.24 NES 10 10.60 9.00 12.00 1.08 0.34
EE* 10 3.55 2.67 4.33 0.57 0.18
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 199
bin specimens that were previously evaluated as
Asaccus elisae (Böhme, 1973; Baran and Gru-
ber, 1982; Tok, Atatür and Tskavak, 1997; Tok,
Atatür and Cihan, 2002) in Anatolia must be re-
examined in terms of morphological characters
in order to stabilize the distribution of new taxa
in Anatolia.
Sympatric lizard and snake species. Other
reptiles collected or observed at these ruins in-
clude: Acanthodacylus harranensis,Ophisops
elegans,Trachylepis aurata,Eumeces schnei-
deri and Malpolon monspessulanus.
Etymology. The new species is in dedication
to Prof. Dr. ˙
Ibrahim Baran of the University of
Dokuz Eylül, ˙
Izmir, to acknowledge his prolific
and uninterrupted contribution to the herpetol-
ogy of Turkey.
Comparison. Asaccus barani sp. nov. is eas-
ily distinguishable from other Asaccus as fol-
lows. Scansors in A. barani sp. nov. do not ex-
tend beyond claws and tubercles are present on
the arm; this is in contrast to A. gallagheri,A.
platyrhynchus,A. caudivolvulus (Khasab popu-
lation), A. kurdistanensis,A. kermanshahensis,
A. nasrullahi,A. granularis,A. andersoni,A.
griseonotus and A. saffinae (e.g., Arnold and
Gardner, 1994; Torki et al., 2008; Afrasiab and
Mohamad, 2009; Torki, 2009, in press). Knee
and especially elbow scales are globular and
some tubercles present on elbow in A. barani sp.
nov. This is in contrast to A. elisae,A. zagrosi-
cus,A. iranicus,A. tangestanensis (partly), A.
kurdistanensis,A. kermanshahensis,A. nasrul-
lahi,A. granularis,A. andersoni,A. griseono-
tus,A. saffinae,A. gallagheri,A. platyrhynchus
and A. caudivolvulus (e.g., Arnold and Gard-
ner, 1994; Torki and Sharifi, 2007; Torki et al.,
2008, in press; Afrasiab and Mohamad, 2009;
Torki, 2009, in press). A. barani sp. nov. shows
difference with A. caudivolvulus (Jebel popula-
tion) by having claws beyond the scansors. A.
barani sp. nov. has maximum variation in tu-
bercles present on dorsal side in terms of size
and shape, this is in contrast to A. elisae,A.
montanus and other new Asaccus (A. zagrosi-
cus,A. iranicus, and A. tangestanensis) (Arnold
and Gardner, 1994; Gardner, 1994). A. barani
sp. nov. also differs from A. montanus by hav-
ing a large body size (Gardner, 1994).
Evaluation of taxonomic characters of
genus Asaccus
Based on the present study we consider that
genus Asaccus includes 16 species: one species,
A. barani sp. nov. is distributed in the Anatolian
plateau; four species, including A. montanus,A.
gallagheri,A. platyrhynchus and A. caudivolvu-
lus are distributed in the Oman-Emirate Moun-
tains; and eleven species, including A. kurdis-
tanensis,A. kermanshahensis,A. nasrullahi,A.
granularis,A. andersoni,A. griseonotus,A. saf-
finae,A. elisae,A. tangestanensis sp. nov., A.
iranicus sp. nov. and A. zagrosicus sp. nov., are
distributed in the Zagros mountains. The dis-
tribution site of the genus Asaccus is covered
by the Anatolian-Zagros-Oman-Emirate Moun-
tains. These branch mountains were distinctive
of the Persian Gulf. Therefore, Asaccus is dis-
tributed across two branch mountains: (1) the
Anatolian-Zagros branch; and (2) the Oman-
Emirate branch. A. barani sp. nov. is an en-
demic species of Asaccus from Turkey and is
only distributed in the Anatolian plateau, which
is the upper latitude of distribution of Asaccus.
A. barani sp. nov. is different from other species
of Asaccus due to having maximum variation in
tuberculation (type, size and distribution) on the
dorsal body. This character is unique for Anato-
lian Asaccus. The geographic distance from A.
barani to Zagrosian Asaccus is is a greater in-
terval to the other species (the distance between
Harran – type locality of A. barani and Sarv-
abad – type locality of A. kurdistanensis,the
nearest Zagrosian Asaccus to A. barani, is ap-
proximately 680 km). In contrast, a maximum
variation on tuberculation for Asaccus was re-
corded in the Oman-Emirate branch. In this re-
gion it was recorded that minimum tubercula-
tion (A. gallagheri), moderate tuberculation (A.
platyrhynchus) and strong tuberculation Asac-
200 F. Tor k i et al.
cus (A. montanus) occurred in separate species.
The Zagros Mountains are a continuation of
the Anatolian plateau, and in this region twelve
endemic Asaccus are distributed. Finally, four
endemic Asaccus are distributed in the Oman-
Emirate Mountains. Therefore, in this study we
considered that speciation within Asaccus oc-
curred in three hot-spots, as follows: (1) Anato-
lian hot-spot, as a small hot spot; (2) Zagros hot-
spot, as a great hot-spot for speciation; and (3)
Oman-Emirate hot-spot as a moderate hot-spot.
Already, the senior author cited two hot-spots
for Asaccus (Torki et al., 2008; Torki, Heidari
and Khan, 2010): Anatolian-Zagros and Oman-
Emirate. However, based on new material (de-
scription of new Asaccus from the Anatolian
plateau with a special morphological pattern)
we increase this to three hot-spots. Fold moun-
tains are an important element for the speciation
of Asaccus. The Zagros Mountains have great
folds among all hot-spots. Based on our field-
work, Asaccus was distributed on the western
slope of the Zagros Mountains, and all species
of Asaccus were distributed in the folded moun-
tains. Large and small crevices of folded moun-
tains are important shelter sites for Asaccus, and
also for egg laying. Rastegar-Pouyani (2006)
stated that Asaccus is a vicariant taxon and the
most important factor in separation and isola-
tion of the northern (Zagros) populations from
the southern (Arabian Peninsula) populations of
Asaccus was the formation of the Persian Gulf.
According to Rastegar-Pouyani (2006), the cen-
ter of diversification for Asaccus is the Zagros
Mountain because of the species diversity.
Several characters are important for the di-
agnosis of the 16 species. Tubercle type, size
and distribution are particularly important tax-
onomical characters and most species could be
diagnosed based on tuberculation. In this study
we focused on the size, type and distribution
of tubercles all over the dorsal body including
dorsum, neck, upper head and limbs. On the
other hand, the colour patterns of some species
are important taxonomical characters. There-
fore, we briefly discussed important taxonom-
ical characters for all 16 species as follows:
A. gallagheri: distinguished from other Asac-
cus due to a lack of tuberculation on the dor-
sal body; A. kermanshahensis: by four postmen-
tal, tuberculation occurred on elbow and lower
part of arm with dorsal tubercles weakly type a;
A. kurdistanensis: tuberculation occurred all
over the dorsal body (except arms) with dor-
sal scale keeled type a; A. andersoni: 100% of
specimens have dichromatism, and the dorsal
colour in males is silver and in females is or-
ange; A. granularis: simple dorsal tubercles do
not extend onto the neck and upper head; A.
griseonotus: shadow-like crossbands, and more
than 10 rows of tubercles on the dorsal; A. nas-
rullahi: shadow-like crossbands, and 6-8 rows
of tubercles on the dorsal; Asaccus barani sp.
nov.: simple tubercles and intermixed keeled tu-
bercles type a and b; A. zagrosicus sp. nov.: sec-
ondary postmental in 100% of specimens not in
contact with lowerlabials; A. platyrhynchus:tail
tip uncompressed and small tubercles distrib-
uted on the dorsal, neck, upper head and limbs
(except arms); A. khasabensis: tail tip flattened,
scale across midorbit fine, tubercles absent on
arms; A. tangestanensis sp. nov.: strong tubercu-
lation (large and type c) all over the dorsal body
(especially on the dorsum, limbs); A. montanus:
granules covered the dorsal body, strong tuber-
culation; A. iranicus sp. nov.: small body size
and digits (forelimbs), different joint to palm
from other Asaccus;A. elisae: tuberculation oc-
curred all over the dorsal body (including arms),
uniform dorsal tuberculation.
Afrasiab and Mohamad (2009) recently de-
scribedanewAsaccus from Iraq (A. saffinae).
Afrasiab and Mohamad (2009) placed A. saffi-
nae close to A. kermanshahensis, but is it easy
to distinguish that A. saffinae from A. kurdis-
tanensis due to the many differences between
them, such as keeled dorsal tubercles, dorsal tu-
bercles extended into upper head, four postmen-
tals and so on (Torki and Sharifi, 2007; Torki et
al., 2008; Torki, Heidari and Khan, 2010). Torki
et al. (2008) and Torki, Heidari and Khan (2010)
found other variations in all Asaccus inhabitants
Four new Asaccus Dixon and Anderson, 1973 (Reptilia: Phyllodactylidae) from Iran and Turkey 201
of Iran, including A. kermanshahensis.Forex-
ample a dark band on the tail is an not impor-
tant characteristic, because Rastegar-Pouyani
(1996) found that both regenerated tails and
original tails can have more than 7-9 dark bands
(Torki et al., 2008; Torki, Heidari and Khan,
2010). Based on above reasons and all FTHM
materials (n =8: FTHM 002800-002807), it
is clear that A. saffinae is very different to A.
kermanshahensis. Based on our material on A.
griseonotus (n =60: FTHM 003040-003099),
all the characteristics are similar in this popula-
tion. Previously, Torki et al. (2008) and Torki,
Heidari and Khan (2010) had reported this vari-
ation for A. griseonotus. Based on FTHM col-
lections (FTHM 003040-003099) and a recent
report (Torki et al., 2008; Torki, Heidari and
Khan, 2010), in this study we redescribed A.
griseonotus as follows: dorsal tubercles simple
or pointed, extended into neck and occipital (do
not extended into upper head and interorbits); 5-
6 shadow-like bands on dorsal (5 m-shaped, in
Afrasiab and Mohamad, 2009); 1st postmentals
in contact or distinguished by one scale; median
dorsal vertebrate groove present. Other charac-
teristics are the same as previous reports about
A. griseonotus (Dixon and Anderson, 1973; An-
derson, 1999; Torki et al., 2008; Torki, Heidari
and Khan, 2010), such as a nostril surrounded
by five scales, scansore clearly extends beyond
claw, upper arm without tubercles, and so on.
All the characteristics of Asaccus saffinae de-
scribed in Afrasiab and Mohamad (2009) are
very similar to Asaccus griseonotus (Dixon and
Anderson, 1973; Anderson, 1999; Torki et al.,
2008; Torki, Heidari and Khan, 2010). We can
not find any differences between A. saffinae and
the Iranian A. griseonotus,soA. saffinae is a
synonym for A. griseonotus. Therefore in this
study we omitted A. saffinae from the list of
Asaccus.
The first data on molecular phylogenetic
analysis of genus Asaccus reveals high sequence
divergence among the sampled species (Papen-
fuss et al., 2010). Molecular data show that one
population of Iranian sample of A. elisae is more
closely related to A. griseonotus than it is to the
Turkish samples of A. elisae and the concept of
A. elisae may be species complex. In this study
we show morphological differences among four
new species with A. elisae and other Asaccus
from Iranian and Anatolian plateau.
Key to the species of the genus Asaccus
(after Arnold and Gardner, 1994; Anderson,
1999; Afrasiab and Mohamad, 2009; Torki,
Heidari and Khan, 2010)
1a: digits (forelimbs) parallel joint to palm . .
...............Asaccus iranicus sp. nov.
1b: digits (forelimbs) joint to palm different to
above...............................2
2a: dorsal tubercles absent . . . . . . . . . . . . . . . . .
....................Asaccus gallagheri
2b: dorsal tubercles present . . . . . . . . . . . . . . . 3
3a: dorsal tubercles keeled . . . . . . . . . . . . . . . .7
3b: dorsal tubercles simple or weakly keeled4
3c: dorsal tubercles intermixed . . . . . . . . . . . . .
................Asaccus barani sp. nov.
4a: dorsal tubercles strongly extend onto neck
and upper head . . . . . . . . . . . . . . . . . . . . . . .5
4b: dorsal tubercles extend onto neck and
weakly upper head . . . . . . . . . . . . . . . . . . . 6
4c: dorsal tubercles do not extend onto neck
and upper head . . . . . . Asaccus granularis
5a: interorbit scales fine . . . . . . . . . . . . . . . . . . .
.................Asaccus platyrhynchus
5b: interorbit scales coarse . . . . . . . . . . . . . . . . .
.....................Asaccus andersoni
6a: 6-8 tubercles rows on dorsal . . . . . . . . . . . .
....................Asaccus nasrullahi
6b: more than 10 tubercles rows on dorsal .. .
...................Asaccus griseonotus
(A. saffinae Afrasiab and Mohamad, 2009)
7a: tubercles do not extend on arm. . . . . . . . . 8
7b: tubercles strongly extend on arm . . . . . . 10
8a: four postmentals . . . . . . . . . . . . . . . . . . . . . . .
..............Asaccus kermanshahensis
8b: 2-3 postmentals . . . . . . . . . . . . . . . . . . . . . . 9
9a: black ring present on tail . . . . . . . . Asaccus
caudivolvulus (Khasab population)
9b: black ring absent on tail . . . . . . . . . . . . . . . .
.................Asaccus kurdistanensis
202 F. Tor k i et al.
10a: scansor do not clearly extend beyond claw
.................Asaccus caudivolvulus
(Jebel population)
10b: scansor clearly extend beyond claw . . . 11
11a: enlarged and trihedral tubercles all over
dorsal body; body and limbs thin . . . . . . . .
.........Asaccus tangestanensis sp. nov.
11b: large or moderate trihedral tubercles on
dorsal; body and limbs not thin . . . . . . . 12
12a: 2nd postmentals distinct from lowerlabials
.............Asaccus zagrosicus sp. nov.
12b: 2nd postmentals contact lowerlabials . . . .
...................................13
13a: body size large (more than 50 mm). . .. . .
........................Asaccus elisae
13b: body size small (less than 50 mm) . . . . . . .
.....................Asaccus montanus
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... Despite the intrinsic scientific interest surrounding the Asaccus species in the Zagros, research efforts have been limited, and comprehensive studies covering the entire region are scarce. Since 1994, the number of Asaccus species recognized as endemic to the Zagros massif has risen to 10, indicating a growing but still incomplete understanding of their diversity and distribution [4,[8][9][10][11][12][13][14][15][16][17][18][19][20]. This incremental discovery highlights both the richness of the Zagros as a habitat for unique species and the urgent need for further, more expansive genetic and ecological studies to fully comprehend the range and evolutionary dynamics of the Asaccus genus in this biodiverse region. ...
... This may suggest that A. elisae comprises different species with similar morphological characters. The two species of A. iranicus and A. tangestanensis described by Torki et al. [11]. have until now been recorded from their type localities only. ...
... Following nomenclatural priorities, A. nasrullahi was synonymized under A. griseonotus. Also, A. zagrosicus which was recently described from south of Iran [11] was found to be closely related to A. elisae. Although A. zagrosicus is diagnosably different from A. elisae in having secondary postmentals separated from lower labials by 1-3 granules, it was nested within A. elisae (Fig. 2) with high support in the phylogenetic trees, species delimitation results and haplotype networks. ...
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... More recently, molecular studies have shown that reptile diversity in the Hajar Mountains was still underestimated, with some gecko groups of the genera Hemidactylus , Pristurus (Badiane et al., 2014;Garcia-Porta, Sim o-Riudalbas, Robinson, & Carranza, 2017), Ptyodactylus (Metallinou et al., 2015;Sim o-Riudalbas et al., 2017), Trachydactylus (de Pous et al., 2016a) and Asaccus (Carranza, Sim o-Riudalbas, Jayasinghe, Wilms, & Els, 2016;Papenfuss et al., 2010) harbouring several cryptic species and deep lineages, some of them with restricted distributions of just a few kilometres. From all the reptile genera of the Hajar Mountains, Asaccus Dixon & Anderson, 1973 is the one with the highest number of endemics (see below) and a good example of the faunal affinities between the Hajar Mountains in Arabia and the Zagros Mountains in Iran Papenfuss et al., 2010;Torki, Ahmadzadeh, Ilgaz, Avcı, & Kumlutaş, 2011a;Uetz, Goll, & Hallerman, 2017). In a similar way, closely related taxa and populations from various reptile groups are present on both sides of the Gulf of Oman and provide an excellent scenario for reconstructing their evolutionary relationships and to explore the biogeography of the region (Dakhteh, Kami, & Anderson, 2007;de Pous et al., 2016b;Kapli et al., 2008Kapli et al., , 2015Krause, Ahmadzadeh, Moazeni, Wagner, & Wilms, 2013;Metallinou et al., 2012Metallinou et al., , 2014Yousofi, Rastegar-Pouyani, & Hojati, 2015). ...
... All voucher specimens were obtained from S. Carranza's field series housed at the Institute of Evolutionary Biology (IBE), Barcelona, Spain; the Natural History Museum, London, UK (NHMUK) and the Oman Natural History Museum, Muscat, Oman (ONHM) (Tables S1 and S4, see supplemental material online). Variables for the morphological analyses were selected based on previous taxonomic studies of Asaccus (Afrasiab & Mohamad, 2009;Arnold, 1972;Arnold & Gardner, 1994;Carranza et al., 2016;Dixon & Anderson, 1973;Gardner, 1994;Rastegar-Pouyani, 1996;Rastegar-Pouyani, Nilson, & Faizi, 2006;Torki, 2010;Torki et al., 2011a;Torki, Fathinia, Rostami, Gharzi, & Nazari-Serenjeh, 2011b;Werner, 2006). Specimens were sexed looking at two external characters: presence of hemipenal bulges and colour of original tails (yellow in males). ...
... Therefore, we describe the unnamed populations from the Eastern Hajars as a new species endemic to Oman. Data for the comparison with other Asaccus species were obtained from our own morphological dataset (Table 1 and Fig. S4, see supplemental material online) and also from morphological information available from the original descriptions of all 18 species of Asaccus (Afrasiab & Mohamad, 2009;Arnold, 1972;Arnold & Gardner, 1994;Carranza et al., 2016;Dixon & Anderson, 1973;Gardner, 1994;Rastegar-Pouyani, 1996;Rastegar-Pouyani et al., 2006;Torki, 2010;Torki et al., 2011aTorki et al., , 2011bWerner, 2006). ...
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In this study, Asaccus elisae specimens, which were caught from Harran, Vilayet Þanlýurfa in SE Turkey was examined in terms of morphological features, pholidosis properties, coloration and pattern. Specimens collected from Harran were compared with specimens from Birecik (Turkey), Iraq, Syria and Iran with regard to literature.
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