Amphibia-Reptilia 38 (2017): 449-459

Geographic variation in morphology of the Agamura Blanford, 1874 in

Seyyed Saeed Hosseinian Yousefkhani1, Mansour Aliabadian1,2,∗, Eskandar Rastegar-Pouyani3, Jamshid Darvish1,2

Abstract. The genus Agamura was previously known from four species occurring on the Iranian Plateau, but was recently revised as a monotypic genus that excluded three species. In the present study, we examined different populations of Agamura persica morphologically. We found that A. persica shows geographic variation with respect to two groups, with the eastern population considered as A. cruralis and A. persica proposed for the western population. Separation between the two populations of A. persica was verified based upon ANOVA results for many morphological characters, including Head Height (HH), Interorbital distance (IO), Forelimb length (FLL), Number of scales across widest part of abdomen (NSA), Loreal scales (LOS), and Number of scales that separate two adjacent tubercles (NTV). Principal Component Analysis (PCA) and Discriminant Function Analysis (DFA) based on metric and meristic characters confirmed the revision of the genus Agamura. Multivariate analysis indicated that all studied OTUs were assigned to the correct classification and have significantly different morphological characters.

Keywords: Agamura cruralis, cryptic divergence, Cyrtopodion, Iranian Plateau, Rhinogecko.

Introduction and Cyrtopodion hormozganum (Nazarov, Bon- darenko and Rajabizadeh, 2012). In these stud- The Iranian Plateau is one of the richest ar- ies, several characters were given that distin- eas for in southwestern Asia (Rastegar- guish these species from the genus Agamura, Pouyani et al., 2015). Many species groups including the presence of a keel on the dorsal and genera have diversified in this region, tubercles, the presence of dark spots on the dor- including the genera Tropiocolotes and Cyr- sum, the number of scales across the venter, and topodion (Bauer et al., 2013; Krause et al., others. 2013; Freitas et al., 2016). Diversification of The genus Agamura is one of the gekkonid Gekkonids has been studied extensively and genera occurring in Western Asia (on the Ira- several complexes have been identified within nian Plateau). The genus was regarded at one them. Recently, several species were described time as having four species: A. persica, A. agamuroides Cyrtopodion from the group as gastropholis, A. femoralis, and A. misonnei kiabi Cyrtopodion (Ahmadzadeh et al., 2011), (Szczerbak and Golubev, 1996), but Anderson golubevi Cyrtopodion persepolense , (Nazarov, (1999) retained only the type species, Aga- Ananjeva and Rajabizadeh, 2009), Cyrtopodion mura persica. Anderson (1999) mentioned that sistanense (Nazarov and Rajabizadeh, 2007), Rhinogecko differs from Agamura in several characteristics such as mental shields and nasal scales that are sharply swollen or tube-like. 1 - Department of Biology, Faculty of Sciences, Ferdowsi University of , Mashhad, Iran Additionally, Anderson (1999) placed A. gas- 2 - Research Department of Zoological Innovations tropholis in the genus Cyrtopodion, with which (RDZI), Institute of Applied Zoology, Faculty of it shares a pigmented peritoneum. According to Sciences, Ferdowsi University of Mashhad, Mashhad, Macey et al. (2000), the genus Cyrtopodion is a Iran 3 - Department of Biology, Faculty of Science, Hakim monophyletic group. Cervenka, Kratochvil, and Sabzevari University, Sabzevar, Iran Frynta (2008) later confirmed the monophyly ∗Corresponding author; e-mail: [email protected] of Cyrtopodion. Anderson (1999) indicated that Downloaded from Brill.com10/04/2021 09:37:33AM via free access © Koninklijke Brill NV, Leiden, 2017. DOI:10.1163/15685381-00003129 450 S.S. Hosseinian Yousefkhani et al. different populations of A. persica exhibit dif- sized gravel (pers. observation). Endemicity in ferentiation in color pattern. He divided them angular-toed on the Iranian plateau sug- into two subspecies differentiated by patterns gests that there are many isolated populations of transverse bars: A. persica persica (the west- related to this group, but a comprehensive in- ern population in central and eastern Iran) with vestigation is needed. three transverse bars and A. persica cruralis In the present study, we aimed to examine (the eastern population limited to southeast Iran, morphological characteristics of all populations Afghanistan and Pakistan) with five transverse of A. persica from the Iranian Plateau. The main bars. Smith (1935) had earlier synonymized A. goal of the study was to test the validity of the persica and A. cruralis, but found some varia- two OTUs of A. persica in the Iranian Plateau. tion among them. For example, Smith (1935) re- ported the presence of large postmental scales in Material and methods Baluchistan specimens that suggested the prob- able presence of A. p. cruralis. Szczerbak and We examined 30 specimens representing A. persica (Ap- pendix 1). These specimens were collected from May 2011 Golubev (1996) indicated that these variations to July 2016 on the Iranian Plateau (fig. 1; exact locations need more consideration, but could not make are provided in Appendix 1). All specimens were preserved in 75% ethanol and deposited in the Sabzevar University the comparison themselves due to lack of suf- Herpetological Collection (SUHC). Twelve mensural and ficient materials. 15 meristic characters (table 1) were recorded from these In respect to habitat, the region inhabited by specimens. Mensural characters were measured using dig- ital calipers with 0.01 mm accuracy and meristic charac- A. persica is different in central and eastern Iran. ters were measured using an Olympus loupe. Because we A. p. cruralis can be found in very rocky ar- combined all samples to examine geographic variation, we removed sexually dimorphic characters from the analyses. eas in eastern Iran, but the habitat of A. p. per- Based upon t-test analysis of all characters, only preanal sica in is characterized by small- pores were distinguished as a sexually dimorphic character.

Figure 1. Map of Iran and sampling localities of all examined species in this study. Downloaded from Brill.com10/04/2021 09:37:33AM via free access Morphology of the genus Agamura 451

Table 1. All metric and meristic characters examined in this study, including their definitions.

Character Definition

SVL Snout-Vent Length HL Head length (from tympanum to tip of rostrum) HW Head width (maximum distance between tympani) HH Head height (straight-line dorsiventral distance between apex of head and ventral surface of lower jaw) SL Snout length (eyeÐnostril distance) IO Interorbital distance EED Distance from eye to tympanum WPW Widest lateral dimension of abdomen FHD Distance between forelimb and hindlimb FLL Forelimb length HLL Hind limb length NSS Number of supralabial scales NIS Number of infralabial scales ION Number of interorbital scales (except palpebral fold scales) LOS Loreal scales (between eye and nostril, in a single longitudinal row) POS Number of scales from postnasal to occiput (in a single longitudinal row) NPS Number of postmental scales NSA Number of scales across widest part of abdomen NPV Number of scales between postmental scales and vent NTW Number of dorsal tubercles at widest lateral dimension of thorax NTP Number of tubercles in a paravertebral row (from neck to hindlimb) NAT Number of scales around a tubercle (forming a circumference) NTV Number of scales that separate two adjacent tubercles (longitudinal) NTH Number of scales that separate two adjacent tubercles NPP Number of preanal pores SLT Subdigital lamella under fourth toe NUT Number of scales under the tail

Statistical analyses were performed using SPSS 16.0 Results and all data were evaluated for normality prior to anal- ysis. Mensural (parametric) and meristic (non-parametric) Color and pattern characters were evaluated using the Shapiro-Wilks and the Kolmogorov-Smirnov tests, respectively. H0 in both tests Based on data from all examined specimens, is a normal distribution, so the dataset is normally dis- color pattern within A. persica is variable and tributed if the values are not significantly different from H 0 may be dependent upon ground color (pers. (P>0.05). Intraspecific analyses of A. persica were per- formed according to two major operational taxonomic units obs). In the western individuals, the entire body (OTUs). We selected these OTUs according to geographic color is light. However, the dorsal color is dark barriers. in the eastern individuals. The number of dorsal After testing for normality, a one-way analyses of vari- ance (ANOVA) was used to examine the characters between crossbars was five in all examined specimens populations of A. persica to verify the morphological diver- of A. persica from throughout the range and all gence between them and to determine whether they fit An- specimens had tails with five to eight black bars derson’s (1999) taxonomic interpretation. The null hypoth- (fig. 2a, b). esis in an ANOVA is that all data are distributed uniformly, so a significant result (P<0.05) indicates that charac- ters are not distributed equally and are therefore differenti- Scalation ated between groups. Principal Component Analysis (PCA) and Canonical Variate Analysis (CVA) were conducted on Differences between populations of A. persica the significant characters identified using the ANOVA. The on the Iranian plateau were identified for some PCA was used to determine the variation between popula- types of scales. Postmental scales are larger tions based on significant characters. The CVA was used to determine the correct classification based on the population in the eastern OTU than in the western one grouping. (fig. 3a, b). There is a notable keel on the Downloaded from Brill.com10/04/2021 09:37:33AM via free access 452 S.S. Hosseinian Yousefkhani et al.

Figure 2. Color pattern of dorsal and tail in Agamura persica (eastern clade (A); western clade (B)).

Figure 3. Mental and postmental scales in Agamura persica (eastern clade A; western clade B). dorsal tubercles in the eastern form, but the Normality test tubercles in the western group are smooth. The rostral scale was completely divided in both All characters were examined for normality. eastern and western OTUs and was not related All of characters had distributions that did not to geographic distribution. The rostral scale is differ significantly from a normal distribution. not divided in the western OTU. Two main groupings were performed on the Downloaded from Brill.com10/04/2021 09:37:33AM via free access Morphology of the genus Agamura 453

Table 2. Mean, standard deviation, and range of 25 metric and meristic characters measured in specimens of Agamura persica for two OTUs across the entire distribution range on the Iranian Plateau.

Character Eastern population (n = 16) Western population (n = 14) P value Mean ± SD Range Mean ± SD Range

SVL 60.93 ± 9.42 46.49-74.79 57.60 ± 7.09 46.72-69.37 0.289 HL 18.23 ± 2.70 14.36-22.83 17.01 ± 1.53 14.45-20.23 0.148 HW 14.51 ± 3.00 10.75-21.56 12.92 ± 1.32 11.22-15.19 0.078 HH 10.63 ± 2.70 7.25-17.97 8.44 ± 0.95 7.20-10.12 0.008 SL 4.68 ± 0.76 3.34-6.14 4.80 ± 0.57 3.81-5.77 0.647 IO 10.10 ± 1.78 6.60-13.63 8.79 ± 0.98 6.75-10.04 0.022 EED 4.74 ± 0.86 2.94-6.05 4.72 ± 0.69 3.83-6.08 0.969 WPW 14.32 ± 3.61 8.49-19.25 13.23 ± 2.88 9.59-18.43 0.373 FHD 26.02 ± 4.71 16.83-33.25 23.98 ± 4.19 16.22-30.61 0.224 FLL 32.20 ± 4.12 25.43-41.09 29.33 ± 3.44 21.55-34.10 0.050 HLL 43.57 ± 6.93 30.03-56.67 40.35 ± 4.01 33.72-47.51 0.138 NSS 13.75 ± 1.43 11.00-16.00 13.57 ± 0.93 12.00-15.00 0.695 NIS 10.81 ± 1.04 8.00-12.00 11.35 ± 0.63 11.00-13.00 0.102 ION 26.43 ± 2.92 21.00-32.00 26.78 ± 2.91 22.00-31.00 0.747 LOS 13.31 ± 1.25 11.00-16.00 12.07 ± 1.59 10.00-15.00 0.024 NSA 29.68 ± 5.26 21.00-40.00 37.35 ± 4.65 31.00-47.00 0.000 NPV 51.62 ± 4.12 46.00-59.00 53.85 ± 3.52 49.00-59.00 0.125 NTW 8.75 ± 1.39 6.00-11.00 9.28 ± 1.06 7.00-11.00 0.252 NTP 25.12 ± 2.33 21.00-29.00 23.78 ± 2.32 18.00-28.00 0.128 NAT 9.50 ± 0.63 8.00-10.00 9.42 ± 0.85 8.00-11.00 0.795 NTV 3.25 ± 0.57 2.00-4.00 2.71 ± 0.82 2.00-4.00 0.047 NTH 3.00 ± 0.73 2.00-4.00 3.07 ± 0.82 2.00-5.00 0.804 SLT 18.81 ± 1.37 16.00-21.00 18.28 ± 1.38 16.00-21.00 0.306 NUT 46.70 ± 5.90 32.00-52.00 46.60 ± 3.57 45.00-53.00 0.973 TL 53.10 ± 6.52 37.38-60.69 49.43 ± 4.06 43.48-54.74 0.274 samples: for intraspecific variation, A. persica Table 3. Factor loading of the first three principal compo- nents (PCs) from a correlation matrix of six characters for were defined into two populations (western and the two populations of Agamura persica. eastern). Character PC1 PC2 PC3

HH 0.802 −0.410 0.017 Statistical analyses IO 0.957 0.114 0.022 FLL 0.851 0.376 −0.228 Descriptive statistics of characters done for each NSA 0.109 0.817 0.558 OTU and are presented in table 2. According to NTV 0.226 −0.835 0.438 − the descriptive statistics and an ANOVA, there LOS 0.324 0.658 0.513 Eigenvalue 2.347 1.687 0.556 are significant differences between the eastern Accumulated percent 46.936 80.677 91.978 and western A. persica populations in HH, IO, of trace FLL, LOS, NSA and NTV (P<0.05) (table 2). Multivariate analyses were done using two popular approaches: Principal Component have the most weight; 33.74% is explained by Analysis (PCA) and Canonical Variate Analy- PC2, which is mainly attributed to NTV and sis (CVA). These techniques were employed to NSA; and 11.12% is explained by PC3, which is explain the variation between two populations mainly attributed to NSA and LOS (table 3 and of A. persica. In Principal Component Analy- fig. 4). In the next step, discriminant analyses sis, the first three components explain 91.97% were done to provide more accurate discrimi- of the total variation. Of this value, 46.93% nation between populations (within A. persica). is explained by PC1, in which IO and FLL CVA between two populations of A. persica is Downloaded from Brill.com10/04/2021 09:37:33AM via free access 454 S.S. Hosseinian Yousefkhani et al.

Figure 4. PCA plot of intra- and inter-specific variation within the Agamura persica in the Iranian Plateau. highly significant and the first component ex- (such as A. persica), regions with mod- plained 100% of all variance. erate temperature and moisture in submontane areas are the best habitats and areas of great- est population density (Hosseinian Yousefkhani Discussion et al., 2017). Anderson (1999) considered the Iran is a mountainous country in southwest- genus Agamura a monotypic genus. Accord- ern Asia that has a rich herpetofauna (Fath- ingly, A misonnei and A. femoralis were in- nia et al., 2010). Several mountain ranges in cluded within the genus Rhinogecko (R. mison- Iran have played a crucial role in the diver- nei) by having 4 preanal pores, two postmentals sification of reptiles. Additionally, the various in contact, and a row of large scales under the fe- types of habitats in Iran reveal substantial ra- mur. Agamura gastropholis was included in the diation among reptiles, represented by many genus Cyrtopodion on the basis of having a low species complexes in lizards (Nazarov, Anan- number of ventral scales. jeva and Rajabizadeh, 2009; Rastegar-Pouyani Our ANOVA results indicated that six mor- et al., 2010). Speciation was promoted by the phological characters (HH, IO, FLL, LOS, NSA uplift of the Zagros Mountains in western Iran, but the Central Plateau of Iran has different and NTV; four metric and two mensural) were ecological conditions. Ecological factors such significantly different between two A. persica as temperature and moisture are two impor- OTUs; these OTUs also differ in dorsal color tant variables affecting population isolation in and size of the postmental (color in the eastern the central deserts of Iran. For ground dwelling population is darker and postmental scales are Downloaded from Brill.com10/04/2021 09:37:33AM via free access Morphology of the genus Agamura 455 larger than in the western population; addition- populations is distinctive. Accordingly, A. cru- ally, dorsal tubercles in eastern population show ralis can be considered as the eastern popula- a keel-like crest). The presence of A. cruralis in tion in Baluchistan and A. persica as the western the eastern part of the distribution is confirmed population from Central Iran (fig. 1) as distinct (but not according to (Szczerbak and Golubev, species. Because the first specimens of A. cru- 1996)); head height is larger and the number ralis were collected from Bahu Kalat, Baluchis- of ventral scales is fewer in the eastern popula- tan (Blanford, 1874), we considered this as the tion than in the western. Szczerbak and Golubev type locality of A. cruralis. We conclude that the (1996) had a poor sample from the central part decision of Anderson (1999) on the previously of the Iranian Plateau and they evaluated A. per- known Agamura is acceptable and that A. cru- sica according to the specimens from Khorasan. ralis is present in southeastern Iran. Adding more sampling locations from Central Iran, especially from , and Taxonomic implications provinces, can provide insights on geographic Given the morphological differences between variation. Based upon this evidence, we propose the two putative species of the genus Agamura that A. cruralis (the eastern OTU) can be vali- and in concordance with the history of their de- dated as a full species. scription, we assign the species level to both This study confirmed the geographic vari- them (Anderson, 1999). As the result, we sug- ation of A. persica and its monotypic status gest the following nomenclatural and taxonomic cannot be accepted. Dumeril (1856) mentioned actions: we resurrect the species A. cruralis the two species of the genus Agamura from Blanford, 1874 from the synonymy of A. per- Iran as A. persica and A. cruralis based on sica and apply it to the species that occurs in the finding that the size of different charac- SE Iran and southern Pakistan. Some morpho- ters is smaller in A. persica. Blanford (1874) logical characters were measured on all speci- collected several specimens from Baluchistan, mens and are presented in Appendixes 2 and 3. southeast Iran and used different morpholo- gical characters to introduce two species, as did Dumeril (1856). These species were as Key to the genus Agamura Blanford, 1874 A. cruralis (syntypes: BMNH74.11.23.52-54 (British Museum of Natural History), type lo- 1a. Dorsal tubercles sharply keeled; large tu- cality Bahou Kalat, Baluchistan, and A. per- bercles presented top of the thigh; long sica (Syntype: MNHN 6761(Muséum national forelimb; number of ventral scales lower d’histoire naturelle, Paris), type locality from than 32 a cross midbody; head large; Persia. Szczerbak and Golubev (1996) revised number of scales between tubercles 4; the species using more specimens from cen- color dark brown . . . . . Agamura cruralis tral and southeast Iran. Szczerbak and Gol- 1b. Dorsal tubercles smooth; small tubercles ubev (1996) considered the Baluchistan popula- presented top of the thigh; short fore- tion as a subspecies, A. p. cruralis, with differ- limb; number of ventral scales higher ent metric and mensural characters. Anderson than 32; head small; number of scales be- (1999) reconsidered the previous classifications tween tubercles 3; color light brown . . . . and only confirmed the Szczerbak and Golubev ...... Agamura persica (1996) classification, proposing that the genus Agamura is monotypic. However, we examined 30 specimens from throughout the range of dis- Acknowledgements. We thank all of the friends who helped us collect samples from the Iranian Plateau. These tribution in Iran and found that morphological include Hossein Nabizadeh, Iman Ronaghi, Morteza Moa- differentiation between two eastern and western dab, Aghil Keyvanloo, Hamze Oraei, and Azar Khosravani. Downloaded from Brill.com10/04/2021 09:37:33AM via free access 456 S.S. Hosseinian Yousefkhani et al.

We also thank Ann Paterson, Aaron Bauer, and Steven C. Macey, J.R., Ananjeva, N.B., Wang, Y., Papenfuss, Anderson, who edited and commented on an early draft T.J. (2000): Phylogenetic relationships among Asian of the manuscript. This research was funded by a grant gekkonid lizards formerly of the genus Cyrtodactylus (no. 3/39887) from Ferdowsi University of Mashhad. We based on cladistic analyses of allozymic data: mono- also thank from Department of Environment of Iran that phyly of Cyrtopodion and Mediodactylus. J. Herpetol. permitted us to visit all habitats of the species in Iran. 34: 258-265. Nazarov, R., Ananjeva, N., Radjabizadeh, M. (2009): Two new species of angular-toed geckoes (: References ) from South Iran. Russ. J. Herpetol. 16: Ahmadzadeh, F., Flecks, M., Torki, F., Boehme, W. (2011): 311-324. A new species of angular-toed , genus Cyr- Nazarov, R.A., Bondarenko, D.A., Radjabizadeh, M. topodion (Squamata: Gekkonidae), from southern Iran. (2012): A new species of thin-toed geckos Cyrtopodion Zootaxa 2924: 22-32. sensu lato (Squamata: Sauria: Gekkonidae) from Hor- Anderson, S. (1999): The Lizards of Iran. Contributions to mozgan province, south Iran. Russ. J. Herpetol. 19: 292- Herpetology, 15. Society for the Study of Amphibians 298. and Reptiles, Oxford Press, California. Nazarov, R.A., Rajabizadeh, M. (2007): A new species of Bauer, A.M., Masroor, R., Titus-Mcquillan, J., Heinicke, angular-toed gecko of the genus Cyrtopodion (Squa- M.P., Daza, J.D., Jackman, T.R. (2013): A preliminary mata: Sauria: Gekkonidae) from south-east Iran (Sistan- phylogeny of the Palearctic naked-toed geckos (Reptilia: Squamata: Gekkonidae) with taxonomic implications. Baluchistan province). Russ. J. Herpetol. 14: 137-144. Zootaxa 3599: 301-324. Rastegar-Pouyani, E., Rastegar- Pouyani, N., Kazemi Cervenka, J., Kratochvil, L., Frynta, D. (2008): Phylogeny Noureini, S., Joger, U., Wink, M. (2010): Molecular and of the Middle Eastern geckos of the genus phylogeny of the Eremias persica complex of the Ira- Cyrtopodion and their selected relatives. Zootaxa 1931: nian plateau (Reptilia: Lacertidae), based on mtDNA se- 25-36. quences. Zool. J. Linn. Soc. 158: 641-660. Duméril, A.H.A. (1856): Description des reptiles nouveaux Rastegar-Pouyani, N., Gholamifard, A., Karamiani, R., ou imparfaitement connus de la collection du Muséum d’Histoire Naturelle et remarques sur la classification et Bahmani, Z., Mobaraki, A., Abtin, E., Faizi, H., Heidari, les charactères des reptiles. Archives du Muséum Hist. N., Takesh, M., Sayyadi, F., Ahsani, N., Browne, R.K. Nat. Paris 8: 438-588. (2015): Sustainable management of the herpetofauna of Freitas, S., Rocha, S., Campos, J., Ahmadzadeh, F., Corti, the Iranian Plateau and coastal Iran. Amph. Rep. Con- C., Sillero, N., Ilgaz, Ç., Kumluta¸s, Y., Arakelyan, M., serv. 9: 1-15. Harris, D.J. (2016): Parthenogenesis through the ice Smith, M.A. (1935): The Fauna of British India, Including ages: a biogeographic analysis of Caucasian rock lizards Ceylon and Burma. (genus Darevskia). Mol. Phylo. Evol. 102: 117-127. Szczerbak, N., Golubev, M. (1996): Gecko Fauna of the Hosseinian Yousefkhani, S.S., Aliabadian, M., Rastegar- Pouyani, E., Darvish, J. (2017): Predicting the impact of USSR and Contiguous Regions, Contr. to Herptol. 13. change on the distribution pattern of Agamura Soc. Study of Amph. and Rept., Oxford, Ohio. persica (Dumeril, 1856) (Squamata: Gekkonidae) in Iran. Belg. J. Zool., in press. Krause, V., Ahmadzadeh, F., Moazeni, M., Wagner, P., Submitted: December 15, 2016. Final revision received: Wilms, T.M. (2013): A new species of the genus Tro- piocolotes Peters, 1880 from western Iran (Squamata: September 20, 2017. Accepted: September 26, 2017. Sauria: Gekkonidae). Zootaxa 3716: 22-38. Associate Editor: Miguel Carretero.

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Appendix 1. Museum number, species name, and locality of each specimen.

Museum number Species name Locality

SUHC 6135 Agamura persica Iran- province-Kerman SUHC 6139 Agamura persica Iran--Faryab SUHC 6137 Agamura persica Iran-Kerman province-GowdGhool-Sirjan SUHC 6133 Agamura persica Iran-Kerman province-Rabor SUHC 6138 Agamura persica Iran-Kerman province-Bardsir SUHC 6136 Agamura persica Iran-Kerman province-GowdGhool-Sirjan SUHC 6134 Agamura persica Iran-Kerman province-Kerman SUHC 6142 Agamura cruralis Pakistan-Nushki SUHC 6141 Agamura cruralis Pakistan-Nushki SUHC 6140 Agamura cruralis Pakistan-Kharan SUHC 6020 Agamura persica Iran--Nehbandan SUHC 6050 Agamura persica Iran-South Khorasan province-Nehbandan SUHC 4984 Agamura persica Iran--Bayazeh SUHC 5947 Agamura cruralis Iran-Sistan and Balushistanprovinc- SUHC 6065 Agamura persica Iran-Qom province-Qom SUHC 6063 Agamura persica Iran-Qom province-Qom SUHC 6064 Agamura persica Iran-Qom province-Qom SUHC 5948 Agamura cruralis Iran-Sistan and Balushistanprovinc-Zahedan SUHC 6126 Agamura persica Iran--30 km N of MorcheKhort SUHC 4973 Agamura persica Iran-Yazd province-Near ChakChak SUHC 312 Agamura persica Iran-Qom province-KamarKooh SUHC 1589 Agamura persica Iran--Kavir National Park SUHC 1588 Agamura persica Iran-Semnan province-Kavir National Park SUHC 574 Agamura persica Iran-Qom province-Qom-EmamzadehAbdollah SUHC 475 Agamura persica Iran-Qom province-Qom SUHC 573 Agamura persica Iran-Qom province-Qom SUHC 569 Agamura persica Iran-Qom province-Qom SUHC 570 Agamura persica Iran-Qom province-Qom SUHC 1654 Agamura persica Iran--Noghab village SUHC 1697 Agamura persica Iran-Razavi Khorasan province-Noghab village

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Appendix 2. All measurements of Agamura cruralis specimens from southeast Iran (metric characters were measured in mm).

Character SUHC 6142 SUHC 6141 SUHC 6140 SUHC 5947 SUHC 5948

SVL 69.04 55.871.05 65.39 63.51 HL 22.83 16.73 19.47 18.54 19.64 HW 21.56 14.19 16.75 13.94 13.57 HH 17.97 11.54 12.91 11.56 9.31 SL 5.13 4.49 5.58 5.29 4.69 IO 10.22 9.811.48.81 10.35 EED 6.05 4.47 5.21 5.93 4.05 WPW 17.66 14.15 17.58 19.25 12.27 FHD 28.29 22.94 28.71 30.87 27.79 FLL 35.232.51 32.87 33.57 35.35 HLL 49.44 43.71 46.74 46.63 48.31 NSS1412141214 NIS1110111112 ION2927212524 LOS1612151411 NNS33333 NSA2830253027 NPV5346535955 NTW1010798 NTP2925232424 NAT 10 9 8 10 10 NTV33334 NTH34334 NPP00002 SLT1918182120 NUT 45 32 49 52 51 SexffffM TL 56.74 37.38 60.69 55.77 58.88

Downloaded from Brill.com10/04/2021 09:37:33AM via free access Morphology of the genus Agamura 459 47 13 11 29 10 3 40 50 10 24 9 2 3 0 16 Ð F Ð 74 16 12 31 13 3 26 54 8 25 10 3 3 0 18 Ð f Ð 19 1292 11 15 13 31 28 14 13 3 3 36 35 59 56 10 10 23 25 8 10 2 3 3 2 0 0 17 19 53 Ð F 54.74 F Ð 36 12 10 23 13 3 26 50 9 27 10 4 4 0 18 Ð f Ð 53 14 10 27 14 3 37 55 11 29 10 2 2 0 20 51 f 53.9 91 14 11 24 11 3 35 50 11 24 9 2 3 2 19 Ð M Ð 03 11 8 25 13 3 25 49 10 21 9 4 3 0 19 Ð f Ð 55 15 11 25 14 3 42 53 9 24 11 3 3 2 19 45 M 49.21 61 14 12 31 12 3 42 58 9 23 9 2 3 2 21 Ð m Ð 67 15 11 32 13 3 21 46 8 25 9 3 2 2 20 45 m 53.02 69 14 12 25 15 3 34 58 8 22 9 2 4 2 19 Ð M Ð 52 14 11 27 13 3 32 55 10 26 10 4 3 0 21 46 f 51.99 72 14 11 22 10 3 32 49 10 23 10 3 2 0 18 Ð F Ð 71 13 11 24 12 3 47 50 7 25 9 2 3 2 17 45 M 48.82 1 13 12 30 13 3 33 55 9 27 10 4 3 0 18 45 F 50.91 57 14 11 24 12 3 26 50 9 26 9 3 2 0 19 Ð f Ð 51 14 11 29 12 3 35 52 8 23 10 3 3 0 19 Ð F Ð 85 14 10 28 13 3 36 57 8 27 10 4 4 2 17 Ð m Ð 18 13 11 29 15 3 36 46 10 21 10 3 3 0 16 Ð f Ð 89 15 12 26 13 3 40 51 6 26 9 3 3 2 19 51 m 48.45 38 13 11 27 10 3 40 57 10 18 10 4 5 2 20 45 M 43.48 88 14 11 24 12 3 41 56 9 24 8 2 4 0 17 Ð F Ð 07 16 12 25 13 3 30 47 7 24 9 3 2 2 18 45 m 54.26 38 12 11 26 11 3 31 51 10 28 10 4 2 0 17 Ð F Ð ...... 24 40 06 45 24 41 57 44 79 49 04 35 43 30 79 40 68 42 09 56 639 82 47 55 33 23 39 41 40 77 35 147 535 18 36 33 36 64 37 49 36 12 45 73 48 98 39 ...... 46 31 61 34 51 28 39 33 25 32 22 25 125 39 29 54 30 62 41 17 30 81 36 721 37 31 05 30 09 26 19 34 83 29 81 27 53 26 26 27 01 27 26 32 15 33 25 28 ...... 89 30 923 724 59 16 83 23 428 77 25 64 22 96 30 92 22 87 31 55 33 45 20 16 19 43 29 49 19 04 16 71 22 229 77 20 61 22 15 22 77 26 94 26 83 27 ...... 78 13 811 86 12 99 9 45 12 29 14 57 18 89 11 99 11 08 15 17 9 71 18 16 11 83 11 318 94 8 52 9 25 15 95 15 56 10 91 8 08 17 62 14 61 13 02 16 ...... specimens that were collected from central Iran (metric characters were measured in mm). 57 4 75 3 59 4 83 22 5 91 4 66 3 63 5 67 4 45 85 5 27 4 58 3 63 5 13 2 44 4 91 4 98 5 21 4 04 6 49 4 54 4 63 64 76 5 ...... 23 8 81 6 34 8 49 10 65 11 62 9 47 8 11 13 65 8 31 11 14 9 52 7 51 8 89 9 58 9 74 7 33 9 77 9 37 8 37 10 14 8 06 12 04 6 66 9 21 8 ...... 89 5 37 3 74 4 79 5 38 4 51 4 33 5 25 4 85 4 89 6 93 4 75 4 96 4 83 3 95 4 23 4 93 5 12 5 95 4 24 4 24 71 4 56 5 25 3 47 5 ...... Agamura persica 03 8 32 7 81 8 96 12 09 8 92 8 811 15 7 73 9 92 12 22 7 02 7 79 94 8 14 7 74 9 19 8 04 10 97 51 10 97 75 7 76 8 95 7 79 ...... 65 14 45 11 76 17 58 16 58 13 91 11 65 15 69 12 97 13 54 16 54 11 81 12 04 13 48 10 65 11 46 12 23 15 48 15 311 75 14 43 11 27 10 62 13 36 10 41 13 ...... 87 17 72 14 39 22 73 19 02 16 88 20 14 16 97 17 79 19 61 16 36 15 56 16 49 14 69 15 92 15 55 20 37 17 06 15 66 18 44 16 53 15 37 19 07 14 56 17 81 17 ...... All measurements of SUHC 574 61 SUHC 570 46 SUHC 6134 56 SUHC 6020 74 SUHC 4984 57 Appendix 3. Character SVLSUHC 6135 67 HL HW HH SL IO EED WPW FHD FLL HLL NSS NIS ION LOS NNS NSA NPV NTW NTP NAT NTV NTH NPP SLT NUT Sex TL SUHC 475 56 SUHC 6136 63 SUHC 6050 74 SUHC 4973 53 SUHC 6065 63 SUHC 6139 46 SUHC 1654 50 GWD1 48 SUHC 6137 50 SUHC 573 65 SUHC 1589 65 SUHC 6063 51 SUHC 6133 63 SUHC 569 49 SUHC 1697 51 SUHC 1588 69 SUHC 6138 49 SUHC 6064 60 SUHC 6126 57 Downloaded from Brill.com10/04/2021 09:37:33AM via free access