Acta Herpetologica 9(2): 187-202, 2014 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-14384 Geographic variation in the morphology of Macrovipera lebetina (Linnaeus, 1758) (Ophidia: Viperidae) in Iran Naeim Moradi1,2,*, Nasrullah Rastegar-Pouyani1,2, Eskandar Rastegar-Pouyani1,3 1 Iranian Plateau Herpetology Research Group (IPHRG), Faculty of Sciences, Razi University, 6714967346 Kermanshah, Iran. *Corre- sponding author. E-mail: naeim.moradi@yahoo.com 2 Department of Biology, Faculty of Sciences, Razi University, 6714967346 Kermanshah, Iran 3 Department of Biology, Faculty of Sciences, Hakim Sabzevari University, Sabzevar, Iran Submitted on 2014, 14th April; revised on 2014, 10th September; accepted on 2014, 10th September Editor: Marco A.L. Zuffi Abstract. The Levantine viper, Macrovipera lebetina, has an extensive geographical range being distributed in central Asia and the Middle East. The species exhibits high levels of polymorphism, especially in colouration and pattern. Recent studies revealed significant morphological differences between the two subspecies from northeastern and west- ern portions of Iran. However, considering limited geographic samplings, taxonomic status of Iranian Macrovipera are controversial. In this study, uni- and multivariate statistical techniques were used to analyze geographic variation in 31 morphological characters measured in 117 specimens of Macrovipera lebetina covering its entire range in Iran. Sexual dimorphism was obvious in number of scales across the head and subcaudals. Univariate analyses detected substantial geographic variation in several meristic characters. Pholidosis exhibits a general north-south pattern of variation and most scale counts averaged higher in southern regions. Colouration displayed a pattern of strong clinal variation among three broad areas consisting of the combined western and northwestern, northeastern, and southern highland regions. Also, morphometric characters exhibited a general north-south pattern of geographic variation and some characters averaged lower in southern regions. Populations from the southern regions remained clearly distinct in Principal Component, Cluster and Discriminant analyses. In the light of these differences, it is concluded that the southern Iranian populations should not be identified as belonging to Macrovipera lebetina obtusa (Dwigubsky, 1832), which occurs in northwestern and western regions of the Iranian Plateau. Keywords. Viperidae, Macrovipera lebetina, geographic variation, morphology, statistical analyses, Iranian Plateau. INTRODUCTION The genus Macrovipera has been described by Reuss in 1927, and then resurrected by Herrmann et al. (1992). According to Lenk et al. (2001), just two species are known: Macrovipera schweizeri (Werner, 1935), which is endemic to a few islands in the western Cyclades (Nil- son and Andrén, 1988) and Macrovipera lebetina (Lin- naeus, 1758). The Levantine viper, Macrovipera lebetina is distributed from Central Asia to the Middle East (Nil- son and Andrén, 1988; Nilson et al., 1988; Böhme and Wiedl, 1994; Göçmen et al., 1996; David et al., 1999; Atatür and Göçmen, 2001; Ananjeva et al., 2006; Göç- men et al., 2006). Traditionally, up to six distinct sub- species were recognized of M. lebetina: M. l. chernovi (Chikin and Szczerbak, 1992), M. l. lebetina (Linnaeus, 1758), M. l. obtusa (Dwigubsky, 1832), M. l. peilei (Mur- ray, 1892), M. l. transmediterranea (Nilson and Andrén, 1988) and M. l. turanica (Terentiev and Chernov, 1940). Recently, Stümpel and Joger (2009) and Stümpel (2012) in their molecular analysis showed that haplotypes of Macrovipera lebetina are subdivided into five major line- ages which support the validity of the allopatric subspe- cies lebetina, euphratica, obtusa, turanica and chernovi 188 N. Moradi et alii with no samples available from peilei (southern Afghani- stan and Pakistan) and transmediterranea (north Africa). One more lebetina-like taxon from south-central Iran (Jiroft/Baft Mts.) is genetically very distinct from all other Macrovipera lebetina taxa (Stümpel, 2012). Macrovipera lebetina obtusa (Dwigubsky, 1832) has a southernmost range from Turkey south to N Jordan, eastwards through the Caucasus to Kashmir and N India (Sindaco et al., 2013). Macrovipera lebetina chernovi (Chikin and Szczer- bak, 1992) occurs in north-eastern Iran, south Turkmeni- stan, and parts of northern Afghanistan, Tadjikistan, and northern Pakistan (Phelps, 2010; Stümpel, 2012) and is replaced by Macrovipera lebetina turanica in central Asia. The body colour of blunt-nosed vipers can be virtu- ally any shade of brown, even pinkish, dark or light grey. Specimens from Cyprus tend to be paler. The blunt- nosed viper can possess faint or bold markings and many seem uniform in colour. Markings, however distinct, take the form of cross bands, dorsal blotches, and vertical lat- eral bars of a darker hue than the ground colour. Some handsome specimens have a more contrasting colour pattern (Mermer et al., 2012). The ventral surface is usu- ally a paler version of the ground colour with or without darker stippling (Phelps, 2010). In recent years, Iranian researchers such as Afroosheh and Kazemi (2011), Rajabizadeh et al. (2011) and Oraie et al. (2012) have done valuable surveys on the taxonomic status of Macrovipera lebetina in Iran. All of the previous researches have focused on the western and northeastern populations of M. lebetina, whereas south- ern populations were not investigated. In order to clarify their taxonomic status, in this paper geographic variation in morphology of all populations of Macrovipera lebetina across the whole distribution range in Iran is discussed. MATERIALS AND METHODS Specimens and sampling In this study, 117 specimens of M. lebetina were exam- ined across the entire range of species distribution in Iran (see Appendix). Of these, 77 mature and intact individuals (29 males and 41 females) were selected for analysis of met- ric and meristic characters used in morphological studies of vipers, and 99 specimens were investigated in non-parametric analysis of colour pattern. Because of the immaturity or lack of revealed colour pattern, 65 specimens among them were select- ed for parametric analysis. Thirty specimens were captured from June 2012 to late October 2013; more than twenty were released subsequent to measuring morphological characters in the field, whereas the remainders according to Pisani (1973) were preserved in ethanol 96% and deposited at the Razi Uni- versity Zoological Museum (RUZM), Museum of Shahid Baho- nar University of Kerman (ZMSBUK), and Sabzevar Univer- sity Herpetological Collection (SUHC). Additional specimens subject of this study come from the herpetological collections (Table 1). Also, six pictures were selected for analyzing the col- our pattern. Characters Characters used in this research are listed in Table 2. All of the specimens were examined for eight metric, seventeen meristic, and six colour pattern characters (Table 2). Metric characters were evaluated with a digital caliper to the nearest 0.01 mm. Because expressing body measurements as a percent- age of Snout-Vent Length (SVL) does neutralize allometry, we calculated the residual values of each body measurements as a function of SVL (Babocsay, 2003). All mensural characters as compared to SVL were analyzed. Table 1. Number of specimens subject of present study come from the herpetological collections and used for meristic, mensural, and color pattern analysis. Type of analysis Herpetological collections ZCRI ZMSBUK SUHC RUZM SUZM FDOI Meristic 43 4 6 4 2 4 Mensural 43 4 6 4 2 4 Colour pattern 55 7 6 6 3 4 Total 61 8 9 8 3 4 ZCRI: Zoological Collection of the Department of Venomous Animals and Antiserum Production, Razi Vaccine and Serum Research Insti- tute, Hesarak, Karaj ZMSBUK: Zoological Museum of Shahid Bahonar University of Kerman SUHC: Sabzevar University Herpetological Collection RUZM: Razi University Zoological Museum SUZM: Shiraz University Zoological Museum FDOI: Species Diversity Museum of Department of Environment, Fars 189Geographic variation in the Iranian Macrovipera lebetina Statistical analyses The Principal Components Analysis (PCA) based on a correlation matrix of meristic data was used to determine whether distinct geographic units exist within Macrovipera leb- etina. Independent multivariate analyses were conducted on combined sexes. In addition, the Cluster observation method was used to explore the population’s groupings, based on significant charac- ters which had r2 > 50%. Also Discriminant Analysis was used for meristic characters, in order to evaluate the actual degree of discrimination among the a priori groups as well as to pre- dict their group membership. Moreover, generalized squared distances between groups were computed to show the distance between each pair of groups. To describe dispersal pattern among morphological char- acters of different localities, descriptive statistical parameters including Minimum, Maximum, Mean and Standard devia- Table 2. Morphological characters used in this study. Abbreviation Characters Meristic characters PreV Number of preventrals + small gular scales along the midline of the ventral side of the head Ven Number of ventral scales Scd Number of subcaudal scales Dors1 Number of dorsal scale rows, counting across the forepart of the body [one head length behind the posterior end of head]1 Dors2 Number of dorsals, counted across mid-body (at mid-SVL length) Dors3 Number of dorsals, counted across the hind part of the body, one head length before the anal plate1 Ap Number of apical scales Blspl Number of scales between last supralabial scales across the head Can Number of canthal scales Interocular Number of scales across head, between the dorsal parts of two eyes Spl* Number of supralabials scales Ifl* Number of sublabial (infralabial) scales inCir* Number of inner circumocular scales outCir* Number of outer circumocular scales (around eye and supraocular scales) Lor* Number of loreal scales Supraoc Number of supraocular scales bESpl Number of scales between eye and supralabials Mensural characters (mm) SVL Snout-vent length TaL Tail length HL Head length (from anterior tip of snout to the angle of jaws) HW Head width HH Head height DbN Distance between nostrils IOD Inter ocular distance SL Snout length (distance between rostrum to the anterior edge of eye) Colour pattern characters SCN The area of the black or dark pigmentation in and around the dorsal blotch, in number of scales. A scale was counted half if it was pigmented over < 50% and one if > 50% of its area (mean of three blotches, halfway between the rostrum and the anus)2 L/WdbR Length to width ratio for a dorsal blotch at mid-SVL. L/DdbR Length of dorsal blotch to distance between two blotches at mid-SVL L/DlbR Length of lateral blotch to distance between two blotches at mid-SVL DbC Dorsal base colour: G) gray; GB) grayish brown; B) brown; R) reddish brown VentP Ventral pattern: D) scattered dots on the entire ventral surface; DD) ventral surface densely dotted; S) scattered dots on the entire ventral surface with a trapezoid splotch * Meristic characters were recorded from both sides and analyzed separately. 1 Campbell et al., 2004. 2 Babocsay, 2003. 190 N. Moradi et alii tion were employed separately for each group. Data normality was checked by drawing residual plots: Residual versus fits and Main effects plot for each character to carrying out the analyses. The existence of sexual dimorphism was checked in all mature specimens using independent sample t-test. Because some data was missing, the General linear model ANOVA was used to explore the significant differences by gender, but also sexes combined. Statistical analyses for all meristic and colour pattern characters were performed using one-way ANOVA followed by Fisher’s LSD method comparison post hoc test to explore the patterns of morphological variation among combined sexes. Also, considering that DbC and VentP are qualitative charac- ters, non-parametric analyses for these characters were per- formed using Kruskal-Wallis test. The significance level for all the statistical tests was set at P < 0.05. Statistical analyses were performed using the MINITAB® (Ver. 16) for the Windows. RESULTS Descriptive statistics Ranges and Mean ± SD for studied characters in each group are provided in Table 3. The specimens were classified according to the different colour patterns and recent studies (Oraie et al., 2012), and the distribution of each pattern in Iran was mapped (Fig. 1). Specimens were grouped in four geographic samples based on geo- graphic units: I. Khorasan Province (Kopet Dagh Mts.); II. Golestan Province (area between Kopet Dagh Mts. and Alborz Mts.); III. Alborz Mts. along with the north- ern part of Zagros Mts.; IV. Southern part of Zagros Mts. (Latitude 33°N) extended south to the Persian Gulf coast and continuing southeast to Sistan Mts. (Longitude 58°E). Results of ANOVA analyses for meristic characters Analysis of meristic data indicated that nine char- acters in males [Ven, AP, Blspl, Spl(L), Spl(R), Ifl(L), Ifl(R), outCir(L) and Supraoc], ten characters in females [Ven, Scd, Spl(L), Ifl(L), Ifl(R), inCir(L), inCir(R), outCir(L), Lor(R) and Supraoc] and thirteen charac- ters when sexes were combined [PreV, Ven, Scd, Can, Interocular, Spl(L), Spl(R), Ifl(L), Ifl(R), inCir(L), outCir(L), Lor(L) and Lor(R)] showed statistically sig- nificant differences (P < 0.05). Results of ANOVA for meristic characters in com- bined sexes show that PreV in group III was significantly lower (P < 0.05) compared to other groups. The num- ber of Ven in group IV are significantly (P < 0.0001) higher than in other groups, whereas Scd in group II are significantly lower compared to other groups. Char- acters of Ifl(L) and Ifl(R) in group IV show significant higher values (P < 0.0001) in comparison with groups II and III, whereas respective values form group IV over- lap with those of group I. Lor(L) in group II were sig- nificantly higher in comparison with group IV. Scale counts of Lor(R) and Spl(L) in groups I and IV are sig- nificantly increased compared to groups II and III and separated from them, whereas, scale counts of Supraoc in these groups (I and IV) was significantly decreased with respect to groups II and III. Also, scale counts of Spl(R) in group IV shows a significant increase (P < 0.05) in comparison with groups II and III and separated from them, whereas this group overlaps with group I. Results of ANOVA analyses for metric characters Results of ANOVA analyses showed that IOD in males, Tal in females, IOD and DbN in combined sexes showed significant differences (P < 0.05). Results for mensural characters in combined sexes indicated that vipers from eastern populations have broader heads than those from western and northwestern populations. Results of ANOVA and non-parametric analyses for colour patterns Analysis of colouration indicates that three char- acters (SCN, L/WdbR and L/DdbR) in combined sexes show statistically significant differences (P < 0.05) among groups. In four groups, results of ANOVA for colour pat- tern show that SCN in group IV has significantly lower number than in group III, but it overlaps with values from groups I and II. Character of L/WdbR in groups I and IV were significantly decreased (P < 0.05) with respect to groups II and III. Character of L/DdbR in groups I and IV were significantly lower (P < 0.05) with respect to group III. Analysis of two non-parametric characters among 99 individuals indicated that both dorsal base colour (DbC) and ventral pattern (VentP) in combined sexes showed powerful significant differences (P < 0.0001) among groups. The median of DbC among four groups was 74.07% R for group I; 50% B for group II; 72% GB for group III and 75.67% G for group IV. Also, the median of VentP among groups was 100% DD for group I; 70% B for group II; 56% D for group III and 86.48% S for group IV. These results indicated that each geographical group have a particular pattern, however, some individuals were diffi- cult to classify with an ambiguous colour patterns such as uniform ground colour without recognizable pattern. According to these results and our observations, three different colour patterns existed among the Iranian populations (Fig. 2). 191Geographic variation in the Iranian Macrovipera lebetina Table 3. Descriptive statistics of meristic, mensural, and colour pattern characters in four groups of Macrovipera lebetina. All Mensural characters as compared to SVL were analyzed. Character Sex I Mean ± SD (n) (range) II Mean ± SD (n) (range) III Mean ± SD (n) (range) IV Mean ± SD (n) (range) PreV M 4.60 ± 0.54 (5) 4-5 4.50 ± 0.57 (4) 4-5 4.00 ± 0.00 (4) 4-4 4.70 ± 0.58 (18) 3-5 F 4.75 ± 0.50 (4) 4-5 4.71 ± 0.48 (7) 4-5 4.18 ± 0.60 (11) 3-5 4.57 ± 0.60 (19) 3-5 Ven M 170.20 ± 1.79 (5) 168-172 167.25 ± 5.12 (4) 162-173 170.50 ± 2.52 (4) 168-174 173.35 ± 2.00 (18) 171-178 F 168.75 ± 2.63 (4) 165-171 167.29 ± 2.69 (7) 164-172 169.09 ± 2.59 (11) 165-173 174.00 ± 2.98 (19) 169-179 Scd M 46.80 ± 3.11 (5) 44-52 44.5 ± 3.11 (4) 42-49 46.00 ± 2.45 (4) 43-49 47.25 ± 3.99 (18) 35-53 F 44.25 ± 4.03 (4) 40-49 39.86 ± 5.46 (7) 29-46 45.60 ± 3.75 (11) 37-49 45.78 ± 3.06 (19) 42-52 Dors 1 M 24.20 ± 1.09 (5) 23-25 25.00 ± 0.00 (4) 25-25 24.50 ± 1.00 (4) 23-25 24.11 ± 1.23 (18) 21-25 F 25.00 ± 0.00 (4) 25-25 24.14 ± 1.06 (7) 23-25 24.45 ± 0.93 (11) 23-25 23.94 ± 1.54 (19) 21-27 Dors 2 M 25.40 ± 0.89 (5) 25-27 25.00 ± 0.00 (4) 25-25 25.00 ± 0.00 (4) 25-25 25.22 ± 0.64 (18) 25-27 F 25.00 ± 0.00 (4) 25-25 25.00 ± 0.00 (7) 25-25 25.18 ± 0.60 (11) 25-27 25.21 ± 0.63 (19) 25-27 Dors 3 M 19.00 ± 0.00 (5) 19-19 18.50 ± 1.00 (4) 17-19 19.00 ± 0.00 (4) 19-19 18.77 ± 0.64 (18) 17-19 F 19.00 ± 0.00 (4) 19-19 19.00 ± 0.00 (7) 19-19 19.18 ±0.60 (11) 19-21 19.00 ± 0.00 (19) 19-19 Ap M 6.80 ± 0.44 (5) 6-7 7.00 ± 0.00 (4) 7-7 6.00 ± 0.00 (4) 6-6 6.38 ± 0.50 (18) 6-7 F 7.00 ± 0.81 (4) 6-8 6.28 ± 0.48 (7) 6-7 6.81 ± 0.87 (11) 6-9 6.31 ± 0.58 (19) 6-8 Blspl M 23.80 ±0.83 (5) 23-25 25.25 ± 0.50 (4) 25-26 26.00 ± 0.81 (4) 25-27 25.20 ± 1.20 (18) 23-28 F 25.25 ± 1.70 (4) 23-27 25.71 ± 0.75 (7) 25-27 25.36 ± 1.28 (11) 23-27 26.12 ± 1.02 (19) 24-28 Can M 2.20 ± 0.44 (5) 2-3 2.00 ± 0.00 (4) 2-2 2.50 ± 0.57 (4) 2-3 2.18 ± 0.40 (18) 2-3 F 2.25 ± 0.50 (4) 2-3 2.00 ± 0.00 (7) 2-2 2.50 ± 0.52 (11) 2-3 2.15 ± 0.37 (19) 2-3 Interocular M 12.00 ± 0.70 (5) 11-13 10.50 ± 0.57 (4) 10-11 11.75 ± 1.25 (4) 10-13 11.11 ± 0.67 (18) 10-12 F 11.75 ± 0.95 (4) 11-13 11.28 ± 0.75 (7) 10-12 12.09 ± 1.22 (11) 10-14 11.21 ± 0.97 (19) 9-13 Spl(L) M 10.40 ± 0.54 (5) 10-11 10.00 ± 0.81 (4) 9-11 9.75 ± 0.50 (4) 9-10 10.72 ±0.57 (18) 10-12 F 10.50 ± 0.57 (4) 10-11 9.85 ± 0.37 (7) 9-10 10.27 ± 0.64 (11) 9-11 10.73 ± 0.65 (19) 10-12 Spl(R) M 10.60 ± 0.54 (5) 10-11 9.75 ± 0.50 (4) 9-10 9.50 ± 0.57 (4) 9-10 10.83 ± 0.61 (18) 10-12 F 10.50 ± 0.57 (4) 10-11 10.28 ± 0.75 (7) 9-11 10.45 ± 0.52 (11) 10-11 10.52 ± 0.61 (19) 10-12 192 N. Moradi et alii Character Sex I Mean ± SD (n) (range) II Mean ± SD (n) (range) III Mean ± SD (n) (range) IV Mean ± SD (n) (range) Ifl(L) M 13.80 ± 0.44 (5) 13-14 12.25 ± 0.50 (4) 12-13 13.00 ± 0.81 (4) 12-14 14.27 ± 0.89 (18) 13-16 F 13.75 ± 0.50 (4) 13-14 13.28 ± 0.75 (7) 13-15 13.36 ± 0.67 (11) 13-15 14.42 ± 1.01 (19) 13-17 Ifl(R) M 13.80 ± 0.83 (5) 13-15 12.50 ± 0.57 (4) 12-13 13.00 ± 0.00 (4) 13-13 14.16 ± 0.85 (18) 13-16 F 13.50 ± 0.57 (4) 13-14 13.57 ± 0.78 (7) 12-14 13.45 ± 0.68 (11) 13-15 13.42 ±0.83 (19) 13-16 inCir(L) M 15.60 ± 1.51 (5) 14-17 14.50 ± 1.00 (4) 13-15 16.00 ± 1.55 (4) 15-17 15.11 ± 1.07 (18) 14-17 F 14.50 ± 1.29 (4) 13-16 16.14 ± 0.69 (7) 15-17 15.90 ± 1.04 (11) 14-17 14.78 ± 1.08 (19) 13-17 inCir(R) M 15.60 ± 2.70 (5) 13-19 14.25 ± 0.95 (4) 13-15 15.50 ± 1.29 (4) 14-17 15.61 ± 1.09 (18) 14-18 F 14.75 ± 1.25 (4) 13-16 16.71 ± 0.75 (7) 16-18 16.18 ± 1.66 (11) 13-19 14.94 ± 1.12 (19) 13-17 outCir(L) M 23.20 ± 0.83 (5) 22-24 22.25 ± 1.50 (4) 21-24 20.75 ± 1.70 (4) 19-23 22.88 ± 1.27 (18) 21-25 F 20.75 ± 1.70 (4) 19-23 21.71 ± 0.95 (7) 20-23 21.36 ± 1.69 (11) 20-26 22.89 ± 1.15 (19) 21-25 outCir(R) M 22.80 ± 1.30 (5) 21-24 22.00 ± 2.16 (4) 20-25 22.75 ± 2.87 (4) 19-26 22.88 ± 1.67 (18) 20-25 F 21.50 ± 2.38 (4) 19-24 22.14 ± 1.77 (7) 20-25 22.72 ± 2.19 (11) 19-27 22.94 ± 1.90 (19) 18-25 Lor(L) M 16.40 ± 1.51 (5) 14-18 13.00 ± 1.41 (4) 12-14 14.50 ± 0.57 (4) 14-15 15.46 ± 2.29 (18) 12-20 F 14.25 ± 3.20 (4) 11-17 13.14 ± 1.67 (7) 11-16 14.00 ± 2.79 (11) 9-19 15.62 ± 1.74 (19) 12-18 Lor(R) M 16.80 ± 2.16 (5) 14-20 14.00 ± 1.41 (4) 13-15 14.75 ± 1.89 (4) 12-16 14.92 ± 1.97 (18) 12-19 F 15.25 ± 2.75 (4) 12-18 13.42 ± 1.90 (7) 11-16 13.81 ± 2.52 (11) 9-17 15.75 ± 1.43 (19) 13-18 Supraoc M 1.40 ± 0.89 (5) 1-3 2.00 ± 0.81 (4) 1-3 2.75 ± 0.50 (4) 2-3 1.33 ± 0.68 (18) 1-3 F 1.50 ± 1.00 (4) 1-3 2.71 ± 0.48 (7) 2-3 2.81 ± 0.40 (11) 2-3 1.15 ± 0.50 (19) 1-3 bESpl M 3.00 ± 0.00 (5) 3-3 3.00 ± 0.00 (4) 3-3 3.00 ± 0.00 (4) 3-3 2.94 ± 0.23 (18) 2-3 F 3.00 ± 0.00 (4) 3-3 3.00 ± 0.00 (7) 3-3 3.00 ± 0.00 (11) 3-3 2.89 ± 0.31 (19) 2-3 Tal/SVL M 0.13 ± 0.00 (5) 0.13 ± 0.01 (4) 0.13 ± 0.00 (4) 0.13 ± 0.01 (17) F 0.13 ± 0.01 (4) 0.12 ± 0.00 (7) 0.13 ± 0.01 (11) 0.13 ± 0.01 (19) HL/SVL M 0.04 ± 0.00 (5) 0.04 ± 0.00 (4) 0.04 ± 0.00 (4) 0.04 ± 0.00 (15) F 0.04 ± 0.00 (4) 0.04 ± 0.00 (7) 0.04 ± 0.00 (11) 0.04 ± 0.00 (16) HW/SVL M 0.02 ± 0.00 (5) 0.02 ± 0.00 (4) 0.0 ± 0.00 (4) 0.02 ± 0.00 (17) F 0.02 ± 0.00 (4) 0.02 ± 0.00 (7) 0.02 ± 0.00 (11) 0.02 ± 0.00 (19) HH/SVL M 0.01 ± 0.00 (5) 0.01 ± 0.00 (4) 0.01 ± 0.00 (4) 0.01 ± 0.00 (18) F 0.01 ± 0.00 (4) 0.01 ± 0.00 (7) 0.01 ± 0.00 (11) 0.01 ± 0.00 (19) DbN/SVL M 0.01 ± 0.00 (5) 0.00 ± 0.00 (4) 0.00 ± 0.00 (4) 0.00 ± 0.00 (15) F 0.01 ± 0.00 (4) 0.00 ± 0.00 (7) 0.00 ± 0.00 (11) 0.00 ± 0.00 (16) Table 3. (Continued). 193Geographic variation in the Iranian Macrovipera lebetina Character Sex I Mean ± SD (n) (range) II Mean ± SD (n) (range) III Mean ± SD (n) (range) IV Mean ± SD (n) (range) IOD/SVL M 0.01 ± 0.00 (5) 0.01 ± 0.00 (4) 0.01 ± 0.00 (4) 0.01 ± 0.00 (15) F 0.01 ± 0.00 (4) 0.01 ± 0.00 (7) 0.01 ± 0.00 (11) 0.01 ± 0.00 (16) SL/SVL M 0.01 ± 0.00 (5) 0.01 ± 0.00 (4) 0.01 ± 0.00 (4) 0.01 ± 0.00 (15) F 0.01 ± 0.00 (4) 0.01 ± 0.00 (7) 0.01 ± 0.00 (11) 0.01 ± 0.00 (16) SCN 16.66 ± 7.35 (7) 6.33-27.33 17.83 ± 3.62 (4) 12.66-21.00 19.82 ± 4.80 (11) 12.00-25.66 13.47 ± 4.95 (25) 7.00-27.00 L/WdbR 0.37 ± 0.12 (7) 0.21-0.52 0.69 ± 0.11 (4) 0.53-0.75 0.68 ± 0.22 (11) 0.34-1.07 0.37 ± 0.12 (25) 0.10-0.61 L/DdbR 0.86 ± 0.38 (7) 0.36-1.25 1.44 ± 0.22 (4) 1.14-1.67 1.61 ± 0.70 (11) 0.65-3.10 0.78 ± 0.41 (25) 0.32-1.97 L/DlbR 0.82 ± 0.32 (7) 0.48-1.38 0.95 ± 0.31 (4) 0.63-1.24 0.93 ± 0.32 (12) 0.52-1.78 0.64 ± 0.37 (26) 0.31-2.07 Table 3. (Continued). Fig. 1. Distribution map of Macrovipera lebetina in Iran, each coloured region represents a studied group. Each circle on the map represents a location from which at least one snake was examined. 194 N. Moradi et alii Principal Component Analysis Multivariate analysis of the meristic characters was carried out to determine whether distinct geographic units exist within Macrovipera lebetina populations. Five Principal Components (PCs) explained 55.8%, and the first eight Principal Components revealed 73.5% of the total variation. Of this total, 18.9% explained by PC1 in which Supraoc, Ven, and Ifl were mainly responsible for this variation; 31.8% explained by PC2 which is mainly attributed to Interocular and inCir; 41.6% explained by PC3 mainly attributed to outCir; and 49.1% explained by PC4 in which Scd and Dors3 were more important. These results showed that the Supraoc, Ven and Interocu- lar are most important characters in separating popula- tions (Table 4). The magnitude and sign of the loadings on PC1 and PC2 showed significant separation among groups and the high degree of separation of southern populations (group IV) from other groups. Also, group I tends to separate from two residual groups (Fig. 3). Cluster analysis A Pearson distance dendrogram based on two mer- istic characters (Ven and Supraoc) with P < 0.05 and r2 > 50% clusters all four geographic groupings used in this study. Males and females are treated together because there is no sexual significant difference between them in these two characters. In Figure 4, the first major dichoto- my separates the southern populations (cluster IV) with a considerable distance from all other clusters. The second major dichotomy separates the northeastern populations (cluster I) with a distance from clusters II and III, which, in turn, constitute two major branches; one branch encompasses Gorgan populations (cluster II) from north- ern Iran, whereas the other branch is composed of all the other localities in northwest, west and western parts of Alborz Mts., approaching cluster IV in the south. Discriminant analysis The reclassifications were compared to the original cluster designations to determine how well the original model was supported by the Discriminant Analysis. Due to existence of missing values, twelve cases were removed from the multivariate analyses. Discriminant Analysis was performed on the 58 remaining snakes with sexes com- bined. Classification results and predicted group member- ship showed that 98.3% of the original allocations were correctly classified. Though vipers from the northeastern population (group I) were correctly classified to 88.9%, this value is 100% for other three groups. However, classi- fication results with Cross-validation indicated that a total of 72.4% of the original groupings were correctly classi- fied. Proportions correctly classified are 66.7% for group I, 77.8% for group II, 46.2% for group III, and 85.2% for group IV. The generalized squared distances among groups were computed to show the distance between each pair of groups as follow: 17.37 between group II and III; 22.63 between group I and II; 19.69 between group I and IV; 31.32 between group II and IV; 19.49 between group I and III; and 28.03 between group III and IV (Table 5). Fig. 2. Typical pattern of Macrovipera lebetina from northeastern populations (Left); northern, western and northwestern populations (Mid- dle); and southern populations (Right) (Sketch by: Naeim Moradi). 195Geographic variation in the Iranian Macrovipera lebetina Table 4. Factor loadings on the first eight Principal Components extracted from a correlation matrix of 22 meristic characters in Macrovi- pera lebetina from Iran. Characters PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PreV 0.214 -0.009 0.048 -0.298 0.361 0.068 -0.008 0.013 Ven 0.323 -0.127 0.192 0.169 -0.307 0.182 0.156 0.059 Scd 0.172 -0.104 -0.018 0.426 -0.076 -0.216 0.042 -0.263 Dors 1 -0.024 0.196 0.065 0.150 0.085 -0.213 0.101 0.613 Dors 2 0.106 0.269 0.265 0.104 -0.021 -0.368 0.139 -0.300 Dors 3 -0.062 0.292 0.155 0.302 -0.096 0.034 0.097 -0.360 Ap 0.032 0.188 0.095 0.207 0.563 -0.178 0.011 -0.057 Blspl -0.010 0.167 -0.109 0.264 -0.259 0.118 -0.550 0.092 Can -0.093 0.244 0.003 0.322 0.218 0.399 -0.248 -0.066 Interocular 0.028 0.425 -0.033 0.032 0.167 -0.080 0.281 0.154 Spl(L) 0.267 0.056 0.135 0.051 0.096 -0.085 -0.498 0.032 Spl(R) 0.230 0.139 0.023 -0.354 0.138 -0.080 -0.330 -0.049 Ifl(L) 0.304 0.172 0.345 -0.079 -0.189 0.193 0.085 0.155 Ifl(R) 0.304 0.230 0.236 -0.176 -0.146 -0.025 -0.135 0.024 inCir(L) -0.124 0.389 -0.234 -0.209 -0.199 0.015 -0.005 -0.023 inCir(R) -0.112 0.349 -0.083 -0.204 -0.372 -0.115 0.076 -0.074 outCir(L) 0.302 0.022 -0.368 -0.020 -0.018 -0.287 0.030 0.011 outCir(R) 0.175 0.017 -0.494 -0.023 -0.003 -0.194 -0.106 -0.248 Lor(L) 0.340 0.139 -0.253 0.030 0.061 0.277 0.237 0.027 Lor(R) 0.289 0.074 -0.330 0.163 0.065 0.383 0.163 0.048 Supraoc -0.368 0.263 -0.105 -0.030 0.062 0.131 -0.048 0.058 bESpl -0.035 0.044 0.147 -0.283 0.135 0.312 0.080 -0.435 Eigenvalue 4.1650 2.8393 2.1580 1.6449 1.4744 1.4230 1.3280 1.1291 Proportion 0.189 0.129 0.098 0.075 0.067 0.065 0.060 0.051 Cumulative 0.189 0.318 0.416 0.491 0.558 0.623 0.683 0.735 Fig. 3. Scatterplot of individuals on the first two Principle Components. 196 N. Moradi et alii Fig. 4. Pearson distance dendrogram resulting from the Cluster Analysis based on two meristic characters (Ventral and Supraocular scales) of Macrovipera lebetina in Iran. Table 5. Linear Discriminant Function for groups. Characters I II III IV PreV 119 125 120 121 Ven 88 88 88 90 Scd -22 -23 -22 -23 Dors 1 48 49 48 49 Dors 2 61 62 58 59 Dors 3 242 246 243 244 Ap -15 -16 -16 -19 Blspl 78 81 79 80 Can -24 -29 -20 -17 Interocular -41 -43 -40 -43 Spl(L) 72 70 73 72 Spl(R) 64 62 61 66 Ifl(L) -162 -164 -159 -164 Ifl(R) -56 -58 -59 -55 inCir(L) 39 40 41 39 inCir(R) -0 0 -1 -1 outCir(L) 131 133 130 134 outCir(R) -56 -56 -53 -56 Lor(L) -11 -12 -11 -10 Lor(R) -32 -33 -35 -34 Supraoc 39 45 44 38 bESpl 371 384 376 368 Constant -12071 -12320 -12210 -12495 Table 6. Standardized Coefficients for determining differences among four groups of Macrovipera lebetina. Characters Function 1 2 3 PreV 0.070 -0.421 0.447 Ven -0.234 0.755 -0.001 Scd -0.045 0.151 -0.204 Dors 1 -0.001 -0.097 0.175 Dors 2 0.142 -0.261 -0.022 Dors 3 0.135 -0.067 0.001 Ap 0.135 -0.422 -0.490 Can -0.345 0.546 0.049 Interocular 0.121 0.651 -0.567 Spl(L) -0.219 0.061 -0.347 Spl(R) -0.228 0.108 -0.131 Ifl(L) -0.208 0.012 0.077 Ifl(R) -0.369 0.198 0.444 inCir(L) 0.115 0.131 -0.141 inCir(R) 0.098 -0.472 0.188 outCir(L) -0.059 0.179 0.391 outCir(R) 0.163 0.249 0.282 Lor(L) 2.340 7.021 2.551 Lor(R) -2.137 -6.654 -2.474 Supraoc 0.874 0.267 0.195 bESpl 0.418 -0.082 -.295 197Geographic variation in the Iranian Macrovipera lebetina Based on additional Discriminant Function analy- sis (DFA) of meristic characters using SPSS® Ver. 20, the first three functions yielded 100% of total information in which, the first function explained 73% of the total variance with characters Supraoc, Ifl, Spl and inCir hav- ing the highest values. The loadings of each character on a particular function are shown in Table 6. Function 1 is heavily weighted by numbers of supraocular scales (Supraoc) and the number of loreal scales (Lor). Func- tion 2 is weighted by ventral (Ven), canthal (Can), Loreal (Lor) and Interocular scales, while function 3 is weighted by loreal scales count (Lor). Totally, loreal scale count was the most powerful character to separate four groups. Based on this analysis, with the ordination of four groups along the first two functions (Function 1 against Function 2), the centroids of each geographic group is well separated from any other group (Fig. 5). Results of sexual dimorphism analyses Firstly, two sample t-tests using all meristic data showed significant sexual dimorphism (P < 0.05) in Scd and Blspl. This indicates that males have more subcau- dal scales (Scd) and females have more scales across the head. A Principal Component Analysis (PCA) for sexes was performed; the magnitude and sign of the loadings on PC1 and PC2 showed apparent sexual dimorphism pattern between males and females. Although, the scat- terplot did not show a clear separation between the sex- es, but a weak pattern of sexual dimorphism is apparent (Fig. 6). Fig. 5. Patterns of variation expressed by the two Discriminant Functions (Function 1 against Function 2) for four groups of Mac- rovipera lebetina. 210-1-2 3 2 1 0 -1 -2 p2 p 1 F M Sex Scatterplot of p1 vs p2 Fig. 6. Scatterplot of Principal Component Analysis for males and females of Macrovipera lebetina. 198 N. Moradi et alii DISCUSSION Latifi in 1983 assigned the Iranian Macrovipera to the subspecies of Macrovipera lebetina obtusa (Dwigubsky, 1832). However, Chikin and Szczerbak in 1992 described a new subspecies from Khorasan Province. Until now, the Iranian Macrovipera consisted of two subspecies: Mac- rovipera lebetina obtusa (Dwigubsky, 1832) (Fig. 7) dis- tributed in west, northwest, and north of Iran, and Mac- rovipera lebetina chernovi (Chikin and Szczerbak, 1992) (Fig. 8) occurring in northeastern Iran. Our results indi- cated that the separation of the northeastern populations of M. lebetina from the northwestern and western popu- lations could be corroborated by morphological char- acters, as previously suggested by Chikin and Szczerbak (1992), Ananjeva et al. (2006), Rajabizadeh et al. (2011), and Oraie et al. (2012). However, the character expression in southern populations likely amounts to another taxo- nomic status in the Iranian Macrovipera taxa. M. lebetina populations from southern region of the Iranian Plateau showed significant differences to all northern populations regarding several morphologi- cal characters. For example, pholidosis showed a gen- eral north-south pattern of geographic variation for most meristic characters. Ventral, infralabial, outer circumocu- lar and supralabial scales averaged higher, and interocu- lar, inner circumocular and supraoculars averaged lower in the southern regions. Many ophidians display latitudinal pattern of geo- graphic variation in Pholidosis. Christman (1980) exam- ined 15 species of snakes in Florida and founded a lati- tudinal pattern of geographic variation in two characters, ventral and subcaudal scales, both of which increased in number from north to south in most species. Higher scales count in the southern regions could also be the result of selection for larger body sizes, which in turn may reflect longer annual growing periods in southern regions. Substantial geographic variation in colouration was observed among the Iranian Macrovipera as well. Vipers from the southern populations have the narrowest cross- bars on dorsum with high distances between them. They also entail lower scale counts compared to northwestern and northeastern populations. The general pattern of body crossbars and blotches in Macrovipera lebetina appeared to be influenced by latitudinal pattern and elevation. The Alborz Mts. of the northern and northwestern regions combine high latitude and high elevation, and conse- quently annual temperatures there are among the coldest in the species’ range. High number of dark pattern ele- ments would be expected to increase the ability of a snake to thermoregulate at high elevations. Integumentary col- our and reflectance are known to exert a strong influence on the thermal biology of snakes (Peterson et al., 1993). Also, according to non-parametric analysis, there was a clinal difference between northern and southern popu- lations, so that with toward the southern latitudinal, dor- sal body colour become brighter and ventral dots pattern become denser and dots form a trapezoid-shaped spots and belly will have a mottled appearance. Geographic variation in pattern may be strongly related to habitat. The differences between the northeast- ern and western regions may be due to the lower eleva- tions and warmer summers of the northeastern regions and different landscape features in Kopet Dagh Mts. In addition, the southern populations remained clearly distinct in all multivariate analyses and the clus- ter analyses showed that the southern populations were distinguishable with a considerable distance from all the other clusters. Multivariate statistical analyses are the Fig. 7. An adult specimen of Macrovipera lebetina obtusa from Tak- ab, western Azerbaijan Province, Iran (Photograph by Omid Mozaf- fari). Fig. 8. An adult specimen of Macrovipera lebetina chernovi from Khorasan Province, Iran (Photograph by Hasan Moghimi). 199Geographic variation in the Iranian Macrovipera lebetina most comprehensive when applied to morphological data, inasmuch as the rely upon multiple aspects of the pheno- type. The formal recognition of distinct geographic units within Macrovipera lebetina in supported by this study. Gorgan populations (cluster II) in the multivari- ate analyses virtually separated from other northeastern and northern-western populations, however, existence of overlapping in pholidosis and colour pattern in some characters can be a sign of a hybrid zone between two major northeastern and northern-western populations, this confirms the existence of two subspecies of M. l. obtusa and M. l. chernovi. In spite of that cannot be rec- ognized a distinct border between ranges of two northern subspecies, individuals of Gorgan populations have more similar to the northern-western subspecies. Eventually, individual characters often showed clear patterns of variation. In all of multivariate analyses, char- acters of Ventral and Supraocular scales were identified as important and significant variables and were used for mapping geographic variation. Differences in dorsal and ventral scale counts are often diagnostic for different species of Macrovipera (Nil- son and Andrén, 1988; Joger, 1984; Göçmen et al., 1999; Phelps, 2010) and Chikin and Szczerbak, 1992 early reli- ance on supraoculars to delineate subspecies of Iranian Macrovipera lebetina. Most recently, Stümpel (2012) in molecular analyses on mtDNA sequences mentioned that the specimen from southeastern Iran were completely distinct from other specimens of M. lebetina. However, studies of reptiles have demonstrated the early classifica- tion based on a few specimens or single morphological characters often do not agree with more comprehensive molecular and morphological analyses. Nevertheless, we accept, provisionally, considering the some morphological evidences, the hypothesis of the specific status of the southern Iranian populations as Macrovipera sp. (Fig. 9). However, complementing phylo- genetic investigation is required to gain a deeper under- standing of the evolution of this species complex. New combinations of morphological characters are used to device a key for identification of Iranian Macrovi- pera, partly based on diagnostic characteristics gathered from Chikin and Szczerbak (1992), David et al. (1999), Joger (1984) and Oraie et al. (2012). IRANIAN MACROVIPERA IDENTIFICATION KEY 1a- Ventral scales 156-188 (less than 171 in 81.08% of specimens) ............................................................................... 2 1b- Ventral scales 169-179 (more than 172 in 83.78% of specimens), supraocular entire, and infralabials 13-17, dorsum brownish gray with narrow crossbars with distin- guishable distance between them, each crossbar includes 13.47 ± 0.99 pigmented scales at mid snout-vent length. Belly covered with dark blotch (3-5 trapezoid spots in posterior part of each ventral scales). Some southeastern specimens’ uniform glazy black ................. Macrovipera sp. 2a- Supraocular entire or divided and larger than circumoc- ulars, snout dark (one third of anterior part of the head). Dorsum reddish brown or dark with large unfilled blotch which sometimes connected together, each dorsum blotch includes 16.66 ± 2.78 pigmented scales at mid snout-vent length. The ratio of interocular distance to head length is 0.39 ± 0.02. Belly often light and splotched by dark dots ....... Macrovipera lebetina chernovi (Chikin and Szczerbak, 1992) 2b- Three supraoculars present and same as the other cir- cumoculars, head uniform, with narrow strips between the corner of the mouth and the posterior part of the eyes, and also between the eye and fourth supralabial scale. Dorsum brown or dark brown with rectangular dark marking and arrange in zigzag pattern, each dorsum blotch includes 19.29 ± 1.16 pigmented scales at mid snout-vent length. The ratio of interocular distance to head length is 0.36 ± 0.01. Belly often light and splotched with dark dots .............. ......................Macrovipera lebetina obtusa (Dwigubsky, 1832) ACKNOWLEDGEMENTS This study was performed based on official permis- sion number 91/49936 issued by Department of the Envi- ronment of Iran. Also, this project was supported by the Iranian Plateau Herpetology Research Group (IPHRG) of Fig. 9. An adult male, Macrovipera sp. from Masjed Soleyman, Khuzestan Province, Iran (Photograph by Naeim Moradi). 200 N. Moradi et alii the Razi University of Kermanshah. We would also like to thank the A. Zare Mirakabadi, curators of zoological collections in the Department of Venomous Animals and Antisera Production, Razi Vaccine and Serum Research Institute, Hesarak, Karaj, and S. Shafiei and H. R. Esmaei- li for borrowing preserved specimens. Also thank to K. Mebert for helpful comments. Our special thanks dedi- cated for F. Todehdehghan, M. E. Sehati-sabet, K. Rabiei, B. Fathinia, A. Sharifi, O. Mozaffari, H. Oraie and F. Mas- jedi for their helps and cooperation during this study. REFERENCES Afroosheh, M., Kazemi, S.M. (2011): Macrovipera lebeti- na cernovi (Ophidia: Viperidae), a newcomer to Iran. 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Herpetological collections, locality, and collection numbers (No.) of the specimens of Macrovipera lebetina used in present study. Specimens Locality No. ZCRI Kalat, near Mashhad, Khorasan Razavi Prov. 2349 Kalat, near Mashhad, Khorasan Razavi Prov. 2350 Kalat, near Mashhad, Khorasan Razavi Prov. 2363 Kalat, near Mashhad, Khorasan Razavi Prov. 2351 Kalat, near Mashhad, Khorasan Razavi Prov. 2346 Kalat, near Mashhad, Khorasan Razavi Prov. 2342 Kalat, near Mashhad, Khorasan Razavi Prov. 2344 Gorgan, Golestan Prov. 1313 Gorgan, Golestan Prov. 2154 Gonbad e Kavoos, Golestan Prov. 2158 Tang-e-Rah (Gonbad e Kavoos), Golestan Prov. 2260 Tang-e-Rah (Gonbad e Kavoos), Golestan Prov. 2261 Gonbad e Kavoos, Golestan Prov. 2256 Gonbad e Kavoos, Golestan Prov. 2259 Tang-e-Rah (Gonbad e Kavoos), Golestan Prov. 2262 Gorgan, Golestan Prov. 2151 Rahmat Abad (Rasht), Gilan Prov. 2143 Rudbar, Gilan Prov. 2150 Arak, Markazi Prov. 2267 Alajigh, Between Tehran & Saveh 2292 Ashk Island, Orumie Lake 129382 Mahallat, Markazi Prov. 1314 Orumie Lake Island 129381 Rezaeie, West Azarbaijan Prov. 129380 Alajigh, Between Tehran & Saveh 2180 Arezu Island, Orumie Lake 129386 Bijar, Kordistan Prov. 1249 Ashk Island, Orumie Lake 129383 Specimens Locality No. Kermanshah, Kermanshah Prov. 972 Semirom, Esfahan Prov. 2274 Khansar, Esfahan Prov. 2237 Khansar, Esfahan Prov. 2238 Meymahe, Esfahan Prov. 2280 Semirom, Esfahan Prov. 2159 Eghlid (Aso pas), Fars Prov. 2144 Albaji, Ahvaz, Khuzistan Prov. 2309 Tang-e-Karun, Eghlid, Fars Prov. 2210 Albaji, Ahvaz, Khuzistan Prov. 2142 Tang-e-Karun, Eghlid, Fars Prov. 2212 Shushtar, Khuzistan Prov. 2248 Abadeh, Fars Prov. 2298 Albaji, Ahvaz, Khuzistan Prov. 2234 Masjed soleyman, Khuzistan Prov. 2299 Albaji, Ahvaz, Khuzistan Prov. 2195 Aligudarz, Lorestan Prov. 2208 Tang-e-Karun, Eghlid, Fars Prov. 2211 Masjed-Soleyman, Khuzistan Prov. 2170 Mamasani, Fars Prov. 119204 Masjed-Soleyman, Khuzistan Prov. 2168 Ramhormoz, Khuzistan Prov. 121817 Kalat, near Mashhad, Khorasan Razavi Prov. (Picture) 426 Kalat, near Mashhad, Khorasan Razavi Prov. (Picture) 979 Kalat, near Mashhad, Khorasan Razavi Prov. 969 Mashhad, Khorasan Razavi Prov. 1385 Mashhad, Khorasan Razavi Prov. 1272 Mashhad, Khorasan Razavi Prov. 1273 Dare Gaz, Khorasan Razavi Prov. 608 202 N. Moradi et alii Specimens Locality No. Dare Gaz, Khorasan Razavi Prov. 2228 Dare Gaz, Khorasan Razavi Prov. 1304 Masjed-Soleyman, Khuzistan Prov. 2171 Masjed-Soleyman, Khuzistan Prov. 2166 SUHC Khorasan Razavi Prov. ERP 986 Khorasan Razavi Prov. ERP 987 Babmaran, Kerman Prov. ERP 143 Bahrame gur Protected area, Fars Province ERP 1531 Bakhtegan National park, Fars Prov. ERP 1518 Pariz, Kerman Prov. ERP 3732 Balvard, Sirjan, Kerman Prov. ERP 1941 Sabzevar, Khorasan Razavi Prov. ERP 666 Shimbar Protected area, Khuzistan Prov. ERP 1741 RUZM Kermanshah, Kermanshah Prov. VV.31.3 Kermanshah, Kermanshah Prov. VV.31.1 Kermanshah, Kermanshah Prov. VV.31.2 Kermanshah, Kermanshah Prov. VV.31.5 Asmari Mts, Masjed soleyman, Khuzistan Prov VV.31.6 Asmari Mts, Masjed soleyman, Khuzistan Prov VV.31.7 Abdanan, Ilam Prov. unlabeled Abdanan, Ilam Prov. unlabeled ZMSBUK Khabr National Park, Kerman Prov. A3 Khabr National Park, Kerman Prov. A18 Khabr National Park, Kerman Prov. A12 Sirjan, Kerman Prov. A46 Sirjan, Kerman Prov. A57 Khabr National Park, Kerman Prov. A58 Pariz, Kerman Prov. A110 Baft, Kerman Prov. A32 SUZM Shiraz, Fars Prov. unlabeled Specimens Locality No. Bamoo National Park, Fars Prov. unlabeled Bamoo National Park, Fars Prov. unlabeled FDOI Sarvestan, Fars Prov. unlabeled Seyf Abad, Fars Prov. unlabeled Seyf Abad, Fars Prov. unlabeled Sarvestan, Fars Prov. unlabeled Picture Abdanan, Ilam Prov. -- Khojir protected area, Tahran Prov. -- Bazman, Siatan Prov. -- Tangestan, Bushehr Prov. -- Capture & Release Golestan National Park, Golestan Prov. -- Golestan National Park, Golestan Prov. -- Golestan National Park, Golestan Prov. -- Damghan, Semnan Prov. -- Shimbar Protected area, Khuzistan Prov. -- Shimbar Protected area, Khuzistan Prov. -- Golgir, Masjid-Soleyman, Khuzistan Prov. -- Balvard, Sirjan, Kerman Prov. -- Balvard, Sirjan, Kerman Prov. -- Balvard, Sirjan, Kerman Prov. -- Pariz, Kerman Prov. -- Pariz, Kerman Prov. -- Ghatruye National park, Fars Prov. -- Bamoo National Park, Fars Prov. -- Bamoo National Park, Fars Prov. -- Gorm protected area, Jahrom, Fars Prov. -- Turan National Park, Semnan Prov. -- Shahrud, Semnan Prov. -- Shahrud, Semnan Prov. -- Rudbarak Protected area, Semnan Prov. -- ZCRI: Zoological Collection of the Department of Venomous Animals and Antiserum Production, Razi Vaccine and Serum Research Institute, Hesarak, Karaj ZMSBUK: Zoological Museum of Shahid Bahonar University of Kerman SUHC: Sabzevar University Herpetological Collection RUZM: Razi University Zoological Museum SUZM: Shiraz University Zoological Museum FDOI: Species Diversity Museum of Department of Environment, Fars