Turkish Journal of Zoology Turk J Zool (2013) 37: 348-356 http://journals.tubitak.gov.tr/zoology/ © TÜBİTAK Research Article doi:10.3906/zoo-1205-1

Reevaluation of the taxonomic status of sand boas of the genus Eryx (Daudin, 1803) (Serpentes: Boidae) in northeastern Iran

1, 1 2 1 Naeimeh ESKANDARZADEH *, Jamshid DARVISH , Eskandar RASTEGAR-POUYANI , Fereshteh GHASSEMZADEH 1 Department of Biology, Faculty of Science, Ferdowsi University of Mashhad, Mashhad, Iran 2 Department of Biology, Faculty of Science, Hakim Sabzevari University, Sabzevar, Iran

Received: 02.05.2012 Accepted: 21.01.2013 Published Online: 29.04.2013 Printed: 29.05.2013

Abstract: Based on the most comprehensive work on Iranian snakes to date, 4 species of the boid snakes of the genus Eryx (Daudin, 1803), E. tataricus (Lichtenstein, 1823), E. miliaris (Pallas, 1773), E. elegans (Gray, 1849), and E. jaculus (Linnaeus, 1758), have been reported from northeastern Iran. In this study, the distribution, morphological characters, and taxonomic status of the genus Eryx in northeastern Iran are reviewed. Four metric and 9 meristic characters were studied in 47 specimens belonging to various populations of Eryx in different parts of Khorasan provinces. Descriptions of the characters of the species of Eryx in northeastern Iran were revised. According to the results, and considering the principle of priority, we concluded that the northeastern populations of E. elegans are conspecific with E. jaculus, and the populations of E. tataricus are conspecific with E. miliaris. In addition, 2 specimens belonging to a population of Eryx in the area were found that differ from all other Iranian species of the genus in several taxonomic characters, but generally were similar to E. miliaris. These specimens need more comprehensive study to clarify their taxonomic status and phylogenetic relationship using more morphology traits, ecology, and molecular studies.

Key words: Eryx, morphological characters, distribution, northeast Iran

1. Introduction E. tataricus (Lichtenstein, 1823), E. miliaris (Pallas, 1773), Species of Boidae are present in the Americas, Africa, E. johnii (Russell, 1801), and E. jayakari (Boulenger, 1888). Europe, Asia, and the southwest Pacific Islands. The Of these, 4 species occur in northeastern Iran: E. elegans, Boidae comprise 3 subfamilies: Boinae, Erycinae, and E. jaculus, E. tataricus, and E. miliaris (Latifi, 1991).E. Ungaliophiinae (Wilcox et al., 2002). Eryx is the only miliaris is known from southern Russia from the western Iranian genus of the Erycinae subfamily and is distributed border of the Caspian Sea (Caucasus) through Kazakhstan, in southwestern Europe, North and East Africa, northern Iran, Afghanistan, Turkmenistan, Uzbekistan, southwestern Asia to India and Sri Lanka, northward to China (Xinjiang through Gansu to western Nei Mongol – Caucasia and Afghanistan, and eastward from Turkestan Inner Mongolia and Ningxia), and Mongolia. E. tataricus to southern Mongolia and western China (Lanza and is distributed along the eastern shore of the Caspian Sea Nistri, 2005). According to Bartlett and Bartlett (1998, from Turkmenistan eastward to eastern Kazakhstan, p. 39), “neither the number of species nor subspecies has Kyrgyzstan, Tajikistan, Uzbekistan, eastern Turkmenistan, ever been agreed on by authorities”. This genus is widely northern Iran, Afghanistan, western Pakistan, western distributed in Iran and can be distinguished by its small China (Xinjiang through Gansu to western Nei Mongol – eyes and indeterminate neck. The head is covered with Inner Mongolia and Ningxia), and southern Mongolia. E. small scales. The tail is short and has one row of subcaudal elegans is found in southern Turkmenistan, northeastern scales. Ventral scales are different from dorsal scales, and Iran, Afghanistan, and northwestern India; the distribution are narrower than the width of the body. They feed on of E. jaculus is southeastern Europe, southwestern lizards, small mammals, and other snakes. The neonates Asia, and North Africa (Terentyev and Chernov, 1965). may sometimes feed on insects (Terentyev and Chernov, According to the osteological studies of Tokar (1989, 1965; Latifi, 1991; Bartlett and Bartlett, 1998; Lanza and 1990), E. miliaris and E. tataricus form a complex together. Nistri, 2005). Until now, 6 species were reported from According to Rastegar-Pouyani et al. (2008) and Venchi Iran: E. elegans (Gray, 1849), E. jaculus (Linnaeus, 1758), and Sindaco (2006), the population of E. tataricus in Iran * Correspondence: [email protected] 348 ESKANDARZADEH et al. / Turk J Zool needs to be clarified as to subspecies. There is no precise variations, we used the ratio of body length to length of identification key for species of Eryx in Iran; this causes tail (LB/LT) and the ratio of interocular space width to misidentification of Iranian species. As yet, no study distance between the posterior edge of eye and corner of has been done on the taxonomic status of Eryx in Iran. mouth (WI/DEM). Discriminant analysis and analysis of The aim of this study is to revise the distribution and variance were performed using SPSS 16.0, with alpha set taxonomic status of Eryx species in northeastern Iran and at 0.05. To survey sexual dimorphism, an independent to determine some diagnostic characters for identification samples t-test was performed. of the species in the area. 3. Results 2. Materials and methods The results of the independent samples t-test showed Forty-seven specimens, including 27 males and 20 that the number of subcaudal scales (ScdS) and the ratio females, were included in the study. The females include of body length to tail length (LB/LT) were significantly 2 samples that differed from all other Iranian species of different between males and females of E. miliaris and the genus in several morphological characters, and so E. tataricus. Number of scales of right eye (RE) was these are indicated as Eryx sp. Detailed information on the significantly different in E. miliaris (P ≤ 0.05). No female specimens, including scientific and common names and was available for either E. elegans or E. jaculus, and so the localities, number of specimens, and deposition places of further analyses were performed separately for males and the samples, are given in Table 1 and Figure 1. females. Descriptive statistics and descriptive characters of Samples were collected by hand, placed within a special the species are shown in Tables 3–6. sack, and transferred to the laboratory for identification As shown in Tables 3 and 5, the metric characters for and complementary studies. Identification of the E. tataricus and E. miliaris show considerable overlap in 10 specimens used in this study was performed according to characters. This is also the case forE. elegans and E. jaculus. Terentyev and Chernov (1965). The only differing character betweenE. tataricus and E. Sex was determined using descriptions presented miliaris is the number of scales posterior to internasal area by Terentyev and Chernov (1965). Metric and meristic (PIN), and that between E. elegans and E. jaculus is the characters used for this study are presented in Table 2. number of dorsal scales (DS). All measurements were log- Metric characters were measured using a digital caliper transformed to normalize. with an accuracy of 0.01 millimeters. A ruler was used for 3.1. Males measuring length of body (LB). Pholidosis features were Twenty-seven male samples, including 6 specimens of counted under a stereomicroscope. The scales of the head E. tataricus, 8 specimens of E. miliaris, 9 specimens of are shown in Figure 2. For considering the allometric E. elegans, and 4 specimens of E. jaculus, were studied.

Table 1. Details of the localities of samples: a) collected samples, b) museum materials included in study. a. Collected samples.

Species No. specimens Locality Sex E. tataricus 3 Fariman, Shahan-Garmab Two females, 1 male E. tataricus 1 Sarakhs Undetermined (because of tail injury) Birjand E. tataricus 1 Female Tehran E. tataricus 1 Female Ghaen E.miliaris 1 Male Fariman, Shahan-Garmab E. miliaris 1 Female Birjand E. elegans 1 Male Kardeh Dam E. elegans 1 Male Kalat E. elegans 1 Male Ghaen E. elegans 3 Male Kaedeh Dam E. jaculus 1 Male

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Table 1. Details of the localities of samples: a) collected samples, b) museum materials included in study. b. Museum materials included in the study.

Species Sample ID number Place of upkeep Locality Sex

ERP 593 E. miliaris ERP 592 Hakim Sabzevari University Female Khorasan ERP 594

E. miliaris ERP 347 Hakim Sabzevari University Dasht Male E. miliaris 1159 Ferdowsi University of Mashhad Jajarm Male

E. miliaris 143 Ferdowsi University of Mashhad - Male

E. miliaris 141 a Ferdowsi University of Mashhad - Male

E. miliaris 317 Azad University of Mashhad Birjand Female

E. miliaris 44 Hakim Sabzevari University - Female Neyshabour Torbat Heidariyeh Road E. miliaris ERP 976 Hakim Sabzevari University Female

ERP 925 E. miliaris ERP 926 Hakim Sabzevari University Joghatai Male ERP 928 Neyshabour Sabzevar Road E. miliaris ERP 806 Hakim Sabzevari University Female

E. tataricus ERP 591 Hakim Sabzevari University Khorasan Female

E. tataricus ERP 929 Hakim Sabzevari University Joghatai Female

E. tataricus ERP 590 Hakim Sabzevari University Torbat Jam Female

E. tataricus ERP 792 Hakim Sabzevari University Sabzevar, Parvand Female

E. tataricus 142 Ferdowsi University of Mashhad - Female

E. tataricus 33 Azad University of Mashhad Torbat Jam Male

22 b E. tataricus Department of the Environment of Kerman Ghaen Male 22 a

E. tataricus ZMGU 1148 Zoological Museum of Golestan University Torbat Jam Male

E. tataricus ZMGU 494 Zoological Museum of Golestan University Neyshabour Male

E. tataricus ZMGU 2001 Zoological Museum of Golestan University Sabzevar Male

E. jaculus ERP 352 Hakim Sabzevari University Dargaz Male

E. jaculus 23 Hakim Sabzevari University Sarigol Male

E. jaculus 215 Azad University of Mashhad Sabzevar Male

E. elegans 120 Azad University of Mashhad North Khorasan Male

E. elegans 27 Hakim Sabzevari University Sarigol Male

E. elegans ZMGU 2091 Zoological Museum of Golestan University Golol Male

Eryx sp. ERP 977 Hakim Sabzevari University Neyshabour Female

Eryx sp. 141 b Ferdowsi University of Mashhad - Female

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Figure 1. Map of Iran and the position of Khorasan provinces, where the specimens of this study were collected. 1- Dasht; 2- Jajarm; 3- Esfraien, Sarigol; 4- Shirvan, Golol; 5- Dargaz; 6- Kalat; 7- Sarakhs; 8- Kardeh Dam; 9- Fariman, Shahan-Garmab; 10- Neyshabour; 11- Neyshabour, Sabzevar Road; 12- Joghatai; 13- Sabzevar; 14- Sabzevar, Parvand; 15- Neyshabour, Torbat Heidariyeh Road; 16- Torbat Jam; 17- Ghaen; 18- Birjand.

Table 2. Description of the most informative metric and meristic characters in Eryx.

Row Variable Abbreviations

1 Distance between posterior edge of eye and corner of mouth DEM

2 Width of interocular space WI

3 Number of scales around right eye RE

4 Number of scales around left eye LE

5 Number of supralabial scales LAB

6 Number of scales posterior to internasal PIN

7 Number of scales between eyes BE

8 Number of dorsal scales DS

9 Number of ventral scales VS

10 Number of subcaudal scales ScdS

11 Body length LB

12 Tail length LT

13 Number of scales between eye and nasal BEN

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Analysis of variance (ANOVA) was performed for meristic and metric characters separately to show significantly variable characters. The result of ANOVA shows that 9 characters (RE, number of scales of left eye [LE], number of supralabial scales [LAB], PIN, number of scales between eyes [BE], DS, ventral scales [VS], number of scales between eyes and nasal area [BEN], and WI/DEM) are significantly variable (P ≤ 0.05). Number of subcaudal scales (ScdS) and the ratio of body length to tail length (LB/LT) with P > 0.05 are not significantly different. 3.1.1. Multivariate analysis Discriminant analysis (DCA) of males was performed, and the first function separated E. tataricus and E. miliaris from E. elegans and E. jaculus (Figure 3). The first and Figure 2. The topography and scalation of the head in Eryx. second discriminant functions (DFs) accounted for 99.7%

Table 3. Descriptive characters of males included in this study.

RE LE LAB BE PIN BEN DS VS ScdS LB/LT WI/DEM E. tataricus N = 6 10–13 10–12 10–13 7–8 3 3–4 44–48 174–187 27–33 6.2–7.6 0.8–1 E. miliaris N = 8 11–13 11–13 11–13 7–8 4 4 46–52 176–188 26–33 6.2–8 0.6–0.9 E. elegans N = 9 8–10 8–9 8–10 6–7 2 3 36–41 159–179 29–45 4.9–8 1.2–1.5 E. jaculus N = 4 8–9 8–10 9–10 6–7 2 3 42–46 165–173 28–45 4.9–7 1.2–1.7

Table 4. Descriptive statistics of males included in this study.

RE LE LAB BE PIN BEN DS VS ScdS LB/LT WI/DEM E. tataricus Mean 12 11 12 7.5 3 3.8 47 179 30 6.8 0.8 Minimum 10 10 10 7 - 3 44 174 27 6 0.8 Maximum 13 12 13 8 - 4 48 187 33 7 1 Std. error 0.5 0.3 0.4 0.2 - 0.2 0.6 2.3 0.9 0.2 0.3 Std. deviation 1.2 0.8 1 0.5 - 0.4 1.5 5.6 2.1 0.5 0.1 E. miliaris Mean 12 12 12 7.5 4 4 49 182 30 7.2 0.7 Minimum 11 11 11 7 - - 46 176 26 6 0.6 Maximum 13 13 13 8 - - 52 188 33 8 0.9 Std. error 0.2 0.2 0.3 0.1 - - 0.8 1.6 0.7 0.2 0.3 Std. deviation 0.6 0.6 0.7 0.4 - - 2.1 4.6 2.1 0.7 0.1 E. elegans Mean 9 8.5 8.5 6.5 2 3 39 169 35 5.8 1.2 Minimum 8 8 8 6 - - 36 159 29 4 1.2 Maximum 10 9 9 7 - - 41 179 45 6 1.5 Std. error 0.3 0.2 0.1 0.2 - - 0.8 2.3 2.3 0.4 0.04 Std. deviation 0.7 0.5 0.4 0.5 - - 2 6.1 6 1 0.1 E. jaculus Mean 8.5 9 9.5 6.5 2 3 43 168 35 5.9 1.3 Minimum 8 8 9 6 - - 42 165 28 4 1.2 Maximum 9 10 10 7 - - 46 173 45 7 1.7 Std. error 0.3 0.4 0.3 0.3 - - 0.9 1.8 3.6 0.5 0.1 Std. deviation 0.5 0.8 0.5 0.5 - - 1.9 3.6 7.3 1 0.3

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Table 5. Descriptive characters of females included in this study.

RE LE LAB BE PIN BEN DS VS ScdS LB/LT WI/DEM E. tataricus N = 10 10–13 10–12 11–13 6–9 3 3–4 43–51 171–193 21–23 8.8–15.6 0.6–1 E. miliaris N = 8 10–13 10–13 11–13 7–9 4 4 44–50 182–188 21–24 8.3–11.3 0.8 Eryx sp. N = 2 12 12 12 7 2 4 41–44 172–187 22–23 9.7–10.9 0.8

Table 6. Descriptive statistics of females included in this study.

RE LE LAB BE PIN BEN DS VS ScdS LB/LT WI/DEM E. tataricus Mean 11 11 12 7 3 4 48 186 22 11.1 0.8 Minimum 10 10 11 6 - - 43 171 21 8.8 0.6 Maximum 13 12 13 9 - - 51 193 23 15.6 1 Std. error 0.3 0.3 0.2 0.3 - - 0.8 2.3 0.3 0.6 0.5 Std. deviation 1 0.9 0.7 0.9 - - 2.4 6.8 0.9 1.9 0.1 E. miliaris Mean 11 11 12 8 4 4 49 185 23 9.9 0.8 Minimum 10 10 11 7 - - 44 182 21 8.3 - Maximum 12 13 13 9 - - 50 188 24 11.3 - Std. error 0.2 0.3 0.2 0.3 - - 0.7 0.9 0.4 0.4 - Std. deviation 0.6 0.9 0.6 0.8 - - 1.9 2.5 1.2 1 - Eryx sp. Mean 12 12 12 12 2 4 43 180 22.5 10.3 0.8 Minimum ------41 172 22 9.7 - Maximum ------44 187 23 10.9 - Std. error ------1.5 7.5 0.5 0.6 - Std. deviation ------2.1 1.1 0.7 0.8 -

Canoncal dscrmnant functons of the variance. It was seen that 97% of the variance was 10 explained by DF1, in which RE, LE, BEN, WI/DEM, and VS had the most effect on its formation; 2.7% of variance was explained by DF2, in which DS had the most effect on 5 its formation; and 0.3% of variance was explained by DF3, in which BE and LAB had the most effect. The eigenvalue 4 of DF1 is 43.09; for DF2, it is 1.18 (Table 7). 3.2. Females 0 1 2 3 Twenty female samples, including 10 E. tataricus, 8

Functon 2 E. miliaris, and 2 Eryx sp., were studied. ANOVA was performed for meristic and metric characters separately code to show significantly variable characters. The results of –5 E. tatarcus E. mlars ANOVA showed that 2 characters were significantly E. elegans different (P ≤ 0.05): DS and PIN. E. jaculus 3.2.1. Multivariate analysis Group centrod –10 DCA of females was performed, and the first function separated E. tataricus and E. miliaris from Eryx sp. (Figure –10 –5 0 5 10 Functon 1 4). The first and second DFs explained 100% of the variance. It was seen that 99% of variance was explained Figure 3. Discriminant analysis of male samples. The first by DF1, in which PIN had the most effect on its formation, function, in which RE, LE, BEN, WI/DEM, and VS have the most and 1% of variance was explained by DF2, in which DS effect, separates E. tataricus and E. miliaris from E. elegans and had the most effect. The eigenvalue of DF1 is 10.66; for E. jaculus. DF2, it is 0.11 (Table 8).

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Table 7. Correlations within groups among discriminant variables and standardized canonical discriminant functions in male samples of Eryx. Variables are ordered by the absolute value of correlation within the function. Asterisk signifies the highest absolute correlation between a variable and the discriminant function. Discriminant function 1 2 3 Eigenvalue 43.09 1.18 0.12 % variance 97 2.7 0.3 Canonical correlation 0.989 0.736 0.336 BEN 0.397* –0.079 –0.100 LE 0.366* 0.189 –0.006 WI/DEM 0.355* 0.113 –0.134 RE 0.302* –0.166 –0.158 VS 0.185* –0.161 –0.122 DS 0.314 0.647* –0.284 BE 0.126 0.243 0.735* LAB 0.322 0.312 –0.544*

Table. 8. Correlations within groups among discriminant variables and standardized canonical discriminant functions in female samples of Eryx. Variables are ordered by the absolute value of correlation within the function. Asterisk signifies the highest absolute correlation between a variable and the discriminant function. Discriminant function 1 2 Eigenvalue 10.66 0.112 % variance 99 1 Canonical correlation 0.956 0.317 PIN 0.952* –0.305 DS 0.246 0.969*

Canoncal dscrmnant functons 4. Discussion 10 As already stated, our results indicate very close relations code and high morphological similarity between E. tataricus E. tatarcus E. mlars and E. miliaris on one hand and E. jaculus and E. elegans on Eryx sp. the other, such that they can hardly be distinguished from 5 Group centrod each other. According to the results, and considering the principle of priority, we believe that in Khorasan provinces the populations of E. elegans are conspecific with E. jaculus, 1 and the populations of E. tataricus are conspecific with E. 0 3 2 miliaris. Dividing the Khorasan populations of Eryx into 4 Functon 2 distinct species, as suggested by other studies (Latifi, 1991), is not supported by the results of our study. This could be –5 due to either our incomplete taxon sampling in the area or misidentification of species in previous studies. We believe that the latter is most likely the case, because we surveyed the area carefully over a long period of time. In addition, –10 2 specimens of Eryx sp. from the area were studied that –10 –5 0 5 10 differed from all other Iranian species of the genus in Functon 1 several morphological characters. They were closest to E. Figure 4. Discriminant analysis of female samples. The first miliaris morphologically, and so we can identify them as function, in which PIN has the most effect, separatesE. tataricus Eryx cf. miliaris. However, the taxonomic status of these and E. miliaris from Eryx sp. specimens remained unclear in our study. We suggest

354 ESKANDARZADEH et al. / Turk J Zool conducting a comprehensive work on this enigmatic genus The second supralabial scale is lower than the third. Three in Khorasan, or, better, in the entire country, to clarify the or 4 scales are located posterior to internasal. Width exact taxonomic status of species and the phylogenetic of interocular space is equal to or smaller than distance relationships within the genus, including this problematic between posterior edge of eye and corner of mouth; eyes population, using molecular markers. Thus, until that directed sideways and somewhat upward. Three or 4 scales time, we consider all the populations of Eryx in Khorasan are present between eyes and nasal area. In most cases, provinces as 3 distinct taxonomic entities at species level: the ventral surface of body is a light cream color with a E. miliaris, E. jaculus, and Eryx cf. miliaris. Diagnostic thick black line that extends along the body closely with features of these taxa are presented below. the head with orange color. Ventral surface is covered 4.1. E. jaculus with some black spots without orange in some specimens. The characters based on the studied samples of Khorasan The undersurface of the head has a few black spots in provinces are as follows. The second supralabial scale is comparison with E. jaculus. On the undersurface of ​​the higher than the third. Two scales posterior to internasal. neck, orange spots with a black middle line are visible. Width of interocular space is more than 1.2 times greater Dense black spots are seen on the lateral surfaces. The than distance between posterior edge of eye and corner of dorsal surface is a mixture of 2 colors, dark brown and mouth; eyes directed sideways. Three scales between eye cream; in most cases, an orange color is seen in the cream and nasal area. The ventral surface is cream-colored with areas. Edge of the brown area is surrounded by a black black spots more or less interconnected; in some cases strip. In some cases, the orange color is not seen (Figure 6). they are seen as linear lines. The undersurface of the head 4.3. Eryx cf. miliaris has scattered spots. Some spots are seen on the lateral We examined only 2 samples. One is from Ferdowsi surfaces. Dorsal surface is an olive or brown color, with University of Mashhad with unclear locality, and the other, scattered black or brown spots that connect as transverse which was collected from Neyshabour, is from Hakim and diagonal lines in most samples (Figure 5). Sabzevari University. Similar to E. miliaris, the second 4.2. E. miliaris supralabial scale is lower than the third, and width of The characters based on the studied samples interocular space is smaller than the distance between of the Khorasan provinces are as follows. posterior edge of eye and corner of mouth. Four scales present between eye and nasal area. Two scales located posterior to internasal. The color and pattern are similar to those of E. miliaris. The characters of this new population are generally similar to E miliaris; a more comprehensive study is needed to clarify

Figure 5. The general view of E. jaculus: a) dorsal and b) ventral Figure 6. The general view of E. miliaris: a) dorsal and b) ventral view. view.

355 ESKANDARZADEH et al. / Turk J Zool its real taxonomic position and phylogenetic relationship 2a. Two scales posterior to internasals …. Eryx cf. miliaris using morphology, ecology, and molecular traits. 2b. Three or 4 scales posterior to internasals….E. miliaris

5. Key to the species of Eryx in northeast Iran Acknowledgments 1a. Width of interocular space more than 1.2 times We gratefully thank Dr Haji Gholi Kami, Soheila Shafiei, greater than distance between posterior edge of eye and members of the Zoological Museum of Ferdowsi and corner of mouth……………………….E. jaculus University of Mashhad, the Zoological Museum of Hakim 1b. Width of interocular space equal or smaller than Sabzevari University, and the Zoological Museum of distance between posterior edge of eye and corner of Golestan University for providing the specimens for this ...………………....……...... ………..………………2 research.

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