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http://journals.tubitak.gov.tr/zoology/

Turkish Journal of Zoology Turk J Zool(2013) 37: 685-692© TÜBİTAKdoi:10.3906/zoo-1204-30

Comparison of geographic variations in Typhlops vermicularis (Merrem, 1820)(Ophidia: Typhlopidae) from the Iranian plateau with Turkey and Turkmenistan

Mohadeseh AFROOSHEH1,*, Nasrullah RASTEGAR-POUYANI1, Seyed Kamran GHOREISHI2, Haji Gholi KAMI3

1Department of Biology, Faculty of Science, Razi University, Kermanshah, Iran2Department of Statistics, Faculty of Science, Razi University, Kermanshah, Iran3Department of Biology, Faculty of Sciences, Golestan University, Golestan, Iran

* Correspondence: m.afroosh@gmail.com

1. IntroductionAbout 2700 snake species are within the suborder Serpentes, which is divided into 2 infraorders, Alethinophidia (typical snakes, true snakes) and Scolecophidia (blindsnakes, wormsnakes, and threadsnakes) (Rage, 1987; Greene, 1997; Pough et al., 1999). Although the scolecophidians are an important group based upon their basal position and number of taxa, most research has centered on taxonomic descriptions, identification keys, and range extensions. The other peculiarities of these snakes, such as diversity, phylogeny, biogeography, and ecology, are poorly known due to their poor dispersal ability, restricted habitat, and very small (for vertebrates) size, all of which reduce their chance of preservation (Wallach, 1998).

Within scolecophidians, blindsnakes (Typhlopidae, approximately 240 species) occur in Africa (south of the Sahara Desert), Madagascar, southern Asia, South and Central America, and Australia, while only 1 representative, Typhlops vermicularis (Greek blindsnake), is found in southeastern Europe (Başoğlu and Baran, 1980; Baran and Atatür, 1998; Cox et al., 2006; Adalsteinsson et al., 2009; Vidal et al., 2010, Kornilios, 2012).

The analysis of geographic variation among populations of some reptiles has already been carried out in the Iranian plateau (e.g., Rastegar-Pouyani et al., 2011). T. vermicularis

is an ideal taxon for such studies, and for testing the effects of changes in topography and ecology on species with poor distributional capabilities.

The Iranian plateau is a connecting crossroad between southeastern Europe and northwestern India and western Pakistan on the one hand, and central Asia and northeastern Africa and Arabia on the other, so zoogeographically it is one of the most important and most complex areas in southwestern Asia. The fauna of Iran, and especially reptile fauna, is the result of interaction between vernacular species of the Iranian plateau and nonvernacular species of North Africa, central and southern Asia, and Europe. These nonvernacular faunal elements have entered the Iranian plateau, and most are widely distributed and occupy various ecological niches (Rastegar-Pouyani, 1991).

Since the last study on this species was that of Latifi (2000), which used old reports from more than 3 decades ago, it seems that this species needs new research to update the older data. The available research studies on T. vermicularis are very few, and, in fact, according to the IUCN, research actions towards its biology, ecology, population numbers, and geographic range are needed.

The main objectives of this study are: 1) analysis of geographic variation in T. vermicularis throughout the

Abstract: Typhlops vermicularis (Merrem, 1820) is one of the cosmopolitan species of the genus Typhlops that occupies a vast range of different localities from North Africa to western Afghanistan. In this study, different populations of T. vermicularis were investigated to clarify if there was any degree of geographic variation in this taxon throughout the studied range. For this purpose, 103 specimens of this taxon belonging to 12 populations from all over Iran, Turkey, and Turkmenistan were studied. Morphometric characters were examined by ANOVA and principal component analysis. Results showed that despite the vast distribution range of this species, there is no significant difference among populations of T. vermicularis in various regions of the Iranian plateau and between populations from within the plateau and outside of it.

Key words: Typhlops vermicularis, geographic variation, morphology, distribution, statistical methods, Iran

Received: 29.04.2012 Accepted: 20.05.2013 Published Online: 04.10.2013 Printed: 04.11.2013

Research Article

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Iranian plateau and comparison of its data with data of the species from Turkey and Turkmenistan; 2) to find any isolated populations of T. vermicularis within its distribution range; and 3) to shed new light on the role of the Zagros and Alborz orogenies in isolation and subsequent evolution of various populations of T. vermicularis.

2. Materials and methodsIn this study, 103 specimens of T. vermicularis were examined (see Appendix for material examined). Of these, 10 specimens were collected during fieldwork from 2008 to 2009 from the western and central regions of Iran. The rest of the specimens were borrowed from the following: Iran National Natural History Museum (INNHM; Tehran, Iran), Razi University Zoological Museum (RUZM; Kermanshah, Iran), Zoology Museum of Gorgan University (ZMGU; Gorgan, Iran), and Göteborgs Naturhistoriska Museum (GNM; Göteborg, Sweden). Collectively, 40 specimens from Golestan, 18 from Kermanshah, 6 from Markazi, 3 from Lorestan, 4 from Razavi Khorasan, 3 from Mazandaran, 2 from West Azerbaijan, 3 from Khuzestan, 3 from Kerman, 2 from Tehran Province, 16 from Turkey, and 3 from Turkmenistan were examined. All specimens from Turkey and Turkmenistan were borrowed from GNM (the material examined is listed in the Appendix) (Figure 1). The measured characters (in millimeters) are as follows: LOA = total length, TL = tail length, MTD =

midtail diameter, ABD = anterior body diameter, MBD = midbody diameter, PBD = posterior body diameter, HW = head width, HL = head length, HW/HL = head width/head length, TL/LOA = tail length/total length, LOA/MBD = total length/midbody diameter, TL/MTD = tail length/midtail diameter, RW = rostral width, RL = rostral length, VL = ventral–rostral length, TSR = total scale row, SC = subcaudals, ASR = anterior scale rows, MSR = midbody scale rows, PSR = posterior scale rows, MPSR = mid- plus posterior-body scale rows, AMPSR = anterior- plus mid- plus posterior-body scale rows, SL = supralabials, IL = infralabials, INS = inferior nasal suture, SIP = supralabial imbrication pattern, Po = postoculars, Pari = parietals, DOR = dorsal pattern (Afroosheh, 2009). Most characters like TSR or HW were measured under a Wild Heerbrugg binocular microscope. All of the specimens were studied without differentiating sex.

Sexual dimorphism in the scolecophidians has not been studied in depth and there was no clear pattern of sexual dimorphism in the few such studies that have been done (Yadgarov, 1971; Dixon and Hendricks, 1979; Perry, 1985). However, sex in typhlopids can be determined through dissection. Since we did not have permission to dissect the borrowed specimens, no sex determination was carried out.

To examine different populations, first, all the studied characters were divided into 2 categories: metric (LOA, TL,

Figure 1. The distribution map of studied localities of Typhlops vermicularis: 1 = Turkey, 2 = West Azerbaijan Prov., 3 = Kermanshah Prov., 4 = Lorestan Prov., 5 = Khuzestan Prov., 6 = Mazandaran Prov., 7 = Golestan Prov., 8 = Turkmenistan, 9 = Razavi Khorasan Prov., 10 = Kerman Prov., 11 = Markazi Prov., 12 = Tehran Prov.

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MTD, ABD, MBD, PBD, HW, HL, RW, RL) and meristic (TSR, SC, ASR, MSR, PSR, SL, MPSR, AMPSR) characters. All characters were standardized and normalized. In this study, 12 localities from different provinces were explored. To discover any degree of geographic variation among various populations, the ANOVA test was employed. Subsequently, a multivariate analysis procedure (including principal component analysis [PCA]) was used to determine the form and pattern of variations. The existence of significant differences among various populations was tested by analysis of variance (ANOVA) followed by the post-hoc least significant difference (LSD) test. The significance level for all of the tests was set at P ≤ 0.05. All statistical analyses were carried out using the SPSS 16 and S-PLUS 8 packages.

3. Results3.1 Results of analyses for metric characters3.1.1. Univariate analysisIn the morphological analysis, descriptive statistical analysis of individuals of 12 populations and then one-way ANOVA (LSD test) were used to explore the patterns of morphological variation among different populations of T. vermicularis for the various morphometric characteristics used in this study.

Results of descriptive and ANOVA pairwise analyses revealed that MTD, ABD, and HL showed statistically significant differences (P < 0.05) among individuals of the 12 studied populations; their means in locality 2 were higher than those of locality 9 (Table 1).

Results of analysis of covariance (ANCOVA) for relative sizes for metric traits among pooled samples of the 12 studied populations of T. vermicularis showed that MTD, ABD, and HL were significantly different among the 12 populations (Table 2), in which MTD was highest (3.70) in locality 9 and was lowest (0.83) in locality 2. For other characters, the mean of ABD was highest (3.89) in locality 9 and lowest (2.11) in locality 2, and the mean of HL was highest (3.62) in locality 9 and lowest (1.87) in locality 2.3.1.2. Multivariate analyses PCA was employed to explore the patterns of geographic variation. The first 3 components accounted for nearly 100% of the total variance, in which the first component showed 98% of the total variance and some characters such as LOA, MBD, and PBD had the highest values. The rate loadings of each character on a particular component for these 3 characters and for all other characters in 3 components are shown in Table 3. Based on this analysis, as shown in Figure 2, in the ordination of 12 populations along the first 3 components, the group centroids of populations are not distinct and the 12 studied populations are not separated from each other significantly.

3.2 Results of analyses for meristic characters3.2.1. Univariate analysesAs with metric characters, based on descriptive and ANOVA analyses, just one meristic character, MPSR, was significantly different among populations (P < 0.05). The means of this character were highest in locality 2 (47.50) and lowest (42.33) in locality 6 (Table 4).

Results of ANCOVA for relative sizes for meristic traits among pooled samples of the 12 studied populations of T. vermicularis showed that MPSR was significantly different among the 12 studied populations (Table 5); its means was highest (47.50 mm) in locality 2 and lowest (42.33 mm) in locality 6.3.2.2. Multivariate analyses PCA was employed to explore the patterns of geographic variation. The first 3 functions explained almost 100% of the total variance, in which the first function showed 98% of the total variance and some characters such as TSR, AMPSR, and MPSR had the highest values. The second component accounted for 1.1%, in which AMPSR, MPSR, and PSR were mainly responsible for this variation. The third function loaded 0.3% of the total variance. The rate of loading of each character in a particular function is shown in Table 6. Ordination of the 12 studied populations along the first 2 components is shown in Figure 3. Obviously, the group centroids of populations are not clear and the 12 studied populations are not separated from each other significantly.

4. DiscussionSemispecies or geographic populations are adaptive aspects of a species in its vast distribution range (Mayer and Ashlock, 1991). Results of this study showed that the one-way ANOVA and ANCOVA analyses did not reveal significant differences among the studied populations at the level of 0.05. Our results show that there is no significant difference among populations of T. vermicularis in various regions of the Iranian plateau and between populations inside and outside of the plateau. In other words, the studied populations were almost homogeneous, not showing distinct patterns of geographic variation.

Patterns of geographic variation in individual characters (i.e. some vary with latitude, some with altitude, and some with longitude) are consistent with the hypothesis that ecogenetically caused geographic variation may result in lower intercharacter congruence than phylogenetically caused geographic variation (Thorpe and Baez, 1993).

According to Kornilios et al. (2012), the clade of T. vermicularis can be split into 2 distinct clades, 1 clade in Jordan/South Syria and 1 in other parts of its distribution (Greece, Turkey, Azerbaijan, Iran). According to the molecular data, populations from western Iran, northeastern Turkey, and Transcaucasia make up a monophyletic group (Kornilios et al., 2011, 2012); the morphometric

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Table 1. Descriptive analysis and one-way ANOVA (LSD test) for metric characters (ranges, mean ± standard deviation) of pooled samples of 12 studied populations of Typhlops vermicularis. 1 = Turkey, 2 = West Azerbaijan Prov., 3 = Kermanshah Prov., 4 = Lorestan Prov., 5 = Khuzestan Prov., 6 = Mazandaran Prov., 7 = Golestan Prov., 8 = Turkmenistan, 9 = Razavi Khorasan Prov., 10 = Kerman Prov., 11 = Markazi Prov., 12 = Tehran Prov.

Characters1 (n = 16) 2 (n = 2) 3 (n = 18) 4 (n = 3)

F-valueP-valueRange Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean± SD

LOA 96.79 –247.92 1.78 ± 38.66 103.11–167.23 1.35 ± 45.33 113.92–236.94 1.83 ± 35.95 202.54–231.37 2.14 ± 15.04 1.67 0.09

TL 1.92–3.84 3.05 ± 0.63 0.60–3.50 2.05 ± 2.05 2.25–4.39 3.35 ± 0.71 3.49–4.27 3.82 ± 0.40 1.23 0.28

MTD 1.82–3.73 2.76 ± 0.52 0.83–1.98 0.83 ± 1.98 1.47–4.48 2.81 ± 0.82 2.88–3.18 3.04 ± 0.15 1.95 0.04

ABD 2.10–4.04 2.85 ± 0.45 1.44–2.78 2.11 ± 0.94 2.12–4.16 3.02 ± 0.63 2.69–4.69 3.60 ± 1.01 2.83 0.00

MBD 2.01–5.45 3.64 ± 0.93 1.80–3.17 2.48 ± 0.96 2.46–7.22 3.77 ± 1.19 3.82–5.32 4.57 ± 0.75 1.82 0.06

PBD 1.68–4.44 2.95 ± 0.72 1.28–2.36 1.82 ± 0.76 1.58–5.11 3.14 ± 0.89 2.29–3.54 2.87 ± 0.62 1.85 0.06

HW 1.92–3.60 2.66 ± 0.40 1.88–2.30 2.09 ± 0.29 2.02–3.47 2.74 ± 0.40 2.77–3.31 3.07 ± 0.27 1.79 0.07

HL 2.14–4.07 2.89 ± 0.48 1.58–2.16 1.87 ± 0.41 2.37–3.95 3.09 ± 0.47 2.50–3.23 2.95 ± 0.39 2.39 0.01

RW 0.75–1.61 1.16 ± 0.20 0.89–1.55 1.16 ± 0.18 0.88–1.50 1.15 ± 0.17 1.14–1.26 1.20 ± 0.08 1.15 0.34

RL 1.41–2.67 1.86 ± 0.33 1.39–2.40 1.89 ± 0.25 1.49–2.42 1.90 ± 0.27 1.80–2.09 1.94 ± 0.20 1.31 0.24

Characters5 (n = 3) 6 (n = 3) 7 (n = 40) 8 (n = 3)

F-valueP-valueRange Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD

LOA 183.3–241.28 2.09 ± 29.31 92.94–196.61 1.28 ± 5 8.63 90.94–277.88 1.75 ± 56.89 205.94–268.11 2.32 ± 26.60 1.67 0.09

TL 2.37–4.11 3.10 ± 0.90 1.65–3.19 2.17 ± 0.88 1.40–5.75 2.93 ± 1.02 3.03–4.54 3.54 ± 0.67 1.23 0.28

MTD 2.75–4.28 3.45 ± 0.77 1.53–3.51 2.27 ± 1.07 1–5.21 2.62 ± 0.99 2.83–3.45 3.22 ± 0.28 1.95 0.04

ABD 2.19–3.80 3.12 ± 0.83 1.47–2.36 1.83 ± 0.46 1.39–4.37 2.79 ± 0.72 3.22–4.40 3.73 ± 0.57 2.83 0.00

MBD 3.16–4.68 4.10 ± 0.82 1.88–4.20 2.79 ± 1.23 1.76–6.39 3.58 ± 1.25 4.35–6.05 4.98 ± 0.76 1.82 0.06

PBD 2.65–4.17 3.49 ± 0.77 1.59–3.25 2.17 ± 0.93 1.19–5.51 2.95 ± 1.07 3.60–4.55 3.99 ± 0.49 1.85 0.06

HW 2.54–3.28 2.96 ± 0.38 1.83–2.75 2.18 ± 0.49 1.73–4.07 2.69 ± 0.58 2.89–3.68 3.21 ± 0.38 1.79 0.07

HL 2.76–3.69 3.28 ± 0.47 2.11–3.22 2.56 ± 0.58 2.05–4.21 2.97 ± 0.58 2.86–4.00 3.45 ± 0.53 2.39 0.01

RW 1.08–1.36 1.24 ± 0.14 0.78–1.20 0.93 ± 0.23 0.78–1.79 1.15 ± 0.24 1.21–1.57 1.39 ± 0.15 1.15 0.34

RL 1.66–2.30 2.07 ± 0.35 1.26–2.06 1.54 ± 0.44 1.25–2.32 1.77 ± 0.32 1.93–2.35 2.13 ±0.22 1.31 0.24

Characters9 (n = 4) 10 (n = 3) 11 (n = 6) 12 (n = 2)

F-valueP-valueRange Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD

LOA 208.61–229.16 2.20 ± 8.88 97.92–189.3 1.49 ± 46.69 112.28–260 1.718 ± 50.20 151.37–185.7 1.685 ± 24.29 1.67 0.09

TL 2.19–4.10 3.11 ± 0.85 2.23–4.37 2.98 ± 1.20 2.06–3.58 2.78 ± 0.49 1.90–2.93 2.41 ± 0.72 1.23 0.28

MTD 3.30–4.42 3.70 ± .49 2.00–2.42 2.18 ± 0.21 1.50–3.31 2.26 ± 0.67 1.97-2.03 2 ± 0.04 1.95 0.04

ABD 3.62–4.25 3.89 ± 0.32 2.08–3.20 2.66 ± 0.56 1.78–4.75 2.94 ± 0.98 2.35–2.67 2.51 ± 0.22 2.83 0.00

MBD 4.43–6.04 5.07 ± 0.72 2.44–3.65 3.13 ± 0.62 1.88–5.46 3.71 ± 1.29 2.52–3.08 2.80 ± 0.39 1.82 0.06

PBD 3.76–5.19 4.40 ± 0.60 2.02–3.08 2.59 ± 0.53 1.46–4.86 2.87 ± 1.20 2.19–2.76 2.47 ± 0.40 1.85 0.06

HW 3.25–3.50 3.37 ± 0.10 2.09–3.05 2.70 ± 0.53 1.99–4.13 2.74 ± 0.72 2.36–2.62 2.49 ± 0.18 1.79 0.07

HL 3.48–3.80 3.62 ± 0.16 2.15–3.18 2.50 ± 0.58 2.54–3.32 2.82 ± 0.313 2.66–2.92 2.79 ± 0.18 2.39 0.01

RW 0.92–1.44 1.24 ± 0.22 0.87–1.11 0.99 ± 0.16 0.92–1.13 1.06 ± 0.08 1.10–1.16 1.13 ± 0.042 1.15 0.34

RL 1.80–2.16 2.03 ± 0.16 1.38–2.04 1.71 ± 0.46 1.66–2.11 1.92 ± 0.17 1.73–2.03 1.88 ± 0.21 1.31 0.24

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data reported here confirm this result. It seems that this diversification started in Anatolia and expanded west to Greece and east to the eastern part of Turkey and Iran. T. vermicularis’ biogeography and divergence were most

probably affected by the glacial periods, during which aridity increased dramatically, since warm and humid interglacials probably had a positive, rather than a negative, effect on its survival.

Table 2. Results of ANCOVA for relative size for metric characters among the pooled samples of 12 studied populations of Typhlops vermicularis.

Variables F-value P-value

LOA 1.67 0.09TL 1.23 0.28MTD 1.95 0.04ABD 2.83 0.00MBD 1.82 0.06PBD 1.85 0.06HW 1.79 0.07HL 2.39 0.01RW 1.15 0.34RL 1.31 0.24

Table 3. The component scores of the coefficient matrix in 12 populations of Typhlops vermicularis used in this study.

Characters Comp. 1 Comp. 2 Comp. 3 Comp. 4

LOA 0.999 –0.034 0.013 0.001TL 0.012 –0.058 –0.989 0.078MTD 0.015 0.395 –0.108 –0.735ABD 0.012 0.298 0.014 0.137MBD 0.020 0.677 –0.027 0.585PBD 0.017 0.509 –0.026 –0.297HW 0.010 0.143 0.036 0.027HL 0.010 0.102 0.058 0.025RW 0.004 0.029 0.048 0.013RL 0.006 0.035 0.022 0.0462

Figure 2. Ordination of PC1 against PC2 for metric characters in different populations of Typhlops vermicularis used in this study. 1 = Turkey, 2 = West Azerbaijan Prov., 3 = Kermanshah Prov., 4 = Lorestan Prov., 5 = Khuzestan Prov., 6 = Mazandaran Prov., 7 = Golestan Prov., 8 = Turkmenistan, 9 = Razavi Khorasan Prov., 10 = Kerman Prov., 11 = Markazi Prov., 12 = Tehran Prov.

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Table 4. Descriptive analysis and one-way ANOVA (LSD test) for meristic characters (ranges, mean ± standard deviation) of pooled samples of 12 studied populations of Typhlops vermicularis. 1 = Turkey, 2 = West Azerbaijan Prov., 3 = Kermanshah Prov., 4 = Lorestan Prov., 5 = Khuzestan Prov., 6 = Mazandaran Prov., 7 = Golestan Prov., 8 = Turkmenistan, 9 = Razavi Khorasan Prov., 10 = Kerman Prov., 11 = Markazi Prov., 12 = Tehran Prov.

Characters1 (n = 16) 2 (n = 2) 3 (n = 18) 4 (n = 3)

F-value P-valueRange Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD

TSR 357–440 3.90 ± 22.72 344–375 3.59 ± 21.92 350–421 3.90 ± 22.98 367– 401 3.83 ± 17.00 1.82 0.06

SC 7–11 9.31 ± 1.07 7–12 8.50 ± 2.12 7–12 9.94 ± 1.43 9–10 9.66 ± 0.58 1.66 0.09

ASR 22–24 23.06 ± 0.99 23–23 23 ± 0 22–24 23.22 ± 1 22–24 23 ± 1 1.31 0.23

MSR 22–24 22.93 ± 0.99 24–24 24 ± 0 22–24 22.77 ± 1 22–24 22.66 ± 1.15 1.35 0.21

PSR 20–24 21.43 ± 1.36 23–24 23.50 ± 0.70 20–24 21.77 ± 0.94 21–23 22 ± 1 1.76 0.07

SL 0.82–1.71 1.14 ± 0.23 0.66–1.60 1.22 ± 0.21 0.95–1.58 1.21 ± 0.17 1.07–1.31 1.19 ± 0.16 1.09 0.38

MPSR 42–48 44.37 ± 1.85 47–48 47.50 ± 0.70 42–48 44.55 ± 1.65 43–47 44.66 ± 2.08 1.90 0.05

AMPSR 64–72 67.44 ± 2.47 70–71 70.50 ± 0.70 64–72 67.78 ± 2.46 66–71 67.67 ± 2.88 1.86 0.06

Characters5 (n = 3) 6 (n = 3) 7 (n = 401) 8 (n = 3)

F-value P-valueRange Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD

TSR 388–428 4.07 ± 20.07 362–369 3.66 ± 23.78 340–436 3.89 ± 23.39 377–431 3.94 ± 24.83 1.82 0.06

SC 7–10 8.33 ± 1.52 7–10 8.33 ± 1.52 7–12 8.65 ± 1.40 7–11 8.65 ± 1.40 1.66 0.09

ASR 22–24 23.33 ± 1.15 22–22 22 ± 0 22–24 23.29 ± 0.92 22–24 23 ± 1.15 1.31 0.23

MSR 22–24 22.66 ± 1.15 21–22 21.66 ± 0.57 21–24 22.90 ± 1.04 22–24 22.50 ± 1 1.35 0.21

PSR 22–22 22.68 ± 1.04 20–22 20.66 ± 1.15 20–24 21.48 ± 0.97 20–22 21.50 ± 1 1.76 0.07

SL 1.08–1.36 1.24 ± 0.14 0.86–1.19 0.98 ± 0.18 0.68–1.80 1.21 ± 0.248 1.09–1.39 1.27 ± 0.13 1.09 0.38

MPSR 44–46 44.66 ± 1.15 41–44 42.33 ± 1.52 42–46 44.39 ± 1.68 42–46 44 ± 1.63 1.90 0.05

AMPSR 66–70 68 ± 20 63–66 64.33 ± 1.52 64–70 67.68 ± 2.36 66–70 67 ± 2 1.86 0.06

Characters9 (n = 4) 10 (n = 3) 11 (n = 6) 12 (n = 2)

F-value P-valueRange Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD

TSR 341–409 3.82 ± 30.20 351–400 3.80 ± 26.08 374–438 4.05 ± 22.48 433–438 4.35 ± 3.53 1.82 0.06

SC 7–9 8.25 ± 0.95 8–12 10 ± 2 7–11 9.17 ± 1.32 8–8 8 ± 0 1.66 0.09

ASR 22–24 23.25 ± 0.95 24–24 24 ± 0 24–24 24 ± 0 24–24 24 ± 00 1.31 0.23

MSR 22–24 22.50 ± 1 22–24 22.66 ± 1.15 22–24 23.66 ± 0.81 24–24 24 ± 0 1.35 0.21

PSR 22–24 22.50 ± 1 20–24 22 ± 2 22–24 22.66 ± 1.03 22–22 22 ± 0 1.76 0.07

SL 1.19–1.41 1.33 ± 0.10 1–1.08 1.04 ± 0.06 0.85–1.25 1.06 ± 0.14 0.93–1.15 1.04 ± 0.15 1.09 0.38

MPSR 44–46 45 ± 1.15 42–46 44.66 ± 2.30 44–48 46.33 ± 1.50 46–46 46 ± 0 1.90 0.05

AMPSR 341–409 3.82 ± 30.20 351–400 3.80 ± 26.08 374–438 4.05 ± 22.48 433–438 4.35 ± 3.53 1.86 0.06

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Despite the vast distribution range of T. vermicularis and occupation of different localities and habitats (e.g., different altitudes, humidities, or substrates), the high degree of homogeneity in the populations of this species is surprising. Here, a question arises: why is T. vermicularis morphologically so homogeneous and conservative

throughout its range? To answer this question, 2 scenarios are proposed here: 1) T. vermicularis is a conservative taxon in its morphological characters, and despite long-term occupation of different habitats, it has mainly remained a homogeneous taxon throughout the range; 2) after the last glacial period (about 10,000 years ago), the small surviving

Table 5. Results of the ANCOVA in relative size for meristic traits among pooled samples of 12 studied populations of Typhlops vermicularis.

Variables F-value P-value

TSR 1.83 0.06SC 1.66 0.09ASR 1.31 0.23MSR 1.35 0.21PSR 1.76 0.07SL 1.09 0.38MPSR 1.90 0.05AMPSR 1.86 0.06

Table 6. The component scores of coefficient matrix in 12 populations of Typhlops vermicularis used in this study.

Characters Comp. 1 Comp. 2 Comp. 3

TSR 0.997 –0.079 0.022SC –0.020 0.006 0.996ASR 0.020 0.202 0.042MSR 0.028 0.193 –0.061PSR 0.013 0.333 0.028SL 0.001 0.006 –0.003MPSR 0.041 0.526 –0.033AMPSR 0.061 0.727 0.009

Figure 3. Ordination of PC1 against PC2 for meristic characters in different populations of Typhlops vermicularis used in this study. 1 = Turkey, 2 = West Azerbaijan Prov., 3 = Kermanshah Prov., 4 = Lorestan Prov., 5 = Khuzestan Prov., 6 = Mazandaran, 7 = Golestan Prov., 8 = Turkmenistan, 9 = Razavi Khorasan Prov., 10 = Kerman Prov., 11 = Markazi Prov., 12 = Tehran Prov.

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populations in southern refugia (e.g., southern regions of the Iranian plateau) expanded their range northward and covered the whole present distribution range of the taxon, not having enough time for genetic and morphological diversification.

With consideration of the special mode of life in this snake (fossorial, insular, and not opportunistic), we are inclined to accept the first scenario.

AcknowledgmentsWe wish to thank Göran Nilson (Göteborg Natural History Museum, Göteborg, Sweden), Alireza Motesharei (Iran National Natural History Museum [INNHM]), Maysam Mashayekhi, and Behzad Fathinia for loan of material.

Appendix. Material examined. Typhlops vermicularis (n = 103)GNM- Re. ex. 1412, 3005, 6457, 6458, 6459, 6460, 6461, 6462, 6464, 6465 (n = 2), 6466, 6467 (n = 2), 6470 (n =

2): Turkey. 5974 (n = 2), 5975: Turkmenistan. 4420 (n = 2): Golestan Prov. Gorgan. 5741, 5751, 5776, 6618: Kermanshah Prov. Kermanshah. 5750: Kermanshah Prov. Eslamabad-e-Gharb. 6631 (n = 2): Lorestan Prov. Khoramabad. INNHM- 16: Tehran Prov. Kan. 1449: Lorestan Prov. 120, 113: West Azerbaijan Prov. Kaboudan Island. RUZM- TT10-5, -6, -7, -9, -10, -11, -16, -31, -32: Kermanshah Prov. Kermanshah. -21, -22: Kermanshah Prov. Ravansar. -17, -20: Kermanshah Prov. Gahvareh., -25, -26, -27, -28, -30: Markazi Prov. Arak. -14, -23, -29: Khuzestan Prov. Dezful. ZMGU- 1179: Kermanshah Prov. Guilan-e-Gharb. 29, 127, 266, 346, 348, 349, 355, 460, 461, 462, 463, 465, 466, 470, 490, 552, 676, 679, 680, 681, 682, 683, 684, 687, 688, 689, 885, 1023, 1085, 1277, 1519, 1593, 1604, 1606, 1607, 1608, 1621, 1622, 1625: Golestan Prov. Gorgan. 1245: Razavi Khorasan Prov. Neishabour. 2082, 2163: Razavi Khorasan Prov. Chenaran. 38: Razavi Khorasan Prov. Mashhad. 1270, 1271, 1623: Mazandaran Prov. Sari. 1997, 1998, 1999: Kerman Prov. Sirjan.

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