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Geographic Variation in Morphology of the Genus Agamura Blanford, 1874 in Iran

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Geographic Variation in Morphology of the Genus Agamura Blanford, 1874 in Iran Amphibia-Reptilia 38 (2017): 449-459 Geographic variation in morphology of the genus Agamura Blanford, 1874 in Iran Seyyed Saeed Hosseinian Yousefkhani1, Mansour Aliabadian1,2,∗, Eskandar Rastegar-Pouyani3, Jamshid Darvish1,2 Abstract. The genus Agamura was previously known from four species occurring on the Iranian Plateau, but was recently revised as a monotypic genus that excluded three species. In the present study, we examined different populations of Agamura persica morphologically. We found that A. persica shows geographic variation with respect to two groups, with the eastern population considered as A. cruralis and A. persica proposed for the western population. Separation between the two populations of A. persica was verified based upon ANOVA results for many morphological characters, including Head Height (HH), Interorbital distance (IO), Forelimb length (FLL), Number of scales across widest part of abdomen (NSA), Loreal scales (LOS), and Number of scales that separate two adjacent tubercles (NTV). Principal Component Analysis (PCA) and Discriminant Function Analysis (DFA) based on metric and meristic characters confirmed the revision of the genus Agamura. Multivariate analysis indicated that all studied OTUs were assigned to the correct classification and have significantly different morphological characters. Keywords: Agamura cruralis, cryptic divergence, Cyrtopodion, Iranian Plateau, Rhinogecko. Introduction and Cyrtopodion hormozganum (Nazarov, Bon- darenko and Rajabizadeh, 2012). In these stud- The Iranian Plateau is one of the richest ar- ies, several characters were given that distin- eas for reptiles in southwestern Asia (Rastegar- guish these species from the genus Agamura, Pouyani et al., 2015). Many species groups including the presence of a keel on the dorsal and genera have diversified in this region, tubercles, the presence of dark spots on the dor- including the genera Tropiocolotes and Cyr- sum, the number of scales across the venter, and topodion (Bauer et al., 2013; Krause et al., others. 2013; Freitas et al., 2016). Diversification of The genus Agamura is one of the gekkonid Gekkonids has been studied extensively and genera occurring in Western Asia (on the Ira- several complexes have been identified within nian Plateau). The genus was regarded at one them. Recently, several species were described time as having four species: A. persica, A. agamuroides Cyrtopodion from the group as gastropholis, A. femoralis, and A. misonnei kiabi Cyrtopodion (Ahmadzadeh et al., 2011), (Szczerbak and Golubev, 1996), but Anderson golubevi Cyrtopodion persepolense , (Nazarov, (1999) retained only the type species, Aga- Ananjeva and Rajabizadeh, 2009), Cyrtopodion mura persica. Anderson (1999) mentioned that sistanense (Nazarov and Rajabizadeh, 2007), Rhinogecko differs from Agamura in several characteristics such as mental shields and nasal scales that are sharply swollen or tube-like. 1 - Department of Biology, Faculty of Sciences, Ferdowsi University of Mashhad, Mashhad, Iran Additionally, Anderson (1999) placed A. gas- 2 - Research Department of Zoological Innovations tropholis in the genus Cyrtopodion, with which (RDZI), Institute of Applied Zoology, Faculty of it shares a pigmented peritoneum. According to Sciences, Ferdowsi University of Mashhad, Mashhad, Macey et al. (2000), the genus Cyrtopodion is a Iran 3 - Department of Biology, Faculty of Science, Hakim monophyletic group. Cervenka, Kratochvil, and Sabzevari University, Sabzevar, Iran Frynta (2008) later confirmed the monophyly ∗Corresponding author; e-mail: [email protected] of Cyrtopodion. Anderson (1999) indicated that Downloaded from Brill.com10/04/2021 09:37:33AM via free access © Koninklijke Brill NV, Leiden, 2017. DOI:10.1163/15685381-00003129 450 S.S. Hosseinian Yousefkhani et al. different populations of A. persica exhibit dif- sized gravel (pers. observation). Endemicity in ferentiation in color pattern. He divided them angular-toed geckos on the Iranian plateau sug- into two subspecies differentiated by patterns gests that there are many isolated populations of transverse bars: A. persica persica (the west- related to this group, but a comprehensive in- ern population in central and eastern Iran) with vestigation is needed. three transverse bars and A. persica cruralis In the present study, we aimed to examine (the eastern population limited to southeast Iran, morphological characteristics of all populations Afghanistan and Pakistan) with five transverse of A. persica from the Iranian Plateau. The main bars. Smith (1935) had earlier synonymized A. goal of the study was to test the validity of the persica and A. cruralis, but found some varia- two OTUs of A. persica in the Iranian Plateau. tion among them. For example, Smith (1935) re- ported the presence of large postmental scales in Material and methods Baluchistan specimens that suggested the prob- able presence of A. p. cruralis. Szczerbak and We examined 30 specimens representing A. persica (Ap- pendix 1). These specimens were collected from May 2011 Golubev (1996) indicated that these variations to July 2016 on the Iranian Plateau (fig. 1; exact locations need more consideration, but could not make are provided in Appendix 1). All specimens were preserved in 75% ethanol and deposited in the Sabzevar University the comparison themselves due to lack of suf- Herpetological Collection (SUHC). Twelve mensural and ficient materials. 15 meristic characters (table 1) were recorded from these In respect to habitat, the region inhabited by specimens. Mensural characters were measured using dig- ital calipers with 0.01 mm accuracy and meristic charac- A. persica is different in central and eastern Iran. ters were measured using an Olympus loupe. Because we A. p. cruralis can be found in very rocky ar- combined all samples to examine geographic variation, we removed sexually dimorphic characters from the analyses. eas in eastern Iran, but the habitat of A. p. per- Based upon t-test analysis of all characters, only preanal sica in central Iran is characterized by small- pores were distinguished as a sexually dimorphic character. Figure 1. Map of Iran and sampling localities of all examined species in this study. Downloaded from Brill.com10/04/2021 09:37:33AM via free access Morphology of the genus Agamura 451 Table 1. All metric and meristic characters examined in this study, including their definitions. Character Definition SVL Snout-Vent Length HL Head length (from tympanum to tip of rostrum) HW Head width (maximum distance between tympani) HH Head height (straight-line dorsiventral distance between apex of head and ventral surface of lower jaw) SL Snout length (eye–nostril distance) IO Interorbital distance EED Distance from eye to tympanum WPW Widest lateral dimension of abdomen FHD Distance between forelimb and hindlimb FLL Forelimb length HLL Hind limb length NSS Number of supralabial scales NIS Number of infralabial scales ION Number of interorbital scales (except palpebral fold scales) LOS Loreal scales (between eye and nostril, in a single longitudinal row) POS Number of scales from postnasal to occiput (in a single longitudinal row) NPS Number of postmental scales NSA Number of scales across widest part of abdomen NPV Number of scales between postmental scales and vent NTW Number of dorsal tubercles at widest lateral dimension of thorax NTP Number of tubercles in a paravertebral row (from neck to hindlimb) NAT Number of scales around a tubercle (forming a circumference) NTV Number of scales that separate two adjacent tubercles (longitudinal) NTH Number of scales that separate two adjacent tubercles NPP Number of preanal pores SLT Subdigital lamella under fourth toe NUT Number of scales under the tail Statistical analyses were performed using SPSS 16.0 Results and all data were evaluated for normality prior to anal- ysis. Mensural (parametric) and meristic (non-parametric) Color and pattern characters were evaluated using the Shapiro-Wilks and the Kolmogorov-Smirnov tests, respectively. H0 in both tests Based on data from all examined specimens, is a normal distribution, so the dataset is normally dis- color pattern within A. persica is variable and tributed if the values are not significantly different from H 0 may be dependent upon ground color (pers. (P>0.05). Intraspecific analyses of A. persica were per- formed according to two major operational taxonomic units obs). In the western individuals, the entire body (OTUs). We selected these OTUs according to geographic color is light. However, the dorsal color is dark barriers. in the eastern individuals. The number of dorsal After testing for normality, a one-way analyses of vari- ance (ANOVA) was used to examine the characters between crossbars was five in all examined specimens populations of A. persica to verify the morphological diver- of A. persica from throughout the range and all gence between them and to determine whether they fit An- specimens had tails with five to eight black bars derson’s (1999) taxonomic interpretation. The null hypoth- (fig. 2a, b). esis in an ANOVA is that all data are distributed uniformly, so a significant result (P<0.05) indicates that charac- ters are not distributed equally and are therefore differenti- Scalation ated between groups. Principal Component Analysis (PCA) and Canonical Variate Analysis (CVA) were conducted on Differences between populations of A. persica the significant characters identified using the ANOVA. The on the Iranian plateau were identified for some PCA was used to determine the variation between
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