Iran. J. Ichthyol. (March 2019), 6(1): 54-64 Received: December 7, 2018 © 2019 Iranian Society of Ichthyology Accepted: March 12, 2019 P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi: 10.22034/iji.v6i1.404 http://www.ijichthyol.org Research Article Interspecific morphological variation among members of the genus Cyprinion Heckel, 1843 (Teleostei: Cyprinidae) in Iran, using landmark- based geometric morphometric technique Manoochehr NASRI1, Soheil EAGDERI*2, Hamid FARAHMAND2, Hasan NEZHADHEYDARI2 1Department of Fisheries Science and Engineering, Faculty of Agriculture and Natural Resources, Lorestan University, Khorramabad, Iran. 2Department of Fisheries, Faculty of Natural Resources, University of Tehran, Karaj, Iran. *Email: [email protected] Abstract: Morphological variation among Cyprinion species in Iran was studied using landmark- based geometric morphometrics. A total of 848 specimens were caught from five basins throughout Iran during 2013-2015. A total number of 15 landmark-points were defined and digitized on 2D pictures. Despite of intraspecific differences among Cyprinion fishes, CVA divided all populations into three major groups viz. C. macrostomum, C. kais and C. tenuiradius (group X), C. watsoni and C. microphthalmum (group Y) and C. milesi (group Z). According to the results, C. macrostomum group can be distinguished from other groups by having deeper body, least head length and depth, the longest dorsal-fin base and least caudal peduncle length. Cyprinion milesi has the longest anal-fin base, least body depth, deepest caudal peduncle and longest head. Cyprinion watsoni has the longest caudal peduncle and least dorsal-fin base and C. microphthalmum is distinguished by having the least caudal peduncle length and depth. The main reasons of observed intraspecies morphological diversity among the fishes is proposed to be some phenotypic plasticity related to climatic and zoogeographical factors. The results suggest that the most possible factor influencing morphological variations among Iranian Cyprinion species is habitat related differences. Keywords: Cyprinidae, Fish distribution, Freshwater, Phenotypic plasticity, Zoogeography. Citation: Nasri, M.; Eagderi S.; Farahmand, H. & Nezhadheydari, H. 2019. Interspecific morphological variation among members of the genus Cyprinion Heckel, 1843 (Teleostei: Cyprinidae) in Iran, using landmark-based geometric morphometric technique. Iranian Journal of Ichthyology 6(1): 54-64. Introduction C. macrostomum and C. kais in Iraq based on some Iran locates in the southern Palearctic biogeographic morphometric characters. Nasri (2008) revealed region bordering the Oriental and Ethiopian regions some morphological differences between C. macro- and is considered as an exchange region for stomum and C. kais in Iranian part of the Tigris river freshwater fishes (Coad 1996, 2019) with 297 species drainage. Banarescu & Herzig-Straschil (1995) in 109 genera and 30 families (Esmaeili et al. 2018; distinguished C. kais, C. macrostomum and C. tenui- Nasri et al. 2018). Among them, the genus Cyprinion radius based on dorsal-fin rays, mouth form and includes six species viz. C. kais, C. macrostomum, lateral line scales. Nasri et al. (2018) studied the C. microphthalmum, C. milesi, C. tenuiradius and morphological diversity of the Iranian Cyprinion C. watsoni found in exorheic and endorheic basins of using traditional morphometric method. Some other Iran, including the Hormuz, Makran, Mashkid, valuable studies e.g. osteological (Aydin et al. 2008; Tigris, Persis, Maharlu, Sistan, Jazmurian, Kerman Nasri et al. 2013b) and molecular (Daştan et al. 2012) and Lut (Keivany et al. 2016; Esmaeili et al. 2018). works are also performed on some member of the Despite their wide distribution in Iran, there are few genus Cyprinion. morphological studies mainly based on traditional Study of the body shape is a routine method for methods (Nasri et al. 2018). Kafuku (1969) compared understanding many aspects of fish biology such as 54 Nasri et al.- Interspecific morphological variation among Cyprinion Table 1. The sampling stations of Cyprinion fishes used in this study. Num Symbol Species Longitude Latitude Basin Province River ber A C. kais 40 47°13'32"E 33°16'31"N Tigris Ilam Saimare (Talkhab) B1 C. macrostomum 32 45º55'15"E 33º30'12"N Tigris Ilam Godarkhosh B2 C. macrostomum 44 46°40'54"E 33°46'26"N Tigris Ilam Homeil B3 C. macrostomum 44 47°13'08"E 33°43'09"N Tigris Ilam Saimare (Sarcham) B4 C. macrostomum 45 46°56'06"E 33°21'47"N Tigris Ilam Saimare (Kolm) B5 C. macrostomum 54 47°13'32"E 33°16'31"N Tigris Ilam Saimare (Talkhab) B6 C. macrostomum 43 47°44'07"E 33°14'08"N Tigris Lorestan Kashkan (Moorani) B7 C. macrostomum 34 48°19'06"E 32°41'00"N Tigris Khuzestan Dez Kohgiluyeh and C1 C. tenuiradius 34 51°15'01"E 30°19'06"N Tigris Shiv Boyer-ahmad C2 C. tenuiradius 64 51°35'00"E 29°45'00"N Persis Fars Shapoor C3 C. tenuiradius 34 51°27'19"E 29°13'57"N Persis Bushehr Faryab C4 C. tenuiradius 31 51°47'18"E 28°53'18"N Persis Bushehr Dasht-e Palang C5 C. tenuiradius 43 53°09'25"E 28°27'12"N Persis Fars Ghareaghach D1 C. watsoni 30 57°41'33"E 28°41'08"N Persis Kerman Halil Sistan and D2 C. watsoni 37 60°43'02"E 27°12'48"N Jazmooryan Sarzeh Aqueduct Baluchestan Sistan and E C. milesi 32 61°15'35"E 26°37'53"N Makran Sarbaz Baluchestan F1 C. microphthalmum 43 54°40'09"E 27°12'28"N Hormoz Hormozgan Ilood spring F2 C. microphthalmum 35 56°50'29"E 27°32'27"N Hormoz Hormozgan Shamil F3 C. microphthalmum 31 57°25'15"E 26°50'27"N Jazmooryan Hormozgan Kahnuj F4 C. microphthalmum 29 57°44'57"E 26°5'57.3"N Makran Hormozgan Kash Sistan and F5 C. microphthalmum 33 61°15'35"E 26°37'53"N Makran Sarbaz Baluchestan Sistan and F6 C. microphthalmum 36 61°35'27"E 26°50'36"N Makran Nhang Baluchestan resource management, evolution, behavior, and supplementary work of Nasri et al. (2018). ecology (Rohlf & Slice 1990; Marcus et al. 1996). For instance, geometric morphometric methods Materials and Methods revealed some phenotypic plasticity evidences A total of 848 Cyprinion specimens were collected related to dam construction in Capoeta gracilis from 22 stations in five inland water basins of Iran (Heidari et al. 2013). Furthermore, this method were using electrofishing device during 2013-2015 (Table well-utilized for intraspecific morphological 1, Fig. 1). The specimens were preserved in 4% comparison in Alburnus chalcoides (Mohadasi et al. buffered formaldehyde after anesthesia in clove oil 2013), A. filippii (Jalili et al. 2015) and interspecific extract 1%. The fishes were identified based on comparison of the genus Alburnus (Khataminejad et Keivany et al. (2016) and Coad (2019). The left side al. 2013). Since, geometric morphometric method of fishes were photographed using a digital camera can detect little differences due to its higher detection (Kodak EasyShare Z650 with 6 MP resolution), and ability (Bookstein 1991; Rohlf & Marcus 1993; to extract data of the body shape, 15 homologous Adams et al. 2004), this work aimed to study the landmark-points (Fig. 2) were defined and digitized morphological variations and phenotypic plasticity using tpsDig2 (Rohlf 2010). Generalized Procrustes among the Iranian members of the genus Cyprinion Analysis (GPA) was used to remove non-shape data, using landmark-based geometric morphometrics as a including size, position and direction of the body 55 Iran. J. Ichthyol. (March 2019), 6(1): 54-64 Fig.1. Sampling stations of Genus Cyprinion. (A- C. kais; B- C. macrostomum; C- C. tenuiradius; D- C. watsoni; E- C. milesi; F- C. microphthalmum). Fig.2. The 15 homologous landmark points representing fish body shape. (1- snout tip; 2- the junction of the head and trunk; 3- the origin of the dorsal fin; 4- the insertion of the dorsal fin; 5- the upper edge of caudal fin base; 6- posterior body extremity; 7- the lower edge of caudal fin base; 8- the insertion of the anal fin; 9- the origin of the anal fin; 10- the ventral end of the gill slit; 11- the lower margin of the orbit; 12- the center of eye; 13- the upper margin of orbit; 14- the end of the head; 15- dorsal origin of the pectoral fin). shape data. can be assumed as Kendall shape coordinates (Rohlf Correlations between the procrustes and tangent 1998). The data were analyzed using canonical shape distances were calculated using tpsSmall 1.33 variant analysis (CVA) with P-value obtained from (Rohlf 2015) to certify if the procrusted coordinates permutation test with 10,000 replications for 56 Nasri et al.- Interspecific morphological variation among Cyprinion Table 2. The Classification matrix showing the numbers of individuals that were correctly classified (Bold values indicate correct classifications). Fig.3. Scatter plot and shape deformation of the studied Cyprinion populations based on the first two canonical variant functions. classification functions and to assign individual analysis (CA) as a complement to CVA by adopting specimens to putative populations and cluster the Euclidean square distance as a measure of 57 Iran. J. Ichthyol. (March 2019), 6(1): 54-64 Table 3. Pairwise Mahalanobis distances for 22 populations of Cyprinion from Iran. dissimilarity. The Mahalanobis distances were depth. The CV2 is related to the orbital diameter, extracted in CVA analysis to explore distance of predorsal distance, caudal peduncle length, anal-fin morphological differences between the studied base length and body depth. In group X, the orbital populations. The shape differences of the studied diameter and snout length are greater and caudal populations in relation to consensus configuration peduncle depth and length are lower. The members were visualized using deformation grids presented of the group Y are largely overlapped morpho- along with CA.