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 Heckel, 1843 (Teleostei: ) 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. 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. All analysis was performed using logically positioning between the groups X and Z. MorphoJ 1.06d (Klingenberg 2011) and PAST 2.17c The southeastern species, C. milesi (group Z) has (Hammer 2012) softwares. higher head depth, caudal peduncle depth and length, and anal-fin base length than those of western species Results i.e. groups X and Y (Fig. 3). The procrustes and tangent distances were highly The cluster analysis of 22 Cyprinion populations correlated (r2=1 and h=0.9996). The CVA revealed isolated C. watsoni of the Jazmooryan basin 21 CVs that the two first one bearing 59.33% of the population as a new group (Y´) with 83% bootstrap variations. The CVA based on the P-value of the support (Fig. 4). According to the results, the three permutation test showed a significant difference southern and southeastern species (C. watsoni, (P<0.001) in the body shape of the populations C. milesi and C. microphthalmum) were clustered studied (Table 2, Fig. 3). Mahalanobis distances together. All western and south-western species viz. between the studied populations are presented in C. macrostomum, C. kais and C. tenuiradius were Table 3. grouped as a single cluster (with 62% bootstrap) The CVA divided all studied populations in three sharing the longer dorsal-fin base length and shorter groups (Fig. 3). Group X includes C. macrostomum, caudal peduncle. The least inter-species divergence C. kais and C. tenuiradius, group Y consists of was observed in group X. can be C. watsoni and C. microphthalmum, and group Z distinguished from other members of the group X by contains only C. milesi. According to the having longer anal fin-base, and lower caudal deformation grids, the morphological variations peduncle depth, head length, head depth and orbital defined by CV1 are the dorsal-fin base length, diameter. (X2) has the postorbital length, caudal peduncle depth and head highest body depth, longest predorsal length, deepest 58 Nasri et al.- Interspecific morphological variation among Cyprinion

Fig.4. Morphological dendrogram of the studied Cyprinion populations of Iran (the bootstrap support less than 50 were removed).

59 Iran. J. Ichthyol. (March 2019), 6(1): 54-64

caudal peduncle and shortest anal-fin base among the rivers’ flow due to low rainfall (Asghari-Moghadam group X. Cyprinion tenuiradius is separated from 2010) can be considered as main factors to stimulate other members of the group X by having lower such a higher phenotype plasticity in groups Y and Z. caudal peduncle depth and length. There was high Whereas, in south-western and western basins, the morphological overlap between C. tenuiradius and freshwater systems are usually stable and all C. macrostomum (X3a). Cyprinion milesi has the connected to the two recently separated basins i.e. longest anal-fin base, lowest body depth and deepest Tigris-Euphrates and Persis (Nasri 2015). Hence, it caudal peduncle. Cyprinion watsoni has the longest can be proposed that in the allopatric C. macro- caudal peduncle and shortest dorsal-fin base, and stomum, C. kais and C. tenuiradius, stable and C. microphthalmum bears the least caudal peduncle similar ecological conditions have led to minimize length and depth. Cyprinion microphthalmum of the their morphological differences. Nahang and Sarbaz rivers (Y2) showed the least The group Y showed both intra and interspecific orbital diameter. According to the results, the most diversity reflecting their unstable and diverse morphological diversity was observed in the southern environmental conditions. Intraspecific morpho- and southeastern species decreasing this variation logical variation of C. watsoni revealed its potential toward southwestern and western taxa. phenotypic plasticity responding to environmental factors (Nasri et al. 2014). In addition, inter- Discussion population morphological study of C. watsoni The morphological differences can be related to showed that fish living lentic regions of rivers have environment factors revealing some habitat related elevated body shape, but those of lotic or turbulent phenotypic plasticity in fish species (Kerschbaumer rivers have more fusiform body shape with longer & Sturmbauer 2011; Heidari et al. 2013; Mohadasi et head (Nasri et al. 2014). The high level of al. 2013). The members of the genus Cyprinion have intraspecies diversity as reported for Iranian showed some phenotype adaptations in response to Cyprinion (Nasri et al. 2013a, b), can be due to geographical and climatic changes after arriving Iran geographical and climatic variation of their habitats from the east i.e. Oriental region, to survive and in Iran (Banarescu & Coad 1991; Coad 1996; Nasri utilize the various environments (Banarescu & Coad 2015). 1991; Coad 1996; Banister & Clarke 1997). Banarescu & Herzig-Straschil (1995) divided all Cyprinion fishes are known as resistant species Iranian Cyprinion into two groups (C. macrostomum (Coad 1996) occupying difficult environmental or group X and C. watsoni-microphthalmum or group conditions such as thermal streams in Turkey (Daştan Y) but in this study, C. milesi was morphologically et al. 2012) and salty and temporal rivers in southern separated from the C. watsoni-microphthalmum Iran (Nasri et al. 2014). group. Cyprinion milesi is characterized by having Based on the results, the southeastern Cyprinion longest anal-fin base, least body depth, deepest populations (groups Y and Z) show more caudal peduncle and longest head that are all the morphological variations than those of group X i.e. proper morphological characters for inhabiting lotic C. macrostomum, C. kais and C. tenuiradius. The ecosystems (Nasri et al. 2014). In addition, the big intraspecific diversity in species with wide head and a wide oblique mouth in C. milesi can be distribution range is high as reported in Cyprinion considered as its trophic adaptation i.e. carnivorous species (Nikolski 1963; Banarescu & Herzig- feeding mode (Svanbäck et al. 2008; Naples & Straschil 1995; Nasri et al. 2013a). In addition, the McAfee 2012). The main morphological differences unstable climate conditions of the southern and between C. milesi and C. kais with their conspecifics southeastern Iran, including seasonal and temporal is mouth form (Banarescu & Herzig-Straschil 1995), 60 Nasri et al.- Interspecific morphological variation among Cyprinion

that it can be related to their feeding habits (Svanbäck supported by Lorestan University and University of et al. 2008; Naples & McAfee 2012). Tehran. The results of the present study showed that C. macrostomum group (group X) can be References distinguished from other groups by having the greater Adams, D.C.; Rohlf, F.J. & Slice, D.E. 2004. Geometric body depth, shorter head length and depth, longer morphometrics: ten years of progress following the dorsal-fin base and shorter caudal peduncle. ‘revolution’. Italian Journal of Zoology 71: 5-16. Explaining the causes of morphological differences Asghari-Moghadam, M.-R. 2010. climatology of Iran in is very difficult. But, it is possible to relate an altered quaternary glaciation period. Journal of Geography 4(13): 131-149. phenotypic characters to known functions Aydin, R.; Şen, D.; Çalta, M. & Canpolat, Ö. 2008. The

(Langerhans et al. 2004). In addition, the potential amount of calcium in bony structures used for age occupied niche of fishes can be predicted based on determination in Cyprinion macrostomus (Heckel, fish morphology, especially in cyprinids (Douglas & 1843). Aquaculture Research 39: 596-602. Matthews 1992; Wood & Bain 1995). A well- Banarescu, P.M. & Coad, B.W. 1991. Cyprinids of developed dorsal fin is a kind of phenotype plasticity Eurasia. In: Winfield, I.J. & Nelson, J.S. (eds.), response to the permanent rivers with high water Cyprinid Fishes. Springer, The Netherlands, pp. 127- level (Lauder & Drucker 2004; Poulet et al. 2004) as 155. seen in the rivers of the Persis and Tigris-Euphrates Banarescu, P.M. & Herzig-Straschil, B. 1995. A revision basins. Furthermore, the shorter caudal peduncle, of the species of the Cyprinion macrostomus-group deeper body and lower head depth and length are led (Pisces: Cyprinidae). Annalen des Naturhistorischen Museums in Wien 97(B): 411-420. to a fusiform body shape appropriating for streams Banister, K.E. & Clarke, M.A. 1997. The freshwater with higher current velocity or lakes with high water fishes of the Arabian Peninsula. Journal of Oman level (Chan 2001). Almost all rivers in distribution Studies, Special Report 1: 111-154. range of C. macrostomum group are permanent and Bookstein, F.L. 1991. Morphometric Tools for high productive. It is difficult to morphologically Landmark Data: Geometry and Biology. Cambridge distinguish C. macrostomum from C. tenuiradius, University Press, Cambridge, UK. and C. watsoni from C. microphthalmum. A wide Chan, M.D. 2001. Fish ecomorphology: predicting interspecific morphological variation reported habitat preferences of stream fishes from their body among Iraqi Cyprinion (Kafuku 1969) and Iranian shape. PhD. thesis, fisheries and wildlife sciences, species (Nasri et al. 2013a, b, 2014, 2015). Virginia Polytechnic Institute and State University, Based on the results, it can be concluded that the Virginia, USA. Coad, B.W. 1996. Zoogeography of the fishes of the most possible factor influencing morphological Tigris-Euphrates basin. Zoology in the Middle East variations among Iranian Cyprinion fishes is habitat 13(1): 51-70. related differences as the members of this taxa known Coad, B.W. 2019. Freshwater fishes of Iran. to be resistant to various environmental condition and www.briancoad.com. accessed (3 January 2019). distributed in diverse habitats with geographical and Daştan, S.D.; Bardakci, F. & Degerli, N. 2012. Genetic climatic differences. Therefore, they can be adapted Diversity of Cyprinion macrostomus Heckel, 1843 to their habitats by changing their body shape as (Teleostei: Cyprinidae) in Anatolia. Turkish Journal generalist species. of Fisheries and Aquatic Sciences 12: 651-659. Douglas, M.E. & Matthews, W.J. 1992. Does Acknowledgements morphology predict ecology? hypothesis testing We would like to thank P. Jalili for her help in within a freshwater stream fish assemblage. Oikos 65(2): 213-224. laboratory works. This study was financially 61 Iran. J. Ichthyol. (March 2019), 6(1): 54-64

Esmaeili, H.R.; Sayyadzadeh, G.; Eagderi, S. & Abbasi, Mohadasi, M.; Shabanipour, N. & Eagderi, S. 2013. K. 2018. Checklist of freshwater fishes of Iran. Habitat-associated morphological divergence in four FishTaxa 3(3): 1-95. Shemaya, Alburnus chalcoides (: Hammer, Ø. 2012. PAST: Paleontological Statistics. Cyprinidae) populations in the southern Caspian Sea Natural History Museum University of Oslo, Oslo. using geometric morphometrics analysis. Heidari, A.; Mousavi-Sabet, H.; Khoshkholgh, M.; International Journal of Aquatic Biology 1(2): 82-92. Esmaeili, H.R. & Eagderi, S. 2013. The impact of Naples, V.L. & McAfee, R.K. 2012. Reconstruction of Manjil and Tarik dams (Sefidroud River, southern the cranial musculature and masticatory function of Caspian Sea basin) on morphological traits of Siah the Pleistocene panamerican ground sloth Mahi Capoeta gracilis (Pisces: Cyprinidae). Eremotherium laurillardi (Mammalia, Xenarthra, International Journal of Aquatic Biology 1(4): 195- Megatheriidae). Historical Biology 24(2): 187-206. 201. Nasri, M. 2008. of bigmouth lotak (Cyprinion Jalili, P.; Eagderi, S. & Keivany, Y. 2015. Body Shape macrostomum Heckel, 1843) and smallmouth lotak Comparison of Kura Bleak (Alburnus filippii) in Aras (Cyprinion kais Heckel, 1843) in Karkheh River and Ahar-Chai Rivers Using Geometric basin and Godarkhosh River in Ilam province. M.Sc. Morphometric Approach. Research in Zoology 5(1): thesis, Department of Natural Resources, Isfahan 20-24. University of Technology, Isfahan. (In Farsi) Kafuku, T. 1969. Morphological differentiation of Nasri, M. 2015. Phylogeography of Genus Cyprinion in Cyprinion in Iraq. Bulletin of Freshwater Fisheries Iran. Ph.D. thesis, Department of Fisheries, Research Laboratory 19(2): 155-159. University of Tehran, Tehran, Iran. (In Farsi) Keivany, Y.; Nasri, M.; Abbasi, K. & Abdoli, A. 2016. Nasri, M.; Eagderi, S. & Farahmand, H. 2014. Atlas of Inland Water Fishes of Iran, 1. Iran Intraspecific morphological variation of Sabzug, Department of Environment, Tehran, Iran. (In Farsi) Cyprinion watsoni (Day, 1872) from southern and Kerschbaumer, M. & Sturmbauer, C. 2011. The utility of southeastern Iran based on Geometric morphometrics geometricmorphometrics to elucidate pathways of method. Journal of Applied Ichthyological Research cichlid fish evolution. International Journal of 2(2): 1-13. (In Farsi) Evolutionary Biology 1-8. Nasri, M.; Eagderi, S.; Farahmand, H. & Hashemzade- Khataminejad, S.; Mousavi-Sabet, H.; Sattari, M.; SegharLoo, I. 2013a. Body shape comparison of Vatandoust, S. & Eagderi, S. 2013. A comparative Cyprinion macrostomum (Heckel, 1843) and study on body shape of the genus Alburnus Cyprinion watsoni (Day, 1872) using geometric (Rafinesque, 1820) in Iran, using geometric morphometric method. International Journal of morphometric analysis. Caspian Journal of Aquatic Biology 1(5): 240-244. Environmental Sciences 11(2): 205-215. Nasri, M.; Eagderi, S.; Keivany, Y.; Farahmand, H.; Klingenberg, C.P. 2011. MorphoJ: an integrated Dorafshan, S. & Nezhadheydari, H. 2018. software package for geometric morphometrics. Morphological diversity of Cyprinion Heckel, 1843 Molecular Ecology Resources 11: 353-357. species (Teleostei: Cyprinidae) in Iran. Iranian Langerhans, R.B. & DeWitt, T.J. 2004. Shared and Journal of Ichthyology 5(2): 96-108. unique features of evolutionary diversification. Nasri, M.; Keivany, Y. & Dorafshan, S. 2013b. American Naturalist 164(3): 335-349. Comparative osteology of Lotaks, Cyprinion kais and Lauder, G.V. & Drucker, E.G. 2004. Morphology and C. macrostomum (, Cyprinidae), from experimental hydrodynamics of fish fin control Godarkhosh River, Western Iran. Journal of surfaces. Ieee Journal of Oceanic Engineering, 29(3): Ichthyology 53(6): 455-463. 556-571. Nikolski, G.V. 1963. Fish Ecology. Academic Press. Marcus, L.F.; Corti, M.; Loy, A.; Naylor, G.J.P. & Slice, London. Poulet, N.; Berrebi, P.; Crivelli, A.J.; Lek, S. & D.E. 1996. Advances in Morphometries. Springer Argillier, C. 2004. Genetic and morphometric Science+ Business Media, LLC. variations in the pikeperch (Sander lucioperca L.) of

62 Nasri et al.- Interspecific morphological variation among Cyprinion

a fragmented delta. Archiv für Hydrobiologie 159(4): 531-554. Rohlf, F.J. & Marcus, L.F. 1993. A revolution in morphometrics. Trends in Ecology and Evolution 8(4): 129-132. Rohlf, F.J. & Slice, D.E. 1990. Extensions of the procrustes method for the optimal superimposition of landmarks. Systematic Zoology 39(1): 40-59. Rohlf, F.J. 1998. On applications of geometric morphometrics to studies of ontogeny and phylogeny. Systematic Biology 47(1): 147-158. Rohlf, F.J. 2010. TpsDig2–Thin Plate Spline Digitise. 2.16. New York. State University of New York. Rohlf, F.J. 2015. TpsSmall–Thin Plate Spline Small Variation Analysis. 1.33. New York. State University of New York. Svanbäck, R.; Eklöv, P.; Fransson, R. & Holmgren, K. 2008. Intraspecific competition drives multiple species resource polymorphism in fish communities. Oikos 117: 114-124. Wood, B.M. & Bain, M.B. 1995. Morphology and microhabitat use in stream fish. Canadian Journal of Fisheries and Aquatic Sciences 52(7): 1487-1498.

63 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

مقاله پژوهشی تغییرات ریختی بینگونهای در ماهیان جنس Cyprinion Heckel, 1843 )خانواده کپورماهیان( ایران با استفاده از روش ریختسنجی هندسی لندمارکپایه

منوچهر نصری1، سهیل ایگدری*2، حمید فرحمند2، حسن نژادحیدری2

1گروه علوم و مهندسی شیالت، دانشکده کشاورزی و منابع طبیعی دانشگاه لرستان، خرمآباد، ایران 2گروه شیالت، دانشکده منابع طبیعی، دانشگاه تهران، کرج، ایران. چکیده: تغییرات ریختی در بین گونههای جنس Cyprinion ایران با استفاده از روش ریختسنجی هندسی لندمارکپایه مورد مطالعه قرار گرفت. نمونههای مورد مطالعه طی شهریور 1392 تا شهریور 1394 از پنج حوضه آبریز ایران صید شدند. تعداد 15 لندمارک بر روی تصاویر دوبعدی ماهیها تعریف و رقومی شد. عالوه بر تفاوتهای ریختی درونگونهای، آزمون CVA تمامی جمعیتهای مورد مطالعه را در سه گروه اصلی دستهبندی کرد شامل: C. kais ،C. macrostomum و C. tenuiradius )گروه X(، C. watsoni و C. microphthalmum )گروه Y( و C. milesi )گروه Z(. بر اساس نتایج، گروه C. macrostomum را میتوان بر اساس داشتن عمق بدن بیشتر، کمترین طول و ارتفاع سر، بیشترین طول قاعده باله پشتی و کمترین طول ساقه دمی از دو گروه دیگر تشخیص داد. C. milesi دارای بیشترین طول قاعده باله مخرجی، کمترین ارتفاع بدن، بیشترین ارتفاع ساقه دمی و بیشترین طول سر است. Cyprinion watsoni دارای بیشترین طول ساقه دمی و کمترین طول قاعده باله پشتی است و C. microphthalmum را میتوان بر اساس کمترین طول و ارتفاع ساقه دمی از سایرین متمایز کرد. مهمترین علت وجود تفاوتهای ریختی درونگونهای را میتوان انعطافپذیری ریختی مرتبط با عوامل جغرافیای جانوری و اقلیمی دانست. نتایج پیشنهاد میکنند که محتملترین عامل مؤثر بر تنوع ریختی در بین گونههای Cyprinion ایران، تفاوتهای مرتبط با خصوصیات زیستگاهی است. کلماتکلیدی: کپورماهیان، پراکنش ماهیان، آب شیرین، انعطافپذیری ریختی، جغرافیای جانوری.

64