A comparative study on the body shape of Alosa caspia (Teleostei, Clupeidae) populations in the southern Caspian Sea basin using geometric morphometric analysis



Geometric morphometric method was used to examine body shape variations among the three populations of the Caspian shad, Alosa caspia, in the southern Caspian Sea coasts of Iran. A total of 289 A. caspia specimens were caught from three localities, from the west to the east including Guilan (Anzali), Mazandaran (Sari) and Golestan (Miankale), respectively. Shape data was extracted using 15 landmark points on 2-D pictures of specimens. The PCA, DFA, CVA/MANOVA analyses and cluster analysis (CA) based on Euclidean square distances were used to examine shape differences among the three populations. Significant differences were found among the populations of A. caspia in terms of body shape. Cluster analysis showed separation of Mazandaran populations from the other provinces. Results revealed that the studied populations are divided into two categories; a group with large body and head depth and short and wide caudal peduncle and the other one with shallow body, small head and longer caudal peduncle. Observed differences in the head shape of Mazandaran population of A. caspia could be indirectly related to the feeding resources. Changes in head and mouth shapes can be considered as reflective of differences in selection of food items and direction of feeding. The obtained body shape properties can provide a shape-based identification key for Alosa caspia, which is useful for fisheries and stock management or conservation programs.


Shape variation, Discriminant, Clupeiformes, Landmark, Stock, Shad.


Adams, D.C.; Rohlf, F.J. & Slice, D.E. 2004. Geometric morphometrics: ten years of progress following the “revolution.” Italian Journal of Zoology 71: 5-16.

Andersson, C.B.; Hughes, S.O.; Fisher, J.O. & Nicklas, T.A. 2005. Cross-cultural equivalence of feeding beliefs and practices: The psychometric properties of the child feeding questionnaire among Blacks and Hispanics. Preventative Medicine 41: 521-531.

Balon, E.K. 1993. Dynamics of biodiversity and mechanisms of change: a plea for balanced attention to form creation and extinction. Biological Conservation 66: 5-16.

Bektas, Y. & Belduz A.O. 2009. Morphological variation among Atlantic horse mackerel, Trachurus trachurus populations from Turkish coastal waters. Journal of Animal and Veterinary Advances 8: 511-7.

Bookstein, F.L. 1991. Morphometric tools for landmark data. Cambridge University Press, Cambridge, UK. 435 p.

Bookstein, F.L. 1996. Biometrics, biomathematics and the morphometric synthesis. Bulletin of Mathematical Biology 58: 313-365.

Cadrin, S.X. & Friedland, K.D. 1999. The utility of image processing techniques identification. Fisheries Research 43: 129-139.

Cadrin, S.X. & Silva, V.M. 2005. Morphometric variation of yellowtail flounder. ICES Journal of Marine Science 62: 683-694.

Coad, B.W. 2017. Freshwater fishes of Iran. Available at: http://www.briancoad.com (accessed on 17 June 2017).

Costa, C. & Cataudella, S. 2007. Relationship between shape and trophic ecology of selected species of Sparids of the Caprolace coastal lagoon (central Tyrrhenian Sea). Environmental Biology of Fishes 78: 115-123.

Dryden, I.L. & Mardia, K.V. 1998. Statistical Shape Analysis. New York: John Wiley and Sons.

Eschmeyer, W.N. & Fong J.D. 2011. Animal biodiversity: An outline of higher level classification and survey of taxonomic richness. Zootaxa 3148: 26-38.

Esmaeili, H.R.; Coad, B.W.; Mehraban, H.R.; Masoudi, M.; Khaefi, R.; Abbasi, K.; Mostafavi, H. & Vatandoust, S. 2014. An updated checklist of fishes of the Caspian Sea basin of Iran with a note on their zoogeography. Iranian Journal of Ichthyology 1: 152-184.

Esmaeili, H.R.; Mehraban, H.; Abbasi, K.; Keivany, Y. & Coad, B. 2017. Review and updated checklist of freshwater fishes of Iran: Taxonomy, distribution and conservation status. Iranian Journal of Ichthyology 4(Suppl. 1): 1-114

Gatz, A.J., Jr. 1979. Community organization in fishes as indicated by morphological features. Ecology 711- 718.

Guill, J.M.; Hood, C.S. & Heins, D.C. 2003. Body shape variation within and among three species of darters (Perciformes: Percidae). Ecology of Freshwater Fish 12: 134-140.

Klingenberg, C.P. 1998. Heterochrony and allometry: the analysis of evolutionary change in ontogeny. Biological Reviews 73: 79-123.

Kohestan-Eskandari, S.; Anvarifar, H.; Mousavi-Sabet, H.; Yousefi, M. & Khanzade, M.; 2014. A morphology-based hypothesis for homeward migration success and population differentiation in the anadromous Kutum Rutilus kutum (Pisces: Cyprinidae) along the southern Caspian Sea, Iran. Folia Zoologica 63(3): 151-160.

Langerhans, B.R.; Laydon, C.A.; Langerhans, A.K. & Dewitt, T.J. 2003. Habitat-associated morphological divergence in two Neotropical fish species. Biological Journal of the Linnaean Society 80: 689-698.

Loy, A.; Boglione, C. & Cataudella, S. 1999. Geometric morphometrics and morpho-anatomy: a combined tool in the study of sea bream (Sparus aurata) shape. Journal of Applied Ichthyiology 15: 104-110.

Magurran, A.E. 1998. Population differentiation without speciation. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences 353: 275-286.

Matthews, W.J. 1998. Patterns in Freshwater Fish Ecology. New York, Chapman and Hall. 757 p.

Mousavi-Sabet, H. & AnvariFar, H. 2013. Landmark-based morphometric variation between Cobitis keyvani and Cobitis faridpaki (Pisces: Cobitidae), with new habitat for C. faridpaki in the southern Caspian Sea basin. Folia Zoologica 62: 167-175.

Page, L.M. & Swofford, D.L. 1984. Morphological correlates of ecological specialization in darters. Environmental Biology of Fishes 11: 139-159.

Pianka, E.R. 1994. Evolutionary ecology. New York, Harper Collins. 397 p.

Richtsmeier, J.T.; Deleon, V.B. & Lele, S.R. 2002. The promise of geometric morphometrics. The Yearbook of Physical Anthropology 45: 63-91.

Rohlf, F.J. & Marcus, L.F. 1993. “A revolution in morphometrics,”Trends in Ecology and Evolution 8(4): 129-132.

Rohlf, F.J. 2004. TpsDig–Thin Plate Spline Digitize, Version 1.4 [Computer software]. Stony Brook: Dept. of Ecology and Evolution, State Univ. of New York.

Sansom, R.S. 2009. Phylogeny, classification, and character polarity of the Osteostraci (Vertebrata). Journal of Systematic Paleontology 7: 95-115.

Schluter, D. & McPhail, J.D. 1992. Ecological character displacement and speciation in sticklebacks. American Naturalist 140: 85-108.

Slice, D.E. 2007. Geometric Morphometrics. Annual Review of Anthropology 36: 261-281.

Sneath, P.H. & Sokal, R.R. 1973. Numerical taxonomy. The principles and practice of numerical classification.

Small, C. 1996. The statistical theory of shape (p. 227). New York: Springer.

Tzeng, T.D. 2004. Morphological variation between populations of spotted Mackerel Scomberaus tralasicus of Taiwan. Fisheries Research 68: 45-55.

Wimberger, P. H.; 1990. Plasticity of fish body shape: the effects of diet, development, family, and age in two species of Geophagus (Pisces: Cichlidae). Biological Journal of the Linnean Society 45: 197-218.

Winemiller, K.O. 1991. Ecomorphological diversification in lowland freshwater fish as assemblages from five biotic regions. Ecological Monographs 343-365.

Wood, B.M. & Bain, M.B. 1995. Morphology and microhabitat use in stream fish. Canadian Journal of Fisheries and Aquatic Sciences 52: 1487-1498.

Yamamoto, S.; Maekawa, K.; Tamate, T.; Koizumi, I.; Hasegawa, K. & Kubota, H. 2006. Genetic evaluation of translocation in artificially isolated populations of white-spotted charr (Salvelinus leucomaenis). Fisheries Research 78: 352-358.


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