Scientia Marina, Vol 74, No 1 (2010)

Body morphometrics, swimming diversity and niche in demersal sharks: a comparative case study from the Mediterranean Sea

Umberto Scacco
I .S.P.R.A ex I.C.R.A.M, Roma , Italy

Gabriele La Mesa
I .S.P.R.A ex I.C.R.A.M, Roma , Italy

Marino Vacchi
I .S.P.R.A ex I.C.R.A.M, Roma , Italy


Seven demersal shark species belonging to six families, obtained from the southern Mediterranean Sea, were selected as representative of different putative swimming modes, inhabited depths and trophic levels. Caudal fin morphometrics (caudal fin span, area and aspect ratio), hepatosomatic index and muscular-skeletal amount per unit length were estimated and compared within and among species by means of both parametric and non-parametric multivariate and univariate statistical procedures. Intra-specific variation was related to size and, for one species only, also to sex. Two inter-specific patterns of variation were observed: the first pattern was interpreted as a swimming ability gradient between slow benthic and more pelagic species, as a function of caudal morphometry and body characteristics; the second pattern is related to the depth the species inhabit and/or their trophic level. These two hypotheses were tested with the current data and literature information, and discussed in the context of shark swimming modes, habit and environment. Based on observed differences in caudal fin morphometrics, liver and muscular masses, this paper is the first to provide a quantitative and comparative dataset for demersal sharks as a group, and attempts to describe sample variability as a function of expected swimming ability and occupied niche (depth inhabited and/or trophic level).


demersal sharks; swimming ability; tail aspect ratio; hepatosomatic index

Full Text:



Alberch, P., S.J. Gould, G.F. Oster and D.B. Wake. – 1979. Size and shape in ontogeny and phylogeny. Paleobiology, 5(3): 296-317.

Aleev, Y.G. – 1963. Function and gross morphology in fish. Israel: Program for Scientific Translation.

Alexander, R.M. – 1965. The lift produced by the heterocercal tails of Selachii. J. Exp. Biol., 43: 131-138.

Alexander, R.M. –1967. Functional design in fishes. London: Hutchinson University Library.

Alexander, R.M. – 1968. Animal Mechanics. London: Sidgwick and Jackson.

Baldridge, H.D. Jr – 1970. Sinking factors and average densities of Florida sharks as function of liver buoyancy. Copeia, 4: 744-754. doi:10.2307/1442317

Baldridge, H.D. Jr – 1972. Accumulation and function of liver oil in Florida sharks. Copeia, 2: 306-325. doi:10.2307/1442493

Belluscio, A., U. Scacco, F. Colloca, P. Carpentieri and G.D. Ardizzone. – 2000. Feeding strategies of two species of demersal chondrichthyans, Galeus melastomus (Rafinesque, 1810) and Etmopterus spinax (Linnaeus, 1758), in the Central Tyrrhenian Sea. Biol. Mar. Medit., 7: 417-426.

Bertrand, J., L. Gil De Sola, C. Papaconstantinou, G. Relini and A. Souplet. – 2000. Contribution on the distribution of elasmobranchs in the Mediterranean Sea (from the MEDITS surveys). Biol. Mar. Medit., 7(1): 385-399.

Blake, R.W. – 2004. Fish functional design and swimming performance. J. Fish Biol., 65: 1193-1222. doi:10.1111/j.0022-1112.2004.00568.x

Bone, Q. and B.LRoberts. – 1969. The density of elasmobranches. J. Mar. Biol. Ass. UK, 49: 913-937.

Breder, C.M. – 1926. The locomotion of fishes. Zoologica, 4: 159-297.

Budker, P. – 1971. The life of sharks. Columbia University Press, New York.

Carrassón, M., C. Stefanescu and J.E. Cartes. – 1992. Diets and bathymetric distributions of two bathyal sharks of the Catalan deep sea (western Mediterranean). Mar. Ecol. Progr. Ser., 82: 21-30 doi:10.3354/meps082021

Clarke, K.R. and R.M. Warwick. – 2001. Change in marine communities: an approach to statistical analysis and interpretation, 2nd edn. PRIMER-E, Plymouth.

Clarke, M.W., P.L. Connolly and J.J. Braken. – 2001. Aspects of reproduction of the deep water sharks Centroscymnus coelolepis and Centrophorus squamosus from west of Ireland and Scotland. J. Mar. Biol. Ass. UK, 81: 1019-1029.

Cliff, G., S.F.J. Dudley and B. Davis. – 1989. Sharks caught in the protective gill nets off Natal, South Africa. 2. The great white shark Carcharodon carcharias (Linnaeus). S. Afr. J. Mar. Sci., 8: 131-144.

Cognetti, G. and M. Sará. – 1974. Biologia Marina. Calderini, Bologna.

Compagno, L.J.V. – 1984. FAO species catalogue. Vol. 4. Sharks of the world. An annotated and illustrated catalogue of shark species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fisheries Synopsis. (125, Vol. 4, Part 1), 249 pp.

Compagno, L.J.V., D.A. Ebert and M.J. Smale. – 1989. Guide to the sharks and rays of southern Africa. NEMHolland (Publ.) Ltd, London.

Compagno, L.J.V. and V.H. Niem. – 1998 b. Hexanchidae. Cowsharks, sixgill, and seven gill sharks. In: K.E. Carpenter and V.H. Niem (eds.), FAO Identification Guide for Fisheries Purposes. The Living Marine Resources of the Western Central Pacific, pp. 1208-1210. FAO, Rome.

Compagno, L.J.V. – 2002. Sharks. Key to families occurring in the area. In: K.E.Carpenter (ed.), FAO species identification guide for fishery purposes. The living marine resources of the Western Central Atlantic: Introduction, molluscs, crustaceans, hagfishes, sharks, batoid fishes, and chimaeras, Vol. 1, pp. 363-505. FAO, Rome.

Corner, E.D.S., F.R.S. Denton and G.R. Forster. – 1969. On the buoyancy of some deep sea sharks. Proc. Roy. Soc. B-Biol., 171: 415-429.

Cortès, E. – 1999. Standardized diet compositions and trophic levels of shark. ICES J. Mar. Sci., 56: 707-717. doi:10.1006/jmsc.1999.0489

Craik, J.C.A. – 1978. An annual cycle of vitellogenesis in the elasmobranch Scyliorhinus canicula. J. Mar. Biol. Ass. UK, 58: 719-726. doi:10.1017/S0025315400041369

D’Ortensio, F., D. Iudicone, C. de Boyer Montegut, P. Testor, D. Antoine, S. Marullo, R. Santoleri and G. Madec.– 2005. Seasonal variability of the mixed layer depth in the Mediterranean Sea as derived from in situ profiles. Geophys. Res. Lett., 32: L12605. doi:10.1029/2005GL022463

Domenici, P., G. Claireaux and D.J. McKenzie. – 2007. Environmental constraints upon locomotion and predator–prey interactions in aquatic organisms: an introduction. Philos. Trans. Roy. Soc. B, 362: 1929-1936. doi:10.1098/rstb.2007.2078 PMCid:2042526

Donley, J. and R. Shadwick.– 2003. Steady swimming muscle dynamics in the leopard shark Triakis semifasciata. J. Exp. Biol., 206: 1117-1126. doi:10.1242/jeb.00206 PMid:12604571

Ellis, J.R., M.G. Pawson, and S.E. Shackley. – 1996. The comparative feeding ecology of six species of shark and four species of ray (Elasmobranchii) in the north-east Atlantic. J. Mar. Biol. Ass. UK, 76: 89-106. doi:10.1017/S0025315400029039

Ferry, L.A. and G.V. Lauder. – 1996. Heterocercal tail function in leopard sharks: a three-dimensional kinematic analysis of two models. J. Exp. Biol., 199: 2253-2268.

Fischer, W., M.L. Bauchot and M. Schneider. – 1987 Fiches FAO d’identification des espèces pour les besoins de la pêche. Méditerranée et Mer Noire. Zones de pêche 37 Vertébrés. Vol. 2. FAO, Rome.

Fish, F.E. and L.D. Shannahan. - 2000. The role of the pectoral fins in body trim of sharks. J. Fish Biol., 56: 1062-1073. doi:10.1111/j.1095-8649.2000.tb02123.x

Froese, R. and D. Pauly. – 2007. FishBase. World Wide Web electronic publication., version (11/2007).

Gabriel, K.R. – 1971. The biplot graphic display of matrices with application to principal component analysis. Biometrika, 58(3): 453-467. doi:10.1093/biomet/58.3.453

Gabriel, K.R. – 1982. Biplot. In: S. Kotz and N.L. Johnson (eds.), Encyclopedia of Statistical Sciences, vol. 1, pp. 263-271, Wiley, New York.

Gabriel, K.R. and C.L. Odoroff. – 1990. Biplots in biomedical research. Stat Med., 9(5): 469-485. doi:10.1002/sim.4780090502 PMid:2349401

Ghirardelli, E. – 1981. La vita nelle acque. Utet, Torino

Gibson, R.N. and I.A. Ezzi. – 1987. Feeding relationships of a demersal fish assemblage on the west coast of Scotland. J. Fish. Biol., 31: 55-69. doi:10.1111/j.1095-8649.1987.tb05214.x

Gould, S.J. and N. Eldredge. – 1977. Punctuated equilibria: the tempo and mode of evolution reconsidered. Paleobiology, 3: 115-151.

Gower, J.C. and D.J. Hand. – 1996. Biplots. Chapman and Hall, London.

Graffelman, J. and T. Aluja-Banet. - 2003. Optimal representation of supplementary variables in biplots from principal component analysis and correspondence analysis. Biometrical J., 45(4): 491-509. doi:10.1002/bimj.200390027

Hennemann, R.M. – 2001. Sharks and rays: Elasmobranch guide of the world. IKAN- Unterwasserarchiv, Frankfurt.

Higham, T.E. – 2007. The integration of locomotion and prey capture in vertebrates: Morphology, behaviour, and performance. Integr. Comp. Biol., 47(1): 82-95. doi:10.1093/icb/icm021

Imre, I., R.L. McLaughlin and D.L.G. Noakes. – 2002. Phenotypic plasticity in brook charr: changes in caudal fin induced by water flow. J. Fish Biol., 61(5): 1171-1181. doi:10.1111/j.1095-8649.2002.tb02463.x

Kawabe, R., Y. Naito, K. Sato, K. Miyashita and N. Yamashita. – 2004. Direct measurement of the swimming speed, tailbeat, and body angle of Japanese flounder (Paralichthys olivaceus). ICES J. Mar. Sci., 7: 1080-1087. doi:10.1016/j.icesjms.2004.07.014

Kohler, N.E., J.G. Casey and P.A. Turner. – 1996. Length-length and length-weight relationships for 13 Shark Species from the Western North Atlantic. NOAATechnical Memorandum NMFS-NE-110, May 1996.

Koob, T.J. and I.P. Callard. – 1999. Reproductive endocrinology of female elasmobranches: lessons from the little skate (Raja erinacea) and spiny dogfish (Squalus acanthias). J. Exp. Zool., 284: 557-574. doi:10.1002/(SICI)1097-010X(19991001)284:5<557::AID-JEZ12>3.0.CO;2-P PMid:10469994

Kramer, E. – 1960. Zur Form und Funktion des Lokomotionsapparates der Fische. Z. Wiss. Zool. ABT A, 163: 1-63.

Krebs, C.J. – 1989. Ecological Methodology. NEMYork: Harper and Row.

Last, P.R. and J.D. Stevens. – 1994. Sharks and rays of Australia. Australia: CSIRO.

Lauder, G.V. – 2000. Function of the caudal fin during locomotion in fishes: kinematics, flow visualization, and evolutionary patterns. Am. Zool., 40: 101-122. doi:10.1668/0003-1569(2000)040[0101:FOTCFD]2.0.CO;2

Lighthill, M.J. – 1970. Aquatic animal propulsion of high hydromechanical efficiency. J. Fluid Mech., 44: 265-301. doi:10.1017/S0022112070001830

Lindsey, C.C. – 1978. Form, function and locomotory habits in fish. In: W.S. Hoar and D.J. Randall (eds.), Fish Physiology, vol. 7, pp. 1-100. Academic Press, London.

Lingham-Soliar, T. – 2004a. Caudal fin allometry in the white shark Carcharodon carcharias: implications for locomotory performance and ecology. Naturwissenschaften, 92: 231-236. doi:10.1007/s00114-005-0614-4 PMid:15772806

Lingham-Soliar, T. – 2004b. Dorsal Fin in the White Shark, Carcharodon carcharias: ADynamic Stabilizer for Fast Swimming. J. Morph., 260: 1-11.

Lingham-Soliar, T. - 2005. Caudal Fin in the White Shark, Carcharodon carcharias: ADynamic Propeller for Fast, Efficient Swimming. J. Morph., 264: 233-252. doi:10.1002/jmor.10328 PMid:15795938

Lingham-Soliar, T. and G. Plodowski. – 2007. Taphonomic evidence for high-speed adapted fin in thunniform ichthyosaurs. Naturwissenschaften, 94: 65-70. doi:10.1007/s00114-006-0160-8 PMid:17021914

Macpherson, E. – 1979. Relations trophiques des poisons dans la Méditerranée occidentale. Rapp. Comm. Int. Explor. Sci. Mer Méditerr., 25/26: 49-58.

Macpherson, E. and B.A.Roel. – 1987. Trophic relationships in the demersal fish community off Namibia. S. Afr. J. Mar. Sci., 5: 585-596.

Magnuson, J.J. – 1978. Locomotion by scombrid fishes: hydromechanics, morphology and behaviour. In: W.S. Hoar and D.J. Randall (eds.), Fish Physiology, vol. 7, pp. 239-313. Academic Press, London.

Marcil, J., D.P. Swain and J.A. Hutchings. – 2006. Genetic and environmental components of phenotypic variation in body shape among populations of Atlanticcod (Gadus morhua L.). Biol. J. Linn. Soc., 88(3): 351-365. doi:10.1111/j.1095-8312.2006.00656.x

Massutí, E. and J. Moranta. – 2003. Demersal assemblages and depth distribution of elasmobranches from the continental shelf and slope off the Balearic Islands (western Mediterranean). ICES J. Mar. Sci., 60: 753-766. doi:10.1016/S1054-3139(03)00089-4

Newton, G.M. – 1999. The deep-sea environment - Earth’s final frontier. Aust. Mar. Sci. Bull., 147: 17-21.

Notarbartolo Di Sciara, G. and I. Bianchi. – 1998. Guida degli squali e delle razze del Mediterraneo. Franco Muzzio.

Odum, W.E. and E.J. Heald. – 1975. The detritus-based food web of an estuarine mangrove community. In: L.E. Cronin (ed.), Estuarine research, Vol 1, pp. 265-286. Accademic Press, New York.

Olaso, I., F. Velasco, F. Sànchez, A. Serrano, C. Rodrìguez-Cabello and O. Cendrero. – 2004. Trophic relations of lesser-spotted catshark (Scyliorhinus canicula) and blackmouth catshark (Galeus melastomus) in the Cantabrian Sea. J. Northwest Atl. Fish. Soc., 35 upload date: 16 Dec 04.

Pauly, D. – 1986. Asimple method for estimating the food consumption of fish populations from growth data and food conversion experiments. Fish. Bull., 84(4): 827-840.

Perry, C. and B.D. Roitberg. – 2006. Trophic egg laying: hypotheses and tests. Oikos, 112(3): 706-714. doi:10.1111/j.0030-1299.2006.14498.x

Pinardi, N. and E. Masetti. – 2000. Variability of the large scale general circulation of the Mediterranean Sea from observations and modelling: a review. Palaeogeogr. Palaeoclimatol. Palaeoecol., 158: 153-173. doi:10.1016/S0031-0182(00)00048-1

Richardson, A.J., G. Maharaj, L.J.V. Compagno, R.W. Leslie, D.A. Ebert and M.J. Gibbons. – 2000. Abundance, distribution, morphometrics, reproduction and diet of the Izak catshark. J. Fish Biol., 56: 552-576. doi:10.1111/j.1095-8649.2000.tb00755.x

Sambilay Jr., V.C. – 1990. Interrelationships between swimming speed, caudal fin aspect ratio and body length of fishes. Fishbyte, 8: 16-20.

Sandoval-Castillo, J., A. Rocha-Olivares, C. Villavicencio-Garayzar and E. Balart. – 2004. Cryptic isolation of Gulf of California shovelnose guitarfish evidenced by mitochondrial DNA. Mar. Biol., 145(5): 983-988. doi:10.1007/s00227-004-1378-7

Serena, F. – 2005. Field identification guide to the sharks and rays of the Mediterranean and Black Sea. FAO, Rome.

Siegel, S. and N.J.Jr. Castellan. – 1992. Statistica non parametrica. Mc Graw Hill, Milano.

Sokal, R.R. and F.J. Rohlf. – 1995. Biometry. Freeman, W.H. and Co, New York.

STATISTICA6.0 (Electronic Version): StatSoft, Inc. – 2006. Electronic Statistics Textbook. Tulsa, OK: StatSoft. WEB: (Printed Version): Hill, T., Lewicki, P. 2006). STATISTICS Methods and Applications. StatSoft: Tulsa, OK.

Stergiou, K.I. and V.S. Karpouzi. – 2002. Feeding habits and trophic levels of Mediterranean fish. Rev. Fish Biol. Fisher., 11: 217-254. doi:10.1023/A:1020556722822

Sulikowski, J.A., W.B. IIIDriggers, T.S. Ford, R.K. Boonstra and J.K. Carlson. – 2007. Reproductive cycle of the black nose shark Carcharhinus acronotus in the Gulf of Mexico. J. Fish. Biol., 70(2): 428-440. doi:10.1111/j.1095-8649.2007.01314.x

Thomson, K.S. – 1976. On the heterocercal tail in sharks. Paleobiology, 2: 19-38.

Thomson, K.S. and D.E. Simanek. – 1977. Body form and locomotion in sharks. Am. Zool., 17: 343-354.

Tortonese, E. – 1956. Fauna d’Italia. Leptocardia, Ciclostomata, Selachii. Calderini, Bologna.

Underwood, A.J. – 1997. Experiments in ecology: their logical design and interpretation using analysis of variance. Cambridge University Press, Cambridge, U.K.

Webb, P.W. – 1978. Hydrodynamics: nonscombroid fish. In: W.S. Hoar and D.J Randall (eds.), Fish Physiology, vol. 7, pp.189-237. Academic Press, London.

Webb, P.W. – 1984. Form and function in fish swimming. Sci. Am., 251: 58-68.

Webb, P.W. and R.S. Keyes. – 1982. Swimming kinematics of sharks. Fish. B.-NOAA, 80: 803-812.

Webb, P.W. and R.W. Blake. – 1985. Swimming. In: M. Hildebrand, D.M. Bramble, K.F. Liem and D.B. Wake (eds), Functional vertebrate Morphology, pp. 111-128. Belknap Press, Harward.

Wilga, C.D. and G.V. Lauder. – 2000. Three-dimensional kinematics and wake structure of the pectoral fins during locomotion in leopard sharks, Triakis semifasciata. J. Exp. Biol., 203: 2261-2278.

Wilga, C.D. and G.V. Lauder. – 2001. Functional morphology of the pectoral fins in bamboo sharks, Chiloscyllium plagiosum: benthic versus pelagic station holding. J. Morph., 249: 195-209. doi:10.1002/jmor.1049 PMid:11517464

Wilga, C.D. and G.V. Lauder. – 2002. Function of the heterocercal tail in sharks: quantitative wake dynamics during steady horizontal swimming and vertical manoeuvring. J. Exp. Biol., 205: 2365-2374.

Wilga, C.D. and G.V. Lauder. – 2004 a. Biomechanics: hydrodynamic function of the shark’s tail. Nature, 430: 850. doi:10.1038/430850a PMid:15318211

Wilga, C.D. and G.V. Lauder. – 2004 b. Biomechanics of locomotion in sharks, rays, and chimeras. In M. Hildebrand, D.MBramble, K.F. Liem and D.B. Wake (eds.), Biology of sharks and their relatives, pp. 139-164. CRCPress: LLC.

Yano, K. – 1995. Reproductive biology of the black dogfish, Centroscyllium fabricii, collected from waters off western Greenland. J. Mar. Biol. Ass. UK, 75: 285-310. doi:10.1017/S002531540001818X

Copyright (c) 2010 Consejo Superior de Investigaciones Científicas (CSIC)

Creative Commons License
This work is licensed under a Creative Commons Attribution 4.0 International License.

Contact us

Technical support