Morphology of Vipera palaestinae: Intraspecific Variability and Sexual Dimorphism

Stanislav Volynchik

Abstract


I present new data on intraspecific variations in external morphology and sexual dimorphism in different characteristics of Vipers, Vipera palaestinae in Israel. Corporal proportions and scalation pattern of museum specimens of this snake, the most common viperid species in Israel, were recorded and analyzed. Strong sexual dimorphism was found in tail measurements (its relative length, and number of subcaudal scales), head proportions and in the number of ventral scales. The sexual differences were evident in the head scalation – females have significantly more supralabial and sublabial scales; males, in turn exceed females in circumorbital scale counts. There is no sexual size dimorphism (SSD) in average body weight, snout-vent and total length. The sexes display a similar allometry between SVL/head and SVL/tail length; between tail length and the number of subcaudal scales (weak, but significant relationship). There is no correlation between SVL and the number of ventral or dorsal scales. The morphological plasticity caused by reproduction requirements is well registered and was evident in relating to tail measurements and body scalation, although the possible causes and selective forces of the pholidosis intersexual distinctions can be cleared up through future ecological studies.

Keywords


Vipera palaestinae; snakes; scalation; intraspecific variability; sexual dimorphism; Viperidae

Full Text:

PDF

References


Alexander A. A. and Gans C. (1966), «The pattern of dermal- vertebral correlation in snakes and amphisbaenians», Zool. Mededelingen, 41, 171 – 190.

Allon N. and Kochva E. (1974), «The quantities of venom injected into prey of different size by Vipera palaestinae in a single bite», J. Exp. Zool., 188, 71 – 76.

Andrén C. and Nilson G. (1981), «Reproductive success and risk of predation normal and melanistic colour morphs of the adder, Vipera berus», Biol. J. Linn. Soc., 15, 235 – 246.

Andrews R. M. (1982), «Patterns of growth in reptiles», in: C. Gans and F. H. Pough (eds.), Biology of the Reptilia. Vol. 13, Acad. Press, New York, pp. 273 – 320.

Arnold S. J. (1988), «Quantitative genetics and selection in natural populations: microevolution of vertebral numbers in the garter snake Thamnophis elegans», in: B. S. Weir, E. J. Eisen, M. M. Goodman, and G. Namkoong (eds.), Proc. of the 2nd Int. Conf. on Quantitative Genetics, Sinauer, Sunderland, MA, pp. 619 – 636.

Arnold S. J. and Bennett A. F. (1988), «Behavioural variation in natural populations. V. Morphological correlates of locomotion in the garter snake (Thamnophis radix)», Biol. J. Linn. Soc., 34, 175 – 190.

Arnold S. J. (1993), «Foraging theory and prey-size predator-size relations in snakes», in: R. A. Seigel and J. T. Collins (eds.), Snakes. Ecology and Behavior, McGraw-Hill, New York, pp. 87 – 116.

Aubret F., Bonnet X., Harris M., and Maumelat S. (2005), «Sex differences in body size and ectoparasite load in the ball python, Python regius», J. Herpetol., 39, 315 – 320.

Babocsay G. (2006), Geographic Variation of Morphological Characters in the Venomous Snake, Echis coloratus: Environmental correlates, Taxonomic and Practical Implications. Ph. D. Dissertation, Hebrew Univ. of Jerusalem, Israel.

Barash A. and Hoofien J. (1956), Reptiles of Israel [in Hebrew], Hakibutz Hameuchad Ltd.

Bonnet X., Shine R., Naulleau G., and Thiburce C. (2001), «Plastic vipers: influence of food intake on the size and shape of Gaboon vipers (Bitis gabonica)», J. Zool., 255, 341 – 351.

Boulenger G. A. (1913), The Snakes of Europe, Methuen & Co., London.

Bouskila A. and Amitai P. (2001), Handbook of Amphibians and Reptiles of Israel [in Hebrew], Keter Publ. House Ltd., Jerusalem.

Brito J. C., Santos X., Pleguezuelos J. M., Fahd S., Llorente G., and Parellada X. (2006), «Morphological variability of the Lataste’s viper (Vipera latastei) and the Atlas dwarf viper (Vipera monticola): patterns of biogeographical distribution and taxonomy», Amphibia–Reptilia, 27, 219 – 240.

Disi A., Modrý D., Nečas P., and Rifai L. (2001), Amphibians and Reptiles of the Hashemite Kingdom of Jordan. An Atlas and Field Guide, Edition Chimaira, Frankfurt am Main.

Faiman R., Razzeti E., Seligman H., and Werner Y. L. (2005), «Sexual dimorphism in snakes», Proc. of the 55th Ann. Meet. of the Zool. Soc. of Israel. Isr. J. Zool., 51, 63 – 64.

Fitch H. S. (1970), «Reproductive cycles in lizards and snakes», Publ. Mus. Nat. Hist. Univ. Kansas, 52, 1 – 247.

Fitch H. S. (1981), «Sexual size differences in reptiles», Publ. Mus. Nat. Hist. Univ. Kansas, 70, 1 – 72.

Ford N. B. and Seigel R. A. (1989), «Relationships among body size, clutch size, and egg size in three species of oviparous snakes», Herpetologica, 45, 75 – 83.

Forsman A. (1991a), «Variation in sexual size dimorphism and maximum body size among adder populations: effects of prey size», J. Anim. Ecol., 60, 253 – 267.

Forsman A. (1991b), «Adaptive variation in head size in Vipera berus L. populations», Biol. J. Linn. Soc., 43, 281 – 296.

Fox W., Gordon C., and Fox M. H. (1961), «Morphological effects of low temperature during the embryonic development of the garter snake, Thamnophis elegans», Zoologica, 46, 57 – 71.

Garrigues T., Dauga C., Ferquel E., Choumet V., and Failloux A.-B. (2005), «Molecular phylogeny of Vipera Laurenti, 1768 and the related genera Macrovipera (Reuss, 1927) and Daboia (Gray, 1842), with comments about neurotoxic Vipera aspis aspis populations», Mol. Phylogen. Evol., 35, 35 – 47.

Gibbons J. W. (1972), «Reproduction, growth and sexual dimorphism in the canebrake rattlesnake (Crotalus horridus atricaudatus)», Copeia, 1972, 222 – 226.

Gibbons J. W. and Lovich J. E. (1990), «Sexual dimorphism in turtles with emphasis on the slider turtle (Trachemys scripta)», Herpetol. Monogr., 4, 1 – 29.

Gitter S. (1962), «Toxicology and biochemistry of the venoms of some Israeli snakes», Folia Med., 21, 12 – 14.

Groombridge B. (1980), A Phyletic Analysis of Viperine Snakes. Ph. D. Dissertation, Polytechnic College, London.

Groombridge B. (1986), «Phyletic relationships among viperine snakes», in: Z. Roček (ed.), Studies in Herpetology, Charles Univ., Prague, pp. 219 – 222.

Hasegawa M. and Mori A. (2008), «Does a gigantic insular snake grow faster or live longer to be gigantic? Evidence from a long-term field study», S. Am. J. Herpetol., 3, 145 – 154.

Houston D. and Shine R. (1993), «Sexual dimorphism and niche divergence: feeding habits of the Arafura filesnake», J. Anim. Ecol., 62, 737 – 748.

Jayne B. C. and Bennett A. F. (1990), «Selection on locomotor performance capacity in a natural population of garter snakes», Evolution, 44, 1204 – 1229.

King W. (1959), «Vertebra duplication, an osteological anomaly widespread in snakes», Herpetologica, 15, 87 – 88.

King R. B. (1989), «Sexual dimorphism in snake tail length: sexual selection, natural selection, or morphological constraint?» Biol. J. Linn. Soc., 38, 133 – 154.

King R. B. (1997), «Variation in brown snake (Storeria dekayi) morphology and scalation: sex, family, and microgeographic differences», J. Herpetol., 31, 335 – 346.

King R. B., Bittner T. D., Queral-Regil A., and Cline J. H. (1999), «Sexual dimorphism in neonate and adult snakes», J. Zool., 247, 19 – 28.

Klauber L. M. (1943), «Tail length differences in snakes with note on sexual dimorphism and the coefficient of divergence», Bull. Zool. Soc. San Diego, 18, 1 – 60.

Klauber L. M. (1945), «Herpetological correlations. 1. Correlations in homogeneous populations», Bull. Zool. Soc. San Diego, 21, 5 – 101.

Klauber L. M. (1956), Rattlesnakes. Their Habits, Life Histories and Influence on Mankind, 1st Edition, Univ. of California Press, Berkeley, California.

Kochva E. (1960), «A quantitative study of venom secretion by Vipera palaestinae», Am. J. Trop. Med. Hyg., 9, 381 – 390.

Kochva E. and Gans C. (1965), «The venom gland of Vipera palaestinae with comments on the glands of some other viperines», Acta Anat., 62, 365 – 401.

Kochva E. (2004), Venomous Snakes and Snake Venoms [in Hebrew], Teva HaDvarim, Tel-Aviv Univ., Israel.

Kramer E. (1961), «Variation, Sexual dimorphismus, Wachstum und Taxonomie von Vipera ursinii (Bonaparte, 1835) und Vipera kaznakovi Nikolskij, 1909», Rev. suisse Zool., 68, 627 – 725.

Krause M. A., Burghardt G. M., and Gillingham J. C. (2003), «Body size plasticity and local variation of relative head and body size sexual dimorphism In Garter Snakes (Thamnophis sirtalis)», J. Zool., 261, 399 – 407.

Lenk P., Kalyabina S., Wink M., and Joger U. (2001), «Evolutionary relationships among the true vipers (Viperinae) inferred from mitochondrial DNA sequences», Mol. Phylogen. Evol., 19, 94 – 104.

Lindell L. E., Forsman A., and Merilä J. (1993), «Variation in number of ventral scales in snakes: effects on body size, growth rate and survival in the adder, Vipera berus», J. Zool., 230, 101 – 115.

Madsen T. and Shine R. (1993a), «Phenotypic plasticity in body sizes and sexual size dimorphism in European grass snakes», Evolution, 47, 321 – 325.

Madsen T. and Shine R. (1993b), «Male mating success and body size in European grass snake», Copeia, 1993, 561 – 564.

Madsen T. and Shine R. (1994), «Costs of reproduction influence the evolution of sexual size dimorphism in snakes», Evolution, 48, 1389 – 1397.

Madsen T., Shine R., Loman J., and Hakansson T. (1993), «Determinants of mating success in male adders, Vipera berus», Anim. Behav., 45, 491 – 499.

Mallow D., Ludwig D., and Nilson G. (2003), True Vipers: Natural History and Toxinology of Old World Vipers, Krieger Publ. Comp., Malabar, Florida.

Mendelssohn H. (1963), «On the biology of the venomous snakes in Israel. Part I», Isr. J. Zool., 12, 143 – 170.

Merilä J., Forsman A., and Lindell L. E. (1992), «High frequency of ventral scales anomalies in Vipera berus populations», Copeia, 1992, 1127 – 1130.

Mertens R. (1952), «Türkiye amfibi ve reptilleri hakkinda. Amphibien und Reptilien aus der Türkei», Istanbul Üniv. Fakült. Mecmuasi, 17, 41 – 75.

Monney J.-C., Luiselli L., and Capula M. (1995), «Correlates of melanism in a population of adders (Vipera berus) from the Swiss Alps and comparisons with other alpine populations», Amphibia–Reptilia, 16, 323 – 330.

Nilson G. and Andrén C. (1986), The Mountain Vipers of the Middle East — The Vipera xanthina Complex, Bonn. Zool. Monogr. 20, Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn.

Nilson G. and Andrén C. (2001), «The meadow and steppe vipers of Europe and Asia — the Vipera (Acridophaga) ursinii complex», Acta Zool. Acad. Sci. Hung., 47, 87 – 267.

Osgood D. W. (1978), «Effects of temperature on the development of meristic characters in Natrix fasciata», Copeia, 1978, 33 – 47.

Pearson D., Shine R., and How R. (2002), «Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)», Biol. J. Linn. Soc., 77, 113 – 125.

Queral-Regil A. and King R. B. (1998), «Evidence for phenotypic plasticity in snake body size and relative head dimensions in response to amount and size of prey», Copeia, 1998, 423 – 429.

Rivas J. A. and Burghardt G. M. (2001), «Understanding sexual size dimorphism in snakes: wearing the snake’s shoes», Anim. Behav., 62, F1 – F6.

Saint-Girons H. (1978), «Morphologie externe comparée et systematique des Viperes d’Europe (Reptilia, Viperidae)», Rev. suisse Zool., 85, 565 – 595.

Santos X., Gonzalez-Solis J., and Llorente G. A. (2000), «Variation in the diet of the viperine snake Natrix maura in relation to prey availability», Ecography, 23, 185 – 192.

Shine R. (1978), «Sexual size dimorphism and male combat in snakes», Oecologia, 33, 269 – 277.

Shine R. (1988), «The evolution of large body size in females: a critique of Darwin’s «fecundity advantage» model», Am. Naturalist, 131, 124 – 131.

Shine R. (1989), «Ecological causes for the evolution of sexual dimorphism: a review of the evidence», Quart. Rev. Biol., 64, 419 – 461.

Shine R. (1991), «Intersexual dietary divergence and the evolution of sexual dimorphism in snakes», Am. Naturalist, 138, 103 – 122.

Shine R. (1993), «Sexual dimorphism in snakes», in: R. A. Seigel and J. T. Collins (eds.), Snakes: Ecology and Behavior, McGraw-Hill, New York, pp. 49 – 86.

Shine R. (1994), «Sexual size dimorphism in snakes revisited», Copeia, 1994, 326 – 346.

Shine R. (2000), «Vertebral numbers in male and female snakes: the roles of natural, sexual and fecundity selection», J. Evol. Biol., 13, 455 – 465.

Shine R. and Seigel R. A. (1996), «A neglected life-history trait: clutch-size variance in snakes», J. Zool., 239, 209 – 223.

Shine R., Branch W. R., Harlow P. S., and Webb J. K. (1996), «Sexual dimorphism, reproductive biology, and food habits of two species of African filesnakes (Mehelya, Colubridae)», J. Zool., 240, 327 – 340.

Shine R., Olsson M. M., Moore I. T., LeMaster M. P., and Mason R. T. (1999), «Why do male snakes have longer tails than females?» Proc. R. Soc. B, 266, 2147 – 2151.

Shine R., Olsson M. M., Moore I. T., LeMaster M. P., Greene M., and Mason R. T. (2000), «Body size enhances mating success in male garter snakes», Anim. Behav., 59, F4 – F11.

Simmons J. E. (2002), Herpetological Collecting and Collections Management, Revised Edition, Soc. Study Amph. Rept., Herpetol. Circ., 31.

Stuart J. N. (1995), «Observations on formalin-inducted darkening of herpetological specimens», SPNHC Coll. Forum, 11, 39 – 45.

Stümpel N. and Joger U. (2009), «Recent advances in phylogeny and taxonomy of Near and Middle Eastern Vipers — an update», ZooKeys, 31, 179 – 191.

Theodor O. (1955), On Poisonous Snakes and Snake Bite in Israel [in Hebrew], The Israel Scientific Press, Jerusalem.

Thorpe R. S. (1989), «Pattern and function of sexual dimorphism: A biometric study of character variation in the grass snake (Natrix natrix, Colubridae) due to sex and its interaction with geography», Copeia, 1989, 53 – 63.

Tomović L., Radojičić J., Džukić G., and Kalezić M. (2002), «Sexual dimorphism of the Sand Viper (Vipera ammodytes L.) from the central part of Balkan peninsula», Russ. J. Herpetol., 9(1), 69 – 76.

Vincent S. E., Herrel A., and Irschick D. J. (2004), «Sexual dimorphism in head shape and diet in the cottonmouth snake (Agkistrodon piscivorus)», J. Zool., 264, 53 – 59.

Volynchik S. (2001), «Biology and ecology of Vipera berus L. and Natrix natrix L. in Southern Transuralia», Proc. of the 1st Conf. of Nikolsky Herpetol. Soc. [in Russian with English summary], Moscow, pp. 58 – 61.

Volynchik S. (2002), Ecology of Snakes (Serpentes) of Southern Transuralia. Ph. D. Dissertation [in Russian], Izd. SGU, Surgut, Russia.

Weatherhead P. J., Barry F. E., Brown G. P., and Forbes M. R. (1995), «Sex ratios, mating behavior and sexual size dimorphism of the northern water snake, Nerodia sipedon», Behav. Ecol. Sociobiol., 36, 301 – 311.

Werner F. (1938), «Eine verkannte Viper (Vipera palaestinae n. sp.)», Zool. Anz., 122, 313 – 318.

Werner Y. L. (1994), «Head size variation in Cerastes (Ophidia: Viperidae) parallels body size variation in potential prey (Rodentia: Gerbillinae)», The Snake, 26, 57 – 60.

Werner Y. L. and Seifan T. (2006), «Eye size in geckos: asymmetry, allometry, sexual dimorphism, and behavioral correlates», J. Morphol., 267, 1486 – 1500.

Wolf M. and Werner Y. L. (1994), «The striped color pattern and striped/non-striped polymorphism in snakes (Reptilia: Ophidia)», Biol. Rev., 69, 599 – 610.

Zuffi M. A. L. and Bonnet X. (1999), «Italian subspecies of the asp viper, Vipera aspis: patterns of variability and distribution», Ital. J. Zool., 66, 87 – 95.


Refbacks

  • There are currently no refbacks.