Climatic gradients impose clinal selection on animal ecological and physiological performance, often promoting geographic body size clines. Bergmann’s rule predicts that body size increases with decreasing environmental temperatures given the need to retain body-heat through adjustments of body-mass-to-surface-area ratio. This prediction generally holds for endotherms, but remains controversial for ectotherms. An alternative interpretation, the ‘resource rule’, suggests that food abundance, primary productivity and precipitation (which, unlike temperature, do not necessarily correlate with geography), drive body size clines. We investigate geographic variation in body size within 65 species of lizards and snakes (squamates) based on an intercontinental dataset (6,500+ specimens belonging to 56 Israeli species, and multiple populations of nine Liolaemus species from Argentina and Chile). Bergmann’s rule is only rarely supported by our data (in four species, 6 %), whereas six species (9 %) follow its converse (hence, it is unsupported in 94 % of cases). Similarly, size increases with resource abundance in only 12 species (18 %). Therefore, although neither of the rules is supported, factors suggested by the resource rule are better predictors of body size than temperature. Surprisingly, we show that some measures of the extent of a species’ climatic envelope do not affect the likelihood of it showing a size-climate relationship. We conclude that negative size-temperature associations are an exception rather than a generality among squamates.
This is a preview of subscription content, log in to check access.
Buy single article
Instant access to the full article PDF.
Price includes VAT for USA
Subscribe to journal
Immediate online access to all issues from 2019. Subscription will auto renew annually.
This is the net price. Taxes to be calculated in checkout.
Adams, D. C., & Church, J. O. (2008). Amphibians do not follow Bergmann’s rule. Evolution, 62, 413–420.
Amarello, M., Nowak, E. M., Taylor, E. N., Schuett, G. W., Repp, R. A., Rosen, P. C., et al. (2010). Potential environmental influences on variation in body size and sexual size dimorphism among Arizona populations of the western diamond-backed rattlesnake (Crotalus atrox). Journal of Arid Environments, 74, 1443–1449.
Ashton, K. G., & Feldman, C. R. (2003). Bergmann’s rule in nonavian reptiles: Turtles follow it, lizards and snakes reverse it. Evolution, 57, 1151–1163.
Ashton, K. G., Tracy, M. C., & de Queiroz, A. (2000). Is Bergmann’s rule valid for mammals? American Naturalist, 156, 390–415.
Bar, A., & Haimovitch, G. (2011). A field guide to reptiles and amphibians of Israel. Israel: Pazbar.
Bergmann, C. (1847). Ueber die Verhaltnisse der warmeokonomie der thiere zu ihrer grosse. Gottinger Studien, 3, 595–708.
Blackburn, T. M., & Gaston, K. J. (2003). Macroecology. Concepts and consequences. Oxford: Blackwell.
Blackburn, T. M., Gaston, K. J., & Loder, N. (1999). Geographic gradients in body size: A clarification of Bergmann’s rule. Diversity and Distributions, 5, 165–174.
Blackburn, T. M., & Ruggiero, A. (2001). Latitude, elevation and body mass variation in Andean passerine birds. Global Ecology and Biogeography, 10, 245–259.
Blanckenhorn, W. U., & Demont, M. (2004). Bergmann and converse Bergmann latitudinal clines in arthropods: Two ends of a continuum? Integrative and Comparative Biology, 44, 413–424.
Cei, J. M. (1993). Reptiles del noroeste, nordeste y este de la Argentina. Herpetofauna de las selvas subtropicales, puna y pampas (p. 947). Torino: Museo Regionale di Scienze Naturali di Torino.
Costa, G. C., Vitt, L. G., Pianka, E. R., Mesquita, D. O., & Colli, G. R. (2008). Optimal foraging constrains macroecological patterns: Body size and dietary niche breadth in lizards. Global Ecology and Biogeography, 17, 670–677.
Cruz, F. B., Fitzgerald, L. A., Espinoza, R. E., & Schulte, J. A. (2005). The importance of phylogenetic scale in tests of Bergmann’s and Rapoport’s rules: Lessons from a clade of South American lizards. Journal of Evolutionary Biology, 18, 1559–1574.
Dayan, T., Tchernov, E., Yom-Tov, Y., & Simberloff, D. (1989). Ecological character displacement in Saharo-Arabian Vulpes: Outfoxing Bergmann’s rule. Oikos, 55, 263–272.
Dillon, M. E., Frazier, M. R., & Dudley, R. (2006). Into thin air: Physiology and evolution of alpine insects. Integrative and Comparative Biology, 46, 49–61.
Espinoza, R. E., Wiens, J. J., & Tracy, C. R. (2004). Recurrent evolution of herbivory in small, cold-climate lizards: Breaking the ecophysiological rules of reptilian herbivory. Proceedings of the National Academy of Sciences, USA, 101, 16819–16824.
Feldman, A. & Meiri, S. (2013). Length-mass allometry in snakes. Biological Journal of the Linnean Society (In Press).
Frankenberg, E., & Werner, Y. L. (1992). Egg, clutch and maternal sizes in lizards: Intra- and interspecific relations in near-eastern Agamidae and Lacertidae. Herpetological Journal, 2, 7–18.
Goldberg, S. R. (2012a). Reproduction in Kotschy’s gecko Mediodactylus kotschyi (Squamata: Gekkonidae) from the Greek Islands and Israel. Herpetological Bulletin, 119, 15–18.
Goldberg, S. R. (2012b). Reproduction in the desert lacerta, Mesalina guttulata, from Israel (Squamata: Lacertidae). Zoology in the Middle East, 56, 27–38.
Gur, H., & Gur, M. K. (2012). Is spatial variation in food availability an explanation for a Bergmannian size pattern in a North American hibernating, burrowing mammal? An information-theoretic approach. Journal of Zoology, 287, 104–114.
Hijmans, R. J., Cameron, S. E., Parra, J. L., Jones, P. G., & Jarvis, A. (2005). Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25, 1965–1978.
Huston, M. A., & Wolverton, S. (2011). Regulation of animal size by NPP, Bergmann’s rule, and related phenomena. Ecological Monographs, 81, 349–405.
Imhoff, M. L., Bounoua, L., Ricketts, T., Loucks, C., Harriss, R., & Lawrence, W. T. (2004). Global patterns in human consumption of net primary production. Nature, 429, 870–873.
Iverson, J. B. (1982). Adaptations to herbivory in Iguanine lizards. In G. M. Burghardt, & A. S. Rand (Eds.), Iguanas of the world. Their behaviour, ecology and conservation. New Jersey: Noyes Publications. (pp. 60–76).
James, F. C. (1970). Geographic size variations in birds and its relationship with climate. Ecology, 51, 365–390.
King, G. (1996). Reptiles and herbivory. New York: Chapman and Hall.
Labra, A., Pienaar, J., & Hansen, T. F. (2009). Evolution of thermal physiology in Liolaemus lizards: Adaptation, phylogenetic inertia, and niche tracking. American Naturalist, 174, 204–220.
Lindsey, C. C. (1966). Body sizes of poikilotherm vertebrates at different latitudes. Evolution, 20, 456–465.
Lovegrove, B. G. (2000). The zoogeography of mammalian basal metabolic rate. American Naturalist, 156, 201–219.
Mayr, E. (1956). Geographical character gradients and climatic adaptation. Evolution, 10, 105–108.
McNab, B. K. (2010). Geographic and temporal correlations of mammalian size reconsidered: A resource rule. Oecologia, 164, 13–23.
Meiri, S. (2008). Evolution and ecology of lizard body sizes. Global Ecology and Biogeography, 17, 724–734.
Meiri, S. (2010). Length-weight allometries in lizards. Journal of Zoology, 281, 218–226.
Meiri, S. (2011). Bergmann’s rule: What’s in a name? Global Ecology and Biogeography, 20, 203–207.
Meiri, S., Bauer, A. M., Chirio, L., Colli, G. R., Das, I. Doan, T. M., Feldman, A., Castro-Herrera, F., Novosolov, M., Pafilis, P., Pincheira-Donoso, D., Powney, G., Torres-Carvajal, O., Uetz, P., & Van Damme, R. (2013). Are lizards feeling the heat? A tale of ecology and evolution under two temperatures. Global Ecology and Biogeography (In Press). doi:10.1111/geb.12053.
Meiri, S., Brown, J. H., & Sibly, R. M. (2012). The ecology of lizard reproductive output. Global Ecology and Biogeography, 21, 592–602.
Meiri, S., & Dayan, T. (2003). On the validity of Bergmann’s rule. Journal of Biogeography, 30, 331–351.
Meiri, S., Meijaard, E., Wich, S., Groves, C., & Helgen, K. (2008). Mammals of Borneo—small size on a large island. Journal of Biogeography, 35, 1087–1094.
Meiri, S., & Thomas, G. H. (2007). The geography of body size—challenges of the interspecific approach. Global Ecology and Biogeography, 16, 689–693.
Meiri, S., Yom-Tov, Y., & Geffen, E. (2007). What determines conformity to Bergmann’s rule? Global Ecology and Biogeography, 16, 788–794.
Mendelssohn, H. (1963). On the biology of the venomous snakes in Israel, Part I. Israel Journal of Zoology, 12, 143–170.
O’Brien, R. M. (2007). A caution regarding rules of thumb for variance inflation factors. Quality and Quantity, 41, 673–690.
Olalla-Tarraga, M. A. (2011). “Nullius in Bergmann” or the pluralistic approach to ecogeographical rules: A reply to Watt et al. (2010). Oikos, 120, 1441–1444.
Olalla-Tarraga, M. A., Rodriguez, M. A., & Hawkins, B. A. (2006). Broad-scale patterns of body size in squamate reptiles of Europe and North America. Journal of Biogeography, 33, 781–793.
Oufiero, C. E., Adolph, S. C., Gartner, G. E. A., & Garland, T. (2011). Latitudinal and climatic variation in body size and dorsal scale counts in Sceloporus lizards: A phylogenetic perspective. Evolution, 65, 3590–3607.
Partridge, L., & Coyne, J. A. (1997). Bergmann’s rule in ectotherms: Is it adaptive? Evolution, 51, 632–635.
Peters, R. H. (1983). The ecological implications of body size. Cambridge: Cambridge University Press.
Pincheira-Donoso, D. (2010). The balance between predictions and evidence and the search for universal macroecological patterns: Taking Bergmann’s rule back to its endothermic origin. Theory in Biosciences, 129, 247–253.
Pincheira-Donoso, D. (2011). Predictable variation of range-sizes across an extreme environmental gradient in a lizard adaptive radiation: Evolutionary and ecological inferences. PLoS One, 6, e28942.
Pincheira-Donoso, D., Bauer, A. M., Meiri, S., & Uetz, P. (2013). Global taxonomic diversity of living reptiles. PLoS One, 8, e59741.
Pincheira-Donoso, D., Fox, S. F., Scolaro, J. A., Ibargüengoytía, N., Acosta, J. C., Corbalán, V., et al. (2011). Body size dimensions in lizard ecological and evolutionary research: Exploring the predictive power of mass estimation equations in two Liolaemidae radiations. Herpetological Journal, 21, 35–42.
Pincheira-Donoso, D., Hodgson, D. J., & Tregenza, T. (2008a). The evolution of body size under environmental gradients in ectotherms: Why should Bergmann’s rule apply to lizards? BMC Evolutionary Biology, 8, 68.
Pincheira-Donoso, D., & Núñez, H. (2005). Las especies chilenas del género Liolaemus. Taxonomía, sistemática y evolución. Publicación Ocasional del Museo Nacional de Historia Natural de Chile, 59, 1–487.
Pincheira-Donoso, D., Scolaro, J. A., & Sura, P. (2008b). A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania). Zootaxa, 1800, 1–85.
Pincheira-Donoso, D., Tregenza, T., & Hodgson, D. J. (2007). Body size evolution in South American Liolaemus lizards of the boulengeri clade: A contrasting reassessment. Journal of Evolutionary Biology, 20, 2067–2071.
Raia, P., & Meiri, S. (2006). The island rule in large mammals: Paleontology meets ecology. Evolution, 60, 1731–1742.
Roitberg, E. S., Orlova, V. F., Kuranova, V. N., Bulakhova, N. A., Zinenko, O. I., Ljubisavljevic, K., et al. (2011). Inter-observer and intra-observer differences in measuring body length: A test in the common lizard, Zootoca vivipara. Amphibia-Reptilia, 32, 477–484.
Rosenzweig, M. L. (1968). The strategy of body size in mammalian carnivores. American Midland Naturalist, 80, 299–315.
Schmidt-Nielsen, K. (1984). Scaling. Why is animal size so important?. Cambridge: Cambridge University Press.
Sears, M. W., & Angilletta, M. J. (2004). Body size clines in Sceloporus lizards: Proximate mechanisms and demographic constraints. Integrative and Comparative Biology, 44, 433–442.
Tracy, C. R. (1999). Differences in body size among chuckwalla (Sauromalus obesus) populations. Ecology, 80, 259–271.
Uetz, P. (2012). The Reptile Database. Available at http://www.reptile-database.org. Accessed 31 Mar 2012.
Vezina, A. F. (1985). Empirical relationships between predator and prey size among terrestrial vertebrate predators. Oecologia, 67, 555–565.
Volynchik, S. (2012). Morphological variability in Vipera palaestinae along an environmental gradient. Asian Herpetological Research, 3, 227–239.
Waldron, A. (2007). Null models of geographic range size evolution reaffirm its heritability. American Naturalist, 170, 221–231.
Watt, C., Mitchell, S., & Salewski, V. (2010). Bergmann’s rule; a concept cluster? Oikos, 119, 89–100.
Webb, T. J., & Gaston, K. J. (2003). On the heritability of geographic range sizes. American Naturalist, 166, 129–135.
Wilson, A. B. (2009). Fecundity selection predicts Bergmann’s rule in syngnathid fishes. Molecular Ecology, 18, 1263–1272.
Yom-Tov, Y. (2003). Body sizes of carnivores commensal with humans have increased over the past 50 years. Functional Ecology, 17, 323–327.
Yom-Tov, Y., & Geffen, E. (2006). The determination of mammal body size: Ambient temperature or food? Oecologia, 148, 213–218.
We thank Erez Maza for invaluable help resolving the correct geographic origin of species in the TAUM, and Anat Feldman and Stanislav Volynchic for valuable discussion. We are also grateful to referee’s and editor’s comments that greatly improved our manuscript. D.P.-D. thanks the Leverhulme Trust and a University of Lincoln Faculty Starting Funding for financial support. D.P.D. dedicates this paper to Natalia Feltrin, a young Argentinean ecologist, and a good friend, who devoted her career to the study of Liolaemus lizards, and who recently passed away in a road accident. Her life was interrupted on her way to her Ph.D viva. Natalia’s work will be greatly remembered and appreciated.
Daniel Pincheira-Donoso and Shai Meiri contributed equally.
About this article
Cite this article
Pincheira-Donoso, D., Meiri, S. An Intercontinental Analysis of Climate-Driven Body Size Clines in Reptiles: No Support for Patterns, No Signals of Processes. Evol Biol 40, 562–578 (2013). https://doi.org/10.1007/s11692-013-9232-9
- Bergmann’s rule
- Resource rule
- Climatic variability
- Geographic variation in body size