Geographic variation of body size in New World anurans: energy and water in a balance

The validity of Bergmann's rule, perhaps the best known ecogeographical rule, has been questioned for ectothermic species. Here, we explore the interspecific version of the rule documenting body size gradients for anurans across the whole New World and evaluating which environmental variables best explain the observed patterns. We assembled a dataset of body sizes for 2761 anuran species of the Western Hemisphere and conducted assemblage‐based and cross‐species analyses that consider the spatial and phylogenetic structure in the data. In accordance with heat and water‐related explanations for body size clines, we found a consistent association of median body size and potential evapotranspiration across the New World. A relevant role of water availability also emerges, suggesting the joint importance of body size for thermoregulation and hydroregulation in anurans. Anurans do not follow a simple Bergmannian pattern of increasing size towards high latitudes. Consistent with previous regional findings, our Hemisphere‐wide analyses detect that the geographic variation in anuran body sizes is highly dependent on a trade‐off between heat and water balance. The observed size‐climate relationships possibly emerge from the interplay between thermoregulatory abilities and the benefits inherent to reduced surface‐to‐volume ratios in larger species, which decrease the rates of evaporative water loss and favour heat retention. Our results also show how temperature becomes important for species that are directly in contact with the substrate and water, like burrowing and terrestrial anurans, while arboreal species exhibit a body size cline linked with potential evapotranspiration.     

Environment and morphology in Australian Aborigines: a re-analysis of the Birdsell database

Pursuant to his major research interest in the cultural ecology of hunter-gatherers, Birdsell collected an unparalleled body of phenotypic data on Aboriginal Australians during the mid twentieth century. Birdsell did not explicitly relate the geographic patterning in his data to Australia's climatic variation, instead arguing that the observable differences between groups reflect multiple origins of Australian Aborigines. In this article, bivariate correlation and multivariate analyses demonstrate statistically significant associations between climatic variables and the body build of Australians that are consistent with the theoretical expectations of Bergmann's and Allen's rules. While Australian Aborigines in comparison to Eurasian and New World populations can be generally described as long-headed, linear in build, and characterized by elongated distal limbs, the variation in this morphological pattern across the continent evidently reflects biological adaptation to local Holocene climates. These results add to a growing body of evidence for the role of environmental selection in the development of modern human variation.     

Deep phylogeny,net primary productivity,and global body size gradient in birds

Body size is evolutionarily constrained, but the influence of phylogenetic relationships on global body size (i.e. body mass) gradients is unexplored. We quantify and map the family‐level phylogenetic and non‐phylogenetic structure of the global gradient of birds, evaluating the extent to which it is influenced by phylogenetic inertia in contrast to heat conservation, resource availability, starvation resistance, niche conservatism, or interspecific competition. Phylogenetic eigenvector regression (PVR) partitioned the global bird body size gradient into phylogenetically autocorrelated (PA) and phylogenetically independent (PI) components. Simple, piecewise, and partial regressions were used to investigate associations between the PA and PI components of body size and environmental correlates, and to quantify independent and overlapping contributions of environment, phylogenetic autocorrelation, and species richness to the body size gradient. Two‐thirds of the geographic variation in bird body size can be explained by phylogenetic relationships at the family level. The global variation in body size, independent of phylogenetic relationships, is most strongly associated with net primary productivity, which is consistent with ‘starvation resistance’. However, the New and Old worlds have very different patterns. We found no independent association of species richness with body size. Despite major unresolved regional differences, deep phylogenetic relationships, heat conservation, and starvation resistance probably operate in concert in shaping the global bird body size gradient in different parts of the world. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Seasonality rather than average temperature explains body size variation in a Neotropical fruit-eating bat

We investigated variation in body size of the widely distributed Neotropical bat Chiroderma villosum across its entire range. Our objective was to verify if the size-related geographic variation in the species is related to environmental variables. We took 13 measurements of 410 specimens from 198 localities in Mesoamerica and South America, and collected information on latitude, longitude, altitude, precipitation, and temperature, totalling 22 variables. We detected clinal variation in size related to latitude and longitude, with a pattern that conforms to the Bergmann's rule. Clinal variation of size along longitude was influenced by the taxonomic component, with subspecies C. v. jesupi being smaller than C. v. villosum. In contrast the latitudinal cline was explained by temperature seasonality and precipitation, with a 14% increase in size between the north and south extremes of the range. In other words, size of individuals is larger in areas with more seasonal oscillations in temperature and with lower precipitation. Our results support the notion that low temperatures alone do not explain large size of mammals in high latitudes. One hypothesis is that large size is favoured in more seasonal climates because somatic growth is faster when resources are abundant, and also larger animals can endure food scarcity better than small ones. We also postulate that pressures related to interspecific competition and resource use may be more intense in more areas marked by seasonal climatic variations. Specifically, a larger size in seasonal areas may allow individuals to explore a wider niche. We suggest that future approaches, refining regional variation in the diet of C. villosum may serve as a further test to this hypothesis.     

Moth body size increases with elevation along a complete tropical elevational gradient for two hyperdiverse clades

The body size of an animal is probably its most important functional trait. For arthropods, environmental drivers of body size variation are still poorly documented and understood, especially in tropical regions. We use a unique dataset for two species‐rich, phylogenetically independent moth taxa (Lepidoptera: Geometridae; Arctiinae), collected along an extensive tropical elevational gradient in Costa Rica, to investigate the correlates and possible causes of body‐size variation. We studied 15 047 specimens (794 species) of Geometridae and 4167 specimens (308 species) of Arctiinae to test the following hypotheses: 1) body size increases with decreasing ambient temperature, as predicted by the temperature–size rule; 2) body size increases with increasing rainfall and primary productivity, as predicted from considerations of starvation resistance; and 3) body size scales allometrically with wing area, as elevation increases, such that wing loading (the ratio of body size to wing area) decreases with increasing elevation to compensate for lower air density. To test these hypotheses, we examined forewing length as a proxy for body size in relation to ambient temperature, rainfall, vegetation index and elevation as explanatory variables in linear and polynomial spatial regression models. We analysed our data separately for males and females using two principal approaches: mean forewing length of species at each site, and mean forewing length of complete local assemblages, weighted by abundance. Body size consistently increased with elevation in both taxa, both approaches, both sexes, and also within species. Temperature was the best predictor for this pattern (–0.98 < r < –0.74), whereas body size was uncorrelated or weakly correlated with rainfall and enhanced vegetation index. Wing loading increased with elevation. Our results support the temperature–size rule as an important mechanism for body size variation in arthropods along tropical elevational gradients, whereas starvation resistance and optimization of flight mechanics seem to be of minor importance.     

Global patterns of body size evolution are driven by precipitation in legless amphibians

Body size shapes ecological interactions across and within species, ultimately influencing the evolution of large‐scale biodiversity patterns. Therefore, macroecological studies of body size provide a link between spatial variation in selection regimes and the evolution of animal assemblages through space. Multiple hypotheses have been formulated to explain the evolution of spatial gradients of animal body size, predominantly driven by thermal (Bergmann's rule), humidity (‘water conservation hypothesis’) and resource constraints (‘resource rule’, ‘seasonality rule’) on physiological homeostasis. However, while integrative tests of all four hypotheses combined are needed, the focus of such empirical efforts needs to move beyond the traditional endotherm–ectotherm dichotomy, to instead interrogate the role that variation in lifestyles within major lineages (e.g. classes) play in creating neglected scenarios of selection via analyses of largely overlooked environment–body size interactions. Here, we test all four rules above using a global database spanning 99% of modern species of an entire Order of legless, predominantly underground‐dwelling amphibians (Gymnophiona, or caecilians). We found a consistent effect of increasing precipitation (and resource abundance) on body size reductions (supporting the water conservation hypothesis), while Bergmann's, the seasonality and resource rules are rejected. We argue that subterranean lifestyles minimize the effects of aboveground selection agents, making humidity a dominant selection pressure – aridity promotes larger body sizes that reduce risk of evaporative dehydration, while smaller sizes occur in wetter environments where dehydration constraints are relaxed. We discuss the links between these principles with the physiological constraints that may have influenced the tropically‐restricted global radiation of caecilians.     

Selection does not favor larger body size at lower temperature in a seed-feeding beetle

Body size of many animals increases with increasing latitude, a phenomenon known as Bergmann's rule (Bergmann clines). Latitudinal gradients in mean temperature are frequently assumed to be the underlying cause of this pattern because temperature covaries systematically with latitude, but whether and how temperature mediates selection on body size is unclear. To test the hypothesis that the "relative" advantage of being larger is greatest at cooler temperatures we compare the fitness of replicate lines of the seed beetle, Stator limbatus, for which body size was manipulated via artificial selection ("Large,"Control," and "Small" lines), when raised at low (22 degrees C) and high (34 degrees C) temperatures. Large-bodied beetles (Large lines) took the longest to develop but had the highest lifetime fecundity, and highest fitness (r(C)), at both low and high temperatures. However, the relative difference between the Large and Small lines did not change with temperature (replicate 2) or was greatest at high temperature (replicate 1), contrary to the prediction that the fitness advantage of being large relative to being small will decline with increasing temperature. Our results are consistent with two previous studies of this seed beetle, but inconsistent with prior studies that suggest that temperature-mediated selection on body size is a major contributor to the production of Bergmann clines. We conclude that other environmental and ecological variables that covary with latitude are more likely to produce the gradient in natural selection responsible for generating Bergmann clines.     

Ecological factors affecting body size of Japanese adolescents

Prefectural data of 17-year-old Japanese adolescents' body size were excerpted from the annual series of Reports on School Health Statistics (Ministry of Education) for the last 35 years, and were correlated with the mean annual temperature of prefectural capitals. A negative correlation was found for both height and weight. Thus, it can be stated that children in colder climates conform to Bergmann's rule that they are larger in their height and weight than children growing up in warmer climates. Partial correlation coefficients indicate that body weight is primarily related to the temperature. Further, the results of a multiple regression analysis, using temperature, daily energy intake (National Nutrition Surveys), and annual per capita income (Annual Reports of Prefectural Accounts) as the independent variables among the 12 districts (groups of prefectures) for the last 25 years, suggest that the observed Bergmann's phenomenon is related to geographical differences in nutrition; people in colder regions consume more calories than those in warmer regions. © 1993 Wiley-Liss, Inc.     

Macroecology and macroevolution of body size in Anolis lizards

Body size is one of the most influential traits affecting many ecological and physiological processes across animal and plant taxa. Studies of the environmental factors shaping body size patterns may evaluate either temporal or spatial dimensions. Here, we analyzed body size evolution in the radiation of Anolis lizards across both geographical and temporal dimensions. We used a set of macroecological and macroevolutionary methods to test current and past environmental effects on geographical gradients of body size and its evolutionary rates. First, we test whether a set of current ecological/physiological hypotheses (heat balance, productivity and seasonality) explains spatial body size gradients. Second, we evaluate how tempo (i.e. evolutionary rates) and mode (i.e. evolutionary process) of body size evolution changed through time and the role of paleo-temperatures on rates of body size evolution during the Cenozoic. We did not find a signature of current environmental variables driving spatial body size gradients. By contrast, we found strong support for a correlation between temperature changes (i.e. climate cooling) during the Cenozoic and rates of body size evolution (i.e. body size diversification). We suggest that patterns of body size evolution in Anolis lizards might be influenced by thermoregulatory behavior across clades and regions.     

Length of activity season drives geographic variation in body size of a widely distributed lizard

Understanding the factors that drive geographic variation in life history is an important challenge in evolutionary ecology. Here, we analyze what predicts geographic variation in life‐history traits of the common lizard, Zootoca vivipara, which has the globally largest distribution range of all terrestrial reptile species. Variation in body size was predicted by differences in the length of activity season, while we found no effects of environmental temperature per se. Females experiencing relatively short activity season mature at a larger size and remain larger on average than females in populations with relatively long activity seasons. Interpopulation variation in fecundity was largely explained by mean body size of females and reproductive mode, with viviparous populations having larger clutch size than oviparous populations. Finally, body size‐fecundity relationship differs between viviparous and oviparous populations, with relatively lower reproductive investment for a given body size in oviparous populations. While the phylogenetic signal was weak overall, the patterns of variation showed spatial effects, perhaps reflecting genetic divergence or geographic variation in additional biotic and abiotic factors. Our findings emphasize that time constraints imposed by the environment rather than ambient temperature play a major role in shaping life histories in the common lizard. This might be attributed to the fact that lizards can attain their preferred body temperature via behavioral thermoregulation across different thermal environments. Length of activity season, defining the maximum time available for lizards to maintain optimal performance, is thus the main environmental factor constraining growth rate and annual rates of mortality. Our results suggest that this factor may partly explain variation in the extent to which different taxa follow ecogeographic rules.     

Cope's rule and the adaptive landscape of dinosaur body size evolution

The largest known dinosaurs weighed at least 20 million times as much as the smallest, indicating exceptional phenotypic divergence. Previous studies have focused on extreme giant sizes, tests of Cope's rule, and miniaturization on the line leading to birds. We use non‐uniform macroevolutionary models based on Ornstein–Uhlenbeck and trend processes to unify these observations, asking: what patterns of evolutionary rates, directionality and constraint explain the diversification of dinosaur body mass? We find that dinosaur evolution is constrained by attraction to discrete body size optima that undergo rare, but abrupt, evolutionary shifts. This model explains both the rarity of multi‐lineage directional trends, and the occurrence of abrupt directional excursions during the origins of groups such as tiny pygostylian birds and giant sauropods. Most expansion of trait space results from rare, constraint‐breaking innovations in just a small number of lineages. These lineages shifted rapidly into novel regions of trait space, occasionally to small sizes, but most often to large or giant sizes. As with Cenozoic mammals, intermediate body sizes were typically attained only transiently by lineages on a trajectory from small to large size. This demonstrates that bimodality in the macroevolutionary adaptive landscape for land vertebrates has existed for more than 200 million years.     

Shifting latitudinal clines in avian body size correlate with global warming in Australian passerines

Intraspecific latitudinal clines in the body size of terrestrial vertebrates, where members of the same species are larger at higher latitudes, are widely interpreted as evidence for natural selection and adaptation to local climate. These clines are predicted to shift in response to climate change. We used museum specimens to measure changes in the body size of eight passerine bird species from south-eastern Australia over approximately the last 100 years. Four species showed significant decreases in body size (1.8–3.6% of wing length) and a shift in latitudinal cline over that period, and a meta-analysis demonstrated a consistent trend across all eight species. Southern high-latitude populations now display the body sizes typical of more northern populations pre-1950, equivalent to a 7° shift in latitude. Using ptilochronology, we found no evidence that these morphological changes were a plastic response to changes in nutrition, a likely non-genetic mechanism for the pattern observed. Our results demonstrate a generalized response by eight avian species to some major environmental change over the last 100 years or so, probably global warming.     

Size matters for lice on birds: Coevolutionary allometry of host and parasite body size

Body size is one of the most fundamental characteristics of all organisms. It influences physiology, morphology, behavior, and even interspecific interactions such as those between parasites and their hosts. Host body size influences the magnitude and variability of parasite size according to Harrison's rule (HR: positive relationship between host and parasite body sizes) and Poulin's Increasing Variance Hypothesis (PIVH: positive relationship between host body size and the variability of parasite body size). We analyzed parasite–host body size allometry for 581 species of avian lice (～15% of known diversity) and their hosts. We applied phylogenetic generalized least squares (PGLS) methods to account for phylogenetic nonindependence controlling for host and parasite phylogenies separately and variance heterogeneity. We tested HR and PIVH for the major families of avian lice (Ricinidae, Menoponidae, Philopteridae), and for distinct ecological guilds within Philopteridae. Our data indicate that most families and guilds of avian lice follow both HR and PIVH; however, ricinids did not follow PIVH and the “body lice” guild of philopterid lice did not follow HR or PIVH. We discuss mathematical and ecological factors that may be responsible for these patterns, and we discuss the potential pervasiveness of these relationships among all parasites on Earth.     

Age and annual growth rate cause spatial variation in body size in Phrynocephalus przewalskii (Agamid)

Whether or not biogeographic rules dealing with spatial patterns of animal body sizes are valid for ectotherms is controversial. As the ectotherms grow all their lives, we explored the role of age and annual growth rate in body size variation in Phrynocephalus przewalskii in northern China. Morphological data were collected from 11 populations across a broad geographic gradient. Correlations between age, sex, climatic factors, and body size were analyzed using generalized linear model (GLM) and generalized linear mixed model (GLMM). GLM analysis indicated that the general body size of both sexes and the appendage size of females increased significantly with increasing temperature; however, the coefficient of determination was very small. GLMM analysis indicated that body size only correlated with age, whereas appendage size was affected by age, temperature, rainfall, and sunshine. Annual growth rates were positively correlated with temperature. We concluded that body size variation was mainly caused by age structure and plasticity of the growth rate in P. przewalskii and did not follow Bergmann''s rule; however, females followed Allen''s rule. Future studies to investigate the effect of energy restriction are needed to further understand the relationship between growth rate and body size. We also suggest that further studies on thermal advantage and sexual selection may be helpful to understand appendage size variation in P. przewalskii.     

Spatial patterns in the size of Chinese lizards are driven by multiple factors

BackgroundFor almost two centuries, ecologists have examined geographical patterns in the evolution of body size and the associated determinants. During that time, one of the most common patterns to have emerged is the increase in body size with increasing latitude (referred to as Bergmann''s rule). Typically, this pattern is explained in terms of an evolutionary response that serves to minimize heat loss in colder climates, mostly in endotherms. In contrast, however, this rule rarely explains geographical patterns in the evolution of body size among ectotherms (e.g., reptiles).LocationChina.AimIn this study, we assembled a dataset comprising the maximum sizes of 211 lizard species in China and examined the geographical patterns in body size evolution and its determinants. Specifically, we assessed the relationship between body size and climate among all lizard species and within four major groups at both assemblage and interspecific levels.ResultsAlthough we found that the body size of Chinese lizards was larger in warmer regions, we established that at the assemblage level, size was correlated with multiple climatic factors, and that body size–climate correlations differed within the four major groups. Phylogenetic analysis at the species level revealed that no single climatic factor was associated with body size, with the exception of agamids, for which size was found to be positively correlated with temperature.Main conclusionsGeographical patterns in Chinese lizard body size are driven by multiple factors, and overall patterns are probably influenced by those of the major groups. We suggest that our analyses at two different levels may have contributed to the inconsistent results obtained in this study. Further studies investigating the effects of altitude and ecological factors are needed to gain a more comprehensive understanding of the evolution of ectotherm body size.     

Interrelations of global macroecological patterns in wing and thorax size,sexual size dimorphism,and range size of the Drosophilidae

Support for macroecological rules in insects is mixed, with potential confounding interrelations between patterns rarely studied. We here investigate global patterns in body and wing size, sexual size dimorphism and range size in common fruit flies (Diptera: Drosophilidae) and explore potential interrelations and the predictive power of Allen's, Bergmann's, Rensch's and Rapoport's rules. We found that thorax length (r² = 0.05) and wing size (r² = 0.09) increased with latitude, supporting Bergmann's rule. Contrary to patterns often found in endothermic vertebrates, relative wing size increased towards the poles (r² = 0.12), a pattern against Allen's rule, which we attribute to selection for increased flight capacity in the cold. Sexual size dimorphism decreased with size, evincing Rensch's rule across the family (r² = 0.14). Yet, this pattern was largely driven by the virilis–repleta radiation. Finally, range size did not correlate with latitude, although a positive relationship was present in a subset of the species investigated, providing no convincing evidence for Rapoport's rule. We further found little support for confounding interrelations between body size, wing loading and range size in this taxon. Nevertheless, we demonstrate that studying several traits simultaneously at minimum permits better interpretation in case of multiple, potentially conflicting trends or hypotheses concerning the macroecology of insects.     

Geographic variation in body size and its relationship with environmental gradients in the Oriental Garden Lizard,Calotes versicolor

Patterns of geographic variation in body size are predicted to evolve as adaptations to local environmental gradients. However, many of these clinal patterns in body size, such as Bergmann's rule, are controversial and require further investigation into ectotherms such as reptiles on a regional scale. To examine the environmental variables (temperature, precipitation, topography and primary productivity) that shaped patterns of geographic variation in body size in the reptile Calotes versicolor, we sampled 180 adult specimens (91 males and 89 females) at 40 locations across the species range in China. The MANOVA results suggest significant sexual size dimorphism in C. versicolor (F_23,124 = 11.32, p < .001). Our results showed that C. versicolor failed to fit the Bergmann's rule. We found that the most important predictors of variation in body size of C. versicolor differed for males and females, but mechanisms related to heat balance and water availability hypotheses were involved in both sexes. Temperature seasonality, precipitation of the driest month, precipitation seasonality, and precipitation of the driest quarter were the most important predictors of variation in body size in males, whereas mean precipitation of the warmest quarter, mean temperature of the wettest quarter, precipitation seasonality, and precipitation of the wettest month were most important for body size variation in females. The discrepancy between patterns of association between the sexes suggested that different selection pressures may be acting in males and females.     

Body size evolution in South American Liolaemus lizards of the boulengeri clade: a contrasting reassessment

Bergmann's rule predicts larger body sizes in species living in higher latitudes and altitudes. This rule appears to be valid for endotherms, but its relevance to ectotherm vertebrates has largely been debated. In squamate reptiles (lizards and snakes), only one study, based on Liolaemus species of the boulengeri clade, has provided phylogenetic evidence in favour of Bergmann's clines. We reassessed this model in the same lizard clade, using a more representative measure of species body size and including a larger number of taxa in the sample. We found no evidence to support Bergmann's rule in this lineage. However, these non-significant results appear to be explained only by the inclusion of further species rather than by a different estimation of body size. Analyses conducted on the 16 species included in the previous study always revealed significant relationships between body size and latitude-altitude, whereas, the enlarged sample always rejected the pattern predicted by Bergmann's rule.