On the validity of Bergmann's rule

Aim We reviewed the occurrence of Bergmann's rule in birds (ninety‐four species) and mammals (149 species), using only studies where statistical significance of the results was tested. We also tested whether studies using different characters as surrogates of body size have a different tendency to conform to Bergmann's rule, whether body size and nest type (in birds) have an influence on the tendency to conform to the rule, and whether sedentary birds conform to the rule more than migratory birds. Location Worldwide. Methods We reviewed published data on geographic and temporal variation in body size, using only studies where the statistical significance of the results was tested. We asked how many species conform to the rule out of all species studied in each order and family. Results Over 72% of the birds and 65% of the mammal species follow Bergmann's rule. An overall tendency to follow the rule occurs also within orders and families. Studies using body mass in mammals show the greatest tendency to adhere to Bergmann's rule (linear measurements and dental measurements show a weaker tendency); while in birds, studies using body mass and other surrogates (linear measurements and egg size) show a similar tendency. Birds of different body mass categories exhibit a similar tendency to follow Bergmann's rule, while in mammals the lower body size categories (4–50 and 50–500 g) show a significantly lower tendency to conform to the rule. Sedentary birds tend to conform to Bergmann's rule more than migratory species. Nest type does not affect the tendency to conform to Bergmann's rule. Main conclusions Bergmann's rule is a valid ecological generalization for birds and mammals.     

Morphological variations in a widely distributed Eastern Asian passerine cannot be consistently explained by ecogeographic rules

Ecogeographic rules that describe quantitative relationships between morphologies and climate might help us predict how morphometrics of animals was shaped by local temperature or humidity. Although the ecogeographic rules had been widely tested in animals of Europe and North America, they had not been fully validated for species in regions that are less studied. Here, we investigate the morphometric variation of a widely distributed East Asian passerine, the vinous‐throated parrotbill (Sinosuthora webbiana), to test whether its morphological variation conforms to the prediction of Bergmann''s rule, Allen''s rules, and Gloger''s rule. We at first described the climatic niche of S. webbiana from occurrence records (n = 7838) and specimen records (n = 290). The results of analysis of covariance (ANCOVA) suggested that the plumage coloration of these parrotbills was darker in wetter/warmer environments following Gloger''s rule. However, their appendage size (culmen length, beak volume, tarsi length) was larger in colder environments, the opposite of the predictions of Allen''s rule. Similarly, their body size (wing length) was larger in warmer environments, the opposite of the predictions of Bergmann''s rule. Such disconformity to both Bergmann''s rule and Allen''s rule suggests that the evolution of morphological variations is likely governed by multiple selection forces rather than dominated by thermoregulation. Our results suggest that these ecogeographic rules should be validated prior to forecasting biological responses to climate change especially for species in less‐studied regions.     

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.     

A global assessment of Bergmann's rule in mammals and birds

Bergmann's rule states that endotherms have a large body size in high latitudes and cold climates. However, previous empirical studies have reported mixed evidence on the relationships between body size and latitude, raising the question of why some clades of endotherms follow Bergmann's rule, whereas others do not. Here, we synthesized the interspecific relationships between body size and latitude among 16,187 endothermic species (5422 mammals and 10,765 birds) using Bayesian phylogenetic generalized linear mixed models to examine the strength and magnitude of Bergmann's rule. We further assessed the effect of biological and ecological factors (i.e., body mass categories, dietary guild, winter activity, habitat openness, and climate zone) on the variations in the body mass–latitude relationships by adding an interaction term in the models. Our results revealed a generally weak but significant adherence to Bergmann's rule among all endotherms at the global scale. Despite taxonomic variation in the strength of Bergmann's rule, the body mass of species within most animal orders showed an increasing trend toward high latitudes. Generally, large-bodied, temperate species, non-hibernating mammals, and migratory and open-habitat birds tend to conform to Bergmann's rule more than their relatives do. Our results suggest that whether Bergmann's rule applies to a particular taxon is mediated by not only geographic and biological features, but also potential alternate strategies that species might have for thermoregulation. Future studies could explore the potential of integrating comprehensive trait data into phylogenetic comparative analysis to re-assess the classic ecogeographic rules on a global scale.     

Bergmann's rule: a biophysiological rule examined in birds

The most studied ecogeographic rule is Bergmann's rule, but aspects of the original paper are often presented incorrectly even though Bergmann (1847) is explicitly cited. The goal of this paper is to 1) summarize the contents of Bergmann's paper, supported by direct translations, and 2) to discuss the main issues surrounding Bergmann's rule based on Bergmann's intentions and early definitions of the rule. Although Bergmann himself never formulated an explicit rule, based on Bergmann's (1847) intentions and early definitions of Bergmann's rule, Bergmann's rule is: “Within species and amongst closely related species of homeothermic animals a larger size is often achieved in colder climates than in warmer ones, which is linked to the temperature budget of these animals.” Bergmann (1847) assumed that the surface area of an animal is a measure for heat dissipation and an animal's volume a measure of its heat production. As body size increases, an animal's surface area increases less than its volume; however, modifications in morphology and behaviour will also influence the temperature budget. Bergmann hypothesized that when everything but size is equal, the smaller animals should live in warmer areas. This was supported by empirical data on > 300 bird species belonging to 86 genera. Recommendations for use of the term Bergmann's rule include 1) inclusion of a thermoregulatory mechanism, 2) application only to homoeothermic animals, 3) but to any taxonomic group, 4) tests of the rule should test the assumption that larger animals have to produce less heat to increase body temperatures, and 5) future authors should either go back to the original publication (Bergmann 1847) when referring to it or simply not cite it at all. Synthesis Based on Bergmann's (1847) intentions and early definitions, Bergmann's rule is: “Within species and amongst closely related species of homeothermic animals a larger size is often achieved in colder climates than in warmer ones, which is linked to the temperature budget of these animals.” Recommendations for use of the term Bergmann's rule include 1) inclusion of a thermoregulatory mechanism, 2) application only to homoeothermic animals, 3) and to any taxonomic group, 4) tests of the rule should examine whether larger animals have to produce less heat to increase body temperatures, and 5) authors should go back to the original publication (Bergmann 1847) when referring to it.     

Body size but not colony size increases with altitude in the holarctic ant,Leptothorax acervorum

1. Bergmann's rule states that organisms inhabiting colder environments show an increase in body size or mass in comparison to their conspecifics living in warmer climates. Although originally proposed for homoeothermic vertebrates, this rule was later extended to ectotherms. In social insects, only a few studies have tested this rule and the results were ambiguous. Here, ‘body size’ can be considered at two different levels (the size of the individual workers or the size of the colony). 2. In this study, data from 53 nests collected along altitudinal gradients in the Alps were used to test the hypotheses that the worker body size and colony size of the ant Leptothorax acervorum increase with increasing altitude and therefore follow Bergmann's rule. 3. The results show that the body size of workers but not the colony size increases with altitude. Whether this pattern is driven by starvation resistance or other mechanisms remains to be investigated.     

Patterns of within-species body size variation of birds: strong evidence for Bergmann's rule

Aim The aim of this study is to test whether Bergmann's rule, a general intraspecific tendency towards larger body size in cooler areas and at higher latitudes, holds for birds throughout the world. Location This study includes information on species of birds from throughout the world. Methods I gathered data on body size variation from the literature and used two general meta‐analytical procedures to test the validity of Bergmann's rule in birds: a modified vote‐counting approach and calculation of overall effect sizes. Related species may show similar body size trends, thus I performed all analyses using nonphylogenetic and phylogenetic methods. I used tests of phylogenetic signal for each data set to decide which type of statistical analysis (nonphylogenetic or phylogenetic) was more appropriate. Results The majority of species of birds (76 of 100 species) are larger at higher latitudes, and in cooler areas (20 of 22 species). Birds show a grand mean correlation coefficient of +0.32 for body size and latitude, and ?0.81 for body size and temperature, both significant trends. Sedentary species show stronger body size trends in some, but not all, analyses. Neither males nor females consistently have stronger body size trends. Additionally, the strength of body size trends does not vary with latitude or body mass. Conclusions Bergmann's rule holds for birds throughout the world, regardless of whether temperature or latitude (as a proxy) is used. Previous studies have suggested that Bergmann's rule is stronger for sedentary than migratory species, males than females and temperate than tropical taxa. I did not find strong support for any of these as general themes for birds, although few studies of tropical taxa have been conducted. The processes responsible for Bergmann's rule remain somewhat of a black box; however, fasting endurance is probably a more important factor than the traditional hypothesis of heat conservation.     

Rule reversal: Ecogeographical patterns of body size variation in the common treeshrew (Mammalia,Scandentia)

There are a number of ecogeographical “rules” that describe patterns of geographical variation among organisms. The island rule predicts that populations of larger mammals on islands evolve smaller mean body size than their mainland counterparts, whereas smaller‐bodied mammals evolve larger size. Bergmann's rule predicts that populations of a species in colder climates (generally at higher latitudes) have larger mean body sizes than conspecifics in warmer climates (at lower latitudes). These two rules are rarely tested together and neither has been rigorously tested in treeshrews, a clade of small‐bodied mammals in their own order (Scandentia) broadly distributed in mainland Southeast Asia and on islands throughout much of the Sunda Shelf. The common treeshrew, Tupaia glis, is an excellent candidate for study and was used to test these two rules simultaneously for the first time in treeshrews. This species is distributed on the Malay Peninsula and several offshore islands east, west, and south of the mainland. Using craniodental dimensions as a proxy for body size, we investigated how island size, distance from the mainland, and maximum sea depth between the mainland and the islands relate to body size of 13 insular T. glis populations while also controlling for latitude and correlation among variables. We found a strong negative effect of latitude on body size in the common treeshrew, indicating the inverse of Bergmann's rule. We did not detect any overall difference in body size between the island and mainland populations. However, there was an effect of island area and maximum sea depth on body size among island populations. Although there is a strong latitudinal effect on body size, neither Bergmann's rule nor the island rule applies to the common treeshrew. The results of our analyses demonstrate the necessity of assessing multiple variables simultaneously in studies of ecogeographical rules.     

Geographical variation of human gut microbial composition

Although we know there is considerable variation in gut microbial composition within host species, little is known about how this variation is shaped and why such variation exists. In humans, obesity is associated with the relative abundance of two dominant bacterial phyla: an increase in the proportion of Firmicutes and a decrease in the proportion of Bacteroidetes. As there is evidence that humans have adapted to colder climates by increasing their body mass (e.g. Bergmann''s rule), we tested whether Firmicutes increase and Bacteroidetes decrease with latitude, using 1020 healthy individuals drawn from 23 populations and six published studies. We found a positive correlation between Firmicutes and latitude and a negative correlation between Bacteroidetes and latitude. The overall pattern appears robust to sex, age and bacterial detection methods. Comparisons between African Americans and native Africans and between European Americans and native Europeans suggest no evidence of host genotype explaining the observed patterns. The variation of gut microbial composition described here is consistent with the pattern expected by Bergmann''s rule. This surprising link between large-scale geography and human gut microbial composition merits further investigation.     

A Reassessment of Bergmann's Rule in Modern Humans

It is widely accepted that modern humans conform to Bergmann''s rule, which holds that body size in endothermic species will increase as temperature decreases. However, there are reasons to question the reliability of the findings on which this consensus is based. One of these is that the main studies that have reported that modern humans conform to Bergmann''s rule have employed samples that contain a disproportionately large number of warm-climate and northern hemisphere groups. With this in mind, we used latitudinally-stratified and hemisphere-specific samples to re-assess the relationship between modern human body size and temperature. We found that when groups from north and south of the equator were analyzed together, Bergmann''s rule was supported. However, when groups were separated by hemisphere, Bergmann''s rule was only supported in the northern hemisphere. In the course of exploring these results further, we found that the difference between our northern and southern hemisphere subsamples is due to the limited latitudinal and temperature range in the latter subsample. Thus, our study suggests that modern humans do conform to Bergmann''s rule but only when there are major differences in latitude and temperature among groups. Specifically, groups must span more than 50 degrees of latitude and/or more than 30°C for it to hold. This finding has important implications for work on regional variation in human body size and its relationship to temperature.     

Patterns of size variation in bees at a continental scale: does Bergmann's rule apply?

Body size latitudinal clines have been widley explained by the Bergmann's rule in homeothermic vertebrates. However, there is no general consensus in poikilotherms organisms in particular in insects that represent the large majority of wildlife. Among them, bees are a highly diverse pollinators group with high economic and ecological value. Nevertheless, no comprehensive studies of species assemblages at a phylogenetically larger scale have been carried out even if they could identify the traits and the ecological conditions that generate different patterns of latitudinal size variation. We aimed to test Bergmann's rule for wild bees by assessing relationships between body size and latitude at continental and community levels. We tested our hypotheses for bees showing different life history traits (i.e. sociality and nesting behaviour). We used 142 008 distribution records of 615 bee species at 50 × 50 km (CGRS) grids across the West Palearctic. We then applied generalized least squares fitted linear model (GLS) to assess the relationship between latitude and mean body size of bees, taking into account spatial autocorrelation. For all bee species grouped, mean body size increased with higher latitudes, and so followed Bergmann's rule. However, considering bee genera separately, four genera were consistent with Bergmann's rule, while three showed a converse trend, and three showed no significant cline. All life history traits used here (i.e. solitary, social and parasitic behaviour; ground and stem nesting behaviour) displayed a Bergmann's cline. In general there is a main trend for larger bees in colder habitats, which is likely to be related to their thermoregulatory abilities and partial endothermy, even if a ‘season length effect’ (i.e. shorter foraging season) is a potential driver of the converse Bergmann's cline particularly in bumblebees.     

Body size phenology in a regional bee fauna: a temporal extension of Bergmann's rule

Bergmann's rule originally described a positive relationship between body size and latitude in warm‐blooded animals. Larger animals, with a smaller surface/volume ratio, are better enabled to conserve heat in cooler climates (thermoregulatory hypothesis). Studies on endothermic vertebrates have provided support for Bergmann's rule, whereas studies on ectotherms have yielded conflicting results. If the thermoregulatory hypothesis is correct, negative relationships between body size and temperature should occur in temporal in addition to geographical gradients. To explore this possibility, we analysed seasonal activity patterns in a bee fauna comprising 245 species. In agreement with our hypothesis of a different relationship for large (endothermic) and small (ectothermic) species, we found that species larger than 27.81 mg (dry weight) followed Bergmann's rule, whereas species below this threshold did not. Our results represent a temporal extension of Bergmann's rule and indicate that body size and thermal physiology play an important role in structuring community phenology.     

Urban living influences the nesting success of Darwin’s finches in the Galápagos Islands

Urbanization is expanding worldwide with major consequences for organisms. Anthropogenic factors can reduce the fitness of animals but may have benefits, such as consistent human food availability. Understanding anthropogenic trade‐offs is critical in environments with variable levels of natural food availability, such as the Galápagos Islands, an area of rapid urbanization. For example, during dry years, the reproductive success of bird species, such as Darwin''s finches, is low because reduced precipitation impacts food availability. Urban areas provide supplemental human food to finches, which could improve their reproductive success during years with low natural food availability. However, urban finches might face trade‐offs, such as the incorporation of anthropogenic debris (e.g., string, plastic) into their nests, which may increase mortality. In our study, we determined the effect of urbanization on the nesting success of small ground finches (Geospiza fuliginosa; a species of Darwin''s finch) during a dry year on San Cristóbal Island. We quantified nest building, egg laying and hatching, and fledging in an urban and nonurban area and characterized the anthropogenic debris in nests. We also documented mortalities including nest trash‐related deaths and whether anthropogenic materials directly led to entanglement‐ or ingestion‐related nest mortalities. Overall, urban finches built more nests, laid more eggs, and produced more fledglings than nonurban finches. However, every nest in the urban area contained anthropogenic material, which resulted in 18% nestling mortality while nonurban nests had no anthropogenic debris. Our study showed that urban living has trade‐offs: urban birds have overall higher nesting success during a dry year than nonurban birds, but urban birds can suffer mortality from anthropogenic‐related nest‐materials. These results suggest that despite potential costs, finches benefit overall from urban living and urbanization may buffer the effects of limited resource availability in the Galápagos Islands.     

Population level differences in adult body mass emerge in infancy and early childhood: Evidence from a global sample of low and lower‐income countries

Many studies have linked measures of adult body shape and mass in ancient and contemporary populations to ecogeographical variables such as temperature and latitude. These results tend to support Bergmann's rule, which posits that bodies will be relatively less slender for their height in colder climates and more slender in warmer climates. Less well explored is the ontogeny of these population‐level differences. Here we use data on infants and children from 46 low and lower income countries to test whether children's weight for height is associated with measures of temperature and latitude. We also test the hypothesis that children living in areas with greater pathogen prevalence will be lighter for their height because of life history trade‐offs between investment in immune function and growth. Finally, we test whether population specific adult body mass predicts infant and child body mass, and whether this is independent of ecogeographical variables. Our results show that maximum monthly temperature explains 17% of children's weight for height while adult population‐level body mass explains ～44% (Table 5 ). The measures of pathogen prevalence explain little of the variation in children's body shape (8%; P > 0.05). Our results suggest that population differences are consistent with Bergmann's rule but parental body shape explains more variance. Moreover, these population‐level differences arise early in development, suggesting that any possible environmental influences occur in utero and/or result from epigenetic or population genetic differences. Am J Phys Anthropol 154:232–238, 2014. © 2014 Wiley Periodicals, Inc.     

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.     

Higher temperatures during development reduce body size in the zebra finch in the laboratory and in the wild

The most commonly documented morphological response across many taxa to climatic variation across their range follows Bergmann's rule, which predicts larger body size in colder climates. In observational data from wild zebra finches breeding across a range of temperatures in the spring and summer, we show that this relationship appears to be driven by the negative effect of high temperatures during development. This idea was then experimentally tested on zebra finches breeding in temperature‐controlled climates in the laboratory. These experiments confirmed that those individualso produced in a hot environment (30 °C) were smaller than those produced in cool conditions (18 °C). Our results suggest a proximate causal link between temperature and body size and suggest that a hotter climate during breeding periods could drive significant changes in morphology within and between populations. This effect could account for much of the variation in body size that drives the well‐observed patterns first described by Bergmann and that is still largely attributed to selection on adult body size during cold winters. The climate‐dependent developmental plasticity that we have demonstrated is an important component in understanding how endotherms may be affected by climate change.     

Bergmann's rule does not apply to geometrid moths along an elevational gradient in an Andean montane rain forest

Aim Bergmann's rule generally predicts larger animal body sizes with colder climates. We tested whether Bergmann's rule at the interspecific level applies to moths (Lepidoptera: Geometridae) along an extended elevational gradient in the Ecuadorian Andes. Location Moths were sampled at 22 sites in the province Zamora‐Chinchipe in southern Ecuador in forest habitats ranging from 1040 m to 2677 m above sea level. Methods Wingspans of 2282 male geometrid moths representing 953 species were measured and analysed at the level of the family Geometridae, as well as for the subfamily Ennominae with the tribes Boarmiini and Ourapterygini, and the subfamily Larentiinae with the genera Eois, Eupithecia and Psaliodes. Results Bergmann's rule was not supported since the average wingspan of geometrid moths was negatively correlated with altitude (r = ?0.59, P < 0.005). The relationship between body size and altitude in Geometridae appears to be spurious because species of the subfamily Larentiinae are significantly smaller than species of the subfamily Ennominae and simultaneously increase in their proportion along the gradient. A significant decrease of wingspan was also found in the ennomine tribe Ourapterygini, but no consistent body size patterns were found in the other six taxa studied. In most taxa, body size variation increases with altitude, suggesting that factors acting to constrain body size might be weaker at high elevations. Main conclusions The results are in accordance with previous studies that could not detect consistent body size patterns in insects at the interspecific level along climatic gradients.     

Body size explains interspecific variation in size–latitude relationships in geographically widespread beetle species

1. In most birds and mammals, larger individuals of the same species tend to be found at higher latitudes, but in insects, body size–latitude relationships are highly variable. 2. Recent studies have shown that larger‐bodied insect species are more likely to decrease in size when reared at increased temperature, compared with smaller‐sized species. These findings have led to the prediction that a positive relationship between body size and latitude should be more prevalent in larger‐bodied insect species. 3. This study measured the body size of > 4000 beetle specimens (12 species) collected throughout North America. Some beetle species increased in size with latitude, while others decreased. Importantly, mean species body size explained c. 30% of the interspecific variation in the size–latitude response. 4. As predicted, larger‐bodied beetle species were more likely to show a positive relationship between body size and latitude (Bergmann's rule), and smaller‐bodied species were more likely to show a negative body size–latitude relationship (inverse Bergmann's rule). 5. These body size–latitude patterns suggest that size‐specific responses to temperature may underlie global latitudinal distributions of body size in Coleoptera, as well as other insects.     

Bergmann's rule in shrews: geographical variation of body size in Palearctic Sorex species

According to Bergmann's rule, individuals of a given species tend to be larger in colder (northern) climates. Traditional explanation points to the relatively lower surface‐to‐volume ratio in larger animals and, consequently, relatively lower costs of thermoregulation. We examined intraspecific covariation of body size with geographical location and climate in five species of Sorex shrews, animals that are among the smallest extant mammals. The condylobasal length of skull (CBL), compiled from literature data and measured on museum specimens, was used as an indicator of the overall body size of shrews. Surprisingly, in three out of five shrew species, the CBL was negatively correlated with latitude, and the same trend, although not statistically significant, was found in the fourth species. In general, shrews were smaller in colder areas, as evidenced by the positive correlations between the CBL and temperature. In two species, these positive correlations appeared when the effect of longitude was held constant in the partial correlation analysis. Characteristically, the strongest negative correlation with latitude and positive with temperatures was found in S. minutus, the smallest species under study. Shrews were in general larger in environments with high actual evapotranspiration. Body mass reviewed in S. araneus paralleled the pattern found in the CBL variation in this species, i.e. it decreased northward, both in summer‐ and winter‐caught animals. In addition, contrary to the widely accepted ? but not rigorously tested ? belief, body mass recession from summer to winter (the Dehnel Effect) did not correlate with latitude. We concluded that shrews followed the converse to Bergmann's rule, and hypothesize that part of their body size variation along the west‐east axis may be explained by character displacement. We also hypothesize that scarcity of food, especially in winter, is a major factor selecting for small body size in shrews in northern areas, as smaller individuals should require less food. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78 , 365–381.     

Functional adaptation rather than ecogeographical rules determine body-size metrics along a thermal cline with elevation in the Chinese pygmy dormouse (Typhlomys cinereus)

Phenotypic plasticity is crucial for how organisms respond to variation in their environment, affecting their diversity and distribution, especially in the light of rapid environmental change. Ecogeographical rules predict an association between specific adaptive morphological and physiological traits with cooler conditions due to higher latitude, elevation, or climate change. Such ecogeographical effects are often most evident in ancient species due to continuous selective adaptation occurring over long periods of time. Here, we use the suitably ancient Chinese pygmy dormouse (Typhlomys cinereus) to test whether body-size, appendage length and heart size vary in accordance with Bergmann's, Allen's and Hesse's rule, respectively. Based on a sample of 67 adult individuals (female, n = 29; male n = 38) trapped at 37 sites transcending an elevational range from 414 to 1757 m, we tested for trait concordance with Bergmann's rule (body mass, length and SMI), Allen's rule (length of tail, foot, ear, snout), and Hesse's rule (wet and dry heart mass). Effects of elevation (and thus temperature lapse rate; calculated as 0.61 °C per 100 m) on body size, appendage length and heart size, were tested by fitting Standardized Major Axis (SMA) models. We observed substantial heterogeneity in morphometric traits allowing for the detection of ecogeographical clines. However, none conformed with Bergmann's, Allen's (except ear size), or Hesse's rule. However, our results indicate some support for Geist's rule of net primary productivity. We conclude that pervasive functional life-history adaptations in this blind, arboreal, echolocating ancient species exceeded selection for morphological energy efficiency constraints, with the notable exception of reduced ear pinnae size at colder, elevated sites. This is an important consideration for predicting how species, and populations in general, may adapt to human induced rapid environmental change, contrary to expectations of warming driving selection for smaller body-size.