Latitudinal and altitudinal body size variation among north‐west European land snail species

Bergmann's rule states that, among conspecific populations, individuals are larger in cooler than in warmer environments as a consequence of selection related to heat conservation. Many of the most comprehensive assessments of Bergmann's rule to date have examined clinal patterns in body size among species assemblages. Our study is a more direct test of Bergmann's rule because we examine the pattern within a single, widely distributed species. We examined geographic variation in body and cell size in the spotted turtle ( Clemmys guttata). Our analysis of 818 turtles collected from the entire range (45-28 degrees N), indicated that body size increased with latitude; however, the relationship was driven by a population of large turtles at the northern extreme of the species' range. When the northern population was removed from the analyses, Bergmann's rule was not supported, and the smallest turtles occurred near the central part of the species' distribution. Recent literature has suggested that latitudinal clines in body size may simply be a physiological byproduct of the effects of temperature on cell division, resulting in larger cells, and hence larger organisms, from cooler temperatures. Measurements of the diameter of skin cells did not support the hypothesis that cell size increases with latitude and decreases with temperature in the spotted turtle, nor was there a significant relationship between body size and cell size. Our study suggests that neither Bergmann's rule nor cell size variation sufficiently explain the body size cline observed in the spotted turtle. We hypothesize that patterns in body size are related to variation in female size at maturity and reproductive cycles.

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.

Using published distributions of 65 species from the British Isles and northern Europe, we show that ant assemblages change with latitude in two ways. First, as commonly found for many types of organisms, the number of ant species decreased significantly with increasing latitude. For Ireland and Great Britain, species richness also increased significantly with region area. Second, although rarely demonstrated for ectotherms, the body size of ant species, as measured by worker length, increased significantly with increasing latitude. We found that this body-size pattern existed in the subfamily Formicinae and, to a lesser extent, in the Myrmicinae, which together comprised 95% of the ant species in our study area. There was a trend for formicines to increase in size with latitude faster than myrmicines. We also show that the pattern of increasing body size was due primarily to the ranges of ant species shifting to higher latitudes as their body sizes increased, with larger formicines becoming less represented at southerly latitudes and larger myrmicines becoming more represented at northerly latitudes. We conclude by discussing five potential mechanisms for generating the observed body-size patterns: the heat-conservation hypothesis, two hypotheses concerning phylogenetic history, the migration-ability hypothesis, and the starvation-resistance hypothesis.

Spatial patterns of body size are associated with environmental factors like temperature, seasonality, precipitation, resources, and altitude, among the most relevant. Latitudinal size patterns have been extensively studied. Studies on altitudinal gradients of body size, however, are relatively few. Besides, it is still unclear whether there is variation in the relationship between environmental factors and body size of species, or correlations within different altitudes. China provides a good opportunity to assess the altitudinal gradients of body size, because the entire terrain of China changes from high altitude in the west to low altitude in the east, with three natural terrain stages (low: less than 660 m.a.s.l.; medium: 660–2600 m.a.s.l.; and high: more than 2600 m.a.s.l.). Similarly, lizards are distributed across a wide altitudinal range (from −154.31 m to ~6000 m above sea level). We examined 211 lizard species in China and assessed the altitudinal gradients of Chinese lizard body sizes at assemblage and interspecific levels. Piecewise structural equation models (pSEM) were used to investigate the direct and indirect relationships between body size and environmental factors within different terrains (low, medium, and high altitudes). Overall, we found that body size in Chinese lizards decreases with increasing altitude. However, there were variations in the altitudinal gradients and drivers within the three terrain stages. Altitude has both direct (medium and low stages) and indirect (high stages) effects in shaping lizard size. We established that body size increased with increasing altitudes in the low and medium terrain stages. Seasonality had different effects on body size in medium (positively) and low (negatively) stages. However, among the three terrain stages, temperature was consistently negatively correlated with lizard size. Overall, we identified complex patterns and drivers of altitudinal gradients of body size within the three terrain stages, probably due to the different environmental conditions across altitudes.

The evolution of body size within and among species is predicted to be influenced by multifarious environmental factors. However, the specific drivers of body size variation have remained difficult to understand because of the wide range of proximate factors that covary with ectotherm body sizes across populations with varying local environmental conditions. Here, we used female Eremias argus lizards collected from different populations across their wide range in China, and constructed linear mixed models to assess how climatic conditions and/or available resources at different altitudes shape the geographical patterns of lizard body size across altitude. Lizard populations showed significant differences in body size across altitudes. Furthermore, we found that climatic and seasonal changes along the altitudinal gradient also explained variations in body size among populations. Specifically, body size decreased with colder and drier environmental conditions at high altitudes, reversing Bergmann's rule. Limited resources at high altitudes, measured by the low vegetative index, may also constrain body size. Therefore, our study demonstrates that multifarious environmental factors could strongly influence the intraspecific variation in organisms' body size.

Widespread butterflies follow inconsistent trends in the Bergmann's rule and flight morphometry: Implications for conservation in the Western Himalaya

AimIn general, within temperate latitudes, races of endotherms tend to show larger body size at higher latitudes, while smaller bodied ones tend to be found at lower latitudes, a pattern known as Bergmann's rule (or James's rule for intra‐specific patterns). The adaptive basis of this geographical variation in body size is explained mostly by the heat conservation hypothesis. Application of Bergmann's rule has been confirmed mostly among terrestrial taxa and studies of marine taxa are rare. Our aim was to examine whether the intra‐specific variation in body size of a pelagic seabird species (streaked shearwaterCalonectris leucomelas) follows Bergmann's/James's rule.LocationEight different breeding colonies of the streaked shearwater,ranging from 24–39° N in latitude (approximately the full latitudinal range of the species) and 123–142° E in longitude.MethodsWe measured morphological traits of 454 adult streaked shearwaters in total. Principal components analysis was used to create a composite body size index for each sex from each colony using the measured traits, and the value of first principal component was compared. The relationships between body size index and latitude, longitude and the averaged mean air temperature were examined using generalized linear models.ResultsBody size was positively correlated with latitude and longitude, and negatively correlated with the mean air temperature. Latitude and longitude were highly correlated with air temperature. Nonetheless, between colonies, populations that are considered to forage over large areas appeared to exhibit smaller body size than those of lower latitude populations.Main conclusionThe overall trend of the geographical variations in body size of streaked shearwaters followed Bergmann's/James's rule, suggesting that this rule may be applicable to seabirds. However, in addition to thermoregulatory adaptation, intra‐specific differences in foraging characteristic in relation to local marine environment might also affect the body size in highly mobile seabirds.

Geographic patterns in body size are often associated with latitude, elevation, or environmental and climatic variables. This study investigated patterns of body size and cell size of the green anole lizard, Anolis carolinensis, and potential associations with geography or climatic variables. Lizards were sampled from 19 populations across the native range, and body size, red blood cell size and size and number of muscle cells were measured. Climatic data from local weather stations and latitude and longitude were entered into model selection with Akaike's information criterion to explain patterns in cell and body sizes. Climatic variables did not drive any major patterns in cell size or body size; rather, latitude and longitude were the best predictors of cell and body size. In general, smaller body and cell sizes in Florida anoles drove geographic patterns in A. carolinensis. Small size in Florida may be attributable to the geological history of the peninsular state or the unique ecological factors in this area, including a recently introduced congener. In contrast to previous studies, we found that A. carolinensis does not follow Bergmann's rule when the influence of Florida is excluded. Rather, the opposite pattern of larger lizards in southern populations is evident in the absence of Florida populations, and mirrors the general pattern in squamates. Muscle cell size was negatively related to latitude and red blood cell size showed no latitudinal trend outside of Florida. Different patterns in the sizes of the 2 cell types confirm the importance of examining multiple cell types when studying geographic variation in cell size.

Theoretical mechanisms describing species abundance distributions should also underpin geographic variation in life-history traits. However, recent studies suggest that abun- dance and trait patterns may not co-vary and may respond differently to abiotic conditions acting at different spatial scales. We examined patterns in abundance and body size of 2 estuarine mol- luscs, the arkshell Anadara trapezia and the mudsnail Batillaria australis, across their wide distri- butions in eastern Australia. We related abundance and body size patterns to abiotic variables including water temperature, pH, salinity, sediment redox and dissolved oxygen content at multi- ple spatial scales. Two hypotheses were tested: (1) geographic patterns in abundance and body size do not co-vary, and (2) patterns in abundance are more strongly influenced by abiotic condi- tions occurring at a large spatial scale (e.g. across latitudinal gradients) whereas body size is more strongly influenced by variation in abiotic conditions occurring at smaller scales. The influence of spatial scale and associated abiotic variables on abundance and body size distributions was deter- mined using multiple linear regression, ANOVA and variance component analyses. Geographic variation in abundance and body size were independent of each other in both species. Abiotic variation across latitudinal gradients was the strongest predictor of abundance, but factors that varied substantially at local scales (e.g. dissolved oxygen and sediment redox) were the strongest predictors of body size. Our data indicate that geographic patterns in body size and abundance can be disconnected from each other, most likely due to differential responses to abiotic variation acting at different spatial scales.

Hypotheses that explain geographic variation in body size were examined using cranial measurements of 950 bobcats (Lynx rufus) from western North America. Bobcats were divided into 25 geographic localities of similar habitats and landform (based on ecoregions). Principal component analyses were used to derive a single estimate of size from scores on the first principal component. Males and females were examined separately, because they were significantly dimorphic in body size and because sex and locality exhibited a significant interaction. We expected that female body size would best reflect environmental influences, because male size may be influenced by sexual selection. We found significant geographic variation in bobcat body size, with about 44% of the variation in males and 47% of the variation in females accounted for by comparison among the localities. We also found that variation in body size was associated with Bergmann's rule, as indicated by significant multiple regression of body size of males (R2 = 0.426) and females (R2 = 0.480) on latitude and elevation. Using correlation and regression analyses, we examined the association of body size with selected environmental variables that represent the classical physiological explanation of Bergmann's rule, James' moisture-humidity modification of Bergmann's rule, Rosenzweig's productivity hypothesis, and Boyce's seasonality hypothesis. Only the productivity hypothesis was not supported. The relative strengths of associations suggested, however, that James' modification was better supported than the classical explanation for Bergmann's rule. Path analyses permitted further discrimination of hypotheses, and only the seasonality hypothesis received significant support. As expected, this support was only evident for females. Path analysis may provide a tool for evaluating relative strengths of competing but correlated explanations of geographic variation.

Intraspecific variability and species turnover drive variations in Collembola body size along a temperate-boreal elevation gradient

ABSTRACTAim To describe the geographical pattern of mean body size of the non‐volant mammals of the Nearctic and Neotropics and evaluate the influence of five environmental variables that are likely to affect body size gradients.Location The Western Hemisphere.Methods We calculated mean body size (average log mass) values in 110 × 110 km cells covering the continental Nearctic and Neotropics. We also generated cell averages for mean annual temperature, range in elevation, their interaction, actual evapotranspiration, and the global vegetation index and its coefficient of variation. Associations between mean body size and environmental variables were tested with simple correlations and ordinary least squares multiple regression, complemented with spatial autocorrelation analyses and split‐line regression. We evaluated the relative support for each multiple‐regression model using AIC.Results Mean body size increases to the north in the Nearctic and is negatively correlated with temperature. In contrast, across the Neotropics mammals are largest in the tropical and subtropical lowlands and smaller in the Andes, generating a positive correlation with temperature. Finally, body size and temperature are nonlinearly related in both regions, and split‐line linear regression found temperature thresholds marking clear shifts in these relationships (Nearctic 10.9 °C; Neotropics 12.6 °C). The increase in body sizes with decreasing temperature is strongest in the northern Nearctic, whereas a decrease in body size in mountains dominates the body size gradients in the warmer parts of both regions.Main conclusions We confirm previous work finding strong broad‐scale Bergmann trends in cold macroclimates but not in warmer areas. For the latter regions (i.e. the southern Nearctic and the Neotropics), our analyses also suggest that both local and broad‐scale patterns of mammal body size variation are influenced in part by the strong mesoscale climatic gradients existing in mountainous areas. A likely explanation is that reduced habitat sizes in mountains limit the presence of larger‐sized mammals.

ABSTRACTAimTo evaluate (1) whether three migratory nightjar species (Family Caprimulgidae) adhere to Bergmann's rule, (2) whether environmental factors on the breeding or wintering grounds determine body size, and (3) which mechanistic hypotheses best explain Bergmannian patterns in body size.LocationNorth and South America; Europe and Africa.TaxonEastern whip‐poor‐will (Antrostomus vociferus), Common nighthawk (Chordeiles minor) and European nightjar (Caprimulgus europaeus).MethodsWe used GPS tracking and morphometric data to assess competing hypotheses explaining variation in body size for each species, based on their breeding (n = 3388) and wintering (n = 189) locations.ResultsAll three species exhibited Bergmannian patterns in body size, providing the first evidence that nightjars conform to Bergmann's rule despite adaptations to severe environmental conditions. Environmental and geographic variables at breeding sites were stronger predictors of body size than wintering‐site variables. Although we found partial support for Bergmann's temperature regulation hypothesis, geographic variables, rather than specific environmental factors, emerged as the strongest predictors of body size variation.Main ConclusionsLatitude and longitude correlated strongly with environmental variables and migratory distance; thus, these geographical variables likely encompass many factors that influence body size in nightjars. The present study is among the first to use tracking data from individual birds to understand how environmental pressures across the annual cycle are related to body size. Our findings highlight the critical role of geographic breeding‐ground factors in shaping Bergmannian patterns, offering robust evidence to support nearly two centuries of research since Bergmann's rule was first described in 1847.

AimWhether intraspecific spatial patterns in body size are generalizable across species remains contentious, as well as the mechanisms underlying these patterns. Here we test several hypotheses explaining within‐species body size variation in terrestrial vertebrates including the heat balance, seasonality, resource availability and water conservation hypotheses for ectotherms, and the heat conservation, heat dissipation, starvation resistance and resource availability hypotheses for endotherms.LocationGlobal.Time period1970–2016.Major taxa studiedAmphibians, reptiles, birds and mammals.MethodsWe collected 235,905 body size records for 2,229 species (amphibians = 36; reptiles = 81; birds = 1,545; mammals = 567) and performed a phylogenetic meta‐analysis of intraspecific correlations between body size and environmental variables. We further tested whether correlations differ between migratory and non‐migratory bird and mammal species, and between thermoregulating and thermoconforming ectotherms.ResultsFor bird species, smaller intraspecific body size was associated with higher mean and maximum temperatures and lower resource seasonality. Size–environment relationships followed a similar pattern in resident and migratory birds, but the effect of resource availability on body size was slightly positive only for non‐migratory birds. For mammals, we found that intraspecific body size was smaller with lower resource availability and seasonality, with this pattern being more evident in sedentary than migratory species. No clear size–environment relationships were found for reptiles and amphibians.Main conclusionsWithin‐species body size variation across endotherms is explained by disparate underlying mechanisms for birds and mammals. Heat conservation (Bergmann's rule) and heat dissipation are the dominant processes explaining biogeographic intraspecific body size variation in birds, whereas in mammals, body size clines are mostly explained by the starvation resistance and resource availability hypotheses. Our findings contribute to a better understanding of the mechanisms behind species adaptations to the environment across their geographic distributions.

Patterns of abundance across space and time, and intraspecific variation in body size, are two species attributes known to influence diet breadth and the structure of interaction networks. Yet, the relative influence of these attributes on diet breadth is often assumed to be equal among taxonomic groups, and the relationship between intraspecific variation in body size on interaction patterns is frequently neglected. We observed bee–flower interactions in multiple locations across Montana, USA, for two growing seasons and measured spatial and temporal patterns of abundance, along with interspecific and intraspecific variation in body size for prevalent species. We predicted that the association between spatial and temporal patterns of abundance and intraspecific variation in body size, and diet breadth, would be stronger for bumble bee compared to non-bumble bee species, because species with flexible diets and long activity periods can interact with more food items. Bumble bees had higher local abundance, occurred in many local communities, more intraspecific variation in body size, and longer phenophases compared to non-bumble bee species, but only local abundance and phenophase duration had a stronger positive association with the diet breadth of bumble bee compared to non-bumble bee species. Communities with a higher proportion of bumble bees also had higher intraspecific variation in body size at the network-level, and network-level intraspecific variation in body size was positively correlated with diet generalization. Our findings highlight that the association between species attributes and diet breadth changes depending on the taxonomic group, with implications for the structure of interaction networks.