Biogeography of body size in terrestrial isopods (Crustacea: Oniscidea)

This study tries to unveil the contribution of climatic shift in shaping the extreme body size diversity in terrestrial isopods (Oniscidea). Trying to explain size variation at an interspecific level, we test five hypotheses: (1) Bergmann's Rule and the temperature‐size rule postulate large size in cold areas; (2) The metabolic cold adaptation theory postulates small animal sizes in cold environments; (3) The primary productivity hypothesis predicts size increase in resource‐rich areas; (4) The aridity resistance hypothesis predicts large size in arid regions; and (5). The acidosis hypothesis predicts smaller size with decreasing soil pH. Globally, Bergmann's rule and the aridity hypothesis are weakly supported. Among families and genera, results are variable and idiosyncratic. Conglobating species sizes provide weak support for the acidosis hypothesis. Overall, size is strongly affected by familial affiliation. Isopod size evolution seems to be mainly affected by phylogenetically constrained life‐history traits.     

Recent spatial and temporal changes in body size of terrestrial vertebrates: probable causes and pitfalls

Geographical and temporal variations in body size are common phenomena among organisms and may evolve within a few years. We argue that body size acts much like a barometer, fluctuating in parallel with changes in the relevant key predictor(s), and that geographical and temporal changes in body size are actually manifestations of the same drivers. Frequently, the principal predictors of body size are food availability during the period of growth and ambient temperature, which often affects food availability. Food availability depends on net primary productivity that, in turn, is determined by climate and weather (mainly temperature and precipitation), and these depend mainly on solar radiation and other solar activities. When the above predictors are related to latitude the changes have often been interpreted as conforming to Bergmann's rule, but in many cases such interpretations should be viewed with caution due to the interrelationships among various environmental predictors. Recent temporal changes in body size have often been related to global warming. However, in many cases the above key predictors are not related to either latitude and/or year, and it is the task of the researcher to determine which particular environmental predictor is the one that determines food availability and, in turn, body size. The chance of discerning a significant change in body size depends to a large extent on sample size (specimens/year). The most recent changes in body size are probably phenotypic, but there are some cases in which they are partly genetic.     

Variation in body size and sexual size dimorphism in the most widely ranging lizard: testing the effects of reproductive mode and climate

Reproductive mode, ancestry, and climate are hypothesized to determine body size variation in reptiles but their effects have rarely been estimated simultaneously, especially at the intraspecific level. The common lizard (Zootoca vivipara) occupies almost the entire Northern Eurasia and includes viviparous and oviparous lineages, thus representing an excellent model for such studies. Using body length data for >10,000 individuals from 72 geographically distinct populations over the species' range, we analyzed how sex‐specific adult body size and sexual size dimorphism (SSD) is associated with reproductive mode, lineage identity, and several climatic variables. Variation in male size was low and poorly explained by our predictors. In contrast, female size and SSD varied considerably, demonstrating significant effects of reproductive mode and particularly seasonality. Populations of the western oviparous lineage (northern Spain, south‐western France) exhibited a smaller female size and less female‐biased SSD than those of the western viviparous (France to Eastern Europe) and the eastern viviparous (Eastern Europe to Far East) lineages; this pattern persisted even after controlling for climatic effects. The phenotypic response to seasonality was complex: across the lineages, as well as within the eastern viviparous lineage, female size and SSD increase with increasing seasonality, whereas the western viviparous lineage followed the opposing trends. Altogether, viviparous populations seem to follow a saw‐tooth geographic cline, which might reflect the nonmonotonic relationship of body size at maturity in females with the length of activity season. This relationship is predicted to arise in perennial ectotherms as a response to environmental constraints caused by seasonality of growth and reproduction. The SSD allometry followed the converse of Rensch's rule, a rare pattern for amniotes. Our results provide the first evidence of opposing body size—climate relationships in intraspecific units.     

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.     

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.     

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.     

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.     

Life on the edge: carnivore body size variation is all over the place

Evolutionary biologists have long been fascinated by both the ways in which species respond to ecological conditions at the edges of their geographic ranges and the way that species'' body sizes evolve across their ranges. Surprisingly, though, the relationship between these two phenomena is rarely studied. Here, we examine whether carnivore body size changes from the interior of their geographic range towards the range edges. We find that within species, body size often varies strongly with distance from the range edge. However, there is no general tendency across species for size to be either larger or smaller towards the edge. There is some evidence that the smallest guild members increase in size towards their range edges, but results for the largest guild members are equivocal. Whether individuals vary in relation to the distance from the range edges often depends on the way edge and interior are defined. Neither geographic range size nor absolute body size influences the tendency of size to vary with distance from the range edge. Therefore, we suggest that the frequent significant association between body size and the position of individuals along the edge-core continuum reflects the prevalence of geographic size variation and that the distance to range edge per se does not influence size evolution in a consistent way.     

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.     

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.     

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, ?? , ??–??.     

Latitude, elevation and body mass variation in Andean passerine birds

• 1 Relationships between body mass and latitude, and body mass and elevation are examined in the assemblage of Andean passerine birds.
• 2 Across species, body mass is positively correlated with the mid‐point of the species elevational distribution, but there is no significant relationship between body mass and latitudinal range mid‐point.
• 3 When the assemblage is separated into Andean endemic and non‐endemic species, the former group shows a significant positive relationship between body mass and elevation, and the latter a significant positive relationship between body mass and latitude (‘Bergmann’s rule’). Andean endemic species exhibit Bergmann’s rule once elevation is controlled for using multiple regression.
• 4 These relationships are not a consequence of the phylogenetic non‐independence of species. All the effects shown are very weak, with latitude and elevation explaining only a few per cent of the variation in body mass. Relationships are strongest when phylogenetically controlled analyses are performed just within genera.
• 5 The implications of these results for the mechanistic understanding of Bergmann’s rule are discussed.

     

Beyond latitude: Temperature,productivity and thermal niche conservatism drive global body size variation in Odonata

Aim

So far, latitudinal body size clines have been discussed primarily in the context of thermoregulation, sensu Bergmann. However, body size patterns are ambiguous in ectotherms, and this heterogeneity remains poorly understood. We tested whether Bergmann's rule and the resource availability rule, which states that energetic requirements determine species body size, apply to damselflies and dragonflies (Odonata). Furthermore, we hypothesized that the contrasting effects of thermoregulation and resource availability (e.g., productivity) can obscure the overall gradient in body size variation.

Location

Global.

Time period

Contemporary.

Major taxa studied

Odonata.

Methods

Using data for 43% of all odonate species described so far, we tested our hypotheses in phylogenetically and spatially comparative analyses at assemblage and species levels. For the distribution data, we integrated expert range maps and ecoregional ranges based on all available occurrence records. To distinguish between long-term and evolutionarily recent responses of environmental drivers in body size, we constructed a phylogenetically informed classification of all odonate species and decomposed the body size into its phylogenetic and specific components for our subset of species.

Results

We documented a weak positive relationship between body length and latitude but found strong and contrasting effects for temperature between dragonflies and damselflies and consistent positive effects for productivity that explained 35–57% of body size variation. Moreover, we showed a strong phylogenetic signal in sized-based thermoregulation that shaped the distribution of dragonflies, but not of damselflies.

Main conclusions

We concluded that temperature, productivity and conservatism in size-based thermoregulation synergistically determine the distribution of ectotherms, while the taxon-specific importance of these factors can lead to contrasting and weak latitude–size relationships. Our results reinforce the importance of body size as a determinant of species distributions and responses to climate change.     

Developmental variation in ecogeographic body proportions

While ecogeographic variation in adult human body proportions has been extensively explored, relatively less attention has been paid to the effect of Bergmann's and Allen's rules on human body shape during growth. The relationship between climate and immature body form is particularly important, as immature mortality is high, mechanisms of thermoregulation differ between young and mature humans, and immature body proportions fluctuate due to basic parameters of growth. This study explores changes in immature ecogeographic body proportions via analyses of anthropometric data from children included in Eveleth and Tanner's (1976) Worldwide Variation in Human Growth, as well as limb proportion measurements in eight different skeletal samples. Moderate to strong correlations exist between climatic data and immature stature, weight, BMI, and bi-iliac breadth; these relationships are as strong, if not stronger, in immature individuals as they are in adults. Correlations between climate and trunk height relative to stature are weak or nonexistent. Altitude also has significant effects on immature body form, with children from higher altitudes displaying smaller statures and lower body weights. Brachial and crural indices remain constant over the course of growth and display consistent, moderate correlations with latitude across ontogeny that are just as high as those detected in adults. The results of this study suggest that while some features of immature body form, such as bi-iliac breadth and intralimb indices, are strongly dictated by ecogeographic principles, other characteristics of immature body proportions are influenced by intrinsic and extrinsic factors such as nutrition and basic constraints of growth.     

Cranial ontogeny, diet, and ecogeographic variation in African lorises

A series of 22 craniodental measurements were obtained for the three subspecies of potto (Perodicticus) and angwantibos (Arctocebus). To describe patterns of variation in Perodicticus, a discriminant function analysis (DFA) was performed with adult data. To investigate the ecogeographic correlates of size variation in Perodicticus, adult cranial dimensions were compared with field data on latitudinal and longitudinal coordinates for available specimens as well as altitudinal data for a more limited sample. Ontogenetic series for larger-bodied Perodicticus and smaller-bodied Arctocebus were compared to test the hypothesis that inter- and intrageneric variation in skull form results from the differential extension/truncation of shared patterns of relative growth, and to assess morphological variation in the masticatory complex of sister taxa with differing dietary habits. Analyses of relative growth indicate that skull proportions in Perodicticus subspecies are largely ontogenetically scaled. In comparisons between Perodicticus and Arctocebus, most facial dimensions also are ontogenetically scaled, with all but one of the seven divergent comparisons (interorbital breadth) representing a feature of the masticatory apparatus. The DFA provided independent support for prior classifications of Perodicticus into three taxa. Size differentiation in African lorises appears to be correlated with altitudinal variation (Bergmann's Rule) as well as character displacement. The smallest pottos, P. p. potto, occupy low-lying coastal habitats in western Africa, whereas the larger, eastern forms inhabit higher, presumably colder elevations. The largest potto, P. p. edwardsi, is sympatric throughout most of its range with the smallest and most insectivorous African lorises (Arctocebus). A basis for intrageneric taxonomic variation in Perodicticus is supported by such nonclinal size variation, as well as divergences in the ontogeny of masticatory proportions corresponding to interspecific variation in dietary proclivities.     

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.     

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.     

Ecological and evolutionary drivers of range size in Coenagrion damselflies

Geographic range size is a key ecological and evolutionary characteristic of a species, yet the causal basis of variation in range size among species remains largely unresolved. One major reason for this is that several ecological and evolutionary traits may jointly shape species' differences in range size. We here present an integrated study of the contribution of ecological (dispersal capacity, body size and latitudinal position) and macroevolutionary (species' age) traits in shaping variation in species' range size in Coenagrion damselflies. We reconstructed the phylogenetic tree of this genus to account for evolutionary history when assessing the contribution of the ecological traits and to evaluate the role of the macroevolutionary trait (species' age). The genus invaded the Nearctic twice independently from the Palearctic, yet this was not associated with the evolution of larger range sizes or dispersal capacity. Body size and species' age did not explain variation in range size. There is higher flight ability (as measured by wing aspect ratio) at higher latitudes. Species with a larger wing aspect ratio had a larger range size, also after correcting for phylogeny, suggesting a role for dispersal capacity in shaping the species' ranges. More northern species had a larger species' range, consistent with Rapoport's rule, possibly related to niche width. Our results underscore the importance of integrating macroecology and macroevolution when explaining range size variation among species.