Geographic Variation In Body Size Research Articles

Viviparity is associated with larger female size and higher sexual size dimorphism in a reproductively bimodal lizard.

Squamate reptiles are central for studying phenotypic correlates of evolutionary transitions from oviparity to viviparity because these transitions are numerous, with many of them being recent. Several models of life-history theory predict that viviparity is associated with increased female size, and thus more female-biased sexual size dimorphism (SSD). Yet, the corresponding empirical evidence is overall weak and inconsistent. The lizard Zootoca vivipara, which occupies a major part of Northern Eurasia and includes four viviparous and two non-sister oviparous lineages, represents an excellent model for testing these predictions. We analysed how sex-specific body size and SSD is associated with parity mode, using body length data for nearly 14,000 adult individuals from 97 geographically distinct populations, which cover almost the entire species' range and represent all six lineages. Our analyses controlled for lineage identity, climatic seasonality (the strongest predictor of geographic body size variation in previous studies of this species) and several aspects of data heterogeneity. Parity mode, lineage and seasonality are significantly associated with female size and SSD; the first two predictors accounted for 14%-26% of the total variation each, while seasonality explained 5%-7%. Viviparous populations exhibited a larger female size than oviparous populations, with no concomitant differences in male size. The variation of male size was overall low and poorly explained by our predictors. Albeit fully expected from theory, the strong female bias of the body size differences between oviparous and viviparous populations found in Z. vivipara is not evident from available data on three other lizard systems of closely related lineages differing in parity mode. We confront this pattern with the data on female reproductive traits in the considered systems and the frequencies of evolutionary changes of parity mode in the corresponding lizard families and speculate why the life-history correlates of live-bearing in Z. vivipara are distinct. Comparing conspecific populations, our study provides the most direct evidence for the predicted effect of parity mode on adult body size but also demonstrates that the revealed pattern may not be general. This might explain why across squamates, viviparity is only weakly associated with larger size.

Geographic body size variation of a Plateau anuran: evidence supporting the water availability and hibernation hypotheses

Geographic body size variation of a Plateau anuran: evidence supporting the water availability and hibernation hypotheses

Geographic Variation in Body Size and Hematology of the Slate-Colored Coot (Fulica ardesiaca) along the Andes of Peru

Geographic Variation in Body Size and Hematology of the Slate-Colored Coot (Fulica ardesiaca) along the Andes of Peru

Can neutral genetic differentiation explain geographical variation in body size of the natterjack toad, Epidalea calamita?

Population genetic studies are crucial for evolutionary biologists because the population is the basic substrate on which evolution is forged. However little empirical evidence has been able to demonstrate the role that isolation and gene flow play in maintaining differentiation in populations at short geographic scales. Epidalea calamita exhibits a steep variation in body size and reproductive traits in southwestern Spain, associated with changes in the geological substrate. This implies a decrease of 70.9% of body mass and 28.5% in snout-vent length, on a micro-geographic scale of only 60 km. Previous results from both metamorphic and juvenile common garden experiments showed that genetic differentiation may be a causal determinant of geographic variation in adult. This study tested whether neutral genetic differentiation can explain the geographical variation in the body size observed in E. calamita. It was addressed analyzing the level of genetic structuring and gene flow among populations along the cline, comparing the genetic diversity between and within populations, as well as between ecological environments. The study showed that the geographic variation in body size observed in E. calamita has evolved in absence of geographic isolation, with moderate gene flow connecting the populations. Thus, neutral genetic differentiation cannot explain the geographical variation observed. Future studies are needed on the interaction between the genetic component with the environmental factors and will be necessary to analyze the contribution of the maternal effects in the origin and evolution of the geographical variation in the body size observed in E. calamita from southern Spain.

Geographical Variation in Body Size in the Asian Common Toad (Duttaphrynus melanostictus).

The geographic variation in life-history traits of organisms and the mechanisms underlying adaptation are interesting ideas in evolutionary biology. This study investigated age and body size of the Asian common toad (Duttaphrynus melanostictus) among five populations along a geographical gradient. We found that geographical variation in age was non-significant among populations but there was a significant and positive correlation between mean age and body size. Although the body size values at 1043 m are quite different from other sites, after controlling for age effects, there was a significant positive correlation between altitude and body size. Our findings followed the predictions of Bergmann's rule, suggesting that the body size of D. melanostictus is potentially influenced by the low air temperatures at higher altitudes.

Geographical Variation in Body Size and the Bergmann's Rule in Andrew's Toad (Bufo andrewsi).

Environmental variation likely modifies the life-history traits of vertebrates. As ectothermic vertebrates, it is possible that the body size of amphibians is impacted by environmental conditions. Here, we firstly quantified age and body size variation in the Andrew's toad (Bufo andrewsi) across the Hengduan Mountains. Then, we examined the environmental correlates of this variation based on the literature and our unpublished data on the age and body size of the Andrew's toad from 31 populations distributed in southwestern China. Although our analysis revealed significant variations in age and body size across B. andrewsi populations, neither latitude nor altitude correlated with this variability in age and body size. We found that age at sexual maturity, mean age, and longevity increased with decreasing annual mean temperature, whereas age at sexual maturity increased with decreasing temperature seasonality, implying that temperature was a crucial habitat characteristic that modulated age structure traits. Moreover, we revealed positive associations between age structure and UV-B seasonality, and negative relationships between both mean age and longevity and precipitation seasonality. We also found that body size increased with increasing precipitation in the driest month and UV-B seasonality. However, body size did not covary with temperature, signifying no support for Bergmann's rule. These findings help us to understand amphibians' abilities to adapt to environmental variation, which is particularly important in order to provide a theorical basis for their conservation.

Acoustic clue: bringing echolocation call data into the distribution dilemma of Pteronotus (Chiroptera: Mormoopidae) complexes in Central America

Abstract In Central America, the distributional limits and the contact zones of some Pteronotus species, such as the naked-backed bats and the lesser mustached bats, are unclear. To elucidate the distributional dilemma of the species groups Pteronotus fulvus + Pteronotus davyi and Pteronotus psilotis + Pteronotus personatus in Central America, we studied the acoustic variation of their echolocation calls along the range of possible contact zones and the existence of distinct phonic groups. We performed hierarchical k-means clustering on principal components (HCPC) using acoustic samples from Mexico, Honduras, El Salvador, Nicaragua and Costa Rica to describe the global acoustic diversity, possibly overlooking differences between species groups. We assessed whether those acoustic differences were related to the geographical distance and geographical location. We found evidence of sympatry for three phonic groups within each species complex, without a clear correspondence to the known distribution of species. The changes in frequency of their echolocation calls seem to follow a similar pattern to the geographical variation in body size of the species. Future studies in Central America should include an integrative sampling of individually captured, tagged and recorded bats to help in resolution of the distribution dilemma raised here.

Increasing Body Size after Exploitation in a Population of Timber Rattlesnakes (Crotalus horridus)

At a field site in northeastern New York during a 40-yr study, we measured the body sizes (total length, snout–vent length [SVL], mass) in three intrinsic groups of adult Crotalus horridus: nongravid (N) females, gravid (G) females, and males. We argue that a bounty system in effect for 75 yr caused a probable decline of the largest animals. After this lengthy period of exploitation, we observed that body sizes began to increase. Across 4 decades, the three intrinsic groups were significantly larger in the final decade (2009–18) than in the first decade (1979–88), stabilizing at a mean body mass of »710 g in N females, »870 g in G females, and »1,200 g in males. Mean SVLs of N females (94.1 cm) and males (111.0 cm) in the final 3 decades were significantly larger than their SVLs in the first decade. The body condition index (BCI) increased over time, and the intrinsic groups differed significantly in mean BCI: N females (-0.0764), G females (0.0146), and males (0.0899). Among recaptured snakes, wide fluctuations of body mass were not uncommon as most snakes experienced annual losses as well as gains. Geographic variation in body size of C. horridus is illustrated by comparing our New York population with a northern population (Massachusetts) and a southern population (Virginia). Across the species' range, we suggest that body size could be associated primarily with prey size. Because many populations of C. horridus are slow growing and long lived, correlated effects of exploitation on body size may negatively affect the snakes' reproductive biology and longevity.

Geographic variation in body size and plumage colour according to diet composition in a nocturnal raptor

Predator–prey interactions are amongst the strongest selective forces that promote the evolution of local phenotypes in both predators and prey. However, intraspecific spatial covariation in phenotypic traits between predators and prey has been rarely investigated, especially at a large geographic scale. Here, we studied the covariation between prey composition and some phenotypic traits, such as wing length, bill length and plumage colour, of a widely‐distributed nocturnal predator, the western barn owl Tyto alba. By using 3100 specimens collected across its entire range of distribution, spanning from Europe to Middle East and Africa, we showed that wing length positively covaries with prey size, but not with taxonomic composition. This finding suggests that larger prey might have selected for larger body size and/or that larger individuals might be more selective in hunting large prey. In addition, we also found that paler‐plumaged populations generally hunt larger prey. Paler barn owls might be thus better specialized in capturing averagely larger prey and/or mainly hunt in habitats where larger prey are more abundant. In addition, considering that paler individuals are generally larger than brownish ones, it is possible that paler plumage colour might have evolved as a by‐product of selection towards a large body size, which in turn have emerged in response to prey size composition. However, irrespectively of the direction of causality and the phenotypic target of selection, we showed that predator–prey interactions can affect spatial phenotypic variation by promoting the evolution of local adaptations.

Can age and growth patterns explain the geographical variation in the body size of two toad species?

Determining both the age structure and growth pattern allows to establish the causal factors, environmental and/or genetic, that eventually may be responsible for the observed pattern of divergence. We examined the variation in age structure and growth pattern across populations of two toad species, Pelobates cultripes and Epidalea calamita that exhibit a geographic variation in body size in southern Spain. For both species, populations differed in mean age but age structure did not correlate with body size variation across populations. Although the population with the youngest females found for E. calamita was the smallest in body size, the oldest males for both species were found in a small body size population. The growth pattern fit well to a von Bertalanffy growth model and interdemic divergence were found for both the asymptotic body size (Sm ) and the growth coefficients (k). As expected, Large-Bodied populations of both species attained higher Sm but, Small-Bodied population had higher, although non significantly different, k growth coefficients. Also, the Small-Bodied population attained sexual maturity sooner but had also high longevity. The observed pattern may reflect both environmental variations in resources availability affecting body size observed across populations, but also different growth and maturity pathways that may respond to contrasting selective pressures.

Geographic variation in body size among French populations of the European pond turtle

Abstract The processes underlying macroecological gradients in body size are widely debated, in part because their intraspecific variability remains poorly described even in well-studied taxa such as vertebrates. In this study, we investigated how climate, habitat, genetic lineage and sex explain body size variations in French populations of the European pond turtle (Emys orbicularis). We measured 7016 adult individuals captured in 41 populations, covering most of the species’ distribution in metropolitan France, including Corsica. Body size variation in our sample was wide and comparable to that found across the species’ worldwide range. Variation was similar in magnitude at regional and local levels, suggesting that body size is influenced by local factors as much as by regional factors such as climate or genetic lineage. Smaller sizes were associated with Mediterranean or altered oceanic climates, and with two lineages (E. o. galloitalica and E. o. galloitalica/E. o. orbicularis), while larger sizes were associated with northern environments and the orbicularis lineage. Body size variations recorded at local level reflect an adaptive response to environmental constraints, suggesting that habitat is also an important factor in understanding size variation.

Temperature drives caste-specific morphological clines in ants.

The morphology of organisms relates to most aspects of their life history and autecology. As such, elucidating the drivers of morphological variation along environmental gradients might give insight into processes limiting species distributions. In eusocial organisms, the concept of morphology is more complex than in solitary organisms. Eusocial insects such as ants exhibit drastic morphological differences between reproductive and worker castes. How environmental selection operates on the morphology of each caste, and whether caste-specific selection has fitness consequences is largely unknown, but is potentially crucial to understand what limits ant species' distributions. Here we aimed to examine whether ant shape and body size covaries with climate at the scale of an entire continent, and whether such relationship might be caste specific. We used 26,472 georeferenced morphometric measurements from 2,206 individual ants belonging to 32 closely related North American species in the genus Formica to assess how ant morphology relates to geographic variation in the abiotic environment. Although precipitation and seasonality explained some of the geographic variation in morphology, temperature was the best predictor. Specifically, geographic variation in body size was positively related to temperature, meaning that ants are smaller in cold than in warm environments. Moreover, the strength of the relationship between size and temperature was stronger for the reproductive castes (i.e. queens and males) than for the worker caste. The shape of workers and males also varied along these large-scale abiotic gradients. Specifically, the relative length of workers' legs, thoraxes and antennae positively related to temperature, meaning that they had shorter appendages in cold environments. In contrast, males had smaller heads, but larger thoraxes in more seasonal environments. Overall, our results suggest that geographic variation in ambient temperature influences the morphology of ants, but that the strength of this effect is caste specific. In conclusion, whereas ant ecology has traditionally focused on workers, our study shows that considering the ecology of the reproductive castes is imperative to move forward in this field.

Spatial predictors of genomic and phenotypic variation differ in a lowland Middle American bird (Icterus gularis).

Spatial patterns of intraspecific variation are shaped by geographical distance among populations, historical changes in gene flow and interactions with local environments. Although these factors are not mutually exclusive and operate on both genomic and phenotypic variation, it is unclear how they affect these two axes of variation. We address this question by exploring the predictors of genomic and phenotypic divergence in Icterus gularis, a broadly distributed Middle American bird that exhibits marked geographical variation in body size across its range. We combined a comprehensive single nucleotide polymorphism and phenotypic data set to test whether genome-wide genetic and phenotypic differentiation are best explained by (i) isolation by distance, (ii) isolation by history or (iii) isolation by environment. We find that the pronounced genetic and phenotypic variation in I. gularis are only partially correlated and differ regarding spatial predictors. Whereas genomic variation is largely explained by historical barriers to gene flow, phenotypic diversity can be best predicted by contemporary environmental heterogeneity. Our genomic analyses reveal strong phylogeographical structure coinciding with the Chivela Pass at the Isthmus of Tehuantepec that was formed during the Pleistocene, when populations were isolated in north-south refugia. In contrast, we found a strong association between body size and environmental variables, such as temperature and precipitation. The relationship between body size and local climate is consistent with a pattern produced by either natural selection or environmental plasticity. Overall, these results provide empirical evidence for why phenotypic and genomic data are often in conflict in taxonomic and phylogeographical studies.

Geographic variation in body size of five Australian marsupials supports Bergmann’s thermoregulation hypothesis

Abstract Despite a large body of research, little agreement has been reached on the ultimate driver(s) of geographic variation in body size (mass and/or length). Here we use skull length measurements (as a surrogate for body mass) from five Australian marsupial species to test the primary hypotheses of geographic variation in body size (relating to ambient temperature, productivity, and seasonality). We used a revised articulation of Bergmann’s rule, wherein evidence for thermoregulation (heat dissipation or heat conservation) is considered supportive of Bergmann’s rule. We modeled the skull lengths of four Petaurid glider species and the common brushtail possum (Trichosurus vulpecula) as a function of indices of ambient temperature, productivity, and seasonality. The skull length of Petaurus ariel, P. notatus, and the squirrel glider (P. norfolcensis), increased with increasing winter minimum temperature, while that of T. vulpecula decreased with increasing summer maximum temperature. The skull length of P. ariel decreased with indices of productivity, falsifying the productivity hypothesis. Only P. ariel met the hypothesis of seasonality, as skull length increased with increasing seasonality. Thermoregulation was the most consistently supported driver of geographic variation in body size, as we found evidence of either heat conservation or heat dissipation in four of the five species examined. We found the geographic range of the individual species and the climate space in which the species occurred was integral to understanding the species’ responses to climate variables. Future studies should use specimens that are representative of a species’ entire geographic range, encompass a variety of climatic regions, and use consistent methodologies and terminology when testing drivers of geographic variation in body size.

Ambient temperature correlates with geographic variation in body size of least horseshoe bats.

Geographic variation in body size is common within many animal species. The causes of this pattern, however, remain largely unexplored in most vertebrate groups. Bats are widely distributed globally owing to their ability of powered flight. Most bat species encounter a variety of climatic conditions across their distribution range, making them an ideal taxon for the study of ecogeographic patterns in body size. Here, we used adult least horseshoe bats, Rhinolophus pusillus, to test whether geographic variation in body size was determined by heat conservation, heat dissipation, climatic seasonality, or primary productivity. We measured body mass and head-body length for 246 adult bats from 12 allopatric colonies in China. We quantified the ecological conditions inhabited by each colony, including mean maximum temperature of the warmest month, mean minimum temperature of the coldest month, temperature seasonality, precipitation seasonality, and annual net primary productivity (ANPP). Body mass and head-body length, 2 of the most reliable indicators of body size, exhibited marked differences between colonies. After controlling for spatial autocorrelation, the mean minimum temperature of the coldest month explained most of the variation in body size among colonies, regardless of sex. The mean maximum temperature, climatic seasonality, and ANPP had limited power in predicting body size of males or females in comparison with mean minimum temperature. These results support the heat conservation hypothesis and suggest adaptive responses of body size to cold climates in cave-dwelling bats.

Geographic body size variation of a tropical anuran: effects of water deficit and precipitation seasonality on Asian common toad from southern Asia

BackgroundTwo previous studies on interspecific body size variation of anurans found that the key drivers of variation are the species’ lifestyles and the environments that they live in. To examine whether those findings apply at the intraspecific level, we conducted a study of the Asian common toad (Duttaphrynus melanostictus), a terrestrial anuran distributed in tropical regions. The body size of toads from 15 locations, covering the majority of their geographic range, and local environmental data were summarized from published literature. We used a model selection process based on an information-theoretic approach to examine the relationship between toad body size and those environmental parameters.ResultsWe found a positive correlation between the body size of the Asian common toad and the water deficit gradient, but no linkage between body size and temperature-related parameters. Furthermore, there was a positive correlation between the seasonality of precipitation and body size of females from different sampled populations.ConclusionsAs a terrestrial anuran, the Asian common toad should experience greater pressure from environmental fluctuations than aquatic species. It is mainly distributed in tropical regions where temperatures are generally warm and stable, but water availability fluctuates. Therefore, while thermal gradients are not strong enough to generate selection pressure on body size, the moisture gradient is strong enough to select for larger size in both males and females in dryer regions. Larger body size supports more efficient water conservation, a pattern in accordance with the prediction that lifestyles of different species and their local habitats determine the relationship between body size and environment. In addition, larger females occur in regions with greater seasonality in precipitation, which may happen because larger females can afford greater reproductive output in a limited reproductive season.

The patterns and possible causes of global geographical variation in the body size of the greater horseshoe bat (Rhinolophus ferrumequinum)

AbstractAimGeographical variations in endotherm body size (e.g. Bergmann's rule/James's rule and Allen's rule) have long been tested. However, the patterns and causes of geographical variation in body size within bat species, especially within widespread hibernating species, are little known. Here, we evaluated the possible causes of geographical size variation patterns in the greater horseshoe bat (Rhinolophus ferrumequinum), a bat species widely distributed across the Palearctic.LocationPalearctic.MethodsWe collected body size data (body mass and forearm length) from 1,172 adult bats via direct measurements or from the literature. We used high‐resolution environmental data for the sampled sites. We applied multiple linear regressions and an information‐theoretic approach on separate female and male datasets to estimate the support for various hypotheses.ResultsTemperature and latitude failed to predict body mass in both females and males. The best model showed that populations with longer forearms in both sexes were associated with higher mean temperatures (in the warmest quarter) and lower mean temperatures (in the coldest quarter). The independent contribution of these temperature parameters to forearm length in both sexes was higher than that of all other variables.Main conclusionThe observed relationship between temperature and body mass was not consistent with James's rule, and thus failed to support the heat conservation hypothesis. Our results are consistent with the predictions of Allen's rule and indicated that direct heat dissipation from the wing may be the most likely mechanism underlying geographical variation in forearm length. We suggest that the evaluation of the relative importance of multiple causal mechanisms may improve our understanding of patterns of geographical variation in endotherms.

How has the environment shaped geographical patterns of insect body sizes? A test of hypotheses using sphingid moths

AbstractAimWe mapped the geographical pattern of body sizes in sphingid moths and investigated latitudinal clines. We tested hypotheses concerning their possible environmental control, that is, effects of temperature (negative: temperature size rule or Bergmann's rule; positive: converse Bergmann rule), food availability, robustness to starvation during extreme weather and seasonality.LocationOld World and Australia/Pacific region.MethodsBody size data of 950 sphingid species were compiled and related to their distribution maps. Focusing on body length, we mapped the median and maximum size of all species occurring in 100 km grid cells. In a comparative approach, we tested the predictions from explanatory hypotheses by correlating species' size to the average environmental conditions encountered throughout their range, under univariate and multivariate models. We accounted for phylogeny by stepwise inclusion of phylogenetically informed taxonomic classifications into hierarchical random‐intercept mixed models.ResultsMedian body sizes showed a distinctive geographical pattern, with large species in the Middle East and the Asian tropics, and smaller species in temperate regions and the Afrotropics. Absolute latitude explained very little body size variation, but there was a latitudinal cline of maximum size. Species' median size was correlated with net primary productivity, supporting the food availability hypothesis, whereas support for other hypotheses was weak. Environmental correlations contributed much less (i.e. <10%) to explaining overall size variation than phylogeny (inclusion of which led to models explaining >70% of variability).Main conclusionThe intuitive impression of larger species in the tropics is shaped by larger size maxima. Median body sizes are only very weakly related to latitude. Most of the geographical variation in body size in sphingid moths is explained by their phylogenetic past. NPP and forest cover correlate positively with the body size, which supports the idea that food availability allowed the evolution of larger sizes.

Changes in the Body Size of Carabid Beetles Along Elevational Gradients: A Multispecies Study of Between- and Within-Population Variation.

Geographic variation in body size has fascinated biologists since the 19th century as it can provide insight into the evolution of the body size of various organisms. In this study, we investigated body size variation in eight carabid species/subspecies (Coleoptera: Carabidae) along elevational gradients in six Central European mountain ranges. First, we examined elevational variation in body size and whether female and male body sizes differed in their responses to elevation. Second, we examined intrapopulation variation in body size along an elevational gradient, and we compared the degrees of intrapopulation variation between males and females. The investigated species either followed a converse Bergmann's cline (Carabus auronitens auronitens Fabricius 1792; Carabus linnei Panzer 1810; Pterostichus melanarius (Illiger, 1798); Pterostichus pilosus (Host, 1789)) or their size was unaffected by elevation (Carabus auronitens escheri Palliardi, 1825; Carabus sylvestris sylvestris Panzer, 1796; Carabus sylvestris transsylvanicus Dejean, 1826; Pterostichus burmeisteri Heer, 1838). Females were the larger sex in all the investigated species, but the degree of sexual size dimorphism differed between species. In general, the degree of sexual size dimorphism showed no change with elevation. The degree of intrapopulation variation in body size slightly increased with elevation in C. sylvestris sylvestris and P. pilosus. Overall, the intrapopulation variation in body size significantly differed among the investigated carabid species. The existing literature on intrapopulation variation in the body size of insects is limited, but further investigation of this issue could provide a better understanding of the mechanisms that generate geographical clines.

Microgeographic variation in body condition of three Mexican garter snakes in central Mexico.

BackgroundGeographic variation in body size and condition can reveal differential local adaptation to resource availability or climatic factors. Body size and condition are related to fitness in garter snakes (Thamnophis), thus good body condition may increase survival, fecundity in females, and mating success in males. Phylogenetically related species in sympatry are predicted to exhibit similar body condition when they experience similar environmental conditions. We focused on interspecific and geographical variation in body size and condition in three sympatric Mexican garter snakes from the highlands of Central Mexico.MethodsWe assessed SVL, mass, and body condition (obtained from Major axis linear regression of ln-transformed body mass on ln-transformed SVL) in adults and juveniles of both sexes of Thamnophis eques, T. melanogaster, and T. scalaris sampled at different locations and ranges from 3–11 years over a 20-year period.ResultsWe provide a heterogeneous pattern of sexual and ontogenic reproductive status variations of body size and condition among local populations. Each garter snake species shows locations with good and poor body condition; juvenile snakes show similar body condition between populations, adults show varying body condition between populations, and adults also show sex differences in body condition. We discuss variations in body condition as possibly related to the snakes’ life cycle differences.