Geographic variation of wing morphology of great fruit‐eating bats (Artibeus lituratus): environmental,genetic and spatial correlates of phenotypic differences

Responses of species to environmental gradients are important and frequent determinants of geographic phenotypic variation that can drive adaptive processes. Nonetheless, random genetic processes such as drift can also result in geographic variation in phenotypes, and should be evaluated before implicating selection as the process driving phenotypic change. We examined geographic variation in wing morphology of Artibeus lituratus among 18 different sites distributed across interior Atlantic Forest of Paraguay and Argentina. Moreover, we contrasted geographic variation with environmental, spatial, and genetic variation to test hypotheses related to selection and drift and their impacts on wing morphology. For A. lituratus distributed across interior Atlantic Forest, significant differences among sites characterized variation in wing morphology. Geographic variation was significantly related to climatic variables but not spatial or genetic distances. Such a pattern suggests that phenotypic variation is related to selection for particular environmental regimes, and not genetic drift. Four significant dimensions of phenotypic variation were determined. Three dimensions were related to variation among individuals in terms of wing tips, whereas one was related to overall body size. Wing tips are important for manoeuverability during flight and differences among sites likely reflect differences in forest and vegetation structure that must be managed during foraging. Although climate provides good surrogates for environmental variation, it is probably only an indirect cue of selection regimes that determine variation in wing morphology. Future studies should evaluate more direct environmental measures such as vegetation structure when attempting to interpret geographical variation in wing morphology.     

The geographic structure of morphological variation in eight species of fiddler crabs (Ocypodidae: genus Uca) from the eastern United States and Mexico

Species with larger geographic distributions are more likely to encounter a greater variety of environmental conditions and barriers to gene flow than geographically‐restricted species. Thus, even closely‐related species with similar life‐history strategies might vary in degree and geographic structure of variation if they differ in geographic range size. In the present study, we investigated this using samples collected across the geographic ranges of eight species of fiddler crabs (Crustacea: Uca) from the Atlantic and Gulf coasts of North America. Morphological variation in the carapace was assessed using geometric morphometric analysis of 945 specimens. Although the eight Uca species exhibit different degrees of intraspecific variation, widespread species do not necessarily exhibit more intraspecific or geographic variation in carapace morphology. Instead, species with more intraspecific variation show stronger morphological divergence among populations. This morphological divergence is partly a result of allometric growth coupled with differences in maximum body size among populations. On average, 10% of total within‐species variation is attributable to allometry. Possible drivers of the remaining morphological differences among populations include gene flow mediated by ocean currents and plastic responses to various environmental stimuli, with isolation‐by‐distance playing a less important role. The results obtained indicate that morphological divergence among populations can occur over shorter distances than expected based on dispersal potential. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 248–270.     

Prey‐associated head‐size variation in an invasive lizard in the Hawaiian Islands

Biological invasions are recognized as a primary driver of large‐scale changes in global ecosystems. This study addresses ecomorphological variation in head size within and among populations of an ecologically destructive invasive predator, and evaluates the potential roles of environmental components in phenotypic differentiation. We used four size‐corrected measurements of head morphology in Jackson's chameleons, Trioceros jacksonii xantholophus (N = 319), collected from multiple Hawaiian Islands to assess phenotypic variation among and within islands. Results of analysis of variance (ANOVA) comparing chameleon head size (PC1) among islands revealed significant differences (mean difference > 5%) associated with variation in both rainfall and diet composition using Mann–Whitney U‐tests and chi‐squared analyses. These results suggest that morphological differentiation among populations from different islands has occurred over a relatively short ecological timescale, and is likely the result of ecomorphological adaptation to differences in exploited prey hardness. Intra‐island allopatric population variation, however, was also detected in this study. Although we might expect that genetic change is the more likely explanation for differences between islands than within, and that plasticity may be more likely an explanation for the within‐ than the between‐island differences, it is also possible that both within‐ and between‐island patterns are the results of genetic change, or of plasticity.     

Congruence between environmental parameters, morphology and genetic structure in Australia’s most widely distributed eucalypt, Eucalyptus camaldulensis

Eucalyptus camaldulensis is one of the most widely utilised eucalypts. It is also the only eucalypt that occurs across the Australian continent, playing a key ecological role as fauna habitat and in riverbank stabilisation. Despite its ecological and economic importance, uncertainty remains regarding the delineation of genetic and morphological variants. Nine hundred and ninety trees from 97 populations, representing the species’ geographic range were genotyped using 15 microsatellite loci and patterns of diversity compared with restriction fragment length polymorphisms in 29 of these populations. Both markers showed that despite having a riverine distribution, downstream seed dispersal has had less influence than geographic distance on dispersal patterns. Spatial patterns in the distribution of microsatellite genotypes were compared with environmental parameters and boundaries defined by river systems, drainage basins and proposed subspecies. Significant genetic differences among populations within river systems indicated that rivers should not be treated as a single genetic entity in conservation or breeding programmes. Strong geographic trends were evident with 40% of variation in genetic diversity explained by latitude and moisture index. Isolation by distance and significant correlations between genetic distance and environmental parameters for most loci suggest historical factors have had more influence than selection on current patterns of distribution of genetic diversity. Geographic structuring of molecular variation, together with congruence between genetic and morphological variation indicate that E. camaldulensis should be treated as a number of subspecies rather than a single variable taxon. High levels of genetic diversity and geographic trends in the distribution of variation provide a firm basis for further exploration of the species’ genetic resources.     

Integrating large‐scale geographic patterns in flight morphology,flight characteristics and sexual selection in a range‐expanding damselfly

While geographic trait variation along environmental clines is widespread, associated patterns in sexual selection remain largely unexplored. Geographic patterns in sexual selection may be expected if 1) phenotypes vary geographically and sexual selection is dependent on the local phenotypes in the population, and if 2) sexual selection is influenced by geographically structured environmental conditions. We quantified geographic variation in flight‐related traits and flight performance in mated and unmated males and tested for geographic variation in sexual selection on these traits in the poleward range‐expanding damselfly Coenagrion scitulum across a set of eleven core and edge populations ordered along thermal gradients in the larval and in the adult stage. We found little support for trait differentiation between core and edge populations, instead we found considerable geographic trait variation along the larval and adult thermal gradients. As expected under time constraints, body mass decreased with shorter larval growth seasons. Lower temperatures during the adult flight period were associated with a higher body mass, a higher flight speed and a higher fat content; these traits likely evolved to buffer flight ability at suboptimal temperatures and to optimize starvation resistance. Across the large geographic scale, we found a consistent higher flight duration in mated males. Instead, sexual selection for higher fat content was stronger in populations with lower adult flight temperatures and sexual selection for lower body mass acted only in edge populations. Our results indicate sexual selection on flight performance to be consistent over a large geographic scale and this despite the clear geographic patterns in sexual selection on the underlying morphological traits. Our results highlight that to fully understand the fitness implications of geographically changing trait patterns, researchers should consider the entire phenotype–performance–fitness axis and incorporate effects of geographically structured life‐stage specific environmental conditions on this axis.     

Geographic patterns of song variation in four species of Malurus fairy‐wrens

Geographic variation in song is widespread among birds, particularly in species that learn vocalizations. The relationship between geographic distance and song variation is likely related to the degree of isolation between populations. To assess this effect of geographic isolation on song divergence, we examined patterns of geographic song variation in four species of Australian fairy‐wrens (Malurus), two with suspected histories of geographic isolation and two without. Song variation in all four species was consistent with patterns of isolation by distance, and allopatric subspecies in two species were more divergent in song than predicted by distance alone. Each species’ pattern was unique, and some interspecific variation could not be explained by geographic distance. These results indicate that patterns of geographic variation can be influenced by more than geographic distance and historical isolation alone. We suggest that morphological constraints, environmental influences, and sexual selection may all contribute to the variation observed for each species.     

Lean‐season primary productivity and heat dissipation as key drivers of geographic body‐size variation in a widespread marsupial

Geographic body‐size variation characterises many mammal species. Hypotheses centring around heat conservation, heat dissipation, primary productivity and seasonality have been advanced to explain geographic body‐size patterns. However, identification of the primary body‐size drivers has often been hampered by a paucity of data for broadly distributed species and the application of regression models that have not explicitly accounted for the spatial clustering inherent in such datasets. We used Australia's most widespread marsupial, the common brushtail possum Trichosurus vulpecula, as a model species with which to test five proposed drivers of geographic body‐size variation. Using geo‐referenced skull measurements from 588 specimens and a suite of putative environmental covariates, we employed spatial simultaneous autoregressive models, together with information criteria, to evaluate these different hypotheses. Our analysis identified a strong, positive relationship between possum body size and primary productivity during the least productive season, whereas the relationship with mean annual productivity received less support. Consistent with the heat‐dissipation hypothesis, T. vulpecula body size also decreased with increasing mean summer maximum temperature. Spatial autoregression coefficients estimated from the simultaneous autoregressive models were always high, suggesting that additional abiotic or biotic factors might contribute to the spatial patterns observed. We argue that the analysis of geographic body‐size variation should consider multi‐causal possibilities rather than treating the numerous hypotheses as competing, mutually exclusive alternatives.     

Large‐scale geographic variation in iridescent structural ornaments of a long‐distance migratory bird

Iridescent colours produced during moult likely play an important role in pair formation in birds. We sought to quantify geographic variation in such colouration in a duck species, Eurasian teal Anas crecca, in winter (when mating occurs) to evaluate whether this variation reflects birds’ breeding origins or differential individual migration strategies in both males and females. We combined information on feather production region and individual attributes (body size, sex and age) of Eurasian teal from 82 wintering sites in France. Feather production region (moult site or natal origin) was inferred using feather deuterium values (δD_f). We performed spectral measurements to evaluate speculum colour and brightness contrasts for 1052 teal collected over four years. Colouration differed strongly among wintering regions, with birds wintering in eastern France exhibiting higher colour contrast than those wintering in the west. Body size and colouration were positively related. There were no differences in cohort‐specific δD_f values between separate wintering regions in France, indicating that within a winter quarter teal originated from areas across the entire breeding range. Overall, patterns of spatial variation in feather colouration were related most closely to body size which was consistent with predictions of a differential migration hypothesis, with larger and more colour‐contrasting birds wintering closer to their breeding grounds. Because moult speed is also known to affect colour production, early breeders or individuals that skipped reproduction may have invested more or earlier in their feather quality to gain potential advantages in monopolizing future mates.     

Local adaptation versus historical isolation as sources of melanin‐based coloration in the white‐throated thrush Turdus assimilis

Local adaptation seems to be one of the causes of variation in melanin‐based colors in bird plumages, related mainly to the heterogeneity of the environmental conditions along the distribution of a species. Based on comparisons of genetic (mtDNA sequences), ecological (niche models), and quantitative colorimetric data, we explored variation in plumage coloration of the white‐throated thrush Turdus assimilis, a Mesoamerican species whose dorsal color varies from brown (northern and central Mexico) to dark gray (southern Mexico and Central America). Our results suggest the existence of two major patterns of coloration in this bird, which are congruent with the genetic structure, and comparisons of ecological niche models showed that population's niches were more similar than expected by chance, suggesting that color variation in plumage of T. assimilis is not consequence of local adaptation to different environmental conditions. Our results also showed that a greater geographic distance between populations is correlated with greater colorimetric differences, suggesting that color variation in T. assimilis may be consequence of historical isolation.     

Geographical variation in sperm morphology in the red-winged blackbird (<Emphasis Type="Italic">Agelaius phoeniceus</Emphasis>)

Many species differ genetically, physiologically, and morphologically between geographically distinct populations, typically in response to variation in ecological and climatic variables. Little is known, however, about geographical variation in sperm morphology. Sperm morphology is under strong sexual selection, has been shown to evolve rapidly, and often co-varies with other reproductive traits (e.g., testis size or mating system) that differ between populations in some species. The aim of this study was to establish whether sperm morphology varies between populations of the red-winged blackbird (Agelaius phoeniceus), a species with an enormous breeding range and marked inter-population variation in both body size and mating system. We found (1) highly significant variation in sperm morphology among study sites, (2) a gradual increase in sperm length from the southwest to the northeast of the breeding range, and (3) a strong negative association between sperm length and body size. However, the relationship with the mating system remains unclear. Several hypotheses to explain these patterns are proposed.     

Patterns of morphological variation among native and non-native pumpkinseed (<Emphasis Type="Italic">Lepomis gibbosus</Emphasis>) populations: shared and unique aspects of diversification

Contemporary patterns of morphological variation among populations reflects the interplay between historic and contemporary processes that result from selection and constraint. Using the pumpkinseed (Lepomis gibbosus), a species native to North America and introduced to Europe, we assessed the shared and unique aspects of morphological divergence in lentic and lotic environments among native and non-native populations. Ten native and thirteen non-native pumpkinseed populations were collected between 2003 and 2010 from lakes, rivers and reservoirs within the Iberian Peninsula and east-central North America. Fifteen linear external measurements among homologous landmarks that pertain to body size, fin position and fin size were taken from all sampled individuals. Eleven of these measurements were used to test for morphological differences among populations. Pumpkinseed found in lotic water bodies exhibited a more anterior placement of pectoral and pelvic fins and a deeper caudal peduncle and body than those found in lentic water bodies from the same geographic region. However, pumpkinseed also showed morphological differences between geographic origins: pumpkinseed from native populations exhibit a more posterior placement of pectoral and pelvic fins, a narrower anterior caudal peduncle and a more slender body than pumpkinseed from non-native populations. In addition, unique responses of populations to waterbodies within geographic origins revealed a shift between water body types that was opposite in direction for native and non-native populations. Native populations exhibited shorter and deeper caudal peduncles and deeper bodies in lotic habitats, whereas non-native populations showed longer and slender caudal peduncles and more slender bodies in the same type of habitat. Our study demonstrates that contemporary patterns of morphological variation among native and non-native pumpkinseed populations can be explained by contemporary selection and/or a common plastic developmental response among water bodies, historical effects related to geographic origin and unique responses of populations to habitats within geographic origin, and that the effects of history and the interaction between history and contemporary habitat were larger than contemporary processes in explaining morphological variation at this large spatial scale.     

Geographic variation in the Japanese white-toothed shrew<Emphasis Type="Italic">Crocidura dsinezumi</Emphasis>

I investigated geographic variation in the Japanese white-toothed shrewCrocidura dsinezumi (Temminck, 1842) (Insectivora, Soricidae) from the Japanese Archipelago and the northern part of the Ryukyu Archipelago. Univariate and multivariate analyses were conducted based on 11 cranial measurements from 14 sampling localities. Overall size variation seemed to follow clinal changes correlated with latitude and longitude of localities, such that northeastern populations were smaller than southwestern ones. The Hokkaido population and the Kuchinoerabujima population were larger than expected based on estimated values, likely due to changes reflecting environmental factors. In contrast, the Tanegashima population was smaller than its estimated value. The northern Ryukyu populations were divergent from the Kyushu population and from each other. Based on patterns of geographic variation, I suggest that all previously described subspecies ofC. dsinezumi are junior synonyms ofC. dsinezumi; and that the Kuchinoerabujima population and the Nakanoshima population likely represent un- described subspecies.     

Craniofacial variation,body size and ecological factors in aboriginal populations from central Patagonia (2000–200 years B.P.)

Previous studies have shown that ecological factors had a significant role in shaping the patterns of craniofacial variation among South American populations. Here, we evaluate whether temperature and diet contributed to facial diversification in small geographic areas. Facial size and shape of 9 osteological samples from central Patagonia (Argentina) were described using 2D landmarks and semilandmarks. Data on mean annual temperature, diet composition (δ¹³C and δ¹⁵N values) and femoral head maximum breadth, used as a proxy of body mass, were obtained for each sample. We then tested the association of body mass and the ecological variables with facial morphology using spatial regression techniques and a model selection approach. Akaike Information Criterion produced disparate results for both components of facial morphology. The best model for facial size included temperature and body mass proxy, and accounted for more than 80% of variation in size. Lower temperatures were related to larger facial sizes. Body mass was negatively associated with facial size and showed no relationship with the temperature. This suggests a relatively independent variation of cranial traits and body mass at the spatial scale studied here. Facial shape was not associated with the temperature or diet composition, contrasting with the patterns observed at larger spatial scales. Our results point out that the effect of climatic variables on cranial traits might be a source of morphological differentiation not only at large scales but also in small geographic areas, and that size and shape display a differential preservation of environmental signals.     

Variation in morphology and functional performance across distinct evolutionary lineages of the Moorish gecko (Tarentola mauritanica) from the Iberian Peninsula

Deciphering the mechanisms that underlie morphological and functional diversity is essential for understanding how organisms adapt to their environment. Interestingly, phenotypic divergence does not necessarily correspond to the geographic and genetic separation between populations. Here, we explored the morphological and functional divergence among populations of two genetically differentiated clades of the Moorish gecko, Tarentola mauritanica. We used linear and geometric morphometrics to quantify morphological variation and investigated how it translates into biting and CLIMBING PERFORMANCE, to better understand the mechanisms potentially underlying population and lineage divergence. We found marked morphological differences between clades, both in body size and head shape. However, much of this differentiation is more strongly related to local variation between populations of the same clade, suggesting that recent ecological events may be more influential than deep evolutionary history in shaping diversity patterns in this group. Despite a lack of association between morphology and functional diversification in the locomotor system of the Moorish gecko, straightforward links are observed between head morphology and biting performance, providing more hints on the possible underlying causes. Indeed, variation in bite force is mostly determined by size variation and sexual dimorphism, and differences between the two clades concern how sexual variation is expressed, reinforcing the idea that both social and ecological factors contribute in shaping differentiation. Interestingly, the individuals from the islets off the coast of Murcia exhibit particular morphological and functional traits, which suggests that the ecological conditions related to insularity may drive the phenotypic differentiation of this population.     

Seeing the forest for the trees: partitioning ecological and phylogenetic components of Bergmann's rule in European Carnivora

Comparative methods have commonly been applied in macroecological research. However, few methods exist to map and analyze phylogenetic variation in geographical space. Here we develop a general analytical framework to partition the phylogenetic and ecological structures of macroecological patterns in geographic space. As an example, we apply the framework to evaluate interspecific patterns of body size geographic variation (Bergmann's rule) in European Carnivora. We model the components of variance attributable to ecological and phylogenetic effects, and to the shared influence of both factors. Spatial patterns in the ecological component are stronger than those in the original body size data. More importantly, the magnitude of intraspecific body size patterns (as measured by the correlation coefficient between body size and latitude) is significantly correlated with the ecological component across species, providing a unified interpretation for Bergmann's rule at multiple levels of biological hierarchy. This approach provides a better understanding of patterns in macroecological traits and allows improved understanding of their underlying ecological and evolutionary mechanisms.     

The effect of seed traits on geographic variation in body size and sexual size dimorphism of the seed‐feeding beetle Acanthoscelides macrophthalmus

Explaining large‐scale patterns of variation in body size has been considered a central question in ecology and evolutionary biology because several life‐history traits are directly linked to body size. For ectothermic organisms, little is known about what processes influence geographic variation in body size. Changes in body size and sexual size dimorphism (SSD) have been associated with environmental variables, particularly for Bruchinae insects, which feed exclusively on seeds during the larval stage. However, the effect of important seed traits on body size variation has rarely been investigated, and whether SSD varies substantially among populations within bruchine species is poorly known. Using the seed‐feeding beetle Acanthoscelides macrophthalmus infesting its host plant Leucaena leucocephala, we investigated whether specific seed traits (hardness, size, water content, carbon/nitrogen ratio, and phenolic content) were determinant in generating geographic variation in body size and SSD of A. macrophthalmus. We also examined the relationships between body size and SSD with latitude and altitude. The body size of both sexes combined was not related to latitude, altitude, and any of the physical and chemical seed traits. However, the female body size tended to vary more in size than the males, generating significant variation in SSD in relation to latitude and altitude. The females were the larger sex at higher latitudes and at lower altitudes, precisely where seed water content was greater. Therefore, our results suggest that water content was the most important seed trait, most severely affecting the females, promoting geographic variation in SSD of A. macrophthalmus.     

Plasticity and genetic effects contribute to different axes of neural divergence in a community of mimetic Heliconius butterflies

Changes in ecological preference, often driven by spatial and temporal variation in resource distribution, can expose populations to environments with divergent information content. This can lead to adaptive changes in the degree to which individuals invest in sensory systems and downstream processes, to optimize behavioural performance in different contexts. At the same time, environmental conditions can produce plastic responses in nervous system development and maturation, providing an alternative route to integrating neural and ecological variation. Here, we explore how these two processes play out across a community of Heliconius butterflies. Heliconius communities exhibit multiple Mullerian mimicry rings, associated with habitat partitioning across environmental gradients. These environmental differences have previously been linked to heritable divergence in brain morphology in parapatric species pairs. They also exhibit a unique dietary adaptation, known as pollen feeding, that relies heavily on learning foraging routes, or trap-lines, between resources, which implies an important environmental influence on behavioural development. By comparing brain morphology across 133 wild-caught and insectary-reared individuals from seven Heliconius species, we find strong evidence for interspecific variation in patterns of neural investment. These largely fall into two distinct patterns of variation; first, we find consistent patterns of divergence in the size of visual brain components across both wild and insectary-reared individuals, suggesting genetically encoded divergence in the visual pathway. Second, we find interspecific differences in mushroom body size, a central component of learning and memory systems, but only among wild caught individuals. The lack of this effect in common-garden individuals suggests an extensive role for developmental plasticity in interspecific variation in the wild. Finally, we illustrate the impact of relatively small-scale spatial effects on mushroom body plasticity by performing experiments altering the cage size and structure experienced by individual H. hecale. Our data provide a comprehensive survey of community level variation in brain structure, and demonstrate that genetic effects and developmental plasticity contribute to different axes of interspecific neural variation.     

History matters more when explaining genetic diversity within the context of the core–periphery hypothesis

The core–periphery hypothesis (CPH) predicts that populations located at the periphery of a species' range should have lower levels of genetic variation than those at the centre of the range. However, most of the research on the CPH focuses on geographic distance and not on ecological distance, or uses categorical definitions of core and periphery to explain the distribution of genetic diversity. We use current climate data and historical climate data from the last glacial maxima to develop quantitative estimates of contemporary and historical ecological suitability using ecological niche models. We analysed genetic diversity using 12 polymorphic microsatellites to estimate changes in heterozygosity, allelic richness and population differentiation in 31 populations of the wood frog (Lithobates sylvaticus) spanning the species’ entire eastern clade (33^o to 45^o latitude) from Alabama, USA, to Nova Scotia, Canada. Our data support predictions based on the CPH. Populations showed significant differences in genetic diversity across the range, with lower levels of genetic variation at the geographic range edge and in areas with lower levels of historical and contemporary ecological suitability. However, history and geography (not current ecological suitability) best explain the patterns. This study highlights the importance of examining more than just geography when assessing the CPH, and the importance of historical ecological suitability in the maintenance of genetic diversity and population differentiation.