Are social media reports useful for assessing small ape occurrence? A pilot study from Peninsular Malaysia

Citizen science-based research has been used effectively to estimate animal abundance and breeding patterns, to monitor animal movement, and for biodiversity conservation and education. Here, we evaluate the feasibility of using social media observations to assess the distribution of small apes in Peninsular Malaysia. We searched for reports of small ape observations in Peninsular Malaysia on social media (e.g., blogs, Facebook, Instagram, Twitter, YouTube, iNaturalist, etc.), and also used online, radio, print messaging, and word of mouth to invite citizen scientists such as birders, amateur naturalists, hikers, and other members of the public to provide information about small ape observations made during their activities. These reports provided new information about the occurrence of all three species of small apes (Hylobates agilis, Hylobates lar, and Symphalangus syndactylus) in Peninsular Malaysia. Social media users reported observations of small apes in almost every state. Despite the fact that small apes are believed to occur primarily in the interior of large forested areas, most observations were from fairly small (<100 km²) forests near areas of high traffic and high human population (roads and urban areas). This suggests that most outdoor enthusiasts primarily visit well-traveled and easily accessible areas, which results in biased sampling if only incidental observations reported on social media are used. A more targeted approach specifically soliciting reports from citizen scientists visiting large, less-accessible forests may result in better sampling in these habitats. Social media reports indicated the presence of small apes in at least six habitats where they had not been previously reported. We verified the reported data based on whether reports included a date, location, and uploaded photographs, videos and/or audio recordings. Well-publicized citizen science programs may also build awareness and enthusiasm about the conservation of vulnerable wildlife species.     

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.     

Hybridization and transgressive exploration of colour pattern and wing morphology in Heliconius butterflies

Hybridization can generate novel phenotypes distinct from those of parental lineages, a phenomenon known as transgressive trait variation. Transgressive phenotypes might negatively or positively affect hybrid fitness, and increase available variation. Closely related species of Heliconius butterflies regularly produce hybrids in nature, and hybridization is thought to play a role in the diversification of novel wing colour patterns despite strong stabilizing selection due to interspecific mimicry. Here, we studied wing phenotypes in first‐ and second‐generation hybrids produced by controlled crosses between either two co‐mimetic species of Heliconius or between two nonmimetic species. We quantified wing size, shape and colour pattern variation and asked whether hybrids displayed transgressive wing phenotypes. Discrete traits underlain by major‐effect loci, such as the presence or absence of colour patches, generate novel phenotypes. For quantitative traits, such as wing shape or subtle colour pattern characters, hybrids only exceed the parental range in specific dimensions of the morphological space. Overall, our study addresses some of the challenges in defining and measuring phenotypic transgression for multivariate traits and our data suggest that the extent to which transgressive trait variation in hybrids contributes to phenotypic diversity depends on the complexity and the genetic architecture of the traits.     

Does thermal plasticity align with local adaptation? An interspecific comparison of wing morphology in sepsid flies

Although genetic and plastic responses are sometimes considered as unrelated processes, their phenotypic effects may often align because genetic adaptation is expected to mirror phenotypic plasticity if adaptive, but run counter to it when maladaptive. Because the magnitude and direction of this alignment has further consequences for both the tempo and mode of adaptation, they are relevant for predicting an organisms’ reaction to environmental change. To better understand the interplay between phenotypic plasticity and genetic change in mediating adaptive phenotypic variation to climate variability, we here quantified genetic latitudinal variation and thermal plasticity in wing loading and wing shape in two closely related and widespread sepsid flies. Common garden rearing of 16 geographical populations reared across multiple temperatures revealed that wing loading decreases with latitude in both species. This pattern could be driven by selection for increased dispersal capacity in the cold. However, although allometry, sexual dimorphism, thermal plasticity and latitudinal differentiation in wing shape all show similar patterns in the two species, the relationship between the plastic and genetic responses differed between them. Although latitudinal differentiation (south to north) mirrored thermal plasticity (hot to cold) in Sepsis punctum, there was no relationship in Sepsis fulgens. While this suggests that thermal plasticity may have helped to mediate local adaptation in S. punctum, it also demonstrates that genetic wing shape differentiation and its relation to thermal plasticity may be complex and idiosyncratic, even among ecologically similar and closely related species. Hence, genetic responses can, but do not necessarily, align with phenotypic plasticity induced by changing environmental selection pressures.     

Melanin coloration has temperature‐dependent effects on breeding performance that may maintain phenotypic variation in a passerine bird

Fluctuating selection pressure may maintain phenotypic variation because of different types of individuals being adapted to different environmental conditions. We show that the extensive variation in the coloration of male pied flycatchers (Ficedula hypoleuca) can be maintained through differences in the reproductive success of male phenotypes under different conditions. The effects of weather conditions on the relative success of different male phenotypes varied between different phases of breeding. The reproductive output of black males was the highest when it was cold during egg‐laying but warm during the nestling period, whereas the fledgling production of brown males was highest when it was continuously warm. In addition, male forehead and wing patch sizes had context‐dependent effects on timing of breeding and nestling mortality, respectively. These results indicate that environmental heterogeneity plays a role in maintaining phenotypic variation. As melanin‐based coloration is heritable, climate change may alter phenotype frequencies depending on the patterns of warming.     

Genetic variation in heat-stress tolerance among South American <Emphasis Type="Italic">Drosophila</Emphasis> populations

Spatial or temporal differences in environmental variables, such as temperature, are ubiquitous in nature and impose stress on organisms. This is especially true for organisms that are isothermal with the environment, such as insects. Understanding the means by which insects respond to temperature and how they will react to novel changes in environmental temperature is important for understanding the adaptive capacity of populations and to predict future trajectories of evolutionary change. The organismal response to heat has been identified as an important environmental variable for insects that can dramatically influence life history characters and geographic range. In the current study we surveyed the amount of variation in heat tolerance among Drosophila melanogaster populations collected at diverse sites along a latitudinal gradient in Argentina (24°–38°S). This is the first study to quantify heat tolerance in South American populations and our work demonstrates that most of the populations surveyed have abundant within-population phenotypic variation, while still exhibiting significant variation among populations. The one exception was the most heat tolerant population that comes from a climate exhibiting the warmest annual mean temperature. All together our results suggest there is abundant genetic variation for heat-tolerance phenotypes within and among natural populations of Drosophila and this variation has likely been shaped by environmental temperature.     

Habitat variation and wing coloration affect wing shape evolution in dragonflies

Habitats are spatially and temporally variable, and organisms must be able to track these changes. One potential mechanism for this is dispersal by flight. Therefore, we would expect flying animals to show adaptations in wing shape related to habitat variation. In this work, we explored variation in wing shape in relation to preferred water body (flowing water or standing water with tolerance for temporary conditions) and landscape (forested to open) using 32 species of dragonflies of the genus Trithemis (80% of the known species). We included a potential source of variation linked to sexual selection: the extent of wing coloration on hindwings. We used geometric morphometric methods for studying wing shape. We also explored the phenotypic correlation of wing shape between the sexes. We found that wing shape showed a phylogenetic structure and therefore also ran phylogenetic independent contrasts. After correcting for the phylogenetic effects, we found (i) no significant effect of water body on wing shape; (ii) male forewings and female hindwings differed with regard to landscape, being progressively broader from forested to open habitats; (iii) hindwings showed a wider base in wings with more coloration, especially in males; and (iv) evidence for phenotypic correlation of wing shape between the sexes across species. Hence, our results suggest that natural and sexual selection are acting partially independently on fore‐ and hindwings and with differences between the sexes, despite evidence for phenotypic correlation of wing shape between males and females.     

Morphometric differentiation across House Sparrow Passer domesticus populations in Finland in comparison with the neutral expectation for divergence

To understand the biology of organisms it is important to take into account the evolutionary forces that have acted on their constituent populations. Neutral genetic variation is often assumed to reflect variation in quantitative traits under selection, though with even low neutral divergence there can be substantial differentiation in quantitative genetic variation associated with locally adapted phenotypes. To study the relative roles of natural selection and genetic drift in shaping phenotypic variation, the levels of quantitative divergence based on phenotypes (P_ST) and neutral genetic divergence (F_ST) can be compared. Such a comparison was made between 10 populations of Finnish House Sparrows (n = 238 individuals) collected in 2009 across the whole country. Phenotypic variation in tarsus‐length, wing‐length, bill‐depth, bill‐length and body mass were considered and 13 polymorphic microsatellite loci were analysed to quantify neutral genetic variation. Calculations of P_ST were based on Markov‐Chain Monte Carlo Bayesian estimates of phenotypic variances across and within populations. The robustness of the conclusions of the P_ST–F_ST comparison was evaluated by varying the proportion of variation due to additive genetic effects within and across populations. Our results suggest that body mass is under directional selection, whereas the divergence in other traits does not differ from neutral expectations. These findings suggest candidate traits for considering gene‐based studies of local adaptation. The recognition of locally adapted populations may be of value in the conservation of this declining species.     

Temperature shapes the costs,benefits and geographic diversification of sexual coloration in a dragonfly

The environment shapes the evolution of secondary sexual traits by determining how their costs and benefits vary across the landscape. Given the thermal properties of dark coloration generally, temperature should crucially influence the costs, benefits and geographic diversification of many secondary sexual colour patterns. We tested this hypothesis using sexually selected wing coloration in a dragonfly. We find that greater wing coloration heats males – the magnitude of which improves flight performance under cool conditions but dramatically reduces it under warm conditions. In a colder region of the species’ range, behavioural observations of a wild population show that these thermal effects translate into greater territorial acquisition on thermally variable days. Finally, geo‐referenced photographs taken by citizen scientists reveal that this sexually selected wing coloration is dramatically reduced in the hottest portions of the species’ range. Collectively, our results underscore temperature's capacity to promote and constrain the evolution of sexual coloration.     

Equivalence of citizen science and scientific data for modelling species distribution of birds from a tropical savanna

The Wallacean deficit continues to be a challenge to species distribution modelling. Although some authors have suggested that data collected by citizen scientists can be relevant for a better understanding of biodiversity, to our knowledge, no work has quantitatively tested the equivalence between scientific and citizen science data. Here, we investigate the hypothesis that data collected by citizen scientists can be equivalent to data collected by professional scientists when generating species spatial distribution models. For 42 bird species in the Cerrado region we generated and compared species distribution models based on three data sources: (1) scientific data, (2) citizen science data and (3) sample size corrected citizen science data. To test our hypothesis, we compared the equivalence of these datasets. We rejected the hypothesis of equivalence for about one-third (38%) of the evaluated species, revealing that, for most of the species considered, the models generated were equivalent irrespective of the data set used. The distances between centroids of the models that were equivalent were on average smaller than the distances between non-equivalent models. Also, the direction of change in the models showed no pattern, with no trend towards more populated regions. Our results show that the use of data collected by citizen scientists can be an ally in filling the Wallacean deficit gap. In fact, the lack of use of this wide range of data collected by citizen scientists seems to be an unjustified caution. We indicate the potential of using citizen science data for modelling the distribution of species, mainly due to the large set of data collected, which is impracticable for scientists alone to collect. Conservation measures will be favoured by the union of professional and amateur data, aiming for a better understanding of species distribution and, consequently, biodiversity conservation.     

Evolution of the male genitalia: morphological variation of the aedeagi in a natural population of <Emphasis Type="Italic">Drosophila mediopunctata</Emphasis>

To investigate the size and shape of the aedeagus of Drosophila mediopunctata, we used basic statistics and geometric morphometrics. We estimated the level of phenotypic variation, natural and laboratory heritability as well as the phenotypic correlations between aedeagus and wing measures. The wing was used as an indicator for both body size and shape. Positive significant correlation was obtained for centroid size of aedeagus and wing for field parents and their offspring reared in the laboratory. Many positive significant phenotypic correlations were found among linear measures of both organs. The phenotypic correlations were few for aedeagus and wing shape. Coefficients of variation of the measures were on average larger in the aedeagus than in the wing for offspring reared in laboratory, but not for flies coming from the field. Significant “natural” heritabilities were found for five linear measures of the aedeagus and only one for the wing. Few significant heritabilities were found for aedeagus and wing shape, mostly ones concerning the uniform components. In an exploratory analysis, we found that inversion DS-PC0 is associated with both uniform and nonuniform components of shape, respectively, in the wing and aedeagus. Our results do not support the lock-and-key hypothesis for the male genitalia evolution, but cannot refute the sexual selection and pleiotropy hypotheses.     

Prey responses to fine‐scale variation in predation risk from combined predators

While it is well documented that organisms can express phenotypic plasticity in response to single gradients of environmental variation, our understanding of how organisms integrate information along multiple environmental gradients is limited in many systems. Using the freshwater snail Helisoma trivolvis and two common predators (water bugs Belostoma flumineum and crayfish Orconectes rusticus), we explored how prey integrate information along multiple predation risk gradients (i.e. caged predators fed increasing amounts of prey biomass) that induce opposing phenotypes. When exposed to single predators fed increasing amounts of prey biomass, we detected threshold responses; intermediate amounts of consumed biomass induced phenotypic responses, but higher amounts induced little additional induction. This suggests that additional increases in predator‐induced traits with greater predator risk offer minimal increases in fitness or that a limit in the response magnitude was reached. Additionally, the response thresholds were contingent on the predator and focal trait. For shell width, responses were generally detected at a lower amount of consumed biomass by water bugs compared to crayfish. Within the crayfish treatments, we found that the shell thickness response threshold was lower than the shell width response threshold. When we combined gradients of consumed biomass from both predators, we found that the magnitude of response to one predator was often reduced when the other predator was present. Interestingly, these effects were often detected at consumed biomass levels that were lower than the threshold concentration necessary to elicit a response in the single‐predator treatments. Moreover, our combined predator treatments revealed that snails shifted from discrete responses to more continuous (i.e. graded) responses. Together, our results reveal that organisms experiencing multiple environmental gradients can integrate this information to make phenotypic decisions and demonstrate the novel result that an exposure to multiple species of predators can lower the response threshold of prey.     

Genotype‐environment interactions rule the response of a widespread butterfly to temperature variation

Understanding how organisms adapt to complex environments is a central goal of evolutionary biology and ecology. This issue is of special interest in the current era of rapidly changing climatic conditions. Here, we investigate clinal variation and plastic responses in life history, morphology and physiology in the butterfly Pieris napi along a pan‐European gradient by exposing butterflies raised in captivity to different temperatures. We found clinal variation in body size, growth rates and concomitant development time, wing aspect ratio, wing melanization and heat tolerance. Individuals from warmer environments were more heat‐tolerant and had less melanised wings and a shorter development, but still they were larger than individuals from cooler environments. These findings suggest selection for rapid growth in the warmth and for wing melanization in the cold, and thus fine‐tuned genetic adaptation to local climates. Irrespective of the origin of butterflies, the effects of higher developmental temperature were largely as expected, speeding up development; reducing body size, potential metabolic activity and wing melanization; while increasing heat tolerance. At least in part, these patterns likely reflect adaptive phenotypic plasticity. In summary, our study revealed pronounced plastic and genetic responses, which may indicate high adaptive capacities in our study organism. Whether this may help such species, though, to deal with current climate change needs further investigation, as clinal patterns have typically evolved over long periods.     

Flight morphology along a latitudinal gradient in a butterfly: do geographic clines differ between agricultural and woodland landscapes?

Bergman and converse Bergman rules, amongst others, describe latitudinal variation in size of organisms, including flying ectotherms like butterflies. However, geographic clines in morphological traits of functional significance for flight performance and thermoregulation may also exist, although they have received less attention within a biogeographical context. Variation in flight‐related morphology has often been studied relative to landscape structure. However, the extent to which landscape effects interact with latitudinal clines of phenotypic variation has rarely been tested. Here we address the effect of latitude, landscape type and the interaction effect on body size and flight‐related morphology in the speckled wood butterfly Pararge aegeria. Male adult butterflies were collected from two replicate populations in each agricultural and woodland landscape types along a 700 km cline in six latitudinal zones. Overall size, adult body mass and wing area increased with latitude in line with Bergmann's rule. Forewing length, however, decreased with latitude. As predicted from thermoregulatory needs in ectotherms, the basal wing part was darker to the north. Latitudinal trends for flight‐related morphological traits were opposite to predictions about flight endurance under cooler conditions that were observed in some non‐lepidopteran insects, i.e. wing loading increased and wing aspect ratio decreased with latitude. Opposite trends can, however, be explained by other aspects of butterfly flight performance (i.e. mate‐location behaviour). As predicted from differences in environmental buffering in woodland landscapes along the latitudinal gradient, significant landscape×latitude interaction effects indicated stronger latitudinal clines and stronger phenotypic variation for size and flight morphology in the agricultural landscape compared to the woodland landscape. In agreement with significant interaction effects, morphological differentiation increased with latitude and was higher between population pairs of agricultural landscape than between population pairs of woodland landscape. These results demonstrate that landscape, latitude and their interaction contribute to the understanding of the complex geographic variation in P. aegeria adult phenotypes across Europe.     

Dynamics of phenotypic change: wing length declines in a resident farmland passerine despite survival advantage of longer wings

In many taxa, environmental changes that alter resource availability and energetics, such as climate change and land use change, are associated with changes in body size. We use wing length as a proxy for overall structural body size to examine a paradoxical trend of declining wing length within a Yellowhammer Emberiza citrinella population sampled over 21 years, in which it has been previously shown that longer wings are associated with higher survival rates. Higher temperatures during the previous winter (prior to the moult determining current wing length) explained 23% of wing length decrease within our population, but changes may also be correlated with non‐climatic environmental variation such as changes in farming mechanisms linked to food availability. We found no evidence for within‐individual wing length shrinkage with age, but our data suggested a progressive decline in the sizes of immature birds recruiting to the population. This trend was weaker, although not significantly so, among adults, suggesting that the decline in the sizes of recruits was offset by higher subsequent survival of larger birds post‐recruitment. These data suggest that ecological processes can contribute more than selection to observed phenotypic trends and highlight the importance of long‐term studies for providing longitudinal insights into population processes.     

feedr and animalnexus.ca: A paired R package and user‐friendly Web application for transforming and visualizing animal movement data from static stations

Radio frequency identification (RFID) provides a simple and inexpensive approach for examining the movements of tagged animals, which can provide information on species behavior and ecology, such as habitat/resource use and social interactions. In addition, tracking animal movements is appealing to naturalists, citizen scientists, and the general public and thus represents a tool for public engagement in science and science education. Although a useful tool, the large amount of data collected using RFID may quickly become overwhelming. Here, we present an R package (feedr) we have developed for loading, transforming, and visualizing time‐stamped, georeferenced data, such as RFID data collected from static logger stations. Using our package, data can be transformed from raw RFID data to visits, presence (regular detections by a logger over time), movements between loggers, displacements, and activity patterns. In addition, we provide several conversion functions to allow users to format data for use in functions from other complementary R packages. Data can also be visualized through static or interactive maps or as animations over time. To increase accessibility, data can be transformed and visualized either through R directly, or through the companion site: http://animalnexus.ca , an online, user‐friendly, R‐based Shiny Web application. This system can be used by professional and citizen scientists alike to view and study animal movements. We have designed this package to be flexible and to be able to handle data collected from other stationary sources (e.g., hair traps, static very high frequency (VHF) telemetry loggers, observations of marked individuals in colonies or staging sites), and we hope this framework will become a meeting point for science, education, and community awareness of the movements of animals. We aim to inspire citizen engagement while simultaneously enabling robust scientific analysis.     

Wing trait–inversion associations in Drosophila subobscura can be generalized within continents,but may change through time

Clinal variation is one of the most emblematic examples of the action of natural selection at a wide geographical range. In Drosophila subobscura, parallel clines in body size and inversions, but not in wing shape, were found in Europe and South and North America. Previous work has shown that a bottleneck effect might be largely responsible for differences in wing trait–inversion association between one European and one South American population. One question still unaddressed is whether the associations found before are present across other populations of the European and South American clines. Another open question is whether evolutionary dynamics in a new environment can lead to relevant changes in wing traits–inversion association. To analyse geographical variation in these associations, we characterized three recently laboratory founded D. subobscura populations from both the European and South American latitudinal clines. To address temporal variation, we also characterized the association at a later generation in the European populations. We found that wing size and shape associations can be generalized across populations of the same continent, but may change through time for wing size. The observed temporal changes are probably due to changes in the genetic content of inversions, derived from adaptation to the new, laboratory environment. Finally, we show that it is not possible to predict clinal variation from intrapopulation associations. All in all this suggests that, at least in the present, wing traits–inversion associations are not responsible for the maintenance of the latitudinal clines in wing shape and size.     

Evolution along the Great Rift Valley: phenotypic and genetic differentiation of East African white‐eyes (Aves,Zosteropidae)

The moist and cool cloud forests of East Africa represent a network of isolated habitats that are separated by dry and warm lowland savannah, offering an opportunity to investigate how strikingly different selective regimes affect species diversification. Here, we used the passerine genus Zosterops (white‐eyes) from this region as our model system. Species of the genus occur in contrasting distribution settings, with geographical mountain isolation driving diversification, and savannah interconnectivity preventing differentiation. We analyze (1) patterns of phenotypic and genetic differentiation in high‐ and lowland species (different distribution settings), (2) investigate the potential effects of natural selection and temporal and spatial isolation (evolutionary drivers), and (3) critically review the taxonomy of this species complex. We found strong phenotypic and genetic differentiation among and within the three focal species, both in the highland species complex and in the lowland taxa. Altitude was a stronger predictor of phenotypic patterns than the current taxonomic classification. We found longitudinal and latitudinal phenotypic gradients for all three species. Furthermore, wing length and body weight were significantly correlated with altitude and habitat type in the highland species Z. poliogaster. Genetic and phenotypic divergence showed contrasting inter‐ and intraspecific structures. We suggest that the evolution of phenotypic characters is mainly driven by natural selection due to differences in the two macro‐habitats, cloud forest and savannah. In contrast, patterns of neutral genetic variation appear to be rather driven by geographical isolation of the respective mountain massifs. Populations of the Z. poliogaster complex, as well as Z. senegalensis and Z. abyssinicus, are not monophyletic based on microsatellite data and have higher levels of intraspecific differentiation compared to the currently accepted species.     

Beyond the classroom

After school, college or university, adult education in Britain continues in many spheres. The Field Studies Council provides opportunities for a wide scope of amateur and professional naturalists to learn about various aspects of ecology through first hand experience in the field with like-minded people. The FSC is internationally recognized as the pioneer in environmental education and is still at the forefront of ecological thought. Courses are run at a variety of levels on a wide range of subjects. Within our full programme, there should be something for all at home or overseas.     

Seasonal wing colour plasticity varies dramatically between buckeye butterfly populations in different climatic zones

1. In the present study, how seasonal wing colour plasticity in buckeye butterflies (Junonia coenia Hübner) differs between populations in different climatic regions, and whether these differences are explained by regional environmental differences or reflect genetic divergences in plasticity is addressed. 2. Using museum specimen data, it is shown that buckeye wing colour variation is much greater in populations from the humid and subtropical climates of the eastern U.S.A. than in populations from the desert and mediterranean climates of the western U.S.A. 3. Museum specimen data further show that wing colour in eastern populations is strongly correlated with seasonal variation in day length and temperature, whereas wing colour in western populations is only weakly associated with these cues. 4. Controlled incubator experiments comparing Southern California and North Carolina populations suggest that regional differences in wing pattern variation are attributable to dramatically increased robustness to environmental variation in western populations. 5. Together these experiments show that while phenotypic variation can be influenced by environmental cues, the range of this variation can genetically diverge between populations in different regions.