Rapid differentiation of plasticity in life history and morphology during invasive range expansion and concurrent local adaptation in the horned beetle Onthophagus taurus

Understanding the interplay between genetic differentiation, ancestral plasticity, and the evolution of plasticity during adaptation to environmental variation is critical to predict populations’ responses to environmental change. However, the role of plasticity in rapid adaptation in nature remains poorly understood. We here use the invasion of the horned beetle Onthophagus taurus in the United States during the last half century to study the contribution of ancestral plasticity and post-invasion evolution of plastic responses in rapid population differentiation. We document latitudinal variation in life history and morphology, including genetic compensation in development time and body size, likely adaptive responses to seasonal constraints in the North. However, clinal variation in development time and size was strongly dependent on rearing temperature, suggesting that population differentiation in plasticity played a critical role in successful adaptation on ecological timescales. Clinal variation in wing shape was independent of ancestral plasticity, but correlated with derived plasticity, consistent with evolutionary interdependence. In contrast, clinal variation in tibia shape aligned poorly with thermal plasticity. Overall, this study suggests that post-invasion evolution of plasticity contributed to range expansions and concurrent adaptation to novel climatic conditions.     

Random Amplified Polymorphic DNA Diversity among Surface and Subterranean Populations of Asellus aquaticus (Crustacea: Isopoda)

The ecological and evolutionary processes leading to isolation and adaptation of cave animals compared to their surface ancestors are not yet unequivocally understood. In this study the genetic relations of four cave and three surface population of the freshwater crustacean Asellus aquaticus in the Karst region of SW Slovenia and NE Italy were assessed using RAPDs as genetic markers. The results suggest that specialized populations from two caves invaded their subterranean habitat independently, and that their morphological similarity is a result of convergent evolution. Another, less specialized cave population seems to originate from a later colonization of a cave system already inhabited by a more specialized population, but the two populations do not interbreed. This series of temporally and spatially independent invasions has generated a diversity hotspot of non-interbreeding populations of a ubiquitous freshwatercrustacean, which is uniform over most of its range. Genetic variability estimated by the percentage of polymorphic RAPD fragments was similar (86–91%) in most cave and surface populations. Substantially lower values (as low as 49%) were found in two cave populations affected by heavy pollution. Two a priori groupings of populations, traditional subspecies and hydrologically connected groups, were rejected as not significant by means of nested analysis of molecular variance (AMOVA). On the other hand, groupings revealed by UPGMA clustering displayed a significant component of among-group variance. An analysis of gene flow between populations using estimated migration rates confirmed these findings.     

Adaptation and plasticity in insular evolution of the house mouse mandible

Morphometric methods allow the quantification of directions of phenotypic changes and their statistical comparison in a morphometric space. We applied this approach to investigate several candidate factors to explain changes in mandible shape occurring in house mice (Mus musculus domesticus, Mammalia, Rodentia) in Corsica and a nearby islet. The role of niche widening and of the concomitant change in diet was evaluated by comparing the micro‐evolutionary insular change to the macro‐evolutionary difference between omnivorous and herbivorous rodents. Phenotypic plasticity, which may contribute to rapid insular evolution, was assessed by breeding laboratory mice on hard versus soft food. The related change in mandible shape was compared with differences between continental and insular populations. The role of allometry was evaluated by assessing shape change related to size within the continental population and comparing this direction of change with differences on islands. Finally, evolution may be facilitated along the direction of the greatest phenotypic variance. This hypothesis was tested by computing in wild populations vectors corresponding to this direction and by comparing these vectors with those corresponding to estimates of shape changes related to plasticity, micro‐ and macro‐evolutionary processes. In Corsica, the congruence in directions of macro‐ and micro‐evolutionary phenotypic vectors (Corsican/continental mice versus omnivorous/herbivorous rodents) supports the hypothesis of adaptation in mandible shape evolution. By contrast, on the islet, phenotypic divergence follows directions of plastic response to food consistency as well as within‐population allometry. Thus, results suggest differences in the relative importance of processes which may influence rodent mandibular shape depending on the size of the islands they colonized. Faster evolution and plasticity may be more evident in small and often ephemeral populations living on small islands, whereas micro‐evolutionary processes may have enough time and genetic variability to progressively ‘align’ with macro‐evolutionary trends in large populations from big islands.     

Loci under selection during multiple range expansions of an invasive plant are mostly population specific,but patterns are associated with climate

Identifying the genes underlying rapid evolutionary changes, describing their function and ascertaining the environmental pressures that determine fitness are the central elements needed for understanding of evolutionary processes and phenotypic changes that improve the fitness of populations. It has been hypothesized that rapid adaptive changes in new environments may contribute to the rapid spread and success of invasive plants and animals. As yet, studies of adaptation during invasion are scarce, as is knowledge of the genes underlying adaptation, especially in multiple replicated invasions. Here, we quantified how genotype frequencies change during invasions, resulting in rapid evolution of naturalized populations. We used six fully replicated common garden experiments in Brazil where Pinus taeda (loblolly pine) was introduced at the same time, in the same numbers, from the same seed sources, and has formed naturalized populations expanding outward from the plantations. We used a combination of nonparametric, population genetics and multivariate statistics to detect changes in genotype frequencies along each of the six naturalization gradients and their association with climate as well as shifts in allele frequencies compared to the source populations. Results show 25 genes with significant shifts in genotype frequencies. Six genes had shifts in more than one population. Climate explained 25% of the variation in the groups of genes under selection across all locations, but specific genes under strong selection during invasions did not show climate‐related convergence. In conclusion, we detected rapid evolutionary changes during invasive range expansions, but the particular gene‐level patterns of evolution may be population specific.     

Genetic structure of lake and stream populations in a Pyrenean amphibian (Calotriton asper) reveals evolutionary significant units associated with paedomorphosis

Differences in environmental conditions such as those between lakes and streams can produce phenotypic variation and ultimately promote evolutionary diversification. Some species of newts and salamanders can occupy these habitats and express alternative phenotypes: metamorphs that lose gills at metamorphosis and paedomorphs that retain them at the adult stage. Whereas this process is facultative in some species, it is obligatory in others, thus suggesting that isolation and environmental pressures may have canalized developmental pathways. In this study, we focused our research on the Pyrenean brook newt, Calotriton asper, which is present in both lakes and streams, but whose fully aquatic paedomorphic individuals are only present in lakes. We aimed to determine the genetic structure and differentiation of two paedomorphic populations, including their surrounding stream and lake metamorphic populations, to test whether populations of paedomorphs can constitute evolutionary significant units. Although gene flow was identified between lakes and nearby stream populations, there was a low percentage of dispersers, and the paedomorphic populations were genetically differentiated from the populations of metamorphs. It is likely that the studied lakes have offered peculiar conditions that have allowed the development of a paedomorphic phenotype. These populations and phenotypes therefore constitute good models to understand local adaptations. As each of these populations of paedomorphs can be considered evolutionary significant units that cannot be replaced by other nearby populations in case of a population crash, conservation actions should be focused directly on them.     

Differing evolutionary patterns underlie convergence on elongate morphology in endemic fishes of Lake Waccamaw, North Carolina

An extensive body of research has recently demonstrated patterns of parallel and/or convergent evolution that arise from divergent natural selection pressures exerted across environmental gradients. These studies, although providing some of our best empirical evidence for natural selection, have focused on rather narrow phylogenetic scopes, more often than not comparing patterns of morphological change among closely‐related taxa within a single genus. Organisms in replicated populations in these studies are often assumed to have accomplished convergence via similar underlying processes. However, such assumptions cannot be made when looking at evolution across broader phylogenetic and ecological spectra. In the present study, we assessed morphological change across a much broader scale to test whether similar evolutionary and developmental patterns underlie convergence. Specifically, we studied morphological change that has occurred in a novel lake environment (Lake Waccamaw, North Carolina, USA) where three phylogenetically‐disparate fishes representing different orders have speciated and independently evolved streamlined morphologies relative to their deeper‐bodied progenitors occupying nearby streams and coastal regions. Geometric morphometric analyses revealed that, although the bulk of shape change between environments is similar across taxa, significant species‐specific responses, concordant with differing expectations based on the ecologies of these taxa, were also found. Moreover, allometry analyses indicated that the developmental patterns underlying this change also differ across taxa. The present study provides evidence that, within a common environment, convergence can be achieved by different evolutionary and developmental patterns in phylogenetically‐ and ecologically‐disparate taxa. Finally, these results contradict the commonly‐held hypothesis that fishes should be more streamlined in streams than lakes and emphasize the need to also consider other environmental characteristics, such as water clarity and physical complexity, in studies of divergence. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 636–645.     

Geographical variation in growth form traits in Quercus suber and its relation to population evolutionary history

Differential selection pressures caused by environmental disparities lead to populations to become differentiated as they adapt to local environments. In addition, natural selection during the species past can contribute to the observed differentiation. In this study, we examine the geographic variation in a set of four traits related to growth and plant architecture in cork oak (Quercus suber) and investigate to what extent this variation is the result of the effects of ongoing evolution in current environments and the past evolutionary history of the species. Cork oak saplings at the common garden trial exhibited differences in plant architecture associated to cpDNA lineage. Eastern lineages, exhibited the lowest apical dominance and highest branchiness, consistent with the analyses in other cork oak trials. In contrast, patterns linked to the evolutionary past were less evident in height and diameter. These results suggest that selective pressures after cpDNA divergence can have blurred patterns in some traits closely related to fitness, while conserving the past evolutionary imprints in plant architectural traits. Introgressed populations did not show significant differentiation in architecture, which suggests that allele exchanges via hybridization have had a limited effect on population differentiation in cork oak. Finally, populations within lineages also showed differences in growth and architecture. Correlation between population architecture and temperature patterns were observed indicating that environmental factors such as climate also could result crucial in the evolution of plant architecture of cork oak within lineages.     

Seasonal variation in life history traits in two Drosophila species

Seasonal environmental heterogeneity is cyclic, persistent and geographically widespread. In species that reproduce multiple times annually, environmental changes across seasonal time may create different selection regimes that may shape the population ecology and life history adaptation in these species. Here, we investigate how two closely related species of Drosophila in a temperate orchard respond to environmental changes across seasonal time. Natural populations of Drosophila melanogaster and Drosophila simulans were sampled at four timepoints from June through November to assess seasonal change in fundamental aspects of population dynamics as well as life history traits. D. melanogaster exhibit pronounced change across seasonal time: early in the season, the population is inferred to be uniformly young and potentially represents the early generation following overwintering survivorship. D. melanogaster isofemale lines derived from the early population and reared in a common garden are characterized by high tolerance to a variety of stressors as well as a fast rate of development in the laboratory environment that declines across seasonal time. In contrast, wild D. simulans populations were inferred to be consistently heterogeneous in age distribution across seasonal collections; only starvation tolerance changed predictably over seasonal time in a parallel manner as in D. melanogaster. These results suggest fundamental differences in population and evolutionary dynamics between these two taxa associated with seasonal heterogeneity in environmental parameters and associated selection pressures.     

Rapid,nonparallel genomic evolution of Brassica rapa (field mustard) under experimental drought

While we know that climate change can potentially cause rapid phenotypic evolution, our understanding of the genetic basis and degree of genetic parallelism of rapid evolutionary responses to climate change is limited. In this study, we combined the resurrection approach with an evolve-and-resequence design to examine genome-wide evolutionary changes following drought. We exposed genetically similar replicate populations of the annual plant Brassica rapa derived from a field population in southern California to four generations of experimental drought or watered conditions in a greenhouse. Genome-wide sequencing of ancestral and descendant population pools identified hundreds of SNPs that showed evidence of rapidly evolving in response to drought. Several of these were in stress response genes, and two were identified in a prior study of drought response in this species. However, almost all genetic changes were unique among experimental populations, indicating that the evolutionary changes were largely nonparallel, despite the fact that genetically similar replicates of the same founder population had experienced controlled and consistent selection regimes. This nonparallelism of evolution at the genetic level is potentially because of polygenetic adaptation allowing for multiple different genetic routes to similar phenotypic outcomes. Our findings help to elucidate the relationship between rapid phenotypic and genomic evolution and shed light on the degree of parallelism and predictability of genomic evolution to environmental change.     

A new and morphologically distinct population of cavernicolous Poecilia mexicana (Poeciliidae: Teleostei)

The cave molly, Poecilia mexicana, from the Cueva del Azufre, a sulfur cave in Tabasco, Mexico, ranks among the best-studied cave fishes worldwide, despite being known from a single population only. Here we describe a newly discovered second population of cave-dwelling P. mexicana from a nearby, but mostly non-sulfidic cave (Luna Azufre). Despite apparent similarities between the two populations (such as reduced eye diameter and reduced pigmentation), a geometric morphometric analysis revealed pronounced morphological differentiation between the two cave forms.     

THE EVOLUTION OF FEMALE-BIASED SEXUAL SIZE DIMORPHISM: A POPULATION-LEVEL COMPARATIVE STUDY IN HORNED LIZARDS (PHRYNOSOMA)

Female-biased sexual size dimorphism is uncommon among vertebrates and traditionally has been attributed to asymmetric selective pressures favoring large fecund females (the fecundity-advantage hypothesis) and/or small mobile males (the small-male advantage hypothesis). I use a phylogenetically based comparative method to address these hypotheses for the evolution and maintenance of sexual size dimorphism among populations of three closely related lizard species (Phrynosoma douglasi, P. ditmarsi, and P. hernandezi). With independent contrasts I estimate evolutionary correlations among female body size, male body size, and sexual size dimorphism (SSD) to determine whether males have become small, females have become large, or both sexes have diverged concurrently in body size during the evolutionary Xhistory of this group. Population differences in degree of SSD are inversely correlated with average male body size, but are not correlated with average female body size. Thus, variation in SSD among populations has occurred predominantly through changes in male size, suggesting that selective pressures on small males may affect degree of SSD in this group. I explore three possible evolutionary mechanisms by which the mean male body size in a population could evolve: changes in size at maturity, changes in the variance of male body sizes, and changes in skewness of male body size distributions. Comparative analyses indicate that population differentiation in male body size is achieved by changes in male size at maturity, without changes in the variance or skewness of male and female size distributions. This study demonstrates the potential of comparative methods at lower taxonomic levels (among populations and closely related species) for studying microevolutionary processes that underlie population differentiation.     

Evolutionary responses to global change: lessons from invasive species

Biologists have recently devoted increasing attention to the role of rapid evolution in species' responses to environmental change. However, it is still unclear what evolutionary responses should be expected, at what rates, and whether evolution will save populations at risk of extinction. The potential of biological invasions to provide useful insights has barely been realised, despite the close analogies to species responding to global change, particularly climate change; in both cases, populations encounter novel climatic and biotic selection pressures, with expected evolutionary responses occurring over similar timescales. However, the analogy is not perfect, and invasive species are perhaps best used as an upper bound on expected change. In this article, we review what invasive species can and cannot teach us about likely evolutionary responses to global change and the constraints on those responses. We also discuss the limitations of invasive species as a model and outline directions for future research.     

Standing genetic variation as a potential mechanism of novel cave phenotype evolution in the freshwater isopod,Asellus aquaticus

Novel phenotypes can come about through a variety of mechanisms including standing genetic variation from a founding population. Cave animals are an excellent system in which to study the evolution of novel phenotypes such as loss of pigmentation and eyes. Asellus aquaticus is a freshwater isopod crustacean found in Europe and has both a surface and a cave ecomorph which vary in multiple phenotypic traits. An orange eye phenotype was previously revealed by F₂ crosses and backcrosses to the cave parent within two examined Slovenian cave populations. Complete loss of pigmentation, both in eye and body, is epistatic to the orange eye phenotype and therefore the orange eye phenotype is hidden within the cave populations. Our goal was to investigate the origin of the orange eye alleles within the Slovenian cave populations by examining A. aquaticus individuals from Slovenian and Romanian surface populations and Asellus aquaticus infernus individuals from a Romanian cave population. We found orange eye individuals present in lab raised surface populations of A. aquaticus from both Slovenia and Romania. Using a mapping approach with crosses between individuals of two surface populations, we found that the region known to be responsible for the orange eye phenotype within the two previously examined Slovenian cave populations was also responsible within both the Slovenian and the Romanian surface populations. Complementation crosses between orange eye Slovenian and orange eye Romanian surface individuals suggest that the same gene is responsible for the orange eye phenotype in both surface populations. Additionally, we observed a low frequency phenotype of eye loss in crosses generated between the two surface populations and also in the Romanian surface population. Finally, in a cave population from Romania, A. aquaticus infernus, we found that the same region is also responsible for the orange eye phenotype as the Slovenian cave populations and the Slovenian and Romanian surface populations. Therefore, we present evidence that variation present in the cave populations could originate from standing variation present in the surface populations and/or transgressive hybridization of different surface phylogenetic lineages rather than de novo mutations.     

Dispersal evolution diminishes the negative density dependence in dispersal

In many organisms, dispersal varies with the local population density. Such patterns of density-dependent dispersal (DDD) are expected to shape the dynamics, spatial spread, and invasiveness of populations. Despite their ecological importance, empirical evidence for the evolution of DDD patterns remains extremely scarce. This is especially relevant because rapid evolution of dispersal traits has now been empirically confirmed in several taxa. Changes in DDD of dispersing populations could help clarify not only the role of DDD in dispersal evolution, but also the possible pattern of subsequent range expansion. Here, we investigate the relationship between dispersal evolution and DDD using a long-term experimental evolution study on Drosophila melanogaster. We compared the DDD patterns of four dispersal-selected populations and their non-selected controls. The control populations showed negative DDD, which was stronger in females than in males. In contrast, the dispersal-selected populations showed DDD, where neither males nor females exhibited DDD. We compare our results with previous evolutionary predictions that focused largely on positive DDD, and highlight how the direction of evolutionary change depends on the initial DDD pattern of a population. Finally, we discuss the implications of DDD evolution for spatial ecology and evolution.     

Origins and Relationship of Cave Populations of the Blind Mexican Tetra, Astyanax Fasciatus, in the Sierra De El Abra

The blind morph of Astyanax fasciatus (Pisces: Characidae) has been more thoroughly studied than any other cave inhabiting organism. Most studies of A. fasciatus have used individuals from different caves of the Sierra de El Abra, Mexico, and have assumed that each population independently evolved to live in the cave environment. We analyzed the relationships among several cave populations that delineate the Sierra de El Abra using RAPD markers. The results indicate that all cave populations are more closely related to one another than they are to the surface populations. This suggests that present day cave populations derived from a common ancestral stock, most likely due to a single colonization event, or alternatively, that strong gene flow among cave populations has occurred, raising precaution against considering each cave population as independent.     

Forests are not immune to plant invasions: phenotypic plasticity and local adaptation allow <Emphasis Type="Italic">Prunella vulgaris</Emphasis> to colonize a temperate evergreen rainforest

In the South American temperate evergreen rainforest (Valdivian forest), invasive plants are mainly restricted to open sites, being rare in the shaded understory. This is consistent with the notion of closed-canopy forests as communities relatively resistant to plant invasions. However, alien plants able to develop shade tolerance could be a threat to this unique forest. Phenotypic plasticity and local adaptation are two mechanisms enhancing invasiveness. Phenotypic plasticity can promote local adaptation by facilitating the establishment and persistence of invasive species in novel environments. We investigated the role of these processes in the recent colonization of Valdivian forest understory by the perennial alien herb Prunella vulgaris from nearby populations in open sites. Using reciprocal transplants, we found local adaptation between populations. Field data showed that the shade environment selected for taller plants and greater specific leaf areas. We found population differentiation and within-population genetic variation in both mean values and reaction norms to light variation of several ecophysiological traits in common gardens from seeds collected in sun and shade populations. The colonization of the forest resulted in a reduction of plastic responses to light variation, which is consistent with the occurrence of genetic assimilation and suggests that P. vulgaris individuals adapted to the shade have reduced probabilities to return to open sites. All results taken together confirm the potential for rapid evolution of shade tolerance in P. vulgaris and suggest that this alien species may pose a threat to the native understory flora of Valdivian forest.     

Spatial variation in the evolutionary potential and constraints of basal metabolic rate and body mass in a wild bird

An organism's energy budget is strongly related to resource consumption, performance, and fitness. Hence, understanding the evolution of key energetic traits, such as basal metabolic rate (BMR), in natural populations is central for understanding life-history evolution and ecological processes. Here we used quantitative genetic analyses to study evolutionary potential of BMR in two insular populations of the house sparrow (Passer domesticus). We obtained measurements of BMR and body mass (M_b) from 911 house sparrows on the islands of Leka and Vega along the coast of Norway. These two populations were the source populations for translocations to create an additional third, admixed ‘common garden’ population in 2012. With the use of a novel genetic group animal model concomitant with a genetically determined pedigree, we differentiate genetic and environmental sources of variation, thereby providing insight into the effects of spatial population structure on evolutionary potential. We found that the evolutionary potential of BMR was similar in the two source populations, whereas the Vega population had a somewhat higher evolutionary potential of M_b than the Leka population. BMR was genetically correlated with M_b in both populations, and the conditional evolutionary potential of BMR (independent of body mass) was 41% (Leka) and 53% (Vega) lower than unconditional estimates. Overall, our results show that there is potential for BMR to evolve independently of M_b, but that selection on BMR and/or M_b may have different evolutionary consequences in different populations of the same species.     

Tempo and mode of evolutionary divergence in modern and Holocene Vancouver Island marmots (Marmota vancouverensis) (Mammalia, Rodentia)

Marmota vancouverensis is the only insular species among the 14 species of marmots. The evolutionary history of this species is unresolved. Although M. vancouverensis is strongly differentiated in osteological and other morphological characters, its low genetic divergence suggests recent evolution from an ancestral continental species. We used geometric morphometric techniques to assess the morphology of hemimandibles from 239 modern M. vancouverensis , Marmota caligata , Marmota flaviventris , Marmota olympus and 30 Holocene (9435–735 cal. yr bp) subfossil M. vancouverensis . Our results confirm that the mandible of M. vancouverensis is strongly differentiated in shape from continental marmot species, but similar in size to its mainland sister species M. caligata . Temporal variation in size and shape over the past 2500 years among allochronic samples of M. vancouverensis was minimal suggesting that the morphological divergence of this species occurred in a period of rapid change following its isolation from mainland populations in the late Pleistocene. Selection pressures associated with environmental changes and founder effects and genetic drift resulting from population bottlenecks created by population declines and habitat fragmentation are hypothesized as factors contributing to the morphological divergence of this species.     

Rapid evolutionary adaptation to elevated salt concentrations in pathogenic freshwater bacteria Serratia marcescens

Rapid evolutionary adaptions to new and previously detrimental environmental conditions can increase the risk of invasion by novel pathogens. We tested this hypothesis with a 133‐day‐long evolutionary experiment studying the evolution of the pathogenic Serratia marcescens bacterium at salinity niche boundary and in fluctuating conditions. We found that S. marcescens evolved at harsh (80 g/L) and extreme (100 g/L) salt conditions had clearly improved salt tolerance than those evolved in the other three treatments (ancestral conditions, nonsaline conditions, and fluctuating salt conditions). Evolutionary theories suggest that fastest evolutionary changes could be observed in intermediate selection pressures. Therefore, we originally hypothesized that extreme conditions, such as our 100 g/L salinity treatment, could lead to slower adaptation due to low population sizes. However, no evolutionary differences were observed between populations evolved in harsh and extreme conditions. This suggests that in the study presented here, low population sizes did not prevent evolution in the long run. On the whole, the adaptive potential observed here could be important for the transition of pathogenic S. marcescens bacteria from human‐impacted freshwater environments, such as wastewater treatment plants, to marine habitats, where they are known to infect and kill corals (e.g., through white pox disease).     

The evolutionary consequences of habitat fragmentation: Body morphology and coloration differentiation among brook trout populations of varying size

A reduction in population size due to habitat fragmentation can alter the relative roles of different evolutionary mechanisms in phenotypic trait differentiation. While deterministic (selection) and stochastic (genetic drift) mechanisms are expected to affect trait evolution, genetic drift may be more important than selection in small populations. We examined relationships between mature adult traits and ecological (abiotic and biotic) variables among 14 populations of brook trout. These naturally fragmented populations have shared ancestry but currently exhibit considerable variability in habitat characteristics and population size (49 < N_c < 10,032; 3 < N_b < 567). Body size, shape, and coloration differed among populations, with a tendency for more variation among small populations in both trait means and CV when compared to large populations. Phenotypic differences were more frequently and directly linked to habitat variation or operational sex ratio than to population size, suggesting that selection may overcome genetic drift at small population size. Phenotype–environment associations were also stronger in females than males, suggesting that natural selection due to abiotic conditions may act more strongly on females than males. Our results suggest that natural and sexual‐selective pressures on phenotypic traits change during the process of habitat fragmentation, and that these changes are largely contingent upon existing habitat conditions within isolated fragments. Our study provides an improved understanding of the ecological and evolutionary consequences of habitat fragmentation and lends insight into the ability of some small populations to respond to selection and environmental change.