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Adaptive Divergence (adaptive + divergence)
Selected AbstractsRAPID ADAPTIVE DIVERGENCE IN NEW WORLD ACHILLEA, AN AUTOPOLYPLOID COMPLEX OF ECOLOGICAL RACESEVOLUTION, Issue 3 2008Justin Ramsey Adaptive evolution is often associated with speciation. In plants, however, ecotypic differentiation is common within widespread species, suggesting that climatic and edaphic specialization can outpace cladogenesis and the evolution of postzygotic reproductive isolation. We used cpDNA sequence (5 noncoding regions, 3.5 kb) and amplified fragment length polymorphisms (AFLPs: 4 primer pairs, 1013 loci) to evaluate the history of ecological differentiation in the North American Achillea millefolium, an autopolyploid complex of "ecological races" exhibiting morphological, physiological, and life-history adaptations to diverse environments. Phylogenetic analyses reveal North American A. millefolium to be a monophyletic group distinct from its European and Asian relatives. Based on patterns of sequence divergence, as well as fossil and paleoecological data, colonization of North America appears to have occurred via the Bering Land Bridge during the Pleistocene (1.8 MYA to 11,500 years ago). Population genetic analyses indicate negligible structure within North American A. millefolium associated with varietal identity, geographic distribution, or ploidy level. North American populations, moreover, exhibit the signature of demographic expansion. These results affirm the "ecotype" concept of the North American Achillea advocated by classical research and demonstrate the rapid rate of ecological differentiation that sometimes occurs in plants. [source] REPLICATED EVOLUTION OF INTEGRATED PLASTIC RESPONSES DURING EARLY ADAPTIVE DIVERGENCEEVOLUTION, Issue 4 2006Kevin J. Parsons Abstract Colonization of a novel environment is expected to result in adaptive divergence from the ancestral population when selection favors a new phenotypic optimum. Local adaptation in the new environment occurs through the accumulation and integration of character states that positively affect fitness. The role played by plastic traits in adaptation to a novel environment has generally been ignored, except for variable environments. We propose that if conditions in a relatively stable but novel environment induce phenotypically plastic responses in many traits, and if genetic variation exists in the form of those responses, then selection may initially favor the accumulation and integration of functionally useful plastic responses. Early divergence between ancestral and colonist forms will then occur with respect to their plastic responses across the gradient bounded by ancestral and novel environmental conditions. To test this, we compared the magnitude, integration, and pattern of plastic character responses in external body form induced by shallow versus open water conditions between two sunfish ecomorphs that coexist in four postglacial lakes. The novel sunfish ecomorph is present in the deeper open water habitat, whereas the ancestral ecomorph inhabits the shallow waters along the lake margin. Plastic responses by open water ecomorphs were more correlated than those of their local shallow water ecomorph in two of the populations, whereas equal levels of correlated plastic character responses occurred between ecomorphs in the other two populations. Small but persistent differences occurred between ecomorph pairs in the pattern of their character responses, suggesting a recent divergence. Open water ecomorphs shared some similarities in the covariance among plastic responses to rearing environment. Replication in the form of correlated plastic responses among populations of open water ecomorphs suggests that plastic character states may evolve under selection. Variation between ecomorphs and among lake populations in the covariance of plastic responses suggests the presence of genetic variation in plastic character responses. In three populations, open water ecomorphs also exhibited larger plastic responses to the environmental gradient than the local shallow water ecomorph. This could account for the greater integration of plastic responses in open water ecomorphs in two of the populations. This suggests that the plastic responses of local sunfish ecomorphs can diverge through changes in the magnitude and coordination of plastic responses. Although these results require further investigation, they suggest that early adaptive evolution in a novel environment can include changes to plastic character states. The genetic assimilation of coordinated plastic responses could result in the further, and possibly rapid, divergence of such populations and could also account for the evolution of genes of major effect that contribute to suites of phenotypic differences between divergent populations. [source] Phenotypic evolution in high-elevation populations of western fence lizards (Sceloporus occidentalis) in the Sierra Nevada MountainsBIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY, Issue 3 2010ADAM D. LEACHÉ Adaptive divergence in response to variable habitats, climates, and altitude is often accentuated along elevation gradients. We investigate phenotypic evolution in body size and coloration in the western fence lizard (Sceloporus occidentalis Baird & Girard, 1852) across elevation gradients in Yosemite National Park, California, situated in the Sierra Nevada mountains of Western North America. High-elevation populations occurring above 2100 m a.s.l. are recognized as a separate subspecies (Sceloporus occidentalis taylori Camp, 1916), with a distinctive phenotype characterized by a large body size and extensive blue ventral pigmentation. We sampled S. occidentalis from across elevation gradients in Yosemite National Park, California, and collected phenotypic data (body size and ventral coloration measurements; 410 specimens) and mitochondrial DNA sequence data (complete NADH1 gene; 969 bp, 181 specimens) to infer phylogenetic relationships, and examine the genetic and phenotypic diversity among populations. Populations of S. occidentalis in Yosemite National Park follow Bergmann's rule and exhibit larger body sizes in colder, high-elevation environments. The high-elevation subspecies S. o. taylori is not monophyletic, and the mitochondrial DNA genealogy supports a model of convergent phenotypic evolution among high-elevation populations belonging to different river drainages. The hypothesis that separate populations of S. occidentalis expanded up river drainages after the recession of glaciers is supported by population demographic analyses, and suggest that Bergmann's clines can evolve rapidly along elevation gradients. The distinctive high-elevation phenotype that is attributable to S. o. taylori has evolved independently several times, and includes adaptive phenotypic changes associated with increases in body size and ventral coloration. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 630,641. [source] Adaptive Units for Conservation: Population Distinction and Historic Extinctions in the Island Scrub-JayCONSERVATION BIOLOGY, Issue 2 2005KATHLEEN S. DELANEY Aphelocoma; diversidad genética especie; endémica; genética de conservación; Islas Channel Abstract:,The Island Scrub-Jay (Aphelocoma insularis) is found on Santa Cruz Island, California, and is the only insular bird species in the continental United States. We typed seven microsatellite loci and sequenced a portion of the mitochondrial DNA control region of Island Scrub-Jays and their closest mainland relative, the Western Scrub-Jay (Aphelocoma californica), to assess levels of variability and effective population size and to examine the evolutionary relationship between the two species. The estimated female effective population size, Nef, of the Island Scrub-Jay was 1603 (90% confidence interval: 1481,1738) and was about 7.5% of the size of the mainland species. Island and Western Scrub-Jays have highly divergent control-region sequences, and the value of 3.14 ± 0.09% sequence divergence between the two species suggests a divergence time of approximately 151,000 years ago. Because the four northern Channel Islands were joined as one large island as recently as 11,000 years ago, extinctions must have occurred on the three other northern Channel islands, Santa Rosa, San Miguel, and Anacapa, highlighting the vulnerability of the remaining population. We assessed the evolutionary significance of four island endemics, including the Island Scrub-Jay, based on both genetic and adaptive divergence. Our results show that the Island Scrub-Jay is a distinct species of high conservation value whose history and adaptive potential is not well predicted by study of other island vertebrates. Resumen:, Aphelocoma insularis se encuentra en la Isla Santa Cruz, California, y es la única especie de ave insular en Estados Unidos continental. Clasificamos siete locus microsatelitales y secuenciamos una porción de la región control del ADN mitocondrial de A. insularis y su pariente continental más cercano A. californica para evaluar niveles de variabilidad y tamaño poblacional efectivo y examinar las relaciones evolutivas entre las dos especies. El tamaño poblacional efectivo de hembras, Neh, de A. insularis fue estimado en 1603 (90% CI: 1481-1738) y fue aproximadamente 7.5% del tamaño de la especie continental. Aphelocoma insularis y A. californica tienen secuencias muy divergentes en la región control, y el valor de divergencia secuencial de 3.14 ± 0.09% entre las dos especies sugiere un tiempo de divergencia de aproximadamente 151,000 años. Debido a que las cuatro Islas Channel estuvieron unidas en una sola isla tan recientemente como hace 11,000 años, deben haber ocurrido extinciones en las otras tres islas Channel, Santa Rosa, San Miguel y Anacapa, acentuando la vulnerabilidad de la población remanente. Evaluamos el significado evolutivo de cuatro especies insulares endémicas incluyendo A. insularis con base en la divergencia genética y adaptativa. Nuestros resultados muestran que A. insularis es una especie distinta de alto valor de conservación, cuya historia y potencial adaptativo no es pronosticado correctamente por el estudio de otros vertebrados insulares. [source] THE BIOLOGY OF SPECIATIONEVOLUTION, Issue 2 2010James M. Sobel Since Darwin published the "Origin," great progress has been made in our understanding of speciation mechanisms. The early investigations by Mayr and Dobzhansky linked Darwin's view of speciation by adaptive divergence to the evolution of reproductive isolation, and thus provided a framework for studying the origin of species. However, major controversies and questions remain, including: When is speciation nonecological? Under what conditions does geographic isolation constitute a reproductive isolating barrier? and How do we estimate the "importance" of different isolating barriers? Here, we address these questions, providing historical background and offering some new perspectives. A topic of great recent interest is the role of ecology in speciation. "Ecological speciation" is defined as the case in which divergent selection leads to reproductive isolation, with speciation under uniform selection, polyploid speciation, and speciation by genetic drift defined as "nonecological." We review these proposed cases of nonecological speciation and conclude that speciation by uniform selection and polyploidy normally involve ecological processes. Furthermore, because selection can impart reproductive isolation both directly through traits under selection and indirectly through pleiotropy and linkage, it is much more effective in producing isolation than genetic drift. We thus argue that natural selection is a ubiquitous part of speciation, and given the many ways in which stochastic and deterministic factors may interact during divergence, we question whether the ecological speciation concept is useful. We also suggest that geographic isolation caused by adaptation to different habitats plays a major, and largely neglected, role in speciation. We thus provide a framework for incorporating geographic isolation into the biological species concept (BSC) by separating ecological from historical processes that govern species distributions, allowing for an estimate of geographic isolation based upon genetic differences between taxa. Finally, we suggest that the individual and relative contributions of all potential barriers be estimated for species pairs that have recently achieved species status under the criteria of the BSC. Only in this way will it be possible to distinguish those barriers that have actually contributed to speciation from those that have accumulated after speciation is complete. We conclude that ecological adaptation is the major driver of reproductive isolation, and that the term "biology of speciation," as proposed by Mayr, remains an accurate and useful characterization of the diversity of speciation mechanisms. [source] DOES VARIATION IN SELECTION IMPOSED BY BEARS DRIVE DIVERGENCE AMONG POPULATIONS IN THE SIZE AND SHAPE OF SOCKEYE SALMON?EVOLUTION, Issue 5 2009Stephanie M. Carlson Few studies have determined whether formal estimates of selection explain patterns of trait divergence among populations, yet this is one approach for evaluating whether the populations are in equilibria. If adaptive divergence is complete, directional selection should be absent and stabilizing selection should prevail. We estimated natural selection, due to bear predation, acting on the body size and shape of male salmon in three breeding populations that experience differing predation regimes. Our approach was to (1) estimate selection acting within each population on each trait based on an empirical estimate of reproductive activity, (2) test for trait divergence among populations, and (3) test whether selection coefficients were correlated with trait divergence among populations. Stabilizing selection was never significant, indicating that these populations have yet to attain equilibria. Directional selection varied among populations in a manner consistent with trait divergence, indicating ongoing population differentiation. Specifically, the rank order of the creeks in terms of patterns of selection paralleled the rank order in terms of size and shape. The shortest and least deep-bodied males had the highest reproductive activity in the creek with the most intense predation and longer and deeper-bodied males were favored in the creeks with lower predation risk. [source] CONTEMPORARY ISOLATION-BY-DISTANCE, BUT NOT ISOLATION-BY-TIME, AMONG DEMES OF EUROPEAN GRAYLING (THYMALLUS THYMALLUS, LINNAEUS) WITH RECENT COMMON ANCESTORSEVOLUTION, Issue 2 2009Nicola J. Barson The development of isolation by distance (IBD) and isolation by time (IBT) was contrasted among demes of European grayling (Thymallus thymallus) that have diverged within the last 25 generations following colonization of a lake (Lesjaskogsvatnet). We find low but significant levels of genetic differentiation among spawning tributaries and a pattern of IBD among them. We do not, however, find evidence for IBT despite an up to four-week difference in spawning date between "warm/early" and "cold/late" spawning demes and differences in the incubation temperatures experienced by offspring. It appears that IBD has developed more rapidly than IBT in this system and that adaptive divergence has been initiated in the absence of IBT. Although analysis of selected loci could reveal reduced recombination in parts of the genome associated with temporal divergence, our analysis of neutral genetic data suggests that IBD is a more important isolating mechanism in the early stages of adaptive divergence in European grayling. [source] ADAPTIVE POPULATION DIFFERENTIATION IN PHENOLOGY ACROSS A LATITUDINAL GRADIENT IN EUROPEAN ASPEN (POPULUS TREMULA, L.): A COMPARISON OF NEUTRAL MARKERS, CANDIDATE GENES AND PHENOTYPIC TRAITSEVOLUTION, Issue 12 2007David Hall A correct timing of growth cessation and dormancy induction represents a critical ecological and evolutionary trade-off between survival and growth in most forest trees (Rehfeldt et al. 1999; Horvath et al. 2003; Howe et al. 2003). We have studied the deciduous tree European Aspen (Populus tremula) across a latitudinal gradient and compared genetic differentiation in phenology traits with molecular markers. Trees from 12 different areas covering 10 latitudinal degrees were cloned and planted in two common gardens. Several phenology traits showed strong genetic differentiation and clinal variation across the latitudinal gradient, with QST values generally exceeding 0.5. This is in stark contrast to genetic differentiation at several classes of genetic markers (18 neutral SSRs, 7 SSRs located close to phenology candidate genes and 50 SNPs from five phenology candidate genes) that all showed FST values around 0.015. We thus find strong evidence for adaptive divergence in phenology traits across the latitudinal gradient. However, the strong population structure seen at the quantitative traits is not reflected in underlying candidate genes. This result fit theoretical expectations that suggest that genetic differentiation at candidate loci is better described by FST at neutral loci rather than by QST at the quantitative traits themselves. [source] REPLICATED EVOLUTION OF INTEGRATED PLASTIC RESPONSES DURING EARLY ADAPTIVE DIVERGENCEEVOLUTION, Issue 4 2006Kevin J. Parsons Abstract Colonization of a novel environment is expected to result in adaptive divergence from the ancestral population when selection favors a new phenotypic optimum. Local adaptation in the new environment occurs through the accumulation and integration of character states that positively affect fitness. The role played by plastic traits in adaptation to a novel environment has generally been ignored, except for variable environments. We propose that if conditions in a relatively stable but novel environment induce phenotypically plastic responses in many traits, and if genetic variation exists in the form of those responses, then selection may initially favor the accumulation and integration of functionally useful plastic responses. Early divergence between ancestral and colonist forms will then occur with respect to their plastic responses across the gradient bounded by ancestral and novel environmental conditions. To test this, we compared the magnitude, integration, and pattern of plastic character responses in external body form induced by shallow versus open water conditions between two sunfish ecomorphs that coexist in four postglacial lakes. The novel sunfish ecomorph is present in the deeper open water habitat, whereas the ancestral ecomorph inhabits the shallow waters along the lake margin. Plastic responses by open water ecomorphs were more correlated than those of their local shallow water ecomorph in two of the populations, whereas equal levels of correlated plastic character responses occurred between ecomorphs in the other two populations. Small but persistent differences occurred between ecomorph pairs in the pattern of their character responses, suggesting a recent divergence. Open water ecomorphs shared some similarities in the covariance among plastic responses to rearing environment. Replication in the form of correlated plastic responses among populations of open water ecomorphs suggests that plastic character states may evolve under selection. Variation between ecomorphs and among lake populations in the covariance of plastic responses suggests the presence of genetic variation in plastic character responses. In three populations, open water ecomorphs also exhibited larger plastic responses to the environmental gradient than the local shallow water ecomorph. This could account for the greater integration of plastic responses in open water ecomorphs in two of the populations. This suggests that the plastic responses of local sunfish ecomorphs can diverge through changes in the magnitude and coordination of plastic responses. Although these results require further investigation, they suggest that early adaptive evolution in a novel environment can include changes to plastic character states. The genetic assimilation of coordinated plastic responses could result in the further, and possibly rapid, divergence of such populations and could also account for the evolution of genes of major effect that contribute to suites of phenotypic differences between divergent populations. [source] Morphological variation over ontogeny and environments in resource polymorphic arctic charr (Salvelinus alpinus)EVOLUTION AND DEVELOPMENT, Issue 3 2010Kevin J. Parsons SUMMARY Natural selection requires genetically based phenotypic variation to facilitate its action and cause adaptive evolution. It has become increasingly recognized that morphological development can become canalized likely as a result of selection. However, it is largely unknown how selection may influence canalization over ontogeny and differing environments. Changes in environments or colonization of a novel one is expected to result in adaptive divergence from the ancestral population when selection favors a new phenotypic optimum. In turn, a novel environment may also expose variation previously hidden from natural selection. We tested for changes in phenotypic variation over ontogeny and environments among ecomorphs of Arctic charr (Salvelinus alpinus) from two Icelandic lakes. Populations represented varying degrees of ecological specialization, with one lake population possessing highly specialized ecomorphs exhibiting a large degree of phenotypic divergence, whereas the other displayed more subtle divergence with more ecological overlap. Here we show that ecomorphs hypothesized to be the most specialized in each lake possess significant reductions in shape variation over ontogeny regardless of environmental treatment suggesting canalized development. However, environments did change the amount of shape variation expressed in these ecomorphs, with novel environments slowing the rate at which variation was reduced over ontogeny. Thus, environmental conditions may play an important role in determining the type and amount of genetically based phenotypic variation exposed to natural selection. [source] Paradigm shifts in marine fisheries genetics: ugly hypotheses slain by beautiful factsFISH AND FISHERIES, Issue 4 2008Lorenz Hauser Abstract By providing new approaches to the investigation of demographic and evolutionary dynamics of wild populations, molecular genetics has led to fundamental changes in our understanding of marine ecology. In particular, genetic approaches have revolutionized our understanding in three areas: (i) most importantly, they have contributed to the discovery of extensive genetic population structure in many marine species, overturning the notion of large, essentially homogenous marine populations limiting local adaptation and speciation. (ii) Concomitant differences in ecologically important traits now indicate extensive adaptive differentiation and biocomplexity, potentially increasing the resilience to exploitation and disturbance. Evidence for rapid adaptive change in many populations underlies recent concerns about fisheries-induced evolution affecting life-history traits. (iii) A compilation of recent published research shows estimated effective population sizes that are 2,6 orders of magnitude smaller than census sizes, suggesting more complex recruitment dynamics in marine species than previously assumed. Studies on Atlantic cod are used to illustrate these paradigm shifts. In our synthesis, we emphasize the implications of these discoveries for marine ecology and evolution as well as the management and conservation of exploited marine fish populations. An important implication of genetic structuring and the potential for adaptive divergence is that locally adapted populations are unlikely to be replaced through immigration, with potentially detrimental consequences for the resilience to environmental change , a key consideration for sustainable fisheries management. [source] Adaptive versus non-adaptive phenotypic plasticity and the potential for contemporary adaptation in new environmentsFUNCTIONAL ECOLOGY, Issue 3 2007C. K. GHALAMBOR Summary 1The role of phenotypic plasticity in evolution has historically been a contentious issue because of debate over whether plasticity shields genotypes from selection or generates novel opportunities for selection to act. Because plasticity encompasses diverse adaptive and non-adaptive responses to environmental variation, no single conceptual framework adequately predicts the diverse roles of plasticity in evolutionary change. 2Different types of phenotypic plasticity can uniquely contribute to adaptive evolution when populations are faced with new or altered environments. Adaptive plasticity should promote establishment and persistence in a new environment, but depending on how close the plastic response is to the new favoured phenotypic optimum dictates whether directional selection will cause adaptive divergence between populations. Further, non-adaptive plasticity in response to stressful environments can result in a mean phenotypic response being further away from the favoured optimum or alternatively increase the variance around the mean due to the expression of cryptic genetic variation. The expression of cryptic genetic variation can facilitate adaptive evolution if by chance it results in a fitter phenotype. 3We conclude that adaptive plasticity that places populations close enough to a new phenotypic optimum for directional selection to act is the only plasticity that predictably enhances fitness and is most likely to facilitate adaptive evolution on ecological time-scales in new environments. However, this type of plasticity is likely to be the product of past selection on variation that may have been initially non-adaptive. 4We end with suggestions on how future empirical studies can be designed to better test the importance of different kinds of plasticity to adaptive evolution. [source] Five questions on ecological speciation addressed with individual-based simulationsJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 1 2009X. THIBERT-PLANTE Abstract We use an individual-based simulation model to investigate factors influencing progress toward ecological speciation. We find that environmental differences can quickly lead to the evolution of substantial reproductive barriers between a population colonizing a new environment and the ancestral population in the old environment. Natural selection against immigrants and hybrids was a major contributor to this isolation, but the evolution of sexual preference was also important. Increasing dispersal had both positive and negative effects on population size in the new environment and had positive effects on natural selection against immigrants and hybrids. Genetic divergence at unlinked, neutral genetic markers was low, except when environmental differences were large and sexual preference was present. Our results highlight the importance of divergent selection and adaptive divergence for ecological speciation. At the same time, they reveal several interesting nonlinearities in interactions between environmental differences, sexual preference, dispersal and population size. [source] Neutral theory: a historical perspectiveJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 6 2007E. G. LEIGH JR Abstract To resolve a panselectionist paradox, the population geneticist Kimura invented a neutral theory, where each gene is equally likely to enter the next generation whatever its allelic type. To learn what could be explained without invoking Darwinian adaptive divergence, Hubbell devised a similar neutral theory for forest ecology, assuming each tree is equally likely to reproduce whatever its species. In both theories, some predictions worked; neither theory proved universally true. Simple assumptions allow neutral theorists to treat many subjects still immune to more realistic theory. Ecologists exploit far fewer of these possibilities than population geneticists, focussing instead on species abundance distributions, where their predictions work best, but most closely match non-neutral predictions. Neutral theory cannot explain adaptive divergence or ecosystem function, which ecologists must understand. By addressing new topics and predicting changes in time, however, ecological neutral theory can provide probing null hypotheses and stimulate more realistic theory. [source] Maternal genetic effects on adaptive divergence between anadromous and resident brook charr during early life historyJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 5 2005G. M. L. PERRY Abstract The importance of directional selection relative to neutral evolution may be determined by comparing quantitative genetic variation in phenotype (QST) to variation at neutral molecular markers (FST). Quantitative divergence between salmonid life history types is often considerable, but ontogenetic changes in the significance of major sources of genetic variance during post-hatch development suggest that selective differentiation varies by developmental stage. In this study, we tested the hypothesis that maternal genetic differentiation between anadromous and resident brook charr (Salvelinus fontinalis Mitchill) populations for early quantitative traits (embryonic size/growth, survival, egg number and developmental time) would be greater than neutral genetic differentiation, but that the maternal genetic basis for differentiation would be higher for pre-resorption traits than post-resorption traits. Quantitative genetic divergence between anadromous (seawater migratory) and resident Laval River (Québec) brook charr based on maternal genetic variance was high (QST > 0.4) for embryonic length, yolk sac volume, embryonic growth rate and time to first response to feeding relative to neutral genetic differentiation [FST = 0.153 (0.071,0.214)], with anadromous females having positive genetic coefficients for all of the above characters. However, QST was essentially zero for all traits post-resorption of the yolk sac. Our results indicate that the observed divergence between resident and anadromous brook charr has been driven by directional selection, and may therefore be adaptive. Moreover, they provide among the first evidence that the relative importance of selective differentiation may be highly context-specific, and varies by genetic contributions to phenotype by parental sex at specific points in offspring ontogeny. This in turn suggests that interpretations of QST - FST comparisons may be improved by considering the structure of quantitative genetic architecture by age category and the sex of the parent used in estimation. [source] Testing alternative mechanisms of evolutionary divergence in an African rain forest passerine birdJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 2 2005T. B. Smith Abstract Models of speciation in African rain forests have stressed either the role of isolation or ecological gradients. Here we contrast patterns of morphological and genetic divergence in parapatric and allopatric populations of the Little Greenbul, Andropadus virens, within different and similar habitats. We sampled 263 individuals from 18 sites and four different habitat types in Upper and Lower Guinea. We show that despite relatively high rates of gene flow among populations, A. virens has undergone significant morphological divergence across the savanna,forest ecotone and mountain,forest boundaries. These data support a central component of the divergence-with-gene-flow model of speciation by suggesting that despite large amounts of gene flow, selection is sufficiently intense to cause morphological divergence. Despite evidence of isolation based on neutral genetic markers, we find little evidence of morphological divergence in fitness-related traits between hypothesized refugial areas. Although genetic evidence suggests populations in Upper and Lower Guinea have been isolated for over 2 million years, morphological divergence appears to be driven more by habitat differences than geographic isolation and suggests that selection in parapatry may be more important than geographic isolation in causing adaptive divergence in morphology. [source] Spatial and temporal scales of adaptive divergence in marine fishes and the implications for conservationJOURNAL OF FISH BIOLOGY, Issue 2006D. O. Conover Knowledge of geographic and temporal scales of adaptive genetic variation is crucial to species conservation, yet understanding of these phenomena, particularly in marine systems, is scant. Until recently, the belief has been that because most marine species have highly dispersive or mobile life stages, local adaptation could occur only on broad geographic scales. This view is supported by comparatively low levels of genetic variation among populations as detected by neutral markers. Similarly, the time scale of adaptive divergence has also been assumed to be very long, requiring thousands of generations. Recent studies of a variety of species have challenged these beliefs. First, there is strong evidence of geographically structured local adaptation in physiological and morphological traits. Second, the proportion of quantitative trait variation at the among-population level (QST) is much higher than it is for neutral markers (FST) and these two metrics of genetic variation are poorly correlated. Third, evidence that selection is a potent evolutionary force capable of sustaining adaptive divergence on contemporary time scales is summarized. The differing spatial and temporal scales of adaptive v. neutral genetic divergence call for a new paradigm in thinking about the relationship between phenogeography (the geography of phenotypic variation) and phylogeography (the geography of lineages) in marine species. The idea that contemporary selective processes can cause fine-scale spatial and temporal divergence underscores the need for a new emphasis on Darwinian fishery science. [source] Genome-wide single nucleotide polymorphisms reveal population history and adaptive divergence in wild guppiesMOLECULAR ECOLOGY, Issue 5 2010EVA-MARIA WILLING Abstract Adaptation of guppies (Poecilia reticulata) to contrasting upland and lowland habitats has been extensively studied with respect to behaviour, morphology and life history traits. Yet population history has not been studied at the whole-genome level. Although single nucleotide polymorphisms (SNPs) are the most abundant form of variation in many genomes and consequently very informative for a genome-wide picture of standing natural variation in populations, genome-wide SNP data are rarely available for wild vertebrates. Here we use genetically mapped SNP markers to comprehensively survey genetic variation within and among naturally occurring guppy populations from a wide geographic range in Trinidad and Venezuela. Results from three different clustering methods, Neighbor-net, principal component analysis (PCA) and Bayesian analysis show that the population substructure agrees with geographic separation and largely with previously hypothesized patterns of historical colonization. Within major drainages (Caroni, Oropouche and Northern), populations are genetically similar, but those in different geographic regions are highly divergent from one another, with some indications of ancient shared polymorphisms. Clear genomic signatures of a previous introduction experiment were seen, and we detected additional potential admixture events. Headwater populations were significantly less heterozygous than downstream populations. Pairwise FST values revealed marked differences in allele frequencies among populations from different regions, and also among populations within the same region. FST outlier methods indicated some regions of the genome as being under directional selection. Overall, this study demonstrates the power of a genome-wide SNP data set to inform for studies on natural variation, adaptation and evolution of wild populations [source] Genomic pattern of adaptive divergence in Arabidopsis halleri, a model species for tolerance to heavy metalMOLECULAR ECOLOGY, Issue 9 2009CLAIRE-LISE MEYER Abstract Pollution by heavy metals is one of the strongest environmental constraints in human-altered environments that only a handful of species can cope with. Identifying the genes conferring to those species the ability to grow in polluted areas is a first step towards a global understanding of the evolutionary processes involved and will eventually improve phytoremediation practices. We used a genome-scan approach to detect loci under divergent selection among four populations of Arabidopsis halleri growing on either polluted or nonpolluted habitats. Based on a high density of amplified fragment length polymorphism (AFLP) markers (820 AFLP markers, i.e. ~1 marker per 0.3 Mb), evidence for selection was found for some markers in every sampled population. Four loci departed from neutrality in both metallicolous populations and thus constitute high-quality candidates for general adaptation to pollution. Interestingly, some candidates differed between the two metallicolous populations, suggesting the possibility that different loci may be involved in adaptation in the different metallicolous populations. [source] The transcriptomics of life-history trade-offs in whitefish species pairs (Coregonus sp.)MOLECULAR ECOLOGY, Issue 7 2008J. ST-CYR Abstract Despite the progress achieved in elucidating the ecological mechanisms of adaptive radiation, there has been little focus on documenting the extent of adaptive differentiation in physiological functions during this process. Moreover, a thorough understanding of the genomic basis underlying phenotypic adaptive divergence is still in its infancy. One important evolutionary process for which causal genetic mechanisms are largely unknown pertains to life-history trade-offs. We analysed patterns of gene transcription in liver tissue of sympatric dwarf and normal whitefish from two natural lakes, as well as from populations reared in controlled environments, using a 16 006-gene cDNA microarray in order to: (i) document the extent of physiological adaptive divergence between sympatric dwarf and normal species pairs, and (ii) explore the molecular mechanisms of differential life history trade-offs between growth and survival potentially involved in their adaptive divergence. In the two natural lakes, 6.45% of significantly transcribed genes showed regulation either in parallel fashion (2.39%) or in different directions (4.06%). Among genes showing parallelism in regulation patterns, we observed a higher proportion of over-expressed genes in dwarf relative to normal whitefish (70.6%). Patterns observed in controlled conditions were also generally congruent with those observed in natural populations. Dwarf whitefish consistently showed significant over-expression of genes potentially associated with survival through enhanced activity (energy metabolism, iron homeostasis, lipid metabolism, detoxification), whereas more genes associated with growth (protein synthesis, cell cycle, cell growth) were generally down-regulated in dwarf relative to normal whitefish. Overall, parallelism in patterns of gene transcription, as well as patterns of interindividual variation across controlled and natural environments, provide strong indirect evidence for the role of selection in the evolution of differential regulation of genes involving a vast array of potentially adaptive physiological processes between dwarf and normal whitefish. Our results also provide a first mechanistic, genomic basis for the observed trade-off in life-history traits distinguishing dwarf and normal whitefish species pairs, wherein enhanced survival via more active swimming, necessary for increased foraging and predator avoidance, engages energetic costs that translate into slower growth rate and reduced fecundity in dwarf relative to normal whitefish. [source] Hybridization between mtDNA-defined phylogeographic lineages of black ratsnakes (Pantherophis sp.)MOLECULAR ECOLOGY, Issue 12 2006H. LISLE GIBBS Abstract Phylogeographic analyses using mitochondrial DNA (mtDNA) have revealed many examples of apparently deep historical subdivisions (,phylogroups') within many vertebrates. It remains unclear whether these phylogroups represent independently evolving, adaptively differentiated lineages or groups that show little functional differentiation and, hence, will merge on contact. Here, we use mtDNA sequence data to evaluate the phylogeographic relationships between two of the northernmost populations of black ratsnakes (Pantherophis obsoletus complex) in Ontario, Canada and previously analysed populations in the United States. We then use population-level analyses to evaluate the level of adaptive divergence between previously established mtDNA phylogroups. Phylogenetic analyses show that southern Ontario snakes have mtDNA haplotypes that fall within the Central mtDNA phylogroup, as designated by Burbrink et al. (2000). In contrast, snakes in eastern Ontario carry either Central or Eastern-specific haplotypes. Within the hybrid region, we found highly variable frequencies of mtDNA haplotypes among isolated sub-populations, no association between variation in cytonuclear (mtDNA) and nuclear (microsatellite DNA) markers, no difference in survival or reproductive success among snakes with different mtDNA haplotypes, and no effect of mate similarity in mtDNA on female clutch size. These results argue that the Eastern and Central phylogroups have merged in this region, likely due to a lack of adaptive differentiation between individuals in each lineage. Hence, in these snakes, phylogeographic structure in mtDNA is more a reflection of historical isolation rather than adaptive divergence. The observed reticulation between lineages and lack of evidence for hybrid disgenesis also bears on the classification of these lineages as distinct species. [source] INVITED REVIEW: Using genome scans of DNA polymorphism to infer adaptive population divergenceMOLECULAR ECOLOGY, Issue 3 2005JAY F. STORZ Abstract Elucidating the genetic basis of adaptive population divergence is a goal of central importance in evolutionary biology. In principle, it should be possible to identify chromosomal regions involved in adaptive divergence by screening genome-wide patterns of DNA polymorphism to detect the locus-specific signature of positive directional selection. In the case of spatially separated populations that inhabit different environments or sympatric populations that exploit different ecological niches, it is possible to identify loci that underlie divergently selected traits by comparing relative levels of differentiation among large numbers of unlinked markers. In this review I first address the question of whether diversifying selection on polygenic traits can be expected to produce predictable patterns of allelic variation at the underlying quantitative trait loci (QTL), and whether the locus-specific effects of selection can be reliably detected against the genome-wide backdrop of stochastic variability. I then review different approaches that have been developed to identify loci involved in adaptive population divergence and I discuss the relative merits of model-based approaches that rely on assumptions about population structure vs. model-free approaches that are based on empirical distributions of summary statistics. Finally, I consider the evolutionary and functional insights that might be gained by conducting genome scans for loci involved in adaptive population divergence. [source] Correlated evolution of fruit and leaf size in bird-dispersed plants: species-level variance in fruit traits explained a bit further?OIKOS, Issue 3 2002Carlos M. Herrera The astounding morphological diversity exhibited by the fruits of vertebrate-dispersed plants has been traditionally interpreted as the adaptive outcome of divergent selective pressures exerted on plants by the broad array of frugivorous animals involved in seed dispersal. Although the selective capacity of frugivores provides support to this interpretation, recent studies have challenged it by documenting a strong phylogenetic component associated to interspecific variation in most fruit characteristics. Size-related fruit traits provide a conspicuous exception to this pattern, because they exhibit considerable variation at the between-species level which is largely independent of phylogeny and is correlated with consumption by differently-sized dispersal agents. Substantial species-level variance in size-related traits may reflect genuine disperser-driven diversification, but may also be partly influenced by correlated evolution of fruit size with the size of other plant structures. This latter possibility is tested here for bird-dispersed plants of the Iberian Peninsula using phylogenetically independent contrasts. Results demonstrate the existence of correlated evolution of fruit and leaf size at the species level. As all the plant taxa considered have their fruits eaten, and seeds dispersed, by the same relatively reduced set of frugivorous bird species, results suggest that a significant fraction of the variation in fruit size represented in the species sample may be explained as an indirect consequence of variation in leaf size, rather than being associated with adaptive divergence related to seed dispersal agents. [source] Population structure, genetic variation and morphological diversity in indigenous sheep of EthiopiaANIMAL GENETICS, Issue 6 2007S. Gizaw Summary We investigated genetic and morphological diversity and population structure of 14 traditional sheep populations originating from four ecological zones in Ethiopia (sub-alpine, wet highland, sub-humid lowland and arid lowland). All animals (n = 672) were genotyped for 17 microsatellite markers and scored for 12 morphological characters. The sheep were initially classified as fat-tailed (11 populations), thin-tailed (one population) and fat-rumped sheep (two populations). These classifications are thought to correspond to three consecutive introduction events of sheep from the Near-East into East Africa. For the 14 populations, allelic richness ranged from 5.87 to 7.51 and expected heterozygosity (HE) from 0.66 to 0.75. Genetic differentiations (FST values) between all pairs of populations, except between sub-alpine populations, were significantly different from zero (P < 0.001). Cluster analysis of morphological characters and a dendrogram constructed from genetic distances were broadly consistent with the classification into fat-tailed, thin-tailed and fat-rumped sheep. Bayesian cluster analysis using microsatellite markers indicated that there has been further genetic differentiation after the initial introduction of sheep into Ethiopia. Investigation of factors associated with genetic variation showed that an isolation-by-distance model, independently of other factors, explained most of the observed genetic variation. We also obtained a strong indication of adaptive divergence in morphological characters, patterns of morphological variation being highly associated with ecology even when the effect of neutral genetic divergence (FST) was parcelled out in partial Mantel tests. Using a combination of FST values, Bayesian clustering analysis and morphological divergence, we propose a classification of Ethiopian sheep into six breed groups and nine breeds. [source] |