Genetic variability in wild cherry populations in France. Effects of colonizing processes

 Isoenzymes were used to evaluate gene diversity and genetic differentiation among six populations of wild cherry (Prunus avium L.) in France. We contrast the genetic characteristics of a population resulting from a recent colonization with those of a much older population of the same species. No significant genetic structure was observed among populations; in this respect wild cherry does not differ from other forest trees. No founder effects could be detected in the newly colonized population. To explain the results, we discuss classic explanations for the lack of genetic differentiation among populations, including balancing selection and neutral drift/migration. In order to account for the absence of founder effects, we propose a hypothesis based on the life cycle of forest trees, namely that the length of the juvenile phase reduces the impact of small numbers of initial founders. Received : 26 November 1996 / Accepted : 20 December 1996     

Linking extinction-colonization dynamics to genetic structure in a salamander metapopulation

Theory predicts that founder effects have a primary role in determining metapopulation genetic structure. However, ecological factors that affect extinction-colonization dynamics may also create spatial variation in the strength of genetic drift and migration. We tested the hypothesis that ecological factors underlying extinction-colonization dynamics influenced the genetic structure of a tiger salamander (Ambystoma tigrinum) metapopulation. We used empirical data on metapopulation dynamics to make a priori predictions about the effects of population age and ecological factors on genetic diversity and divergence among 41 populations. Metapopulation dynamics of A. tigrinum depended on wetland area, connectivity and presence of predatory fish. We found that newly colonized populations were more genetically differentiated than established populations, suggesting that founder effects influenced genetic structure. However, ecological drivers of metapopulation dynamics were more important than age in predicting genetic structure. Consistent with demographic predictions from metapopulation theory, genetic diversity and divergence depended on wetland area and connectivity. Divergence was greatest in small, isolated wetlands where genetic diversity was low. Our results show that ecological factors underlying metapopulation dynamics can be key determinants of spatial genetic structure, and that habitat area and isolation may mediate the contributions of drift and migration to divergence and evolution in local populations.     

Ecology,demography, and population genetics of barbary macaques in Algeria

Over a 9-year period from 1982 to 1990 ecological and demographic data were collected on two genetic isolates of Barbary macaques (Macaca sylvanus) in Algeria, from the deciduous oak-forest of Akfadou and from the evergreen cedar-oak forest of the National Park Djurdjura. Macaques at Djurdjura profit from more suitable ecological conditions and have a higher rate of population increase as well as a higher male migration rate than those at Akfadou. Genetic data, gained from 23 genetic markers (blood proteins), proved to be highly influenced by the demographic structure of the groups. The macaque populations of Akfadou and Djurdjura have become genetically differentiated. Group fission, coupled with founder effect (genetic drift) and kin-structured (matrilineal) separation, resulted in a priori genetic diversity between one newly-established group and its parent group.     

Which Extent is Plasticity to Light Involved in the Ecotypic Differentiation of a Tree Species from Savanna and Forest?

Light intensity and heterogeneity are some of the main environmental factors that differ between forest and savanna habitats, and plant species from these habitats form distinct functional types. In this study, we tested the hypothesis that not only differences in morphological and physiological traits but also phenotypic plasticity in response to light are involved in adaptation to forest and savanna habitats by investigating ecotypic differentiation between populations of Plathymenia reticulata (Leguminosae: Mimosoideae), a tree from the Brazilian Atlantic Forest and the Brazilian Cerrado (savanna). Seeds from four natural populations (one from each biome core area and two from ecotonal regions) were grown in a common garden with four light treatments. Fifteen morphological and physiological characteristics were evaluated until individuals reached 6 mo old. Comparisons among populations showed differences for seven traits in at least one light treatment. These differences pointed to local adaptation to different biomes. Populations showed different levels of phenotypic plasticity in response to light in seven traits. Higher plasticity was found either in the forest core population or ecotonal populations; lower values were found in the cerrado core population. Lower plasticity in the cerrado population emphasizes the stress resistant syndrome, as lower plasticity is probably advantageous in a habitat where a conservative resource use is crucial. Higher plasticity in forest individuals suggests higher ability in exploiting the light heterogeneity in this habitat. Also, higher plasticity in ecotonal populations can be important to ensure the maintenance of P. reticulata in these temporally and spatially dynamic areas. Abstract in Portugese is available at http://www.blackwell‐synergy.com/loi/btp .     

The adaptive value of epigenetic mutation: Limited in large but high in small peripheral populations

The fate of populations during range expansions, invasions and environmental changes is largely influenced by their ability to adapt to peripheral habitats. Recent models demonstrate that stable epigenetic modifications of gene expression that occur more frequently than genetic mutations can both help and hinder adaptation in panmictic populations. However, these models do not consider interactions between epimutations and evolutionary forces in peripheral populations. Here, we use mainland–island mathematical models and simulations to explore how the faster rate of epigenetic mutation compared to genetic mutations interacts with migration, selection and genetic drift to affect adaptation in peripheral populations. Our model focuses on cases where epigenetic marks are stably inherited. In a large peripheral population, where the effect of genetic drift is negligible, our analyses suggest that epimutations with random fitness impacts that occur at rates as high as 10^–3 increase local adaptation when migration is strong enough to overwhelm divergent selection. When migration is weak relative to selection and epimutations with random fitness impacts decrease adaptation, we find epigenetic modifications must be highly adaptively biased to enhance adaptation. Finally, in small peripheral populations, where genetic drift is strong, epimutations contribute to adaptation under a wider range of evolutionary conditions. Overall, our results suggest that epimutations can change outcomes of adaptation in peripheral populations, which has implications for understanding conservation and range expansions and contractions, especially of small populations.     

Determinants of Genetic Structure in a Nonequilibrium Metapopulation of the Plant Silene latifolia

Population genetic differentiation will be influenced by the demographic history of populations, opportunities for migration among neighboring demes and founder effects associated with repeated extinction and recolonization. In natural populations, these factors are expected to interact with each other and their magnitudes will vary depending on the spatial distribution and age structure of local demes. Although each of these effects has been individually identified as important in structuring genetic variance, their relative magnitude is seldom estimated in nature. We conducted a population genetic analysis in a metapopulation of the angiosperm, Silene latifolia, from which we had more than 20 years of data on the spatial distribution, demographic history, and extinction and colonization of demes. We used hierarchical Bayesian methods to disentangle which features of the populations contributed to among population variation in allele frequencies, including the magnitude and direction of their effects. We show that population age, long-term size and degree of connectivity all combine to affect the distribution of genetic variance; small, recently-founded, isolated populations contributed most to increase F _ST in the metapopulation. However, the effects of population size and population age are best understood as being modulated through the effects of connectivity to other extant populations, i.e. F _ST diminishes as populations age, but at a rate that depends how isolated the population is. These spatial and temporal correlates of population structure give insight into how migration, founder effect and within-deme genetic drift have combined to enhance and restrict genetic divergence in a natural metapopulation.     

Genetic and phenotypic divergence in an island bird: isolation by distance,by colonization or by adaptation?

Discerning the relative roles of adaptive and nonadaptive processes in generating differences among populations and species, as well as how these processes interact, is a fundamental aim in biology. Both genetic and phenotypic divergence across populations can be the product of limited dispersal and gradual genetic drift across populations (isolation by distance), of colonization history and founder effects (isolation by colonization) or of adaptation to different environments preventing migration between populations (isolation by adaptation). Here, we attempt to differentiate between these processes using island populations of Berthelot's pipit (Anthus berthelotii), a passerine bird endemic to three Atlantic archipelagos. Using microsatellite markers and approximate Bayesian computation, we reveal that the northward colonization of this species ca. 8500 years ago resulted in genetic bottlenecks in the colonized archipelagos. We then show that high levels of genetic structure exist across archipelagos and that these are consistent with a pattern of isolation by colonization, but not with isolation by distance or adaptation. Finally, we show that substantial morphological divergence also exists and that this is strongly concordant with patterns of genetic structure and bottleneck history, but not with environmental differences or geographic distance. Overall, our data suggest that founder effects are responsible for both genetic and phenotypic changes across archipelagos. Our findings provide a rare example of how founder effects can persist over evolutionary timescales and suggest that they may play an important role in the early stages of speciation.     

Regional and genetic variations among the Hindu Gujjars of Northwestern India

Serogenetic investigations among the Hindu Gujjars of Ropar District in Punjab were carried out with special reference to their genetic relationship with populations of other localities. Hindu Gujjars of Northwestern India show intragroup genetic variations, significant especially at the ABO locus. The magnitude of intragroup diversity varies depending upon the physical distance between the two groups. The plausible explanation for their diversity seems to be the founder effect, genetic drift and, their breeding and geographic isolation.     

Does local adaptation to resources explain genetic differentiation among Daphnia populations?

Substantial genetic differentiation is frequently observed among populations of cyclically parthenogenetic zooplankton despite their high dispersal capabilities and potential for gene flow. Local adaptation has been invoked to explain population genetic differentiation despite high dispersal, but several neutral models that account for basic life history features also predict high genetic differentiation. Here, we study genetic differentiation among four populations of Daphnia pulex in east central Illinois. As with other studies of Daphnia, we demonstrate substantial population genetic differentiation despite close geographic proximity (<50 km; mean θ = 0.22). However, we explicitly tested and failed to find evidence for, the hypothesis that local adaptation to food resources occurs in these populations. Recognizing that local adaptation can occur in traits unrelated to resources, we estimated contemporary migration rates (m) and tested for admixture to evaluate the hypothesis that observed genetic differentiation is consistent with local adaptation to other untested ecological factors. Using Bayesian assignment methods, we detected migrants in three of the four study populations including substantial evidence for successful reproduction by immigrants in one pond, allowing us to reject the hypothesis that local adaptation limits gene flow for at least this population. Thus, we suggest that local adaptation does not explain genetic differentiation among these Daphnia populations and that other factors related to extinction/colonization dynamics, a long approach to equilibrium F_ST or substantial genetic drift due to a low number of individuals hatching from the egg bank each season may explain genetic differentiation.     

Age-Related Tooth Wear Differs between Forest and Savanna Primates

Tooth wear in primates is caused by aging and ecological factors. However, comparative data that would allow us to delineate the contribution of each of these factors are lacking. Here, we contrast age-dependent molar tooth wear by scoring percent of dentine exposure (PDE) in two wild African primate populations from Gabonese forest and Kenyan savanna habitats. We found that forest-dwelling mandrills exhibited significantly higher PDE with age than savanna yellow baboons. Mandrills mainly feed on large tough food items, such as hard-shell fruits, and inhabit an ecosystem with a high presence of mineral quartz. By contrast, baboons consume large amounts of exogenous grit that adheres to underground storage organs but the proportion of quartz in the soils where baboons live is low. Our results support the hypothesis that not only age but also physical food properties and soil composition, particularly quartz richness, are factors that significantly impact tooth wear. We further propose that the accelerated dental wear in mandrills resulting in flatter molars with old age may represent an adaptation to process hard food items present in their environment.     

The effects of migration and drift on local adaptation to a heterogeneous environment

Local adaptation experiments are widely used to quantify the levels of adaptation within a heterogeneous environment. However, theoretical studies generally focus on the probability of fixation of alleles or the mean fitness of populations, rather than local adaptation as it is commonly measured experimentally or in field studies. Here, we develop mathematical models and use them to generate analytical predictions for the level of local adaptation as a function of selection, migration and genetic drift. First, we contrast mean fitness and local adaptation measures and show that the latter can be expressed in a simple and general way as a function of the spatial covariance between population mean phenotype and local environmental conditions. Second, we develop several approximations of a population genetics model to show that the system exhibits different behaviours depending on the rate of migration. The main insights are the following: with intermediate migration, both genetic drift and migration decrease local adaptation; with low migration, drift decreases local adaptation but migration speeds up adaptation; with high migration, genetic drift has no effect on local adaptation. Third, we extend this analysis to cases where the trait under selection is continuous using classical quantitative genetics theory. Finally, we discuss these results in the light of recent experimental work on local adaptation.     

Distinct subspecies or phenotypic plasticity? Genetic and morphological differentiation of mountain honey bees in East Africa

Identifying the forces shaping intraspecific phenotypic and genotypic divergence are of key importance in evolutionary biology. Phenotypic divergence may result from local adaptation or, especially in species with strong gene flow, from pronounced phenotypic plasticity. Here, we examine morphological and genetic divergence among populations of the western honey bee Apis mellifera in the topographically heterogeneous East African region. The currently accepted “mountain refugia hypothesis” states that populations living in disjunct montane forests belong to a different lineage than those in savanna habitats surrounding these forests. We obtained microsatellite data, mitochondrial sequences, and morphometric data from worker honey bees collected from feral colonies in three montane forests and corresponding neighboring savanna regions in Kenya. Honey bee colonies from montane forests showed distinct worker morphology compared with colonies in savanna areas. Mitochondrial sequence data did not support the existence of the two currently accepted subspecies. Furthermore, analyses of the microsatellite data with a Bayesian clustering method did not support the existence of two source populations as it would be expected under the mountain refugia scenario. Our findings suggest that phenotypic plasticity rather than distinct ancestry is the leading cause behind the phenotypic divergence observed between montane forest and savanna honey bees. Our study thus corroborates the idea that high gene flow may select for increased plasticity.     

Serial founder effects and genetic differentiation during worldwide range expansion of monarch butterflies

Range expansions can result in founder effects, increasing genetic differentiation between expanding populations and reducing genetic diversity along the expansion front. However, few studies have addressed these effects in long-distance migratory species, for which high dispersal ability might counter the effects of genetic drift. Monarchs (Danaus plexippus) are best known for undertaking a long-distance annual migration in North America, but have also dispersed around the world to form populations that do not migrate or travel only short distances. Here, we used microsatellite markers to assess genetic differentiation among 18 monarch populations and to determine worldwide colonization routes. Our results indicate that North American monarch populations connected by land show limited differentiation, probably because of the monarch''s ability to migrate long distances. Conversely, we found high genetic differentiation between populations separated by large bodies of water. Moreover, we show evidence for serial founder effects across the Pacific, suggesting stepwise dispersal from a North American origin. These findings demonstrate that genetic drift played a major role in shaping allele frequencies and created genetic differentiation among newly formed populations. Thus, range expansion can give rise to genetic differentiation and declines in genetic diversity, even in highly mobile species.     

Drivers of population genetic differentiation in the wild: isolation by dispersal limitation,isolation by adaptation and isolation by colonization

Empirical population genetic studies have been dominated by a neutralist view, according to which gene flow and drift are the main forces driving population genetic structure in nature. The neutralist view in essence describes a process of isolation by dispersal limitation (IBDL) that generally leads to a pattern of isolation by distance (IBD). Recently, however, conceptual frameworks have been put forward that view local genetic adaptation as an important driver of population genetic structure. Isolation by adaptation (IBA) and monopolization (M) posit that gene flow among natural populations is reduced as a consequence of local genetic adaptation. IBA stresses that effective gene flow is reduced among habitats that show dissimilar ecological characteristics, leading to a pattern of isolation by environment. In monopolization, local genetic adaptation of initial colonizing genotypes results in a reduction in gene flow that fosters the persistence of founder effects. Here, we relate these different processes driving landscape genetic structure to patterns of IBD and isolation by environment (IBE). We propose a method to detect whether IBDL, IBA and M shape genetic differentiation in natural landscapes by studying patterns of variation at neutral and non‐neutral markers as well as at ecologically relevant traits. Finally, we reinterpret a representative number of studies from the recent literature by associating patterns to processes and identify patterns associated with local genetic adaptation to be as common as IBDL in structuring regional genetic variation of populations in the wild. Our results point to the importance of quantifying environmental gradients and incorporating ecology in the analysis of population genetics.     

Genetic structure in a fragmented Northern Hemisphere rainforest: large effective sizes and high connectivity among populations of the epiphytic lichen Lobaria pulmonaria

An extraordinary diversity of epiphytic lichens is found in the boreal rainforest of central Norway, the highest-latitude rainforest in the world. These rainforest relicts are located in ravine systems, and clear cutting has increased the distance between remaining patches. We hypothesized that the relatively small lichen populations in the remaining forest stands have suffered a depletion of genetic diversity through bottlenecks and founder events. To test this hypothesis, we assessed genetic diversity and structure in the populations of the tripartite lichen Lobaria pulmonaria using eight SSR loci. We sampled thalli growing on Picea abies branches and propagules deposited in snow at three localities. Contrary to expectations, we found high genetic diversity in lichen and snow samples, and high effective sizes of the studied populations. Also, limited genetic differentiation between populations, high historical migration rates, and a high proportion of first generation immigrants were estimated, implying high connectivity across distances <30km. Almost all genetic variation was attributed to variation within sites; spatial genetic structures within populations were absent or appeared on small scales (5-10m). The high genetic diversity in the remaining old boreal rainforests shows that even relict forest patches might be suitable for conservation of genetic diversity.     

Survival rates in a natural population of the damselfly Ceriagrion tenellum: effects of sex and female phenotype

1. Ceriagrion tenellum females show genetic colour polymorphism. Androchrome (erythrogastrum) females are brightly (male‐like) coloured while gynochrome females (typica and melanogastrum) show cryptic colouration. 2. Several hypotheses have been proposed to explain the existence of more than one female morph in damselfly populations. The reproductive isolation and intraspecific mimicry hypotheses predict greater survival of gynochrome females, while the density dependent hypothesis predicts no differential survival between morphs. 3. Mature males had greater recapture probability than females while the survival probability was similar for both sexes. Survival and recapture rates were similar for androchrome and gynochrome females. 4. Gynochrome females showed greater mortality or migration rate than androchrome females during the pre‐reproductive period. This result is not predicted by the above hypotheses or by the null hypothesis that colour polymorphism is only maintained by random factors: founder effects, genetic drift, and migration.     

Body morphology and the savanna — forest transition: A West african example

Four Cameroonian populations were studied for a large set of anthropometric measurements. Three groups live in the forest (two Bantuspeaking, and one pygmoid tribe) and the fourth one in the savanna plateau. Multivariate analysis of the morphological features showed that the two Bantu groups have very similar body morphology. This is interpreted to be a result of common genetic origin and of the fact that they inhabit the same environment Whereas the Pygmoids segregate apart. The savanna dwellers are morphologically equidistant from the two former groups, and when compared to other savanna peoples living in Burkina-Faso, are seen to be more similar to forest populations. Some climatic influences can be identified, mainly on limb proportions and nose and face dimensions. They seem to play a larger part in body differentiation than do nutritional and/or pathological conditions. Differences between the forest Bantu and the savanna-dwellers are mostly due to a shape component, whereas the Pygmoids display an overall reduction in size. Savanna highlanders also exhibit a size reduction, which results in their being more similar in shape to the Pygmies than are the other groups compared. A fruitful approach to morphological differentiation must take advantage of opportunities to compare peoples sharing the same genetic origin but having migrated to occupy contrasting environments. The hypothesis is raised that a common African stock has differentiated in contrasting ecosystems.     

Genetic Diversity in Introduced Populations with an Allee Effect

A phenomenon that strongly influences the demography of small introduced populations and thereby potentially their genetic diversity is the demographic Allee effect, a reduction in population growth rates at small population sizes. We take a stochastic modeling approach to investigate levels of genetic diversity in populations that successfully overcame either a strong Allee effect, in which populations smaller than a certain critical size are expected to decline, or a weak Allee effect, in which the population growth rate is reduced at small sizes but not negative. Our results indicate that compared to successful populations without an Allee effect, successful populations with a strong Allee effect tend to (1) derive from larger founder population sizes and thus have a higher initial amount of genetic variation, (2) spend fewer generations at small population sizes where genetic drift is particularly strong, and (3) spend more time around the critical population size and thus experience more genetic drift there. In the case of multiple introduction events, there is an additional increase in diversity because Allee-effect populations tend to derive from a larger number of introduction events than other populations. Altogether, a strong Allee effect can either increase or decrease genetic diversity, depending on the average founder population size. By contrast, a weak Allee effect tends to decrease genetic diversity across the entire range of founder population sizes. Finally, we show that it is possible in principle to infer critical population sizes from genetic data, although this would require information from many independently introduced populations.