Abstract: 1. In a context of increasing human impact on ecosystems and species distributions, population restoration (introductions, reintroductions, reinforcements) is an essential management tool, especially for plant species with limited colonization ability. However, detailed demographic surveys following restoration and comparisons of demographic rates between restored and natural populations, although essential for identifying the key factors of restoration success, are lacking. 2. We compared the demography over 10 years of six natural and two experimentally introduced populations of the narrowly endemic, cliff-dwelling, self-incompatible plant species Centaurea corymbosa. We analysed the fate of two cohorts of individuals that emerged simultaneously from seed introduction and natural germination. We then built a matrix model of population dynamics (using 6 years of data) and compared the demographic rates and asymptotic growth rate between the natural and introduced populations. 3. Overall, survival rates were higher in the introduced than in the natural populations, either due to better habitat conditions at the cliff scale or to better conditions in microsites selected for seed introduction compared to those reached by chance following natural seed dispersal. 4. In contrast, introduced populations exhibited lower fecundity than natural populations, probably due to the introduction protocol which led, in combination with self-incompatibility, to severely reduced mate availability. 5. Despite clear differences in population dynamics between introduced and natural populations, no significant difference in the asymptotic growth rates could be detected, because the higher survival compensated for the lower fecundity in introduced populations. 6. Synthesis and applications. Creating new populations of C corymbosa in suitable unoccupied sites seems straightforward, provided that the introduction protocol allows sufficiently high fecundity. This key parameter for restoration success can be optimized by sowing seeds from several sources at high density and in several consecutive years, which should increase mate availability for self-incompatible flowering individuals. We suggest that population introduction might be successful for many (endemic) plant species whose geographical range is mainly limited by low colonization ability, especially in Mediterranean landscapes. We show that the simultaneous monitoring of restored and natural populations enables identification of the key parameters to be targeted for management optimization of restored populations.
Abstract: Genetic information is used extensively to reconstruct the evolutionary and demographic history of organisms. Recently, it has been suggested that genetic information from some parasites can complement genetic data from their hosts. This approach relies upon the hypothesis that such parasites share a common history with their host. In some cases, parasites provide an additional source of information because parasite data can better reconstruct the common history. Here, we discuss which parasite traits are important in determining their usefulness for analysing host history. The key is the matching of the traits of the parasite (e.g. effective population size, generation time, mutation rate and level of host specificity) with the timescales (phylogenetic, phylogeographic and demographic) that are relevant to the issues of concern in host history.
Abstract: Despite its significance in evolutionary and conservation biology, few estimates of effective population size (N-e) are available in plant species. Self-fertilization is expected to affect N-e, through both its effect on homozygosity and population dynamics. Here, we estimated N-e using temporal variation in allele frequencies for two contrasted populations of the selfing annual Medicago truncatula: a large and continuous population and a subdivided population. Estimated N-e values were around 5-10% of the population census size suggesting that other factors than selfing must contribute to variation in allele frequencies. Further comparisons between monolocus allelic variation and changes in the multilocus genotypic composition of the populations show that the local dynamics of inbred lines can play an important role in the fluctuations of allele frequencies. Finally, comparing N-e estimates and levels of genetic variation suggest that H-e is a poor estimator of the contemporaneous variance effective population size.
Abstract: Evolution in a single environment is expected to erode genetic variability, thereby precluding adaptation to novel environments. To test this, a large population of spider mites kept on cucumber for approximately 300 generations was used to establish populations on novel host plants (tomato or pepper), and changes in traits associated to adaptation were measured after 15 generations. Using a half-sib design, we investigated whether trait changes were related to genetic variation in the base population. Juvenile survival and fecundity exhibited genetic variation and increased in experimental populations on novel hosts. Conversely, no variation was detected for host choice and developmental time and these traits did not evolve. Longevity remained unchanged on novel hosts despite the presence of genetic variation, suggesting weak selection for this trait. Hence, patterns of evolutionary changes generally matched those of genetic variation, and changes in some traits were not hindered by long-term evolution in a constant environment.
Abstract: Using nitrogen-fixing Sinorhizobium species that interact with Medicago plants as a model system, we aimed at clarifying how sex has shaped the diversity of bacteria associated with the genus Medicago on the interspecific and intraspecific scales. To gain insights into the diversification of these symbionts, we inferred a topology that includes the different specificity groups which interact with Medicago species, based on sequences of the nodulation gene cluster. Furthermore, 126 bacterial isolates were obtained from two soil samples, using Medicago truncatula and Medicago laciniata as host plants, to study the differentiation between populations of Sinorhizobium medicae, Sinorhizobium meliloti bv. meliloti, and S. meliloti bv. medicaginis. The former two can be associated with M. truncatula (among other species of Medicago), whereas the last organism is the specific symbiont of M. laciniata. These bacteria were characterized using a multilocus sequence analysis of four loci, located on the chromosome and on the two megaplasmids of S. meliloti. The phylogenetic results reveal that several interspecific horizontal gene transfers occurred during the diversification of Medicago symbionts. Within S. meliloti, the analyses show that nod genes specific to different host plants have spread to different genetic backgrounds through homologous recombination, preventing further divergence of the different ecotypes. Thus, specialization to different host plant species does not prevent the occurrence of gene flow among host-specific biovars of S. meliloti, whereas reproductive isolation between S. meliloti bv. meliloti and S. medicae is maintained even though these bacteria can cooccur in sympatry on the same individual host plants.
Abstract: Here we explore life history differences in a set of neighbouring metallicolous and nonmetallicolous populations of the heavy metal tolerant plant Thlaspi caerulescens. We contrasted data from field observations and from a common garden experiment, in which soil zinc (Zn) concentration and light availability were manipulated, and data on microsatellite molecular variation. The two ecotypes showed few differences in life history in the field, but large differences in their response to Zn concentration in the common garden. Soil toxicity affected most characters in nonmetallicolous plants, while it had no effect on metallicolous plants. The two ecotypes responded similarly to light. Genetic differentiation for quantitative characters between ecotypes contrasted with the absence of differentiation for microsatellites. Conversely, populations of the same ecotype showed similar responses to Zn, despite their high differentiation for molecular markers. We conclude that divergent selection related to soil toxicity has had a predominant role in shaping life history differences between ecotypes, gene flow weakly opposing local adaptation despite geographical proximity.
Abstract: We investigate the genetic structure and molecular selection pattern of a sympatric population of Sinorhizobium meliloti and Sinorhizobium medicae. These bacteria fix nitrogen in association with plants of the genus Medicago. A set of 116 isolates were obtained from a soil sample, from root nodules of three groups of plants representing among-species, within-species and intraline diversity in the Medicago genus. Bacteria were characterized by sequencing at seven loci evenly distributed along the genome of both Sinorhizobium species, covering the chromosome and the two megaplasmids. We first test whether the diversity of host plants influence the bacterial diversity recovered. Using the same data set, we then analyse the selective pattern at each locus. There was no relationship between the diversity of Medicago plants that were used for sampling and the diversity of their symbionts. However, we found evidence of selection within each of the two main symbiotic regions, located on the two different megaplasmids. Purifying selection or a selective sweep was found to occur in the nod genomic region, which includes genes involved in nodulation specificity, whereas balancing selection was detected in the exo region, close to genes involved in exopolysaccharide production. Such pattern likely reflects the interaction between host plants and bacterial symbionts, with a possible conflict of interest between plants and cheater bacterial genotypes. Recombination appears to occur preferentially within and among loci located on megaplasmids, rather than within the chromosome. Thus, recombination may play an important role in resolving this conflict by allowing different selection patterns at different loci.
Abstract: In plants, genes may disperse through both pollen and seeds. Here we provide a first theoretical study of the mechanisms and consequences of the joint evolution of pollen and seed dispersal. We focus on hermaphroditic self-compatible species distributed in structured populations, assuming island dispersal of pollen and seeds among small patches of plants within large populations. Three traits are studied: the rate of among-patch seed dispersal, the rate of among-patch pollen dispersal, and the rate of within-patch pollen movement. We first analytically derive the evolutionary equilibrium state of each trait, dissect the pairwise selective interactions, and describe the joint three-trait evolutionary equilibrium under the cost of dispersal and kin competition. These results are then analytically and numerically extended to the case when selfed seeds suffer from depressed competitiveness (inbreeding depression, no heterosis). Finally individual-based simulations are used to account for a more realistic model of inbreeding load. Pollen movement is shown to generate opposite selection pressures on seed dispersal depending on spatial scale: within-patch pollen movement favors seed dispersal, whereas among-patch pollen dispersal inhibits seed dispersal. Seed dispersal selects for short-distance movements of pollen and it selects against long-distance dispersal. These interactions shape the joint evolution of these traits. Kin competition favors among-patch seed dispersal over among-patch pollen dispersal for low costs of within-patch pollen movement (and vice versa for significant costs of within-patch pollen movement). Inbreeding depression favors allogamy through high rates of within- and among-patch pollen movement. Surprisingly, it may select either for or against seed dispersal depending on the cost of among-patch pollen dispersal. Heterosis favors increased among-patch dispersal through pollen and seeds. But because these two stages inhibit each other, their joint evolution might lead to decreased seed dispersal in the presence of heterosis. Of crucial importance are the costs of dispersal.
Abstract: We present a study of the genetic diversity and structure of a tropical tree in an insular system. Santalum austrocaledonicum is endemic to the archipelago of New Caledonia and is exploited for oil extraction from heartwood. A total of 431 individuals over 17 populations were analysed for eight polymorphic microsatellite loci. The number of alleles per locus ranged from 3 to 33 and the observed heterozygosity per population ranged from 0.01 in Mare to 0.74 in Ile des Pins. The genetic diversity was lowest in the most recent islands, the Loyautes, and highest in the oldest island, Grande Terre, as well as the nearby small Ile des Pins. Significant departures from panmixia were observed for some loci-population combinations (per population F-IS = 0-0.03 on Grande-Terre and Ile des Pins, and 0-0.67 on Loyautes). A strong genetic differentiation among all islands was observed (F-ST = 0.22), and the amount of differentiation increased with geographic distance in Iles Loyaute and in Grande Terre. At both population and island levels, island age and isolation seem to be the main factors influencing the amount of genetic diversity. In particular, populations from recent islands had large average F-IS that could not be entirely explained by null alleles or a Wahlund effect. This result suggests that, at least in some populations, selfing occurred extensively. Conclusively, our results indicate a strong influence of insularity on the genetic diversity and structure of Santalum austrocaledonicum.
Abstract: Self-incompatibility (SI) systems are widespread mechanisms that prevent self-fertilization in angiosperms. They are generally encoded by one genome region containing several multiallelic genes, usually called the S-locus. They involve a recognition step between the pollen and the pistil component and pollen is rejected when it shares alleles with the pistil. The direct consequence is that rare alleles are favored, such that the S-alleles are subject to negative frequency-dependent selection. Several theoretical articles have predicted the specific patterns of polymorphism, compared to neutral loci, expected for such genes under balancing selection. For instance, many more alleles should be maintained and populations should be less differentiated than for neutral loci. However, empirical tests of these predictions in natural populations have remained scarce. Here, we compare the genetic structure at the S-locus and microsatellite markers for five natural populations of the rare species Brassica insularis. As in other Brassica species, B. insularis has a sporophytic SI system for which molecular markers are available. Our results match well the theoretical predictions and constitute the first general comparison of S-allele and neutral polymorphism.
Abstract: The existence of genetic variability for dispersal is a crucial issue for organisms facing increased habitat fragmentation and climate change. We study the genetic basis and evolutionary potential for diaspore traits related to dispersal in Centaurea corymbosa. Using diaspores collected in natural conditions in four of the six extant populations of this narrow-endemic plant species and diaspores produced in a common garden experiment, we study the variation for pappus and achene sizes, and diaspore mass. Using a sample of achenes from the common garden experiment, we find that the best predictor of terminal velocity is a linear combination of pappus length, achene width, and achene weight. We find significant differences among populations for all traits in both conditions, as well as significant differences among families within population. Although the differences among populations for some traits are not exactly the same in controlled conditions compared to natural conditions, the ranking of populations according to their mean trait values is consistent in both conditions. Our study is therefore one of the first to show a correlation between phenotypic differentiation for dispersal traits in natural conditions vs. controlled conditions. We also show evidence of genetic variation for traits commonly thought to be involved in dispersal ability, suggesting the potential for evolutionary changes following environmental change and management actions.
Abstract: 1 We use a deterministic model to explore theoretically the ecological and evolutionary relevance of plastic changes in seed dispersal along ecological succession. Our model describes the effect of changing disturbance regime, age structure, density and interspecific competition as the habitat matures, enabling us to seek the evolutionarily stable reaction norm for seed dispersal rate as a function of time elapsed since population foundation. 2 Our model predicts that, in the context of ecological succession, selection should generally favour plastic strategies allowing plants to increase their dispersal rate with population age, contrary to previous predictions of models that have assumed genetically fixed dispersal strategies. 3 More complex patterns can evolve showing periods with high production of dispersing seeds separated by periods of intense local recruitment. These patterns are due to the interaction of individual senescence with change in ecological conditions within sites. 4 Evolution of plastic dispersal strategies affects the patterns of density variation with time since foundation and accelerates successional replacement. An interesting parallel can be drawn between the evolution of age-specific dispersal rates in successional systems and the evolution of senescence in age-structured populations. 5 Seed dispersal plasticity could be a potential mechanism for habitat selection in plants and have implications for range expansion in invasive species because recently founded populations at the advancing front may show different patterns to those in the established range.
Abstract: The legume genus Medicago interacts with soil bacteria commonly referred to as rhizobia, in a nitrogen fixing symbiosis. We analysed the diversity of symbiotic association specificity among the two organisms, and its evolution in the plant genus. Nitrogen fixation tests and molecular phylogenetic reconstructions revealed that the genus Medicago includes more symbiotic specificity groups than previously suggested and that plant specificity is highly unstable and has repeatedly switched along the diversification of this genus. A phylogenetic analysis including geographical data shows that bacterial geographical diversity distribution has a strong influence on the geographic distribution of plant species and their ability to colonize new areas. Multiple other modifications of specificity occurred along the diversification of the genus, presumably due to selection for specialization to a single bacterial biovar. Codivergence between plants and bacteria may also have taken place.
Abstract: In this paper, we re-examine how heterogeneous environments can enable protected polymorphisms. Building on the classical models by Levene and Dempster of dispersal and selection in two habitats, we systematically investigate how the maintenance of polymorphisms is affected by (1) local versus global density regulation and (2) constant versus variable output from habitats to the next generation. We show that, for populations capable of habitat choice, a third independent and fundamental class of models needs to be considered. It is characterized by local density regulation (like Levene's model) and variable habitat output (like Dempster's model). Our results indicate that the conditions determining whether a system allows for protected polymorphisms differ qualitatively in the presence and absence of matching habitat choice (which occurs when individuals prefer the habitat to which they are best adapted). Without such habitat choice, the salient distinction is not between local and global density regulation, but between constant and variable habitat output. With matching habitat choice this situation is reversed. Analysis of the third class of models introduced here suggests that the joint evolution of matching habitat choice and local-adaptation polymorphism is easier than was previously thought.
Abstract: In many annual plant populations, seeds may be dormant for several seasons before they germinate. Here, we investigate the consequences of both conditional (dispersed seeds cannot enter a dormant stage) and unconditional seed dormancy on the amount and the distribution of neutral genetic diversity within and among populations. We present joint demographic and population genetics models for single and subdivided populations and derive the effective size and population differentiation at both local and metapopulation scales. We suggest that a Wahlund effect is unlikely to result from age structure alone. Furthermore, the differentiation among populations is decreased by the presence of seed banks. We also extend these models to describe monocarpic (semelparous) perennial life cycle, where the nonreproductive stages are vegetative rosettes instead of dormant seeds. The main difference between the models relies in the way the density-dependent regulation is acting. The effective size of monocarpic perennial species may be less than the census number of individuals, and among-population differentiation is always larger than in annual species. We discuss our results in the light of recent population genetics surveys of annual plants with seed banks.
Abstract: Similar patterns of dispersal and gene flow between closely associated organisms may promote local adaptation and coevolutionary processes. We compare the genetic structures of the two species of a plant genus (Roridula gorgonias and R. dentata) and their respective obligately associated hemipteran mutualists (Pameridea roridulae and P. marlothi) using allozymes. In addition, we determine whether genetic structure is related to differences in host choice by Pameridea. Allozyme variation was found to be very structured among plant populations but less so among hemipteran populations. Strong genetic structuring among hemipteran populations was only evident when large distances isolated the plant populations on which they live. Although genetic distances among plant populations were correlated with genetic distances among hemipteran populations, genetic distances of both plants and hemipterans were better correlated with geographic distance. Because Roridula and Pameridea have different scales of gene flow, adaptation at the local population level is unlikely. However, the restricted gene flow of both plants and hemipterans could enable adaptation to occur at a regional level. In choice experiments, the hemipteran (Pameridea) has a strong preference for its carnivorous host plant (Roridula) above unrelated host plants. Pameridea also prefers its host species to its closely related sister species. Specialization at the specific level is likely to reinforce cospeciation processes in this mutualism. However, Pameridea does not exhibit intraspecific preferences toward plants from their natal populations above plants from isolated, non-natal populations.
Abstract: The fine-scale pattern of cot-related paternity was characterized within a Population of the narrow-endemic model plant species, Centaurea corymbosa, using microsatellites and natural progeny arrays. We used classical approaches to assess correlated mating within sibships and developed a new method based on pairwise kinship coefficients to assess correlated paternity within and among sibships in a spatio-temporal perspective. We also performed numerical simulations to assess the relative significance of different mechanisms promoting correlated paternity and to compare the statistical properties of different estimators of correlated paternity. Our new approach proved very informative to assess which factors contributed most to correlated paternity and presented good statistical properties. Within progeny arrays, we found that about one-fifth of offspring pairs were full-sibs. This level of correlated mating did not result from correlated pollen dispersal events (i.e., pollen codispersion) but rather from limited mate availability, the latter being due to limited pollen dispersal distances, the heterogeneity of pollen production among plants, phenological heterogeneity and, according to simulations, the self-incompatibility system. We point. Out the close connection between correlated paternity and the "TwoGener" approach recently developed to infer pollen dispersal and discuss the conditions to be met when applying the latter.
Abstract: Pollen dispersal was characterized within a population of the narrowly endemic perennial herb, Centaurea corymbosa, using exclusion-based and likelihood-based paternity analyses carried out on microsatellite data. Data were used to fit a model of pollen dispersal and to estimate the rates of pollen flow and mutation/genotyping error, by developing a new method. Selfing was rare (1.6%). Pollen dispersed isotropically around each flowering plant following a leptokurtic distribution, with 50% of mating pairs separated by less than 11 m, but 22% by more than 40 m. Estimates of pollen flow lacked precision (0-25%), partially because mutations and/or genotyping errors (0.03-1%) could also explain the occurrence of offspring without a compatible candidate father. However, the pollen pool that fertilized these offspring was little differentiated from the adults of the population whereas strongly differentiated from the other populations, suggesting that pollen flow rate among populations was low. Our results suggest that pollen dispersal is too extended to allow differentiation by local adaptation within a population. However, among populations, gene flow might be low enough for such processes to occur.
Abstract: Centaurea corymbosa is an endemic plant species restricted to a 3-km(2) area in southern France. This species is known from only six. small populations that are highly differentiated genetically. Matrix models based on eight years of data (1994-2001) were used to assess the pattern of variation in the demographic vital rates of this species, and to investigate the causes of their variation. Asymptotic growth rates X varied widely between years and populations (0.613-1.424). Randomization tests were developed to test for spatial and temporal variation in the asymptotic growth rates. These tests rely on individual data on both survival and fecundity. As our demographic survey only allowed us to estimate,average fecundities, additional fecundity data collected from 1994 to 1996 were used to assess the distribution of individual fecundity expected within populations under demographic stochasticity or sampling error. Randomization tests showed that asymptotic growth rates were significantly different between populations and between years in C. corymbosa. In contrast, log-linear analysis performed only on transition data (i.e., excluding fecundity) suggested that the observed variations in transition probabilities were mostly explained by sampling error or demographic stochasticity rather than environmental stochasticity. This suggested that variations in fecundity among populations and among years may play a key role to explain temporal and spatial differences in X. Life-table response experiment analysis revealed that variations in fecundity and especially in the number of just-emerged seedlings per plant explained most of the observed variance in X. Spatial and temporal variations were detected for most lower-level vital rates, but causal factors that may account for these patterns a. re still unknown: no effect of genetic diversity was detected on the dynamics of the species, and correlation between lower-level vital rates and climatic data did not reveal any clear trends. Our study emphasizes the need to conduct long-term demographic surveys and to collect individual fecundity data to get more insights into the causes of variation of the demographic behavior of C. corymbosa.
Abstract: Marsilea strigosa (Marsileaceae, Pterydophyta) is a rare water fern found in the Mediterranean basin, in temporary flooded habitats only. We analyzed the level and the distribution of genetic variation at seven microsatellite loci, both at the Mediterranean scale and at a narrower scale within a highly fragmented French metapopulation. Genetic diversity among individuals within each pond suggests that M. strigosa reproduces predominantly through selfing. The very high population differentiation at the Mediterranean scale indicates that gene flow (if any) is highly restricted, Similar differentiation is also found at the scale of a single metapopulation, The distribution of multilocus genotypes suggests that the genetic variation in this species is maintained mainly through the interplay of mutation and low recombination.
Abstract: In order to elucidate the mechanisms underlying the large amount of RAPD polymorphism found in 1990 in a population of the selfing; annual Medicago truncatula GAERTN. (Fabaceae), we have analysed most of the individuals (n = 363) from the same population 6 years later using microsatellite loci. We confirm the result of the earlier study, namely that this population is very polymorphic and highly subdivided, with approximately 37% of the variance distributed among subpopulations, only 50 m apart one from another. We use standard F-statistics analyses, linkage disequilibria, minimum spanning network, multilocus assignment tests and spatial autocorrelation analyses to test the hypotheses that spatial structure and outcrossing events are involved in maintaining the large amount of genetic diversity at the level of each subpopulation. Interestingly, fine-scale spatial structure could be observed in only one subpopulation suggesting that other mechanisms are acting elsewhere. To the best of our knowledge, this is the first study of fine spatial genetic structure in a predominantly selfing species.
Abstract: The viability of a metapopulation of the rare, cliff-dwelling Centaurea corymbosa depends critically on the persistence of extant local populations, as it has a very low colonising ability. The variation in components of seed production in space and time was investigated and compared with the variation of supposedly causal plant and population traits. Temporal variation in the number of capitula per plant and in the number of ovules per capitulum were related to resource limitation between years and flowering periods, respectively. The seed-to-ovule ratio also varied widely, because of variation in fertilization rate, and, to a lesser extent, in abortion rate; predation being negligible. Isolated individuals had lower fertilization rates than individuals within dense patches. But despite a high density, the smallest studied population showed the lowest fertilization rates and highest abortion rates. This Allee effect on seed production represents a possible threat for small and/or low-density populations. Reinforcing populations of low density and supplying the smallest populations with new genotypes, by seed deposition into clefts of rocks, are suggested as possible conservation management strategies for C. corymbosa. (C) 2001 Elsevier Science Ltd. All rights reserved.
Abstract: The evolution of dispersal is investigated in a landscape of many patches with fluctuating carrying capacities and spatial heterogeneity in temporal fluctuations. Although asynchronous temporal fluctuations select for dispersal, spatial heterogeneity in the distribution of fluctuating environmental variables selects against it. We find evolutionary branching in dispersal rate leading to the evolutionarily stable coexistence of a high- and a low-dispersal phenotype. We study how the opposing forces of selection for and against dispersal change with the relative size and the environmental qualities of the source and sink habitats. Our results suggest that the evolution of dispersal dimorphism could be a first step towards speciation and local adaptation.
Abstract: Two experiments were set up to investigate how to maintain or create genetic diversity in artificial or managed populations of plants. Using Arabidopsis thaliana, we established 18 metapopulations of 20 populations each, all with the same initial genetic composition. We tested the effects of the population size, the artificial selection regime and the extinction/recolonisation regime. We report, the results of the first four generations of evolution for a trait under selection (precocity) and for allozyme diversity. As expected, overall diversity decreased in each metapopulation, and differentiation among populations increased. As expected, the differentiation was weaker for larger population sizes and in the treatment with extinction and recolonisation with no bottleneck. Artificial selection was effective because the life cycle duration was much reduced. However, most of the reduction occurred during the first generation. We observed all increase of one allele at the LAP-2 locus in all metapopulations, breaching neutral assumptions for this locus. Finally, the selection regime made little difference for small population sizes, whereas large metapopulations were more differentiated when artificial selection was heterogeneous among populations. Altogether, our results agree with theoretical exportations, and provide some new results, which could not have been anticipated. In particular, the overall decrease in genetic diversity was very large (of the order of 20% in 4 generations) even for metapopulations of 2000 individuals.
Abstract: In order to detect the evolutionary potential of two endangered species, Brassica insularis (Brassicaceae) and Centaurea corymbosa (Asteraceae), within and among-population genetic variation for both quantitative traits and allozymic markers was examined. Four populations of each species were studied, representing a large proportion of extant populations. High values of theta (ST) (0.213 and 0.364 for B. insularis and C. corymbosa respectively) suggested that low amounts of gene flow occur among the study populations. In each species, the genetic distance based on allozymes (estimated by the ratio (theta (ST)/1-theta (ST))) was positively correlated with the geographical distance, indicating isolation by distance. In contrast to previous studies in either outcrossing or selfing plant species, and especially for B. insularis, population differentiation for quantitative traits (theta (ST)) was generally found lower than differentiation for allozymes (theta (ST)), suggesting that the populations studied were experiencing similar selective forces acting upon the quantitative traits measured. Such forces would be strong enough to counteract local genetic drift. Interestingly, for both species theta (ST)'s were statistically independent of geographical distance, in contrast to the marginally significant positive isolation by distance shown by theta (ST). Altogether, these results suggest that theta (ST)'s might not always be used as conservative estimates of Q(ST')s, and might instead overestimate the evolutionary potential of endangered species. This would be especially expected in narrow-endemic species, whose ecological niche is often so restricted that indeed homogeneous selective forces are likely to occur, whereas small population sizes and restricted dispersal are likely to produce strong differentiation for :neutral variation. In fact, knowledge of both neutral and quantitative diversity patterns allows identification of those traits undergoing natural selection, and could be useful in designing reinforcement or reintroduction programs. However, this approach might have limitations too, in the presence of outbreeding depression due to locally coevolved gene complexes, (C) 2001 Elsevier Science Ltd. All rights reserved.
Abstract: Absence of enzymatic polymorphism in the endangered species of fern Marsilea strigosa prompted us to develop microsatellite loci in this species. We used an enriched partial genomic library, from which six polymorphic microsatellite loci were obtained. From a previous attempt to develop microsatellite markers with standard techniques, a single long and polymorphic locus was obtained. This highlights the benefits from using enrichment techniques in those species where microsatellite loci may be rare in the genome.
Abstract: Self-incompatibility (SI) is a widespread mechanism that prevents inbreeding in flowering plants. In many species, Sl is controlled by a single focus (the S locus) where numerous alleles are maintained by negative frequency-dependent selection. Inbreeding depression, the decline in fitness of selfed individual, compared to outcrossed ones, is an essential factor in the evolution of SI systems. Conversely, breeding systems influence levels of inbreeding depression. Little is known about the joint effect of SI and drift on inbreeding depression. Here we studied, using a two-locus model, the effect of SI (frequency-dependent selection) on a locus subject to recurrent deleterious mutations causing inbreeding depression. Simulations were performed to assess the effect of Population size and linkage between the two loci oil the level of inbreeding depression and genetic load. We show that the sheltering of deleterious alleles linked to the S locus strengthens inbreeding depression in small populations. We discuss the implications of our results for the evolution of SI systems.
Abstract: Since the origin of life, new species have kept arising while others were disappearing. The new extinction crisis we are now facing, the sixth one, is mainly due to Humans. Habitat loss, introduction of exotic species, pollution are the most important current causes of extinction. Once a species has started to decline, demographical, genetical or environmental: stochastic processes may trap it in a vortex towards extinction. The study of rare species gives new insights into the dynamics of these processes and the evolutionary processes involved. Indeed, there is also an << evolutionary vortex >> that may lead to extinction. A species which is highly adapted to a particular habitat is more vulnerable to habitat loss than a generalist species. As patches of suitable habitats disappear, natural selection in such species favors genes that decrease dispersal propensity. This, in turn, will usually select for increased local adaptation. Natural selection: might also lead to the maintenance of mating systems such:as self-incompatibility, that are particularly deleterious in small populations of rare species.
Abstract: Centaurea corymbosa Pourret (Asteraceae) is a narrow endemic species known only from six populations located in a 3-km(2) area in the south of France. Earlier field experiments have suggested that pollen and seed dispersal were highly restricted within and among populations. Consistent with the field results, populations were highly differentiated for five allozyme loci and among-population variation fitted an isolation-by-distance model. In the present study, we investigated the genetic structure of C. corymbosa using six microsatellite loci. As with allozymes, microsatellites revealed no within-population structure and a large differentiation among populations. However, allozyme loci were less powerful than microsatellites in detecting the extent of gene flow assessed by assignment tests. The patterns of structuration greatly varied among loci for both types of marker; we suggest that differences in single-locus pattern could mainly be an effect of stochastic variation for allozymes and an effect of variation in mutation rate for microsatellites. In contrast to the multilocus results, the two most polymorphic microsatellite loci did not show any isolation-by-distance pattern. Our results suggest that highly variable loci might not always be the best suited markers to quantify levels of gene flow among populations.
Abstract: The evolutionary consequences of changes in landscape dynamics for the evolution of life history syndromes are studied using a metapopulation model. We consider in turn the long-term effects of a change in the local disturbance rate, in the maximal local population persistence, in habitat productivity, and in habitat fragmentation. We examine the consequences of selective interactions between dispersal and reproductive effort by comparing the outcome of joint evolution to a situation where the species has lost the potential to evolve either its reproductive effort or its dispersal rate. We relax the classical assumption that any occupied site in the metapopulation reaches its carrying capacity immediately after recolonization. Our main conclusions are the following: (1) genetic diversity modifies the range of landscape parameters for which the metapopulation is viable, but it alters very little the qualitative evolutionary trends observed for each trait within this range. Although they are both part of a competition/colonization axis, reproductive effort and dispersal are not substitutable traits: their evolution reflects more directly the change in the landscape dynamics, than a selective interaction among them. (2) no general syndrome of covariation between reproductive effort and dispersal can be predicted: the pattern of association between the two traits depends on the type of change in landscape dynamics and on the saturation level. We review empirical evidence on colonizer syndromes and suggest lines for further empirical work.
Abstract: Earlier models on the evolution of dispersal have suggested that evolutionarily stable dispersal rates should increase with the frequency of local extinctions. Most metapopulation models assume site saturation (i.e., no local population dynamics), yet the majority of species distributed as metapopulations rarely attain carrying capacity in all occupied patches. In this article, we relax this assumption and examine the evolutionarily stable dispersal rate under nonsaturated but still competitive demographic conditions. Contrary to previous predictions, we show that increasing local extinction rates may allow decreasing dispersal rates to evolve.
Abstract: 'Conditions of persistence or extinction of a metapopulation of a colonizing annual species are studied in a heterogeneous landscape, a mixture of two elementary landscapes. An elementary landscape is a landscape whose age-structure is described by only one transition matrix, giving the probability for a site to be disturbed, or to follow the process of succession. We first provide an analytical study of the range of dispersal rates that allow metapopulation persistence in an elementary landscape. Second, conditions for metapopulation persistence in a heterogeneous landscape are derived from results obtained in each elementary landscape. Three cases are distinguished. If the two ranges of dispersal rates defined in each elementary landscape overlap, the metapopulation persists in any mixture of the elementary landscapes. If these two dispersal rates ranges are non-overlapping, either the metapopulation goes extinct for some values of the proportion of the elementary landscapes, or two discontinuous ranges of dispersal rates allow the metapopulation persistence. The consequences of these results are discussed in terms of landscape management. In particular, it is shown that under some conditions, a rapid change in environment (from one elementary landscape to another one) might less often lead to metapopulation extinction than a slower change. (C) 1999 Academic Press.
Abstract: Amounts of genetic variability, genetic differentiation among taxa and populations, and population sizes were studied in five populations of Centaurea maculosa ssp. maculosa (a widespread taxon), all six populations of C. corymbosa (a narrowly endemic species), and the single population of C. maculosa ssp. albida Seventeen isozyme loci were studied, of which nine were polymorphic. Results suggest that C. corymbosa and C, maculosa ssp, albida are likely derived from C. maculosa ssp. maculosa because the former represent a sample of tbe diversity of the latter, The percentage of polymorphic loci and Nei's genetic diversity were positively and significantly correlated with population size over all populations, but not within each taxon. Populations of both the widespread C. maculosa ssp. maculosa and the rare C. corymbosa were strongly differentiated: overall, F-ST values were 0.26 and 0.34, respectively. Differentiation among populations of different taxa was of the same order of magnitude as that observed among populations within taxa. Nevertheless, significant differentiation among the three taxa was found by a hierarchical analysis of variance on allele frequencies. We suggest that bottlenecks or founder effects associated with colonization events and ecological specialization in some populations of C. maculosa ssp. maculosa have led to new taxa such as C. corymbosa and C. maculosa ssp. albida This may be a direct consequence of the particularly strong differentiation among populations of the widespread C, maculosa ssp. maculosa Our study highlights the utility of considering closely related widespread taxa in order to understand the population biology and evolution of rare species, as well as to design proper management programs.
Abstract: We performed a molecular phylogenetic study based on the nuclear ribosomal internal and external transcribed spacer (ITS and ETS). Thirty-one annual Medicago species were included in the study, representing more than half of the genus and 85% of the annuals of the genus. Major incongruences were found between phylogenetic relationships and morphological classification of the genus. Morphological and cytological traits were mapped onto the phylogeny. The most parsimonious reconstruction suggested an ancestral spiny state and a recurrent transition from spiny to spineless state. From the ancestral state of 2n = 16, three loss events of chromosomes must have occurred leading to the same specific number of 14 chromosomes whereas species having 30 chromosomes form a monophyletic clade. (C) 1998 Academic Press.
Abstract: We present a molecular phylogeny including most species of the genus Medicago L. (Fabaceae). Based on the consensus of the 48 most parsimonious trees, life-history and mating-system characters are mapped, and a putative history of the genus is suggested. The most parsimonious reconstruction suggests an ancestral annual and selfing state, and recurrent evolution towards perenniality and outcrossing. Based on theoretical predictions and classical hypotheses of the history of the genus, different assumptions about the ancestral state and different weighting schemes of evolution between the character states are made. Assuming an outcrossing, perennial ancestral state (partly supported by morphological features) does not fundamentally change the reconstruction. To meet theoretical expectations, various weighting schemes favouring evolution towards annuality and selfing are applied. Influence and validity of such weighting schemes are discussed with regard to other studies.
Abstract: We have estimated the potential phylogenetic utility of the ribosomal external transcribed spacer (ETS) from the nuclear ribosomal region. The ETS was sequenced from 13 annual Medicago (Fabaceae) species upstream a highly conserved motive which was found among many different organisms. In the genus Medicago, the ETS was found to evolve 1.5 times faster than the internal transcribed spacer and to be 1.5 times more informative. Reduced ribosomal maturation process constraints on ETS are proposed to explain the different evolutionary rates between the two spacers. Maximal phylogenetic resolution and support was obtained when the two spacers were analyzed together. No incongruence between the two spacers was found and ETS appears to be a valuable source of information for solidifying ITS plant phylogeny. The phylogeny obtained in Medicago suggests that none of the three subsections included in the study is monophyletic.
Abstract: About 40% (alpha = 0.05) of the PCR-derived markers scored in a Medicago truncatula and M. tornata intraspecific cross departed from Mendelian expectations at alpha = 0.05. This proportion is among the highest ever documented in the literature, notably for intraspecific crosses. Estimations of DNA amount were also implemented for the parental genotypes or parental lines, and significant variations were observed. Our results suggest that the parental genotypes have diverged for quite a while, and we propose that the level of distortion we documented is correlated with the genome size difference we measured.
Abstract: Centaurea corymbosa (Asteraceae) is endemic to a small area (less than or equal to 3 km(2)), and <500 individuals reproduce in any given year, Nevertheless, enzyme polymorphism was found within and among the six local extant populations, the most distant at 2.3 km, Levels of gene flow among populations and seed and pollen dispersal data indicated very low dispersal capacity, Rarity of long distance dispersal events coupled with traits such as prolonged juvenile period, monocarpy, and self-incompatibility precludes the establishment of new populations and thus the evolution toward colonization ability through increased dispersal rate, polycarpy, or self-compatibility, The species thus appears to be trapped on an evolutionary dead-end toward extinction, even though, from a preliminary introduction experiment, we conclude that several nearby unoccupied sites would be suitable for the species.
Abstract: Using a metapopulation model, we study how local extinctions, limited population life span, and local demographic disequilibrium affect the evolution of the reproductive effort in a species with overlapping generations but no senescence. We show that in a metapopulation with saturation of all sites and an infinite deme maximal life span (no succession), local extinctions simply constitute an additional source of extrinsic mortality. When either the hypothesis of an infinite deme maximal life span or the saturation hypothesis is relaxed, nontrivial predictions arise. In particular, we find interactions between the evolutionarily stable reproductive effort strategy and the demographic dynamics in the metapopulation. We predict that larger reproductive effort may be selected for in habitats of poorer productivity, contrary to what would be predicted in a single population. Also, we predict that higher dispersal rates should favor selection for lower reproductive efforts. However, metapopulation parameters that favor high dispersal rates also favor larger reproductive efforts. Conflicting selection pressures in the metapopulation also allow maintaining evolutionarily stable polymorphism between a low and high reproductive effort for particular trade-offs between survival and fecundity.
Abstract: In this paper we compare mean values, heritability estimates, coefficient of genetic variation, and genetic correlations among several fitness components of two natural populations of a selfing plant species, Medicago truncatula L. It is shown that the population that had been found most polymorphic for molecular markers in a previous study was also the most variable for quantitative characters. Depending on the traits, the larger heritabilities in this population were due to either larger coefficients of genetic variances or smaller coefficients of environmental variances. Whereas genetic and phenotypic correlation matrices were very similar within each population, they were quite different between populations. In particular, although a positive correlation between age and size at maturity was found in both populations, the correlation between age at maturity and reproductive success was negative in the more variable population (late flowering plant, with a larger size at flowering, produced fewer pods), whereas no correlation was observed in the less variable population. We suggest that while in the less variable population all individuals have a high reproductive effort, several strategies coexist in the more variable population, with some early-flowering genotypes showing a high reproductive effort and other late-flowering genotypes showing a larger competitive ability through increased vegetative growth.
Abstract: Two populations of the selfing annual Medicago truncatula Gaertn. (Leguminoseae), each subdivided into three subpopulations, were studied for both metric traits (quantitative characters) and genetic markers (random amplified polymorphic DNA and one morphological, single-locus marker). Hierarchical analyses of variance components show that (1) populations are more differentiated for quantitative characters than for marker loci, (2) the contribution of both within and among subpopulations components of variance to overall genetic variance of these characters is reduced as compared to markers, and (3) at the population level, within population structure is slightly but not significantly larger for markers than for quantitative traits. Under the hypothesis that most markers are neutral, such comparisons may be used to make hypotheses about the strength and heterogeneity of natural selection in the face of genetic drift and gene flow. We thus suggest that in these populations, quantitative characters are under strong divergent selection among populations, and that gene flow is restricted among populations and subpopulations.
Abstract: Using RAPD markers and one morphological marker, we studied the among- and within-population structure in a selfing annual plant species, Medicago truncatula GAERTN. About 200 individuals, sampled from four populations subdivided into three subpopulations each, were scored for 22 markers. It was found that the within-population variance component accounted for 55% of the total variance, while the among-population variance component accounted for 45%. Eighteen percent of the total variance was due to within-population structure (i.e., among subpopulations). Thus, 37% of the total variance was within subpopulations. Using a multilocus approach, it was found that no multilocus genotype was common to two populations. Two of the four studied populations were composed of few (less than or equal to 6) multilocus genotypes, whereas the other two had many (greater than or equal to 15) multilocus genotypes. In the most polymorphic population (37 genotypes), only one genotype was found to be common to two subpopulations. Resampling experiments show that, depending on the population, three to 16 polymorphic loci were necessary and sufficient to score all multilocus genotypes in the population. When these data are compared to published results, it appears that on some occasions, the number of genotypes per population of selfing species might be larger than would be expected from the sole consideration of effective population size. The large within-subpopulation genetic variance observed in some populations could be explained by either small neighborhood sizes within subpopulations, or by outcrossing following migration through seed and/or pollen.
Abstract: In several reciprocal cross-infection experiments parasites were found to be significantly more adapted to their local host populations than to hosts from distant populations. We developed a metapopulation model, taking explicit account of both population densities and gene frequencies, to determine the influence of ecological and genetical parameters on the local adaptation of the parasites and on the spatial distribution of resistance and virulence genes. Our results point to the predominant effect of ecological parameters such as parasite growth rate and host and parasite migration rates on coevolutionary outcomes. In particular, the parasites are more likely to be adapted to their local host population than to allopatric hosts when the parasite migration rate is larger than the host migration rate. The opposite should be observed whenever hosts migrate more than parasites.
Abstract: A Markovian extinction model that takes into account age structure of local populations allows consideration of the effects of demography and successional dynamics on the evolution of migration. Analytical expressions for the evolutionarily stable (ES) rates of dispersal are given for cases in which newly recolonized sites attain carrying capacity within a single season. Using a low-fecundity numerical model, we find that an increase of the level of site saturation increases the dispersal rate. Ecological successions and unequal local extinction rates between newly colonized sites and established populations strongly affect the ES dispersal rate. The frequency of genetic modifiers that enhance the rate of dispersal evolves negative correlations with deme age, with high-migration genotypes predominant among colonizers while progressively declining in frequency as a deme ages. This suggests that between-deme selection (colonization) favors migrants while within-deme selection favors low dispersers, which allows the coexistence of types with different dispersal rates. Because of the interaction between the two levels of selection, the relation between the ES dispersal rate and the deme maximal lifetime is nonmonotone. We suggest that life-history traits other than dispersal might also experience antagonistic selective forces at the between- and within-deme levels.
Abstract: We have investigated the relationship between phenotypic and genetic correlations among a large number of quantitative traits (36) in three different environments in order to determine their degree of disparity and whether phenotypic correlations could be substituted for their genetic counterparts whatever the environment. We also studied the influence of the environment on genetic and phenotypic correlations. Twenty accessions (full-sib families) of Medicago lupulina were grown in three environments. In two of these two levels of environmental stress were generated by harvesting plants at flowering and by growing plants in competition with barley, respectively. A third environment, with no treatment, was used as a control with no stress. Average values of pod and shoot weight indicate that competition induces the highest level of stress. The genetic and phenotypic correlations among the 36 traits were compared. Significant phenotypic correlations were obtained easily, while there was no genetic variation for 1 or the 2 characters being correlated. The large positive correlation between the genetic and phenotypic correlation matrices indicated a good proportionality between genetic and phenotypic correlations matrices but not their similarity. In a given environment, when only those traits with a significant genetic variance were taken into account, there were still differences between genetic and phenotypic correlations, even when levels of significance for phenotypic correlations were lowered. Consequently, it is dangerous to substitute phenotypic correlations for genetic correlations. The number of traits that showed genetic variability increased with increasing environmental stress, consequently the number of significant genetic correlations also increased with increasing environmental stress. In contrast, the number of significant phenotypic correlations was not influnced by the environment. The structures of both phenotypic and genetic matrices, however, depended on the environment, and not in the same way for both matrices.
Abstract: In this note, we suggest that kin-selection could be the main selective force maintaining nodulation and nitrogen fixation in Rhizobium bacteria. Our model stresses how the self-sacrificing individuals that form nodules gain by increasing the Row of root exudates to their relatives in a more or less continuous rhizospheric population.
Abstract: We find the evolutionarily stable allocation of reproductive effort in both hermaphroditic and dioecious perennial plants. The model focuses on the effect of pollen availability on the allocation of reproductive effort and on sex allocation. The model shows that the effect of pollen limitation depends on the detailed assumptions made about the delivery of pollen to individual plants. We consider two extreme models: the pollen-pool model, which assumes that pollen from different individuals is mixed before being delivered, and the single-donor model, which assumes that pollen delivered to each individual comes from only one pollen donor. In the pollen-pool model, a reduction of pollen availability is equivalent to an overall reduction in fertility, while, in the single-donor model, the results are more complex and depend on the sensitivity of fertility to changes in pollen availability. The results for hermaphroditic and dioecious species are quite similar once the sex ratio of the dioecious species is taken into account.
Abstract: The adaptation to a variable environment has been studied within soft and hard selection frameworks. It is shown that an epistatically determined habitat preference, following a Markovian process, always leads to the maintenance of an adaptive polymorphism, in a soft selection context. Although local mating does not alter the conditions for polymorphism maintenance, it is shown that, in that case, habitat selection also leads to the evolution of isolated reproductive units within each available habitat. Habitat selection, however, cannot evolve in the total absence of adaptive polymorphism. This represents a theoretical problem for all models assuming habitat selection to be an initially fixed trait, and means that within a soft selection framework, all the available habitats will be exploited, even the less favourable ones. On the other hand, polymorphism cannot be maintained when selection is hard, even when all individuals select their habitat. Here, the evolution of habitat selection does not need any prerequisite polymorphism, and always leads to the exploitation of only one habitat by the most specialized genotype. It appears then that hard selection can account for the existence of empty habitat and for an easier evolution of habitat specialization.
Abstract: Parasites reduce the reproductive output of their hosts, limit their growth, and sometimes even castrate or kill them. Under certain conditions however, a parasitized host may be better off than an uninfected one. Such 'nice' parasites have a 'pleiotropic' action on their hosts. Parasites can be pleiotropic either in space (in which case they have a beneficial effect on the host in one environment while being detrimental in another) or in time (the parasite is beneficial at one stage of the host's development and 'costly' at another stage). Such pleiotropic parasites may constitute the intermediate stage between parasitism and mutualism.
Abstract: Using a two-loci multiplicative model of resource allocation, we show how the existence of several levels of resource allocation may affect the sign of the genetic correlations between traits linked by trade-offs. Positive genetic correlations between components of fitness affected by genetic trade-offs may result from different amounts of genetic variability at the pleiotropic loci determining the allocation of resources. Thus positive genetic correlations may be obtained in the absence both of environmental variation and of differences between individuals in resource acquisition. Nevertheless, positive correlations between all components of fitness at the same time cannot be obtained without variability in the acquisition of resources.
