Sexual reproduction, clonal diversity and genetic differentiation in patchily distributed populations of the temperate forest herb Paris quadrifolia (Trilliaceae)

Clonal plant species have been shown to adopt different strategies to persist in heterogeneous environments by changing relative investments in sexual reproduction and clonal propagation. As a result, clonal diversity and genetic variation may be different along environmental gradients. We examined the regional and local population structure of the clonal rhizomatous forest herb Paris quadrifolia in a complex of forest fragments in Voeren (Belgium). Relationships between population size (the number of shoots), shoot density (the number of shoots per m²) and local growth conditions were investigated for 47 populations. Clonal diversity and genetic variation within and among 19 populations were investigated using amplified fragment length polymorphism markers. To assess the importance of sexual reproduction, seed set, seed weight and germination success were determined in 18 populations. As predicted, local growth conditions largely affected population distribution, size and density of P. quadrifolia. Populations occurring in moist and relatively productive sites contained significantly more shoots. Here, shoots were also much more sparsely distributed compared to populations occurring in dry and relatively unproductive sites, where shoots showed a strongly aggregated distribution pattern. Clonal diversity was relatively high, compared with other clonal species (G/N ratio = 0.43 and Simpson’s D=0.81). Clonal diversity significantly (P<0.01) decreased with increasing shoot density while molecular genetic variation was significantly (P<0.01) affected by population size and local environmental conditions. Lack of recruitment and out-competition of less-adapted genotypes may explain the decreased genetic variation in dry sites. Analysis of molecular variance revealed significant genetic variation among populations (Φ _ST=0.42, P<0.001), whereas pairwise genetic distances were not correlated to geographic distances, suggesting that gene flow among populations is limited. Finally, the number of generative shoots, the number of seeds per fruit and seed weight were significantly and positively related to population size and local growth conditions. We conclude that under stressful conditions populations of clonal forest plant species can slowly evolve into remnant populations characterized by low levels of genetic variation and limited sexual reproduction. Conservation of suitable habitat conditions is therefore a prerequisite for effective long-term conservation of clonal forest plant species.     

Analyses of spatial patterns and population processes of clonal plants

The nonrandom spatial structure of terrestrial plants is formed by ecological interactions and reproduction with a limited dispersal range, and in turn this may strongly affect population dynamics and population genetics. The traditional method of modelling in population ecology is either to neglect spatial pattern (e.g. in transition matrix models) or to do straightforward computer simulation. We review here three analytical mothods to deal with plant populations in a lattice-structured habitat, which propagate both by seeds that scatter over the whole habitat and by vegetative reproduction (producing runners, rhizomes, etc.) to neighboring vacant sites. [1]Dynamics of global and local densities: Dynamical equations of population density considering nearest-neighbor correlation (spatial clumping) are developed as the joint dynamics of global average density and local density (comparable to mean crowding) based onpair approximation. If there is a linear trade-off between seed production and vegetative reproduction, the equilibrium abundance of the population may be maximized by engaging both means of reproduction. This result is accurately predicted by the pair approximation method, but not by mean-field approximation (neglect of spatial structure). [2]Cluster size distributions: Using global and local densities obtained by pair approximation, we predicted cluster size distribution, i.e. the number of clusters of occupied sites of various sizes. [3]Clonal identity probability decreasing with distance: Multi-locus measurement of allozymes or other neutral molecular markers tells us whether or not a given pair of individuals belong to the same clone. From the pattern of clonal identity probability decreasing with the distance between ramets, we can estimate the relative importance of two modes of reproduction: vegetative propagation and sexual seed production.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

内蒙古中东部草原不同生境克氏针茅（Stipa krylovii Roshev.）种群的形态差异分析

对内蒙古中东部草原分布的克氏针茅进行了种群内和种群间的形态差异分析。结果表明：（1）种群内不同的形态性状存在不同程度的差异;（2）种群间生殖枝中的一部分形态性状以及营养枝高度和营养枝干重均存在较大差异;（3）每穗小花数、每穗籽粒数、每穗小穗数、生殖枝高、穗干重、生殖枝干重、每穗种子重、种子重/生殖枝重比等性状的变化趋势相同,均表现为随生境条件变差而增加,即在生殖上投入增加以增大适合度;（4）主成分分析表明穗长、种子重/生殖枝重比、每穗小花数、每穗籽粒数、生殖枝高、基盘长、第二芒柱长、营养枝高、千粒重、芒针长是不同种群差异的主要指标。这些形态性状的差异可以看作克氏针茅对不同生境的适应性表现。     

Recurrent fruit harvesting reduces seedling density but increases the frequency of clonal reproduction in a tropical tree

Studies on the ecological impacts of non‐timber forest products (NTFP) harvest reveal that plants are often more resilient to fruit and seed harvest than to bark and root harvest. Several studies indicate that sustainable fruit harvesting limits can be set very high (>80% fruit harvesting intensity). For species with clonal and sexual reproduction, understanding how fruit harvest affects clonal reproduction can shed light on the genetic risks and sustainability of NTFP harvest. We studied 18 populations of a gallery forest tree, Pentadesma butyracea (Clusiaceae), to test the impact of fruits harvest, climate and habitat size (gallery forest width) on the frequency of sexual or clonal recruitment in Benin, West Africa. We sampled populations in two ecological regions (Sudanian and Sudano‐Guinean) and in each region, we selected sites with low, moderate and high fruit harvesting intensities. These populations were selected in gallery forests with varying width to sample the natural variation in P. butyracea habitat size. Heavily harvested populations produced significantly less seedlings but had the highest density and proportion of clonal offspring. Our study suggests that for plant species with dual reproductive strategy (via seeds and clonal), fruit harvesting and associated disturbances that come with it can lead to an increase in the proportion of clonal offspring. This raises the issue that excessive fruit harvest by increasing the proportion of clonal offspring to the detriment of seed originated offspring may lead to a reduction in genetic diversity with consequence on harvested species capability to withstand environmental stochasticity.     

Population size and habitat quality affect genetic diversity and fitness in the clonal herb <Emphasis Type="Italic">Cirsium dissectum</Emphasis>

Remaining populations of plant species in fragmented landscapes are threatened by declining habitat quality and reduced genetic diversity, but the interactions of these major factors are rarely studied together for species conservation. In this study, the interactions between population size, habitat quality, genetic diversity and fitness were investigated in 22 populations of the clonal herb Cirsium dissectum throughout the British Isles. Regression analysis was used to identify significant factors, and a structural equation model was developed to illustrate and integrate these interactions. It was found that smaller populations (measured as the total number of plants) had lower genetic diversity (proportion of polymorphic loci), and that reduced genetic diversity (allelic richness) had a negative impact on the survival of seedlings grown under standard conditions. Habitat quality also had a large effect on C. dissectum. Unmanaged sites with tall vegetation, no bare soil and higher nutrient levels had smaller populations of C. dissectum, but flowering was promoted. Flowering was suppressed in heavily grazed sites with short vegetation. Higher levels of bare soil and phosphorus both had a positive relationship with genetic diversity, but probably for distinctly different reasons: bare soil provides safe sites for establishment, whilst phosphorus may promote flowering and improve seed germination. In order to conserve C. dissectum, management needs to maintain site heterogeneity so that C. dissectum can flower and establishment gaps are still available for seedlings; when either component is reduced, negative feedbacks through reduced genetic diversity and individual fitness can be expected. This study therefore highlights the importance of considering both conservation genetics and habitat quality in the conservation of plant species.     

Phenotypic plasticity influences the size, shape and dynamics of the geographic distribution of an invasive plant

Phenotypic plasticity has long been suspected to allow invasive species to expand their geographic range across large-scale environmental gradients. We tested this possibility in Australia using a continental scale survey of the invasive tree Parkinsonia aculeata (Fabaceae) in twenty-three sites distributed across four climate regions and three habitat types. Using tree-level responses, we detected a trade-off between seed mass and seed number across the moisture gradient. Individual trees plastically and reversibly produced many small seeds at dry sites or years, and few big seeds at wet sites and years. Bigger seeds were positively correlated with higher seed and seedling survival rates. The trade-off, the relation between seed mass, seed and seedling survival, and other fitness components of the plant life-cycle were integrated within a matrix population model. The model confirms that the plastic response resulted in average fitness benefits across the life-cycle. Plasticity resulted in average fitness being positively maintained at the wet and dry range margins where extinction risks would otherwise have been high ("Jack-of-all-Trades" strategy JT), and fitness being maximized at the species range centre where extinction risks were already low ("Master-of-Some" strategy MS). The resulting hybrid "Jack-and-Master" strategy (JM) broadened the geographic range and amplified average fitness in the range centre. Our study provides the first empirical evidence for a JM species. It also confirms mechanistically the importance of phenotypic plasticity in determining the size, the shape and the dynamic of a species distribution. The JM allows rapid and reversible phenotypic responses to new or changing moisture conditions at different scales, providing the species with definite advantages over genetic adaptation when invading diverse and variable environments. Furthermore, natural selection pressure acting on phenotypic plasticity is predicted to result in maintenance of the JT and strengthening of the MS, further enhancing the species invasiveness in its range centre.     

Vegetative reproduction and clonal diversity in Rhytidium rugosum (Rhytidiaceae, Bryopsida) inferred by morpho-anatomical and molecular analyses

Many bryophytes exhibit specific life forms that result in the formation of distinct patches. This is primarily achieved by consequent vegetative multiplication through indeterminate growth and vegetative reproduction. Little, however, is known about the underlying mechanisms and genetic patterns. In this study on vegetative multiplication in bryophytes, we apply morpho-anatomical and molecular (AFLP fingerprinting) techniques to analyze the vegetative reproduction system, clonal diversity, and habitat colonization of a pleurocarpous moss, Rhytidium rugosum (Rhytidiaceae). Morpho-anatomically, three types of vegetative diaspores are identified and illustrated: ramets, separated after decay and disintegration of older shoot parts (clonal reproduction); brood branches/branchlets; and caducous shoot apices (both vegetative reproduction s.str.). The AFLP fingerprinting of 35 samples (from two plots in Thuringia, further German populations, and from France, Russia, and Canada) resulted in the identification of three clones from the two plots, each comprising two to 15 samples with identical fingerprints. Samples from one clone occurred in both plots, thus proving dispersal of vegetative diaspores. The AFLP analysis further revealed close relationships of the plot samples, which suggest a clonal rather than generative (sexual) origin of the population.     

Population size, pollination and reproductive success in two endangered Genista species

Habitat fragmentation often leads to small and isolated plant populations as well as decreased habitat quality. These processes can fundamentally disrupt the interactions between plants and pollinators and decrease reproductive success. This concerns especially self-incompatible, non-clonal species that depend on pollination for successful reproduction.In two rare and endangered heathland plant species, Genista anglica and G. pilosa, we examined pollination and reproduction in relation to population size. Eight populations of G. anglica and ten populations of G. pilosa were surveyed in the vicinity of Bremen, NW-Germany. We counted the visits of pollinators (honeybees, bumblebees, and other insects) and determined the reproductive output of the observed shoots.Contrary to our expectation to find increased pollinator visitation rates in larger populations of both Genista species, the number of flower-visiting insects was unrelated to the number of flowering shoots. Increasing shoot length had a positive and increasing temperature a negative impact on the number of visiting honeybees and bumblebees. Despite the general absence of population size effects on pollinator numbers, the number of fruits and seeds in G. anglica increased with increasing population size. Fruit and seed set in G. pilosa were negatively related to the number of ‘other insects’. Our field observations showed that larger populations of both Genista species flowered earlier than smaller populations and much earlier than reported in the literature. Flowering in large populations therefore tends to coincide less well with pollinator abundance, and this may cause a disruption of the temporal coincidence between flowering phenology and pollinator activity.     

Quantitative genetics of seed size variation inPhlox

Summary Because seed size is often associated with survival and reproduction in plant populations, genetic variation for seed size may be reduced or eliminated by natural selection. To test this hypothesis we assessed genetic sources of variation in seed size in a population ofPhlox drummondii to determine whether genetic differences among seeds influence the size they attain. A diallel cross among 12 plants from a population at Bastrop, Texas, USA allowed us to partition variance in the mass of seeds among several genetic and parental effects. We found no evidence of additive genetic variance or dominance genetic variance for seed mass in the contribution of plants to their offspring. Extranuclear maternal effects accounted for 56% of the variance in seed mass. A small interaction was observed between seed genotype and maternal plant. Results of this study support theory that predicts little genetic variation for traits associated with fitness.     

Effects of habitat fragmentation and soil quality on reproduction in two heathland Genista species

Habitat fragmentation decreases plant population size and increases population isolation, as well as altering patterns of plant–animal interactions, all of which may reduce plant fitness. Here, we studied effects of habitat fragmentation (in terms of population size and isolation) and soil quality on the reproduction of two rare legume species, Genista anglica (13 populations) and Genista pilosa (14 populations), confined to remnants of acidic and nutrient‐poor Calluna heathlands. Single individuals of the Genista plants are impossible to distinguish; population size was therefore estimated according to the area occupied (referred to as population size hereafter). We collected soil samples in all heathland sites to determine content of soil water, C, N, P, Ca, K and Mg. In both species values of soil pH and C/N ratio, as well as content of soil P and base cations, reflected the highly acidic and nutrient‐poor environment of the heathlands. Population sizes were unrelated to soil quality. Although the two Genista species are similar in morphology and ecology, effects of explanatory variables on reproduction were largely inconsistent across species. In G. anglica, population size had a positive impact on all reproductive variables except germination rate, which, in contrast, was the only variable affected positively by population size in G. pilosa. In both species, mean total reproductive output, calculated as the product of total seed mass per shoot and total germination, increased with increasing water content and decreased with increasing P. In G. anglica, we found positive effects of the C/N ratio on all reproductive variables except mean single and total seed mass per shoot. In summary, in both species reproductive success per shoot decreased with increasing soil nutrient availability in the heathland sites. The infestation of two large populations of G. pilosa with the pre‐dispersal, seed‐predating weevil Apion compactum had no significant effect on reproduction of the populations.     

Clonal growth is enhanced in the absence of a mating morph: a comparative study of fertile stylar polymorphic and sterile monomorphic populations of Nymphoides montana (Menyanthaceae)

Background and Aims

Many aquatic species with stylar polymorphisms have the capacity for clonal and sexual reproduction and are sensitive to the balance of the two reproductive modes when there are a limited number of mating morphs within a population. This study asked how the clonal and sexual reproductive modes perform in populations that contain only a single morph and where fitness gain through sexual reproduction is rare. In clonal aquatic Nymphoides montana, polymorphic populations normally contain two mating morphs in equal frequencies. Populations are sexually fertile and appear to be maintained by pollen transfer between the two partners. However, in a monomorphic population of N. montana where mating opportunities are unavailable, female and male function is impaired and clonality maintains the population. Here, the consequences of intraspecific variation in sexuality were explored between monomorphic and polymorphic N. montana populations in eastern Australia.

Methods

Comparative measurements of male and female fertility, total dry mass and genotypic diversity using ISSR markers were made between populations with variable sexuality.

Key Results and Conclusions

Very few seeds were produced in the monomorphic population under natural and glasshouse conditions due to dysfunctional pollen and ovules. Stigma–anther separation was minimal in the monomorphic population, which may be a consequence of the relaxed selective pressures that regulate the maintenance of sexual function. However, clonal reproduction was favoured at the expense of sexual reproduction in the monomorphic population; this may facilitate the establishment of sterility throughout the population via resource reallocation or pleiotropic effects. The ISSR results showed that the monomorphic population was one large, single genotype, unlike the multi-genotypic fertile polymorphic populations. Evolutionary loss of sex in a clonal population in which a mating morph is absent was evident; under these conditions clonal growth may assure reproduction and expand the population via spreading stolons.     

Genetic Allee effects on performance, plasticity and developmental stability in a clonal plant

Negative effects of small population size on fitness, so-called Allee effects, may threaten population persistence even in intact habitat remnants. We studied genotypes of 14 isolated populations of the clonal plant Ranunculus reptans, for which molecular genetic (RAPD-) variability is higher for large than for small populations. In a competition-free greenhouse environment vegetative offspring of genotypes from large populations produced more rosettes and flowers, indicating higher fitness. Within-genotype coefficients of variation in performance traits, indicating developmental instability, were lower for genotypes from populations with higher RAPD-variability. In competition with a taller grass, we found relative reduction in leaf length less pronounced for plants from large populations, suggesting higher adaptive plasticity. Our experimental study of a plant with predominantly vegetative reproduction suggests, that negative genetic effects of recent habitat fragmentation, which so far rather were expected in plants with frequent sexual reproduction, are more severe and more common than previously acknowledged.     

Narrow climate and habitat envelope affect the survival of relict populations of a northern Arnica angustifolia

Many species require certain habitat and climate features, especially at range limits. Because of climate warming, isolated relict southern populations of arctic-alpine species could lose their viability. Arnica angustifolia is a rare, clonal northern plant which has distinct populations far away from its central distribution area. I explored its population viability in relation to habitat properties and local temperature regimes using demographic and temperature data collected during 1996-2009 from six southern populations. The data were analyzed with matrix population modeling. Both projected and stochastic population growth rates showed spatiotemporal variation, and indicated decreasing population trends. The population growth rate was most sensitive to the survival of small sized vegetative rosettes, emphasizing the importance of rosette size for fitness. High temperatures and habitat closure were the main threats to viability of Arnica. First, high temperatures during the previous and current growing seasons had a negative impact on rosette size and survival of ramets. Second, high coverage of deciduous shrubs and tall mosses decreased the survival and size of rosettes. Obviously, the viability of populations can be enhanced by habitat amelioration. Population isolation and restricted recruitment and habitat specificity decrease the possibilities for environmental tracking. Further, overgrowth will decrease the availability of growing space inside the populations. Consequently, the long term survival of southern populations in Arnica largely depends on both the ability of species to adapt consequences of changing temperature regimes and the speed of habitat deterioration.     

Negative effects of habitat degradation and fragmentation on the declining grassland plant Trifolium montanum

Changes in land use have resulted in a strong decline in the plant diversity of nutrient-poor grasslands, but little is known about the combined effects of habitat degradation and fragmentation on populations of individual species. We studied these effects on stage structure, recruitment, reproduction and offspring fitness in populations of the declining perennial grassland plant Trifolium montanum in central Germany. Density and survival probability of juvenile plants decreased with light competition, measured as leaf area index (LAI) above T. montanum plants, resulting in aged populations with few juvenile plants at unmanaged sites with higher LAI. Reproduction of T. montanum was not related to LAI, but increased strongly with local density, suggesting pollinator limitation in fragmented populations with a low density of flowering plants. In the common garden, the survival of sown offspring increased with mean seed size, whereas seed production of offspring decreased with isolation, and in strong contrast to previous studies, also decreased with size and density of the population of origin. This could be due to increased inbreeding because of pollination between closely related neighbouring plants in dense and large populations. Our results indicate that both habitat degradation and fragmentation have negative effects on populations of T. montanum, but affect different phases of the life cycle. In the short term, the effects of habitat degradation are more important than those of fragmentation, and populations of T. montanum are primarily threatened by an increase in light competition in unmanaged sites, which rapidly affects the dynamics of the populations. The observed opposite effects of habitat fragmentation on reproduction and offspring fitness indicate that the effects of population size, density and isolation on plant fitness and population viability may be complex.     

Propagule Limitation,Disparate Habitat Quality,and Variation in Phenotypic Selection at a Local Species Range Boundary

Adaptation to novel conditions beyond current range boundaries requires the presence of suitable sites within dispersal range, but may be impeded when emigrants encounter poor habitat and sharply different selection pressures. We investigated fine-scale spatial heterogeneity in ecological dynamics and selection at a local population boundary of the annual plant Gilia tricolor. In two years, we planted G. tricolor seeds in core habitat, margin habitat at the edge of the local range, and exterior habitat in order to measure spatial and temporal variation in habitat quality, opportunity for selection, and selection on phenotypic traits. We found a striking decline in average habitat quality with distance from the population core, yet some migrant seeds were successful in suitable, unoccupied microsites at and beyond the range boundary. Total and direct selection on four out of five measured phenotypic traits varied across habitat zones, as well as between years. Moreover, the margin habitat often exerted unique selection pressures that were not intermediate between core and exterior habitats. This study reveals that a combination of ecological and evolutionary forces, including propagule limitation, variation in habitat quality and spatial heterogeneity in phenotypic selection may reduce opportunities for adaptive range expansion, even across a very local population boundary.     

Differences in life history traits of relatedEpilobium species: Clonality, seed size and seed number

Small changes in morphology can affect the performance and functions of organisms and hence their ecological success. In modular constructed plants, contrasting growth strategies may be realized by differences in the spatial arrangement and size of shoots. Such differences change the way in which meristems and resources are assigned to various functions during the lifespan of a plant. If such changes include the capacity to spread clonally, sexual reproduction may also be affected. I compare patterns in vegetative growth and sexual reproductive traits in four allopatric species ofEpilobium which are sometimes considered as subspecies of a single polymorphic taxon. The four species differ in the location of the buds which annually renew the aerial shoot system.E. dodonaei andE. steveni do not spread clonally and are characterized by a shrub-like habit.E. fleischeri, a species occurring only in the Alps, andE. colchicum, which occurs in the upper region of the Caucasus mountains, both produce buds on horizontal roots or plagiotropic shoots. Both alpine species exhibiting clonal growth have smaller shoots, fewer fruits and smaller seeds than the lowland species. An intraspecific trade-off between seed number per fruit and seed mass is realized. Both alpine species produce more seeds per fruit at the expense of seed mass. The morphological relationship between the four species and their geographical distribution suggest that clonal growth inE. fleischeri (restricted to the Alps) andE. colchicum (restricted to the Caucasus) is adaptively associated with the stressful conditions of alpine habitats. Our results suggest that clonal growth is not necessarily correlated with reduced reproduction by seeds. The success of plants which are already established may largely depend on clonal spread, but the colonization of new habitats depends on the production of a large number of small seeds with high dispersability.     

Which plant traits predict species loss in calcareous grasslands with extinction debt?

Aim Habitat loss and degradation pose a major threat to biodiversity, which can result in the extinction of habitat characteristic species. However, many species exhibit a delayed response to environmental changes because of the slow intrinsic dynamics of populations, resulting in extinction debt. We assess directly the changes in habitat characteristic species composition by comparing historical (1923) and current inventories in highly fragmented grasslands. We aim to characterize the species that constitute extinction debt in European calcareous grasslands. Location Europe, Estonia, 59–60° N, 24–25° E. Methods We related eleven life‐history traits and selected habitat preferences to local extinctions of populations in grasslands where extinction debt has been largely paid. Traits were chosen to describe species dispersal and persistence abilities and were quantified from databases. Results The studied grasslands have lost 90% of their area and 30% of their characteristic plant populations in 90 years. Species more prone to local population extinction were characterized by shorter life span, self‐pollination, a lack of clonal growth, fewer seeds per shoot, lower average height, lower soil nitrogen preference and higher requirements for light, indicating a limited ability to tolerate the range of changes in biotic and abiotic conditions of the sites. Locally extinct populations were also characterized by wind‐dispersed seeds, lower seed weight and lower terminal velocity of seeds, suggesting that species strategies for long‐distance dispersal are not favoured in highly fragmented landscapes. Thus, both increased habitat isolation and decreased habitat quality are important in determining local population extinction. Main conclusions Populations more prone to local extinction were characterized by a number of life‐history traits, demonstrating a greater extinction risk for species with poorer abilities for local persistence and competition. Our results can be applied to less degraded grasslands where the extinction debt is not yet paid to determine those species most susceptible to future extinction.     

Wrong place,wrong time: climate change‐induced range shift across fragmented habitat causes maladaptation and declined population size in a modelled bird species

Many species are locally adapted to decreased habitat quality at their range margins, and therefore show genetic differences throughout their ranges. Under contemporary climate change, range shifts may affect evolutionary processes at the expanding range margin due to founder events. In addition, populations that are affected by such founder events will, in the course of time, become located in the range centre. Recent studies investigated evolutionary changes at the expanding range margin, but have not assessed eventual effects across the species' range. We explored the possible influence of range shift on the level of adaptation throughout the species' total range. For this we used a spatially explicit, individual‐based simulation model of a woodland bird, parameterized after the middle spotted woodpecker ( Dendrocopos medius) in fragmented habitat. We simulated its range under climate change, and incorporated genetic differences at a single locus that determined the individual's degree of adaptation to optimal temperature conditions. Generalist individuals had a large thermal tolerance, but relatively low overall fitness, whereas climate specialists had high fitness combined with a small thermal tolerance. In equilibrium, the populations in the range centre were comprised of the specialists, whereas the generalists dominated the margins. In contrast, under temperature increase, the generalist numbers increased at the expanding margin and eventually also occupied the centre of the shifting range, whereas the specialists were located in the retracting margins. This was caused by founder events and led to overall maladaptation of the species, which resulted in a reduced metapopulation size and thus impeded the species' persistence. We therefore found no evidence for a complementary effect of local adaptation and range shifts on species' survival. Instead, we showed that founder events can cause local maladaptation which can amplify throughout the species' range, and, as such, hamper the species' persistence under climate change.     

Population consequences of changes in an ant-seed mutualism in Sanguinaria canadensis

Summary Seed dispersal by ants was studied in three populations of the myrmecochore, Sanguinaria canadensis, located in three habitats, each of which showed a different level of disturbance. Frequency of seed removal and the distances seeds were carried by ants were related to plant density, dispersion and the relative proportions of sexual and asexual reproduction in each population. Seeds in the least disturbed habitat were removed frequently and carried, by a wide variety of ants, distances of up to 12 m. Plant density was low and clone size was small. There was a relatively low level of sexual (seed) reproduction but seeds were generally transported well beyond the boundaries of the parent clones. By contrast, at the most disturbed site, plant density was high and clone size was very large. While there was a high level of seed production, seeds were rarely moved by ants and since removal distances were short, the probability of a seed being relocated beyond the limits of the parent clone was miniscule. The third population from a habitat which was intermediate in disturbance yielded data intermediate to the others. The data show that habitat disturbance, in disrupting the ant fauna and hence the ant-seed mutualism, has profound effects upon population density, dispersion and patterns of reproduction. Density-dependent regulation of sexual output predicted, for example, by the Strawberry-Coral model (Williams 1975), is maladaptive when the antseed mutualism is disturbed. We discuss the implications of this for theoretical modeling, the significance of mutualisms and the assessment of disturbance for conservation.