Inbreeding Depression, Environmental Stress, and Population Size Variation in Scarlet Gilia (Ipomopsis aggregata)

Despite a large body of theory, few studies have directly assessed the effects of variation in population size on fitness components in natural populations of plants. We conducted studies on 10 populations of scarlet gilia, Ipomopsis aggregata , to assess the effects of population size and year-to-year variation in size on the relative fitness of plants. We showed that seed size and germination success are significantly reduced in small populations (those 100 flowering plants) of scarlet gilia. Plants from small populations are also more susceptible to environmental stress. When plants from small and large populations were subjected to an imposed stress (combined effects of transplanting and experimental clipping, simulating ungulate herbivory) in a common garden experiment, plants from small populations suffered higher mortality and were ultimately of smaller size than plants from large populations. In addition, experimental evidence indicates that observed fitness reductions are genetic, due to the effects of genetic drift and/or inbreeding depression. When pollen was introduced from distant populations into two small populations, seed mass and percentage of germination were bolstered, while pollen transferred into a large population had no significant effect. Year-to-year variation in population size and its effects on plant fitness are also discussed. In one small population, for example, a substantial increase in size from within did not introduce sufficient new (archived) genetic material to fully overcome the effects of inbreeding depression.     

Using environmental and geographic data to optimize ex situ collections and preserve evolutionary potential

Maintenance of biodiversity through seed banks and botanical gardens, where the wealth of species’ genetic variation may be preserved ex situ, is a major goal of conservation. However, challenges can persist in optimizing ex situ collections if trade-offs exist among cost, effort, and conserving species evolutionary potential, particularly when genetic data are not available. We evaluated the genetic consequences of population preservation informed by geographic (isolation by distance [IBD]) and environmental (isolation by environment [IBE]) distance for ex situ collections for which population provenance is available. We used 19 genetic and genomic data sets from 15 plant species to assess the proportion of population genetic differentiation explained by geographic and environmental factors and to simulate ex situ collections prioritizing source populations based on pairwise geographic distance, environmental distance, or both. Specifically, we tested the impact prioritizing sampling based on these distances may have on the capture of neutral, functional, or putatively adaptive genetic diversity and differentiation. Individually, IBD and IBE explained limited population genetic differences across all 3 genetic marker classes (IBD, 10–16%; IBE, 1–5.5%). Together, they explained a substantial proportion of population genetic differences for functional (45%) and adaptive (71%) variation. Simulated ex situ collections revealed that inclusion of IBD, IBE, or both increased allelic diversity and genetic differentiation captured among populations, particularly for loci that may be important for adaptation. Thus, prioritizing population collections based on environmental and geographic distance data can optimize genetic variation captured ex situ. For the vast majority of plant species for which there is no genetic information, these data are invaluable to conservation because they can guide preservation of genetic variation needed to maintain evolutionary potential within collections.     

Invasive Microstegium populations consistently outperform native range populations across diverse environments

Plant species introduced into novel ranges may become invasive due to evolutionary change, phenotypic plasticity, or other biotic or abiotic mechanisms. Evolution of introduced populations could be the result of founder effects, drift, hybridization, or adaptation to local conditions, which could enhance the invasiveness of introduced species. However, understanding whether the success of invading populations is due to genetic differences between native and introduced populations may be obscured by origin x environment interactions. That is, studies conducted under a limited set of environmental conditions may show inconsistent results if native or introduced populations are differentially adapted to specific conditions. We tested for genetic differences between native and introduced populations, and for origin x environment interactions, between native (China) and introduced (U.S.) populations of the invasive annual grass Microstegium vimineum (stiltgrass) across 22 common gardens spanning a wide range of habitats and environmental conditions. On average, introduced populations produced 46% greater biomass and had 7.4% greater survival, and outperformed native range populations in every common garden. However, we found no evidence that introduced Microstegium exhibited greater phenotypic plasticity than native populations. Biomass of Microstegium was positively correlated with light and resident community richness and biomass across the common gardens. However, these relationships were equivalent for native and introduced populations, suggesting that the greater mean performance of introduced populations is not due to unequal responses to specific environmental parameters. Our data on performance of invasive and native populations suggest that post-introduction evolutionary changes may have enhanced the invasive potential of this species. Further, the ability of Microstegium to survive and grow across the wide variety of environmental conditions demonstrates that few habitats are immune to invasion.     

Genetic sub-structure and intermediate optimal outcrossing distance in the marine angiosperm <Emphasis Type="Italic">Zostera marina</Emphasis>

The spatial distribution of genetic variability depends on the spatial patterns of clonal and sexual reproduction, gene flow, genetic drift and natural selection. Species with restricted dispersal may exhibit genetic structuring within populations with immediate neighbours being close relatives, and may show differentiation among populations. Genetic structuring of a species may have important genetic, evolutionary and ecological consequences including distance-dependent mating success. In this study we used microsatellite markers to show that clones of Zostera marina in a population in the Ria Formosa, Portugal, were aggregated and covered distances of up to 3–4 m. Clones within 4 m of each other exhibited significant and positive coancestry values, reflecting the limited seed dispersal of this species. Hand-pollinations between near (0–10.9 m), intermediate (11–32 m) and far (15 km) individuals resulted in similar levels of seed set, although the near pollinations had higher, although not statistically significant, levels of seed abortion during maturation. Seeds from intermediate-distance pollinations had a significantly higher proportion of seeds germinate and shorter germination time than both the near and far seeds. Similarly, the average number of seedlings produced per pollination, used as an overall estimate of fitness, was significantly greater for the intermediate distance when compared to both near and far pollinations. These results suggest that the genetic structuring observed may result in both inbreeding and outbreeding depression, which gives rise to an intermediate optimal outcrossing distance.     

Fragmentation of Landscape as a Cause for Genetic Subdivision in Bank Voles

Abstract: We studied the barrier effects of various roadways on the genetic subdivision of bank vole (Clethrionomys glareolus) populations. Allele frequencies, genetic variability, and genetic distances of natural populations were calculated based on polymorphism of seven microsatellite markers. We compared bank vole populations in control areas without such barriers with animals from both sides of a country road, a railway, and a highway. Using F and R statistics, we demonstrated significant population subdivision in bank vole populations separated by the highway, but not in populations on either side of the other roadways or in the control area. Correlations between geographic and genetic distances were revealed by an extended method based on a Mantel analysis. This allowed us to measure genetic barrier effects and express them as additional geographic distances. For instance, statistically significant differences in allele frequencies in all seven loci examined existed among populations in southern Germany and Switzerland, which are separated by the Rhine River and Lake Constance. The real geographic distance between bank vole populations in Konstanz and those in Lengwil, Switzerland, was 6 km. According to this analysis the genetic barrier effect of the Rhine could be defined as an additional distance of 7.7 km. This study shows for the first time that not only old geographic barriers but also more recent fragmentation of landscape by, for example, highways has an important effect on gene flow and the genetic substructuring of populations, which should be considered in future environmental impact assessments.     

Flight distance of urban birds, predation, and selection for urban life

Numerous species have adapted to humans, especially invasive species associated with humans in towns and cities. Short flight distances of populations adapted to urban environments reflect changes in behavior and physiology, reflecting phenotypic plasticity or evolution. Here, I tested the hypothesis that the decrease in flight distance to a potential predator (an approaching human) reflected adaptation to urbanization, using a data set of flight distances of 44 common species of European birds in different stages of adaptation to urban environments. Urban populations had consistently shorter flight distances than rural populations of the same species. Variation in relative flight distance of urban populations was predicted by the number of generations since urbanization, as expected by a gradual process of adaptation. Furthermore, species with relatively large populations in urban environments would be an indication of local adaptation to urban environments. Relative flight distance of urban population was shorter for species with large populations in urban compared to rural habitats. Species that had adapted to urban environments as shown by short flight distances were less susceptible to predation by the European sparrowhawk Accipiter nisus than species with relatively long flight distances in urban populations. These findings provide evidence consistent with the hypothesis that recent changes in the tameness of urban birds, as reflected by their relatively short flight distances, is an adaptation to the novel urban environment. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Unprecedented Low Levels of Genetic Variation and Inbreeding Depression in an Island Population of the Black-Footed Rock-Wallaby

Abstract: It has been argued that demographic and environmental factors will cause small, isolated populations to become extinct before genetic factors have a significant negative impact. Islands provide an ideal opportunity to test this hypothesis because they often support small, isolated populations that are highly vulnerable to extinction. To assess the potential negative impact of isolation and small population size, we compared levels of genetic variation and fitness in island and mainland populations of the black-footed rock-wallaby ( Petrogale lateralis [Marsupialia: Macropodidae]). Our results indicate that the Barrow Island population of P. lateralis has unprecedented low levels of genetic variation (  H _e = 0.053, from 10 microsatellite loci) and suffers from inbreeding depression (reduced female fecundity, skewed sex ratio, increased levels of fluctuating asymmetry). Despite a long period of isolation ( ∼ 1600 generations) and small effective population size (  N _e ∼ 15), demographic and environmental factors have not yet driven this population to extinction. Nevertheless, it has been affected significantly by genetic factors. It has lost most of its genetic variation and become highly inbred (  F _e = 0.91), and it exhibits reduced fitness. Because several other island populations of P. lateralis also exhibit exceptionally low levels of genetic variation, this phenomenon may be widespread. Inbreeding in these populations is at a level associated with high rates of extinction in populations of domestic and laboratory species. Genetic factors cannot then be excluded as contributing to the extinction proneness of small, isolated populations.     

Adaptive phenotypic differentiation across the intertidal gradient in the alga Silvetia compressa

Populations of intertidal species span a steep environmental gradient driven by differences in emersion time. In spite of strong differential selection on traits related to this gradient, the small spatial scale over which differences occur may prevent local adaptation, and instead may favor a single intermediate phenotype, or nongenetic mechanisms of differentiation. Here I examine whether a common macroalga, Silvetia compressa, exhibits phenotypic differentiation across the intertidal gradient and evaluate how local adaptation, developmental plasticity, and maternal effects may interact to shape individual phenotypes. Reciprocal transplants of both adults and embryos showed a "home-height advantage" in two of the three populations tested. In laboratory trials, the progeny of upper-limit individuals survived exposure to air significantly better than lower-limit progeny from the same population. I compared the emersion tolerance of full-sib families generated from gametes produced in the field to those produced under common garden conditions. The relative advantage of upper-limit lineages was robust to maternal environment during gametogenesis; this pattern is consistent with genetic differentiation. The possible role of local adaptation has historically been ignored in studies of intertidal zonation. In S. compressa, phenotypic differentiation may have important consequences for vertical range, both within and among sites.     

Genetic Variation of Naturally Colonizing Wolves in the Central Rocky Mountains

Recovery of gray wolf ( Canis lupus ) populations in North America depends on minimizing human-caused mortality and enhancing migration from stable source populations to suitable habitat unoccupied by wolves. We used a combination of field observation and DNA microsatellite genotyping to examine natural wolf colonization of Glacier National Park, Montana, and surrounding lands. We found high genetic variation in the colonizing population, showing that these packs were founded by multiple, unrelated wolves from Canada. High dispersal rates, long dispersal distances, and lack of a founding population bottleneck indicate that wolves in the United States and Canada should be viewed and managed as a single population. Restoration in the United States by artificial transplants from Alberta to Yellowstone National Park and central Idaho began in 1995. The transplanted wolves will likely aid demographic recovery, but permanently retaining the high genetic variation of wolves in the United States will require assuring gene flow throughout the central Rocky Mountains.     

River restoration success depends on the species pool of the immediate surroundings

Previous studies evaluating the success of river restorations have rarely found any consistent effects on benthic invertebrate assemblages. In this study, we analyzed data from 24 river restoration projects in Germany dating back 1 to 12 years and 1231 data sets from adjacent river reaches that lie within 0-5, 5-10, and 10-15 km rings centered on the restored sites. We calculated restoration success and recolonization potential of adjacent river reaches based on stream-type-specific subsets of taxa indicative for good or bad habitat quality. On average, the restorations did not improve the benthic invertebrate community quality. However, we show that restoration success depends on the presence of source populations of desired taxa in the surrounding of restored sites. Only where source populations of additional desired taxa existed within a 0-5 km ring around the restored sites were benthic invertebrate assemblages improved by the restoration. Beyond the 5-km rings, this recolonization effect was no longer detected. We present here the first field results to support the debated argument that a lack of source populations in the areas surrounding restored sites may play an important role in the failure to establish desired invertebrate communities by the means of river restorations. In contrast, long-range dispersal of invertebrates seems to play a subordinate role in the recolonization of restored sites. However, because the surroundings of the restored sites were far from good ecological quality, the potential for improvement of restored sites was limited.     

Patterns of Genetic Diversity and Its Loss in Mammalian Populations

Abstract:  Policy aimed at conserving biodiversity has focused on species diversity. Loss of genetic diversity, however, can affect population persistence, evolutionary potential, and individual fitness. Although mammals are a well-studied taxonomic group, a comprehensive assessment of mammalian genetic diversity based on modern molecular markers is lacking. We examined published microsatellite data from populations of 108 mammalian species to evaluate background patterns of genetic variability across taxa and body masses. We tested for loss of genetic diversity at the population level by asking whether populations that experienced demographic threats exhibited lower levels of genetic diversity. We also evaluated the effect of ascertainment bias (a reduction in variability when microsatellite primers are transferred across species) on our assessment of genetic diversity. Heterozygosity did not vary with body mass across species ranging in size from shrews to whales. Differences across taxonomic groupings were noted at the highest level, between populations of marsupial and placental mammals. We documented consistently lower heterozygosity, however, in populations that had experienced demographic threats across a wide range of mammalian species. We also documented a significant ( p = 0.01) reduction in heterozygosity as a result of ascertainment bias. Our results suggest that populations of both rare and common mammals are currently losing genetic diversity and that conservation efforts focused above the population level may fail to protect the breadth of persisting genetic diversity. Conservation policy makers may need to focus their efforts below the species level to stem further losses of genetic resources.     

Nest-site selection of passerines: effects of geographic scale and public and personal information

Nest-site selection is an important determinant of individual fitness in birds. Understanding what information individuals use to choose nest sites is therefore important for understanding the evolution of nest-site selection, the dynamics of populations, and the conservation of species. We used five years of mark-recapture data for Mountain Bluebirds (Sialia currucoides) to examine how dispersal probability and nest-site selection vary with potential cues of nest-site quality. Dispersal distance between breeding seasons and nest-site selection were modeled as a function of personal reproductive success, conspecific density, conspecific reproductive success, and habitat type. Between years, the dispersal probability was related to personal reproductive success, not conspecific information, and individuals fledging fewer young dispersed longer distances. For dispersing individuals, the probability that a nest site was selected in year i was negatively related to distance from the nest site selected in year i - 1 for all age and sex classes, and positively related to conspecific density and reproductive success in year i - 1 for both second-year (SY) and after-hatch-year (AHY) females. However, nest-site selection in year i was more strongly related to conspecific density in year i- 1 for hatch-year (HY) females and was much more strongly related to the reproductive success of conspecifics in year i - 1 for AHY females. Nest-site selection of HY and AHY males was not consistently related to the metrics of conspecific information, but we suspect that relationships were obscured by competitive interactions. We found no evidence indicating that individuals respond differently to conspecific information at longer distances, suggesting that individuals limit dispersal to areas where they have prior knowledge. We predict that these patterns of nest-site selection will allow birds to loosely track nest-site quality and maintain an ideal free distribution, where average fitness is equal in all habitat types.     

Spontaneous Interspecific Hybridization and Patterns of Pollen Dispersal in Ex Situ Populations of a Tree Species (Sinojackia xylocarpa) that is Extinct in the Wild

Abstract:  For endangered plants interspecific hybridization occurring in ex situ collections may lead to failure of reintroduction actions. We used Sinojackia xylocarpa , a well documented Chinese endemic species that is extinct in the wild, as a model case to address this concern. We used paternity analyses to assess the spontaneous hybridization and patterns of pollen flow between S. xylocarpa and its congener species, S. rehderiana , in conserved populations in Wuhan Botanic Garden. Interspecific hybridization events were detected in seven out of eight maternal trees of S. xylocarpa , and an average of 32.7% seeds collected from maternal trees of S. xylocarpa were hybrids. The paternity of 93 out of 249 seedlings from S. xylocarpa assigned to S. rehderiana provided convincing evidence that spontaneous interspecific hybridization occurred extensively in the living garden collection we studied. Different patterns of pollen dispersal (predominantly short-distance vs. long-distance pollination) were observed between intra- and interspecific hybridization events in the garden. Pollen dispersal within the ex situ populations was not restricted by distance, as evidenced by a lack of significant correlations between the average effective pollen dispersal distance (δ) and the geographic distances (d1 and d2) between maternal and paternal trees. The interspecific pollen-dispersal distance ranged from 10 to 620.1 m (mean 294.4 m). Such extensive hybridization in ex situ collections could jeopardize the genetic integrity of endangered species and irrevocably contaminate the gene pool if such hybrids are used for reintroduction and restoration. We recommend strongly that measures be taken to minimize the genetic risks of this kind of hybridization, including establishing buffer zones in ex situ collections, manipulating flowering phenology, testing seed lots before use in reintroduction programs, and controlling pollination for seed purity.     

Guidelines for genetic monitoring of translocated plant populations

Plant translocation is a useful tool for implementing assisted gene flow in recovery plans of critically endangered plant species. Although it helps to restore genetically viable populations, it is not devoid of genetic risks, such as poor adaptation of transplants and outbreeding depression in the hybrid progeny, which may have negative consequences in terms of demographic growth and plant fitness. Hence, a follow-up genetic monitoring should evaluate whether the translocated populations are genetically viable and self-sustaining in the short and long term. The causes of failure to adjust management responses also need to be identified. Molecular markers and fitness-related quantitative traits can be used to determine whether a plant translocation enhanced genetic diversity, increased fitness, and improved the probability of long-term survival. We devised guidelines and illustrated them with studies from the literature to help practitioners determine the appropriate genetic survey methods so that management practices can better integrate evolutionary processes. These guidelines include methods for sampling and for assessing changes in genetic diversity and differentiation, contemporary gene flow, mode of local recruitment, admixture level, the effects of genetic rescue, inbreeding or outbreeding depression and local adaptation on plant fitness, and long-term genetic changes.     

Are direct developers more locally adapted than planktonic developers?

The hypothesis that populations of direct developers exhibit greater geographic differentiation in life history features than populations of planktonic developers, was tested with two species of grazing snails of the genus Littorina from 1986 to 1987. Littorina sitkana (direct developer) and L. scutulata (planktonic developer) coexist on sun- and wave-sheltered beaches from Alaska to Oregon, USA. Seasonal patterns in growth, survival and reproduction were monitored for samples from four geographically separated populations of each species grown in population cages at a common site, Friday Harbor, Washington, USA. The environmental and population effects on growth in the two species were determined in a four-way reciprocal transplant experiment with the same populations. Both the direct and planktonic developers exhibited geographic differentiation in life history features. Differentiation in the direct developer occurred over distances shorter than 30 km, while differentiation in the planktonic developer occurred over the 500 km distance examined (greater than their larvae would likely travel).     

MORPH—An individual-based model to predict the effect of environmental change on foraging animal populations

This paper describes an individual-based model, MORPH, that has been designed to predict the effect of environmental change on foraging animal populations. The key assumptions of MORPH are that individuals within populations behave in order to maximise their perceived fitness, but that perceived fitness may not always be positively related to the actual chances of survival and reproduction. MORPH has been parameterised for coastal birds on several European sites and predicted the effect of environmental change, caused by factors such as habitat loss, disturbance from humans and sea-level rise, on the survival and body condition of these species. However, MORPH contains a basic framework to describe animal physiology and foraging behaviour, and the distribution and abundance of the resources required by these animals. Therefore, MORPH is not restricted to coastal birds, and is potentially applicable to a wider range of systems. To be applied to a forager system, MORPH requires parameters describing (i) the distribution of the food supply and how food quality and abundance changes through time; (ii) the rate at which foragers consume food given the abundance of food and competitors; (iii) the amount of food the forager must consume each day to survive; (iv) the distribution and seasonal changes in other factors which influence the foraging behaviour and survival of foragers. The purpose of this paper is to (i) describe MORPH, (ii) give examples of its application, (iii) describe the types of systems to which MORPH can be applied, and (iv) publish its source code and a user guide.     

Experimental Evidence for Beneficial Fitness Effects of Gene Flow in Recently Isolated Populations

Abstract: A rich theory has been developed to explain the evolution of populations at equilibrium conditions of gene flow, inbreeding, and selection. There are, however, few empirical examples of the effects of gene flow into recently isolated, small populations under nonequilibrium conditions, such as are expected following population fragmentation. We studied the effects of inbreeding and gene flow in small, experimental populations of the mustard Brassica campestris ( rapa ). Replicate populations of five individuals randomly mated in a growth room received treatments of 0, 1, or 2.5 migrants each generation. Plants from the sixth experimental generation were planted in an outdoor common garden to evaluate the effects of the treatments on fitness and the distribution of phenotypic variation. Regression of six fitness components on inbreeding coefficients indicated a negative effect of inbreeding on fitness for five of these components. The 0-migrant treatment had significantly lower fitness than the migrant treatments for four of six fitness components, but fitness did not differ between the 1-migrant and 2.5-migrant treatments. Phenotypic divergence among populations decreased with an increased number of migrants. These data provide empirical evidence of the beneficial fitness effects of a small number of migrants for recently fragmented populations.     

Efficacy of created and restored nesting sites for the conservation of colonial Laridae in the South of France

By rapidly modifying key habitat components, habitat restoration is at risk of producing attractive cues for animals without providing habitats of sufficient quality. As such, individual fitness components, such as reproduction, could be reduced and restored habitats could become ecological traps. This risk notably appears by using artificial constructions in restoration projects, yet few studies have evaluated their efficacy in a robust way. We investigated this by analyzing 154 islets that were created or restored to improve the conservation status of 7 colonial Laridae species in the South of France. From 2007 to 2016, we compared occupancy dynamics and breeding parameters of these species between the restored sites and 846 unmanaged nesting sites. We also explored species’ preference for different nesting site characteristics and their respective effect on breeding parameters. Restored nesting sites were 2–9 times as attractive as unmanaged sites for all species except the Black-headed Gull (Chroicocephalus ridibundus). Colonization probability was up to 100 times higher in sites already used by other species the previous year and increased with distance to the shore until >0.2 when distance was over 250 m. Abandonment probability was 29–70% lower when breeding was successful the previous year in all species except the Sandwich Tern (Thalasseus sandvicensis). Productivity and breeding success probability were 2 times higher on managed sites. Distance from the shore was an important attractive characteristic of artificial nesting sites in all species. Other nesting site characteristics had species-specific effects on colonization, abandonment, and breeding success. Our results indicate that managed nesting sites are successful conservation tools for colonial Laridae in the Mediterranean and do not act as ecological traps. Our study showed that testing the ecological trap hypothesis is a robust way to evaluate the success of restoration projects of breeding habitats.     

When there is no escape: the effects of natural enemies on native, invasive, and noninvasive plants

An important question in the study of biological invasions is the degree to which successful invasion can be explained by release from control by natural enemies. Natural enemies dominate explanations of two alternate phenomena: that most introduced plants fail to establish viable populations (biotic resistance hypothesis) and that some introduced plants become noxious invaders (natural enemies hypothesis). We used a suite of 18 phylogenetically related native and nonnative clovers (Trifolium and Medicago) and the foliar pathogens and invertebrate herbivores that attack them to answer two questions. Do native species suffer greater attack by natural enemies relative to introduced species at the same site? Are some introduced species excluded from native plant communities because they are susceptible to local natural enemies? We address these questions using three lines of evidence: (1) the frequency of attack and composition of fungal pathogens and herbivores for each clover species in four years of common garden experiments, as well as susceptibility to inoculation with a common pathogen; (2) the degree of leaf damage suffered by each species in common garden experiments; and (3) fitness effects estimated using correlative approaches and pathogen removal experiments. Introduced species showed no evidence of escape from pathogens, being equivalent to native species as a group in terms of infection levels, susceptibility, disease prevalence, disease severity (with more severe damage on introduced species in one year), the influence of disease on mortality, and the effect of fungicide treatment on mortality and biomass. In contrast, invertebrate herbivores caused more damage on native species in two years, although the influence of herbivore attack on mortality did not differ between native and introduced species. Within introduced species, the predictions of the biotic resistance hypothesis were not supported: the most invasive species showed greater infection, greater prevalence and severity of disease, greater prevalence of herbivory, and greater effects of fungicide on biomass and were indistinguishable from noninvasive introduced species in all other respects. Therefore, although herbivores preferred native over introduced species, escape from pest pressure cannot be used to explain why some introduced clovers are common invaders in coastal prairie while others are not.