Developmental Instability in Fragmented Populations of Prairie Phlox: A Cautionary Tale

Abstract: Considerable attention has recently been focused on using levels of developmental instability among members of a population to detect environmental or genetic stresses on animals or plants. It is not yet clear, however, that high developmental instability in a sample of individuals always indicates environmental stress or poor genetic quality. We studied 13 fragmented populations of prairie phlox (   Phlox pilosa L.) to test the hypothesis that developmental instability should decrease with increasing population size—as expected if small populations suffer genetic problems associated with inbreeding or are exposed to more environmental stress than larger populations. We used two different measures of developmental instability, each calculated for two different traits: radial asymmetry of flowers (for petal width and petal length) and modular fluctuating asymmetry of leaves (  for leaf widths at two points along the leaf  ). There were weak but significant correlations among individuals for four of six pairwise combinations of these measures. Surprisingly, three of our four measures of developmental instability showed strong population size effects that were opposite to those expected: developmental instability increased with population size. We conclude that measures of developmental instability cannot be applied uncritically for biomonitoring without considerable knowledge of developmental mechanisms, natural history, and population biology of the species in question.     

Reproductive plasticity,oviposition site selection,and maternal effects in fragmented landscapes

Traditionally, evolutionary ecology and conservation biology have primarily been concerned with how environmental changes affect population size and genetic diversity. Recently, however, there has been a growing realization that phenotypic plasticity can have important consequences for the probability of population persistence, population growth, and evolution during rapid environmental change. Habitat fragmentation due to human activities is dramatically changing the ecological conditions of life for many organisms. In this review, we use examples from the literature to demonstrate that habitat fragmentation has important consequences on oviposition site selection in insects, with carryover effects on offspring survival and, therefore, population dynamics. We argue that plasticity in oviposition site selection and maternal effects on offspring phenotypes may be an important, yet underexplored, mechanism by which environmental conditions have consequences across generations. Without considering the impact of habitat fragmentation on oviposition site selection, it will be difficult to assess the effect of fragmentation on offspring fitness, and ultimately to understand the impact of anthropogenic-induced environmental change on population viability.     

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.     

An experimental test of alternative population augmentation scenarios

Human land use is fragmenting habitats worldwide and inhibiting dispersal among previously connected populations of organisms, often leading to inbreeding depression and reduced evolutionary potential in the face of rapid environmental change. To combat this augmentation of isolated populations with immigrants is sometimes used to facilitate demographic and genetic rescue. Augmentation with immigrants that are genetically and adaptively similar to the target population effectively increases population fitness, but if immigrants are very genetically or adaptively divergent, augmentation can lead to outbreeding depression. Despite well‐cited guidelines for the best practice selection of immigrant sources, often only highly divergent populations remain, and experimental tests of these riskier augmentation scenarios are essentially nonexistent. We conducted a mesocosm experiment with Trinidadian guppies (Poecilia reticulata) to test the multigenerational demographic and genetic effects of augmenting 2 target populations with 3 types of divergent immigrants. We found no evidence of demographic rescue, but we did observe genetic rescue in one population. Divergent immigrant treatments tended to maintain greater genetic diversity, abundance, and hybrid fitness than controls that received immigrants from the source used to seed the mesocosms. In the second population, divergent immigrants had a slightly negative effect in one treatment, and the benefits of augmentation were less apparent overall, likely because this population started with higher genetic diversity and a lower reproductive rate that limited genetic admixture. Our results add to a growing consensus that gene flow can increase population fitness even when immigrants are more highly divergent and may help reduce uncertainty about the use of augmentation in conservation.     

Conservation and Distribution of Genetic Variation in a Polytypic Species, the Cutthroat Trout

Abstract: The cutthroat trout (Salmo clarki) presents a series of unusual and difficult problems in conservation biology. As many as 16 subspecies have been recognized in the recent literature. The genetic distance between subspecies based upon 46 enzyme loci ranges from that usually seen between congeneric species to virtual genetic identity. Subspecies from the western portion of the range of the cutthroat trout are genetically more similar to rainbow trout (Salmo gairdneri) than they are to the other subspecies of cutthroat trout. In addition, much of the genetic variation within the west-slope cutthroat trout (S. c. lewisi) results from alleles found in only one or two local populations, but they often occur at high frequencies in those populations. Thus, preserving the genetic variation in westslope cutthroat trout entails preserving as many local populations as possible.
Captive populations of cutthroat trout present a series of opportunities and genetic problems. A number of management agencies are using captive populations to supplement and reestablish natural populations. Basic genetic principles must be understood and followed in establishing and maintaining captive populations. We describe examples of unsuccessful and successful efforts by management agencies to develop captive populations.
The greatest danger to the conservation of the cutthroat trout is introgressive hybridization among subspecies and with rainbow trout. Several factors make salmonid fishes especially susceptible to problems associated with introgressive hybridization. We conclude that biochemical analysis provides a more reliable and informative means of detecting interbreeding than morphological characters. Interbreeding between westslope and Yellowstone cutthroat trout and nonnative Salmo appears to be common and widespread throughout the natural range of these subspecies.     

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.     

进化毒理学——化学品管理的信息工具？

进化毒理学关注的是污染驱动的种群间遗传分化的驱动因子,机制和结果。焦点问题涉及作为选择压力的化学污染类型,遗传学、表观遗传学和受种群影响的人口统计学,以及可能伴随着快速适应的适宜性代价和交叉抗性。在这一领域,研究人员将环境化学、保护遗传学、种群生物学和毒理学的工具结合起来,以了解受影响群体的稳定性和健康。最近有关进化毒理学的研究展示了多样化的种群适应污染案例（青鳉鱼、绦钩虾属和食蚊鱼）。根据定义,“适应”是通过某些个体和基因型的局部丧失实现的,因此提供了生物多样性丧失的单一证据。化学品管理通常主要通过实验室研究预测评估与化学品暴露相关的风险,进化毒理学则通过提供对种群和群落的影响的直接证据补充了这种方法。最近由环境毒理学和化学学会（SETAC）工作组EVOGENERATE（进化和化学品的多代效应）发起了讨论进化毒理学研究对化学品管理的效用。为了进一步讨论,请学术界、政府和工业界三个部门各派一名代表就下列问题发表意见：[在正文中提供]     

When Are Peripheral Populations Valuable for Conservation?

A great deal of effort is spent protecting geographically peripheral populations of widespread species. We consider under what conditions it is appropriate to expend resources to protect these populations. The conservation value of peripheral populations depends upon their genetic divergence from other conspecific populations. Peripheral populations are expected to diverge from central populations as a result of the interwoven effects of isolation, genetic drift, and natural selection. Available empirical evidence suggests that peripheral populations are often genetically and morphologically divergent from central populations. The long-term conservation of species is likely to depend upon the protection of genetically distinct populations. In addition, peripheral populations are potentially important sites of future speciation events. Under some circumstances, conservation of peripheral populations may be beneficial to the protection of the evolutionary process and the environmental systems that are likely to generate future evolutionary diversity.     

Parasites as conservation tools

Parasite success typically depends on a close relationship with one or more hosts; therefore, attributes of parasitic infection have the potential to provide indirect details of host natural history and are biologically relevant to animal conservation. Characterization of parasite infections has been useful in delineating host populations and has served as a proxy for assessment of environmental quality. In other cases, the utility of parasites is just being explored, for example, as indicators of host connectivity. Innovative studies of parasite biology can provide information to manage major conservation threats by using parasite assemblage, prevalence, or genetic data to provide insights into the host. Overexploitation, habitat loss and fragmentation, invasive species, and climate change are major threats to animal conservation, and all of these can be informed by parasites.     

Relationship between Population Size and Fitness

Abstract:  Long-term effective population size, which determines rates of inbreeding, is correlated with population fitness. Fitness, in turn, influences population persistence. I synthesized data from the literature concerning the effects of population size on population fitness in natural populations of plants to determine how large populations must be to maintain levels of fitness that will provide adequate protection against environmental perturbations that can cause extinction. Integral to this comment on what has been done and what needs to be done, sThe evidence suggests that there is a linear relationship between log population size and population fitness over the range of population sizes examined. More importantly, populations will have to be maintained at sizes of >2000 individuals to maintain population fitness at levels compatible with the conservation goal of long-term persistence. This approach to estimating minimum viable population size provides estimates that are in general agreement with those from numerous other studies and strengthens the argument that conservation efforts should ultimately aim at maintaining populations of several thousand individuals to ensure long-term persistence.     

Stress-induced recombination and the mechanism of evolvability

The concept of evolvability is controversial. To some, it is simply a measure of the standing genetic variation in a population and can be captured by the narrow-sense heritability (h²). To others, evolvability refers to the capacity to generate heritable phenotypic variation. Many scientists, including Darwin, have argued that environmental variation can generate heritable phenotypic variation. However, their theories have been difficult to test. Recent theory on the evolution of sex and recombination provides a much simpler framework for evaluating evolvability. It shows that modifiers of recombination can increase in prevalence whenever low fitness individuals produce proportionately more recombinant offspring. Because recombination can generate heritable variation, stress-induced recombination might be a plausible mechanism of evolvability if populations exhibit a negative relationship between fitness and recombination. Here we use the fruit fly, Drosophila melanogaster, to test for this relationship. We exposed females to mating stress, heat shock or cold shock and measured the temporary changes that occurred in reproductive output and the rate of chromosomal recombination. We found that each stress treatment increased the rate of recombination and that heat shock, but not mating stress or cold shock, generated a negative relationship between reproductive output and recombination rate. The negative relationship was absent in the low-stress controls, which suggests that fitness and recombination may only be associated under stressful conditions. Taken together, these findings suggest that stress-induced recombination might be a mechanism of evolvability.     

Application of Reproduction Technologies to the Conservation of Genetic Resources

Abstract: Maintaining the levels of genetic variability in captive populations of endangered species is an important objective of conservation biology. Because of the generally low sizes of captive populations, genetic drift is the main cause of loss of diversity. Several simple management rules, such as equalization of contributions from parents to the next generation, are recommended for minimizing genetic drift, but it cannot be removed completely because of the unavoidable random segregation of heterozygotes. Recent advances in reproductive technology, particularly developed for mammals, are now making a reality of the possibility of using more than one cell from a single meiosis in reproduction. With this technology it is possible to reduce or even completely cancel the genetic drift caused by segregation of heterozygotes. We evaluated the theoretical benefits of the use of such technologies to conservation biology. The effective population size can be increased enormously and, consequently, the amount of drift can be greatly reduced if manipulations in reproduction are made.     

Emerging prion disease drives host selection in a wildlife population

Infectious diseases are increasingly recognized as an important force driving population dynamics, conservation biology, and natural selection in wildlife populations. Infectious agents have been implicated in the decline of small or endangered populations and may act to constrain population size, distribution, growth rates, or migration patterns. Further, diseases may provide selective pressures that shape the genetic diversity of populations or species. Thus, understanding disease dynamics and selective pressures from pathogens is crucial to understanding population processes, managing wildlife diseases, and conserving biological diversity. There is ample evidence that variation in the prion protein gene (PRNP) impacts host susceptibility to prion diseases. Still, little is known about how genetic differences might influence natural selection within wildlife populations. Here we link genetic variation with differential susceptibility of white-tailed deer to chronic wasting disease (CWD), with implications for fitness and disease-driven genetic selection. We developed a single nucleotide polymorphism (SNP) assay to efficiently genotype deer at the locus of interest (in the 96th codon of the PRNP gene). Then, using a Bayesian modeling approach, we found that the more susceptible genotype had over four times greater risk of CWD infection; and, once infected, deer with the resistant genotype survived 49% longer (8.25 more months). We used these epidemiological parameters in a multi-stage population matrix model to evaluate relative fitness based on genotype-specific population growth rates. The differences in disease infection and mortality rates allowed genetically resistant deer to achieve higher population growth and obtain a long-term fitness advantage, which translated into a selection coefficient of over 1% favoring the CWD-resistant genotype. This selective pressure suggests that the resistant allele could become dominant in the population within an evolutionarily short time frame. Our work provides a rare example of a quantifiable disease-driven selection process in a wildlife population, demonstrating the potential for infectious diseases to alter host populations. This will have direct bearing on the epidemiology, dynamics, and future trends in CWD transmission and spread. Understanding genotype-specific epidemiology will improve predictive models and inform management strategies for CWD-affected cervid populations.     

Use of Substitute Species in Conservation Biology

Abstract:  In conservation biology, researchers often want to study the reasons why an endangered population is faring poorly but are unable to study it directly for logistical or political reasons. Instead they study a species that substitutes for the one of concern in the hope that it will cast light on the conservation problem. Here we outline the assumptions underlying this approach. Substitutes can be different populations or species and may be chosen because they are similar biologically to the target or representatives of a constellation of species of which the target is one. They also may be used to develop a predictive model to which the conservation target can be related. For substitutes to be appropriate, they should share the same key ecological or behavioral traits that make the target sensitive to environmental disturbance and the relationship between population vital rates and level of disturbance should match that of the target. These conditions are unlikely to pertain in most circumstances and the use of substitute species to predict endangered populations' responses to disturbance is questionable.     

Transplantation of the Subshrub Lotus scoparius: Testing the Home-Site Advantage Hypothesis

Abstract: The long-term success of restored populations may be jeopardized by the collection locality of transplants if they are ill matched to their new environment. The home-site advantage hypothesis predicts that the relative success of introduced populations will decrease as their genetic and environmental distance to the local native population increases. We evaluated this hypothesis for a geographically variable shrub, Lotus scoparius , in southern Californian coastal sage scrub by planting two common-garden experiments with seedlings from 12 source populations. The common-garden sites differed in environment and were each home to different source populations of the two taxonomic varieties, L. s. var. scoparius or L. s . var. brevialatus . We used allozyme data from each source population to calculate genetic distances between populations, and a combination of climatic data and soil traits to calculate environmental distances. At the more mesic, coastal common garden, cumulative fitness of transplants (survival × flower production) was inversely related to genetic distance between source and resident populations. At the more xeric, inland common garden, cumulative fitness (survival × size) decreased significantly with both genetic and environmental distance after one taxonomic variety was excluded from analyses. Geographic distance was only weakly correlated with genetic distance and had little value in predicting cumulative fitness of transplants. Our data support the home-site advantage hypothesis and the idea that mis-matching source populations of these genetically differentiated seed sources may result in lowered success of restored or constructed populations. The genetic and environmental similarities of source populations should be considered when source materials are chosen for transplantation.     

Mutation and Conservation

Mutation can critically affect the viability of small populations by causing inbreeding depression, by maintaining potentially adaptive genetic variation in quantitative characters, and through the erosion of fitness by accumulation of mildly detrimental mutations. I review and integrate recent empirical and theoretical work on spontaneous mutation and its role in population viability and conservation planning. I analyze both the maintenance of potentially adaptive genetic variation in quantitative characters and the role of detrimental mutations in increasing the extinction risk of small populations. Recent experiments indicate that the rate of production of quasineutral, potentially adaptive genetic variance in quantitative characters is an order of magnitude smaller than the total mutational variance because mutations with large phenotypic effects tend to be strongly detrimental. This implies that, to maintain normal adaptive potential in quantitative characters under a balance between mutation and random genetic drift (or among mutation, drift, and stabilizing natural selection), the effective population size should be about 5000 rather than 500 (the Franklin-Soulé number). Recent theoretical results suggest that the risk of extinction due to the fixation of mildly detrimental mutations may be comparable in importance to environmental stochasticity and could substantially decrease the long-term viability of populations with effective sizes as large as a few thousand. These findings suggest that current recovery goals for many threatened and endangered species are inadequate to ensure long-term population viability.     

Conservation Implications of the Natural Loss of Lineages in Wild Mammals and Birds

Because populations in zoological parks and nature reserves often are derived from only a few individuals, conservationists have attempted to minimize founder effects by equalizing family group sizes and increasing the reproductive contributions of all individuals. Although such programs reduce potential losses of genetic diversity, information is rarely available about the actual persistence of family groups or genetic lineages in natural populations. In the absence of such data, it can be difficult to weigh the importance of human intervention in the conservation of small populations. Separate long-term studies of two mammals, the North American bison (Bison bison) and the white-nosed coati (Nasua narica), and a bird, the Acorn Woodpecker (Melanerpes formicivorus), demonstrate differential extinction of genetic lineages. Irrespective of the mechanisms affecting population structure, which may range from stochastic environmental events to such behavioral phenomena as poor intrasexual competitive abilities, our results show that lineages can be lost at rapid rates from natural populations. A survey of comparable studies from the literature indicates that the loss of matrilines over the course of the study varies from 3% to 87% in wild mammals and from 30% to 80% in birds, with several small mammals losing approximately 20% of matrilines per year of study. These lineage extinctions were not an artifact of the length of the study or the generation time of the species. Such rapid losses of lineages in less than 20-year periods in natural populations suggest that efforts to maintain maximal genetic diversity within populations may not always reflect processes that occur in the wild. Conservation biologists need to give further thought to the extent to which parity among genetic lines should be a primary goal of management of captive and small wild populations.     

Conservation Implications of Drastic Reductions in the Smallest and Most Isolated Populations of Giant Pandas

Abstract: In conservation biology, understanding the causes of endangerment is a key step to devising effective conservation strategies. We used molecular evidence (coalescent simulations of population changes from microsatellite data) and historical information (habitat and human population changes) to investigate how the most‐isolated populations of giant pandas (Ailuropoda melanoleuca) in the Xiaoxiangling Mountains became highly endangered. These populations experienced a strong, recent demographic reduction (60‐fold), starting approximately 250 years BP. Explosion of the human population and use of non‐native crop species at the peak of the Qing Empire resulted in land‐use changes, deforestation, and habitat fragmentation, which are likely to have led to the drastic reduction of the most‐isolated populations of giant pandas. We predict that demographic, genetic, and environmental factors will lead to extinction of giant pandas in the Xiaoxiangling Mountains in the future if the population remains isolated. Therefore, a targeted conservation action—translocation—has been proposed and is being implemented by the Chinese goverment.     

Genetic diversity and IUCN Red List status

The International Union for Conservation of Nature (IUCN) Red List is an important and widely used tool for conservation assessment. The IUCN uses information about a species’ range, population size, habitat quality and fragmentation levels, and trends in abundance to assess extinction risk. Genetic diversity is not considered, although it affects extinction risk. Declining populations are more strongly affected by genetic drift and higher rates of inbreeding, which can reduce the efficiency of selection, lead to fitness declines, and hinder species’ capacities to adapt to environmental change. Given the importance of conserving genetic diversity, attempts have been made to find relationships between red-list status and genetic diversity. Yet, there is still no consensus on whether genetic diversity is captured by the current IUCN Red List categories in a way that is informative for conservation. To assess the predictive power of correlations between genetic diversity and IUCN Red List status in vertebrates, we synthesized previous work and reanalyzed data sets based on 3 types of genetic data: mitochondrial DNA, microsatellites, and whole genomes. Consistent with previous work, species with higher extinction risk status tended to have lower genetic diversity for all marker types, but these relationships were weak and varied across taxa. Regardless of marker type, genetic diversity did not accurately identify threatened species for any taxonomic group. Our results indicate that red-list status is not a useful metric for informing species-specific decisions about the protection of genetic diversity and that genetic data cannot be used to identify threat status in the absence of demographic data. Thus, there is a need to develop and assess metrics specifically designed to assess genetic diversity and inform conservation policy, including policies recently adopted by the UN's Convention on Biological Diversity Kunming-Montreal Global Biodiversity Framework.