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.     

Genetic Diversity of Lepilemur mustelinus ruficaudatus, a Nocturnal Lemur of Madagascar

The genetic polymorphism of natural populations of Lepilemur mustelinus ruficaudatus was studied by protein electrophoresis. We sampled blood from 72 individuals from four populations separated by geographic or anthropogenic barriers from southwestern Madagascar. Six out of 22 enzyme loci showed genetic variation with a degree of polymorphism of 0.273. The expected and observed degree of genetic heterozygosity over all loci is similar to that of other primates (H_e = 0.058, H_o = 0.036). The F-statistics revealed that the four subpopulations were similar with respect to gene structure (F_ST = 0.065, p = 0.016), but the genotypic structures within subpopulations were inconsistent with random mating. For the total of the four subpopulations the proportion of heterozygous individuals was significantly smaller than expected under random mating (F_IS = 0.373, F_IT = 0.414, p < 0.01). These results correspond closely to what is expected considering the low migration ability of individuals of L. m ruficaudatus leading to small and rather isolated inbred populations.     

Genetic Diversity, Population Size, and Fitness in Central and Peripheral Populations of a Rare Plant Lychnis viscaria

Abstract: Genetic diversity is expected to decrease in small and isolated populations as a consequence of bottlenecks, founder effects, inbreeding, and genetic drift. The genetics and ecology of the rare perennial plant Lychnis viscaria (Caryophyllaceae) were studied in both peripheral and central populations within its distribution area. We aimed to investigate the overall level of genetic diversity, its spatial distribution, and possible differences between peripheral and central populations by examining several populations with electrophoresis. Our results showed that the level of genetic diversity varied substantially among populations (  H _exp = 0.000–0.116) and that the total level of genetic diversity (mean H _exp = 0.056) was low compared to that of other species with similar life-history attributes. The peripheral populations of L. viscaria had less genetic variation (mean H _exp = 0.034) than the central ones (0.114). Analysis of genetic structure suggested limited gene flow (mean F _ST = 0.430) and high differentiation among populations, emphasizing the role of genetic drift (  N _e m = 0.33). Isolation was even higher than expected based on the physical distance among populations. We also focused on the association between population size and genetic diversity and possible effects on fitness of these factors. Population size was positively correlated with genetic diversity. Population size and genetic diversity, however, were not associated with fitness components such as germination rate, seedling mass, or seed yield. There were no differences in the measured fitness components between peripheral and central populations. Even though small and peripheral populations had lower levels of genetic variation, they were as viable as larger populations, which emphasizes their potential value for conservation.     

Application of the One-Migrant-per-Generation Rule to Conservation and Management

Abstract:  Endangered species are commonly found in several (partially) isolated populations dispersed on different fragments of a habitat, natural reserve, or zoo. A certain level of connectivity among such populations is essential for maintaining genetic variation within and between populations to allow local and global adaptation and for preventing inbreeding depression. A rule of thumb widely accepted by the conservation community is that one migrant per generation (OMPG) into a population is the appropriate level of gene flow. This rule is based on Wright's study of his island model under a long list of simplifying assumptions. I examined the robustness of the OMPG rule to the violation of each of the many assumptions, quantifying the effect with population genetics theory. I showed that, when interpreted as one effective migrant per generation, OMPG is generally valid for real populations departing from the ideal model in the discrepancies of actual (  N ) and effective (  N_e  ) population sizes and actual ( m ) and effective ( m_e  ) migration rates. I also addressed the issue of converting the effective number of migrants (  M_e= N_em_e  ) into the actual number of migrants ( M = Nm  ) of a certain age and sex. In particular, N_e < N , a case common for natural populations, did not necessarily require M > M_e to maintain a certain level of differentiation among populations. Rather, translating the elusive M_e into the manageable M depends on the specific causes (e.g., biased sex ratio, reproductive skew) that lead to N_e < N .     

Endangered Greater One-horned Rhinoceros Carry High Levels of Genetic Variation

Abstract: The population of Rhinoceros unicornis in the Chitwan Valley, Nepal, was reduced to an estimated effective population size (N_e of 21–28 individuals (60–80 total animals) in 1962. Protein electrophoresis shows that heterozygosity remains very high in this population (H_o= 9.9%) despite its near extinction. We attribute this high heterozygosity to large N_e's prior to the population bottleneck, the recent occurrence of the bottleneck, and long generation time. These results illustrate the importance of considering historical demography and life history parameters when evaluating the possible genetic effects of bottlenecks in wild populations. They also offer support to recent arguments that the erosion of genetic diversity attributed to bottlenecks may be overemphasized.     

DNA Fingerprinting Reveals a Lack of Genetic Variation in Northern Populations of the Western Pond Turtle (Clemmys marmorata)

I used DNA fingerprinting to provide the first analysis of the genetic composition of western pond turtle ( Clemmys marmorata ) populations in Washington, Oregon, and California. Populations of the western pond turtle in Washington and northern Oregon are rapidly approaching extinction. Genetic similarity within the largest northern populations, which are located inland, is high. An analysis of population substructure (F_st) revealed significant genetic divergence between inland populations, indicating a lack of dispersal and gene flow between sites. In contrast, northern coastal sites are not genetically distinct, but there are few if any viable populations remaining in this region. Genetic variability within southern California populations is a great deal higher than in northern inland sites. Similarly, a low F_st value indicated a lack of genetic differentiation between southern sites. An inter-regional analysis of population substructure (F_st = 0.24) revealed a significant degree of genetic divergence between geographical regions throughout the range. In addition, an estimate of western pond turtle phylogeny showed a genetic break in the species between northern and southern populations. Both population subdivision and phylogenetic analyses suggest a lack of appreciable gene flow between geographical regions for a considerable period of time. Genetic analyses support traditional subdivision based solely on the morphological variation of Clemmys marmorata into two subspecies: northern Clemmys marmorata marmorata and southern Clemmys marmorata pallida . Recovery of dwindling northern populations must combine demographic and genetic considerations. A first step should be to preserve local gene pools while augmenting population numbers, with the goal of preventing the extinction of this genetically and morphologically distinct subspecies.     

Estimating the Effective Population Size of Conserved Populations

Accurate estimation of effective population size is important in attempts to conserve small populations of animals or plants. We review the genetic and ecological methods that have been used to estimate effective population size in the past and suggest that, while genetic methods may often be appropriate for the estimation of N _e, and its monitoring, ecological methods have the advantage of providing data that can help predict the effect of a changed environment on N _e. Estimation of N _e, is particularly complex in populations with overlapping generations, and we summarize previous empirical estimates of N _e that used ecological methods in such populations. Since it is often difficult to assess what parameters and assumptions have been used in previous calculations, we suggest a method that provides a good estimate of N _e, makes clear what assumptions are involved, and yet requires a minimum of information. The method is used to analyze data from 14 studies. In 36% (5) of these studies, our estimate is in excellent agreement with the original, and yet we use significantly less information, in 21% (3) the original estimate is markedly lower, in 43% (6) it is markedly higher. Reasons for the discrepancies are suggested. Two of the underestimates involve a failure in the original to account for a long maturation time, and four of life overestimates involve problems in the original with the correction for overlapping generations.     

Comparisons of Genetic Diversity in the Endangered Adenophora lobophylla and Its Widespread Congener, A. potaninii

Abstract: Starch-gel electrophoresis was used to examine the levels and distribution of genetic diversity in two Adenophora species: the narrow endangered Adenophora lobophylla and its widespread congener, A. potaninii . Based on allozyme variation at 18 putative loci, we measured high levels of genetic variability both in the endangered and the widespread species, with 83.3% of the loci being polymorphic. The mean expected heterozygosity within populations (   H _ep  ) and within species (   H _es  ) were 0.234 and 0.244 for A. potaninii and were as high as 0.210 and 0.211 for A. lobophylla . There was higher differentiation among populations in A. potaninii (   F _ST = 0.155) than in A. lobophylla (   F _ST = 0.071). The high levels of genetic diversity in the present allozyme survey are consistent with the morphological variation observed in these species and may be attributed to high outcrossing rates in the Adenophora species. In addition, A. lobophylla was identified as a distinct species on the basis of Nei's genetic distances and thus should be given a high priority for protection. It is noteworthy that the endangered A. lobophylla maintains much higher genetic diversity than most endemic or narrowly distributed plant species in spite of its restricted distribution. We hypothesize that A. lobophylla has become endangered for ecological and stochastic reasons, including habitat destruction or environmental changes, mud slides, and human disturbance such as grazing and mowing. Consequently, habitat protection is of particular importance for conserving this endangered species.     

Predicting the Deleterious Effects of Mutation Load in Fragmented Populations

Abstract:  Human-induced habitat fragmentation constitutes a major threat to biodiversity. Both genetic and demographic factors combine to drive small and isolated populations into extinction vortices. Nevertheless, the deleterious effects of inbreeding and drift load may depend on population structure, migration patterns, and mating systems and are difficult to predict in the absence of crossing experiments. We performed stochastic individual-based simulations aimed at predicting the effects of deleterious mutations on population fitness (offspring viability and median time to extinction) under a variety of settings (landscape configurations, migration models, and mating systems) on the basis of easy-to-collect demographic and genetic information. Pooling all simulations, a large part (70%) of variance in offspring viability was explained by a combination of genetic structure ( F_ST ) and within-deme heterozygosity ( H_S ). A similar part of variance in median time to extinction was explained by a combination of local population size ( N ) and heterozygosity ( H_S ). In both cases the predictive power increased above 80% when information on mating systems was available. These results provide robust predictive models to evaluate the viability prospects of fragmented populations.     

Genetic Diversity in the Endangered Lily Harperocallis flava and a Close Relative, Tofieldia racemosa

We examined genetic diversity in 464 individuals of the monotypic lily Harperocallis flava in its two habitats (seepage bogs and a roadside right-of-way) and five populations of a co-occurring related lily, Tofieldia racemosa. The endangered H. flava, endemic to the Apalachicola lowlands of the Florida panhandle, was monomorphic for the 22 loci scored. In contrast, T. racemosa had a high proportion of polymorphic loci ( P_s = 68.2%; P_p = 47.7%) with moderate genetic diversity (   H_es = 0.134; H_ep = 0.114). Estimated gene flow was moderately high ( Nm = 2.07) for T. racemosa, with most (93%) of the total genetic diversity found within populations. Despite the low level of genetic divergence, some isolation by distance was detected among T. racemosa populations. Harperocallis flava and other species without discernable genetic variation pose special problems for conservation biologists because genetic criteria are not available for the development of ex situ and in situ conservation and management strategies.     

Population Demographics and Genetic Diversity in Remnant and Translocated Populations of Sea Otters

Abstract: The effects of small population size on genetic diversity and subsequent population recovery are theoretically predicted, but few empirical data are available to describe those relations. We use data from four remnant and three translocated sea otter ( Enhydra lutris ) populations to examine relations among magnitude and duration of minimum population size, population growth rates, and genetic variation. Mitochondrial (mt)DNA haplotype diversity was correlated with the number of years at minimum population size ( r _s = −0.741, p = 0.038) and minimum population size ( r _s = 0.709, p = 0.054). We found no relation between population growth and haplotype diversity, although growth was significantly greater in translocated than in remnant populations. Haplotype diversity in populations established from two sources was higher than in a population established from a single source and was higher than in the respective source populations. Haplotype frequencies in translocated populations of founding sizes of 4 and 28 differed from expected, indicating genetic drift and differential reproduction between source populations, whereas haplotype frequencies in a translocated population with a founding size of 150 did not. Relations between population demographics and genetic characteristics suggest that genetic sampling of source and translocated populations can provide valuable inferences about translocations.     

Lineage Loss in Serengeti Cheetahs: Consequences of High Reproductive Variance and Heritability of Fitness on Effective Population Size

Abstract: In natural populations, many breeders do not leave surviving offspring, and as a result many potential genetic lineages are lost. I examined lineage extinction in Serengeti cheetahs ( Acinonyx jubatus ) and found that 76% of matrilines were lost over a 25-year period. Production of future breeders was nonrandom and generally confined to a few families. Five out of 63 matrilines accounted for 45% of the total cheetah population over the course of the study. Lineage persistence is perhaps best illustrated by the variance in lifetime reproductive success ( LRS) and heritability in this parameter. In female cheetahs, variance in LRS was high, and new data show that this LRS was heritable. Variance in LRS and heritability in LRS have dramatic consequences for effective population size, N _e. I calculated N _e for cheetahs, taking into account fluctuating population size, unequal sex ratio, non-Poisson distribution of reproductive success, and heritability of fitness. The N _e was most strongly affected by variance in reproductive success and especially heritability in reproductive success. The variance N _e was 44% of the actual population size, and the inclusion of heritability further reduced N _e to only 15% of the actual population, a ratio similar to that of a social carnivore with reproductive suppression. The current cheetah population in the Serengeti is below numbers suggested by N _e estimates as sufficient to maintain sufficient genetic diversity.     

Developing a Sampling Strategy for Baptisia arachnifera Based on Allozyme Diversity

We analyzed the amount and distribution of genetic variation in Baptisia arachnifera Duncan to develop a sampling strategy for ex situ research. Baptisia arachnifera is an endangered plant species endemic to the coastal plain of Georgia (U.S.) where all populations are within 16 km of each other. A reduction in numbers of individuals has been observed during the last 50 years. Baptisia arachnifera was polymorphic at 24% of the 37 loci examined with an average of 1.32 alleles per locus. The genetic diversity index was relatively low ( H_e = 0.097) as expected for endemic species. Populations were in Hardy-Weinberg equilibrium, suggesting that the species is outcrossing. Consistent with this conclusion is the observation that the majority (approximately 90%) of the genetic variation present in the species is found within individual populations. Indirect evidence of gene flow between populations was detected (   Nm = 2.35). The close proximity of the populations and the recent reduction in population sizes suggest that the populations surveyed may be fragments of a once more continuous gene pool. Based on the observed distribution of genetic diversity among populations (G_ST = 0.096), sampling two populations would capture 99% of the allozyme diversity surveyed. Allozyme data were used to determine which 2 of the 10 populations surveyed should be sampled to maximize the ex situ conservation of genetic diversity. Although the paper-producing companies that own most of the land where Baptisia arachnifera occurs are modifying their harvesting techniques, the species could become extinct without more effective management and preservation efforts.     

Fitting probability models to population dynamics data

Methods for modeling population dynamics in probability using the generalized point process approach are developed. The life history of these populations is such that seasonal reproduction occurs during a short time. Several models are developed and analyzed. Data about two species: colonial spiders (Stegodyphus dumicola) and a migratory bird (wood thrush, Hylocichla mustelina) are used to estimate model parameters with appropriate log maximum likelihood functions. For the spiders, the model is fitted to provide evolutionary feasible colony size based on maximum likelihood estimates of fecundity and survival data. For the migratory bird species, a maximum likelihood estimates are derived for the fecundity and survival rates of young and adult birds and immigration rate. The presented approach allows computation of quantities of interest such as probability of extinction and average time to extinction.     

Sorption and desorption of pymetrozine on six Chinese soils

Pymetrozine is a selective insecticide with a unique chemical structure and mode to control hemipteran and homopteran. While pymetrozine has brought great benefits to crop production by killing insects, its residues in soil may have a detrimental effect on environment. Therefore, it is of great importance to investigate its behaviors in soil. In this study, the sorption and desorption of pymetrozine on six Chinese soils were investigated using a batch equilibrium approach to understand its mobile behavior in the soils. Both sorption and desorption isotherms of pymetrozine were in good agreement with the Freundlich model. The sorption coefficient K_F varied between 3.37 and 58.32 mL∙g⁻¹ and the sorption isotherms were nonlinear, with 1/n ranging from 0.57 to 0.91. A regression equation was proposed to predict the sorption of pymetrozine on six different soil samples: log K_F = 4.3708 − 4.5709 × log (pH in 0.01mol·L⁻¹ CaCl₂) + 0.4700 × log OC% + 0.0057 × sand (%) + 0.0022 × CEC(clay), with R² = 0.9982. The organic carbon content of soil positively affected the sorption of pymetrozine, but soil pH had a negative effect on the sorption. Additionally, effects of CaCl₂ concentration, soil to solution ratio and pesticide form were investigated. The sorption was promoted with an increase in soil to solution ratio and a decrease in CaCl₂ concentration. The possible variation of the five formulated products of pymetrozine was also investigated.     

一种用粪便样方法估计林麝种群密度的新方法

数量(密度)特征是动物种群的一个重要特征，是定点研究一个野生种群生态的一项非常重要的基础工作．其数量和年龄结构为进一步分析其种群的密度、移动、巢域和核域，尤其是为种群动态提供依据，也是珍稀动物保护和资源动物管理利用的先行工作和重要组成部份．关于野生动物种群数量的具体统计方法，有不少的文章报道，然而从动物的空间分布格局来研究大型脊椎动物却较为少见，这主要是受研究方法的限制．为此，作者对此进行了改进，提出了一种新的研究方法：临时空间样方法，并将其应用到冶勒自然保护区林麝的数量调查研究中去．结果表明：(1)集聚分布指数CA=12．09，说明冶勒自然保护区的林麝有较高的集聚度；(2)林麝粪堆在野外的分布属于聚集分布，并与负二项分布拟合较好；(3)在冶勒自然保护区，生活有62～98只左右的林麝，其密度为：1．0～1．6只／km^2．林麝在冶勒自然保护区呈现聚积分布的原因是由于环境和人类活动因素造成的．图1表2参24     

A comparison of models for predicting population persistence

We consider a range of models that may be used to predict the future persistence of populations, particularly those based on discrete-state Markov processes. While the mathematical theory of such processes is very well-developed, they may be difficult to work with when attempting to estimate parameters or expected times to extinction. Hence, we focus on diffusion and other approximations to these models, presenting new and recent developments in parameter estimation for density dependent processes, and the calculation of extinction times for processes subject to catastrophes. We illustrate these and other methods using data from simulated and real time series. We give particular attention to a procedure, due to Ross et al. [Ross, J.V., Taimre, T., Pollett, P.K. On parameter estimation in population models, Theor. Popul. Biol., in press], for estimating the parameters of the stochastic SIS logistic model, and demonstrate ways in which these parameters may be used to estimate expected extinction times. Although the stochastic SIS logistic model is strictly density dependent and allows only for birth and death events, it nonetheless may be used to predict extinction times with some accuracy even for populations that are only weakly density dependent, or that are subject to catastrophes.     

Fe2O3-CeO2-Bi2O3/γ-Al2O3 catalyst in the catalytic wet air oxidation (CWAO) of cationic red GTL under mild reaction conditions

Fe₂O₃-CeO₂-Bi₂O₃/γ-Al₂O₃, an environmental friendly material, was investigated. The catalyst exhibited good catalytic performance in the CWAO of cationic red GTL. The apparent activation energy for the reaction was 79 kJ·mol⁻¹. HO₂· and O₂·⁻ appeared as the main reactive species in the reaction. The Fe₂O₃-CeO₂-Bi₂O₃/γ-Al₂O₃ catalyst, a novel environmental-friendly material, was used to investigate the catalytic wet air oxidation (CWAO) of cationic red GTL under mild operating conditions in a batch reactor. The catalyst was prepared by wet impregnation, and characterized by special surface area (BET measurement), X-ray diffraction (XRD) and X-ray photoelectron spectroscopy (XPS). The Fe₂O₃-CeO₂-Bi₂O₃/γ-Al₂O₃ catalyst exhibited good catalytic activity and stability in the CWAO under atmosphere pressure. The effect of the reaction conditions (catalyst loading, degradation temperature, solution concentration and initial solution pH value) was studied. The result showed that the decolorization efficiency of cationic red GTL was improved with increasing the initial solution pH value and the degradation temperature. The apparent activation energy for the reaction was 79 kJ·mol⁻¹. Hydroperoxy radicals (HO₂·) and superoxide radicals (O₂⁻·) appeared as the main reactive species upon the CWAO of cationic red GTL.     

Effects of Social Structure and Prey Dynamics on Extinction Risk in Gray Wolves

Extinction models based on diffusion theory generally fail to incorporate two important aspects of population biology—social structure and prey dynamics. We include these aspects in an individual-based extinction model for small, isolated populations of the gray wolf (Canis lupus). Our model predicts mean times to extinction significantly longer than those predicted by more general (diffusion) models. According to our model, an isolated population of 50 wolves has a 95% chance of surviving just 9 years and only a 30% chance of surviving beyond 100 years. Reflecting the influence of social structure, a wolf population initially comprising 50 individuals is expected to persist only a few years longer, on average (71 years), than is a population initially comprising just a single reproductive pair (62 years). In contrast, substantially greater average prey abundance leads to dramatically longer expected persistence times. Autocorrelated prey dynamics result in a more complex distribution of extinction times than predicted by many extinction models. We contend that demographic stochasticity may pose the greatest threat to small, isolated wolf populations, although environmental stochasticity and genetic effects may compound this threat. Our work highlights the importance of considering social structure and resource dynamics in the development of population viability analyses.     

Rapid Loss of Genetic Variation in Large Captive Populations of Drosophila Flies: Implications for the Genetic Management of Captive Populations

Levels of variation in eight large captive populations of D. melanogaster (census sizes ∼ 5000) that had been in captivity for periods from 6 months to 23 years (8 to 365 generations) were estimated from allozyme heterozygosities, lethal frequencies, and inversion heterozygosities and phenotypic variances, additive genetic variances ( V _A), and heritabilities ( h ²) for sternopleural bristle numbers. Correlations between all measures of variation except lethal frequencies were high and significant. All measures of genetic variation declined with time in captivity, with those for average heterozygosities, V _A, and h ² being significant. The effective population size ( N _e) was estimated to be 185–253 in these populations, only 0.037–0.051 of census size (N). Levels of allozyme heterozygosities declined rapidly in two large captive populations founded from another wild stock, being reduced by 86% and 62% within 2.5 years in spite of being maintained at sizes of approximately 1000 and 3500. Estimates of N _e/ N for these populations were only 0.016 and 0.004. Two estimates of N _e/ N for captive populations of D. pseudoobscura from data in the literature were also low at 0.036 and 0.012. Consequently, the rate of loss of genetic variation in captive populations and endangered species may be more rapid than hitherto recognized. Merely maintaining captive populations at large census sizes may not be sufficient to maintain essential genetic variation.