Richard Frankham

Effective population size/adult population size ratios in wildlife: a review

Summary The effective population size is required to predict the rate of inbreeding and loss of genetic variation in wildlife. Since only census population size is normally available, it is critical to know the ratio of effective to actual population size (NJN). Published estimates of NJN (192 from 102 species) were analysed to identify major variables affecting the ratio, and to obtain a comprehensive estimate of the ratio with all relevant variables included. The five most important variables explaining variation among estimates, in order of importance, were fluctuation in population size, variance in family size, form of TV used (adults v. breeders v. total size), taxonomic group and unequal sex-ratio. There were no significant effects on the ratio of high v. low fecundity, demographic r. genetic methods of estimation, or of overlapping v. non-overlapping generations when the same variables were included in estimates. Comprehensive estimates of NJN (that included the effects of fluctuation in population size, variance in family size and unequal sex-ratio) averaged only 0-10—0-11. Wildlife populations have much smaller effective population sizes than previously recognized.

Do island populations have less genetic variation than mainland populations

Island populations are much more prone to extinction than mainland populations. The reasons for this remain controversial. If inbreeding and loss of genetic variation are involved, then genetic variation must be lower on average in island than mainland populations. Published data on levels of genetic variation for allozymes, nuclear DNA markers, mitochondrial DNA, inversions and quantitative characters in island and mainland populations were analysed. A large and highly significant majority of island populations have less allozyme genetic variation than their mainland counterparts (165 of 202 comparisons), the average reduction being 29 per cent. The magnitude of differences was related to dispersal ability. There were related differences for all the other measures. Island endemic species showed lower genetic variation than related mainland species in 34 of 38 cases. The proportionate reduction in genetic variation was significantly greater in island endemic than in nonendemic island populations in mammals and birds, but not in insects. Genetic factors cannot be discounted as a cause of higher extinction rates of island than mainland populations.

How closely correlated are molecular and quantitative measures of genetic variation? A meta-analysis.

Abstract.— The ability of populations to undergo adaptive evolution depends on the presence of quantitative genetic variation for ecologically important traits. Although molecular measures are widely used as surrogates for quantitative genetic variation, there is controversy about the strength of the relationship between the two. To resolve this issue, we carried out a meta‐analysis based on 71 datasets. The mean correlation between molecular and quantitative measures of genetic variation was weak (r = 0.217). Furthermore, there was no significant relationship between the two measures for life‐history traits (r =−0.11) or for the quantitative measure generally considered as the best indicator of adaptive potential, heritability (r =−0.08). Consequently, molecular measures of genetic diversity have only a very limited ability to predict quantitative genetic variability. When information about a populations short‐term evolutionary potential or estimates of local adaptation and population divergence are required, quantitative genetic variation should be measured directly.

PREDICTIVE ACCURACY OF POPULATION VIABILITY ANALYSIS IN CONSERVATION BIOLOGY

Population viability analysis (PVA) is widely applied in conservation biology to predict extinction risks for threatened species and to compare alternative options for their mangement. It can also be used as a basis for listing species as endangered under World Conservation Union criteria. However, there is considerable scepticism regarding the predictive accuracy of PVA, mainly because of a lack of validation in real systems. Here we conducted a retrospective test of PVA based on 21 long-term ecological studies—the first comprehensive and replicated evaluation of the predictive powers of PVA. Parameters were estimated from the first half of each data set and the second half was used to evaluate the performance of the model. Contrary to recent criticisms, we found that PVA predictions were surprisingly accurate. The risk of population decline closely matched observed outcomes, there was no significant bias, and population size projections did not differ significantly from reality. Furthermore, the predictions of the five PVA software packages were highly concordant. We conclude that PVA is a valid and sufficiently accurate tool for categorizing and managing endangered species.

Assessing the benefits and risks of translocations in changing environments: a genetic perspective

Translocations are being increasingly proposed as a way of conserving biodiversity, particularly in the management of threatened and keystone species, with the aims of maintaining biodiversity and ecosystem function under the combined pressures of habitat fragmentation and climate change. Evolutionary genetic considerations should be an important part of translocation strategies, but there is often confusion about concepts and goals. Here, we provide a classification of translocations based on specific genetic goals for both threatened species and ecological restoration, separating targets based on ‘genetic rescue’ of current population fitness from those focused on maintaining adaptive potential. We then provide a framework for assessing the genetic benefits and risks associated with translocations and provide guidelines for managers focused on conserving biodiversity and evolutionary processes. Case studies are developed to illustrate the framework.

Introduction to Conservation Genetics: Frontmatter

The biological diversity of the planet is being rapidly depleted due to the direct and indirect consequences of human activity. As the size of animal and plant populations decreases, loss of genetic diversity reduces their ability to adapt to changes in the environment, with inbreeding and reduced fitness inevitable consequences for most species. This textbook provides a clear and comprehensive introduction to genetic principles and practices involved in conservation. Topics covered include: • evolutionary genetics of natural populations • loss of genetic diversity in small populations • inbreeding and loss of fitness • population fragmentation • resolving taxonomic uncertainties • genetic management of threatened species • contributions of molecular genetics to conservation. The text is presented in an easy-to-follow format, with main points and terms clearly highlighted. Each chapter concludes with a concise summary, which, together with worked examples and problems and answers, illuminates the key principles covered. Text boxes containing interesting case studies and other additional information enrich the content throughout, and over 100 beautiful pen-and-ink drawings help bring the material to life.

Estimates of minimum viable population sizes for vertebrates and factors influencing those estimates

Population size is a major determinant of extinction risk. However, controversy remains as to how large populations need to be to ensure persistence. It is generally believed that minimum viable population sizes (MVPs) would be highly specific, depending on the environmental and life history characteristics of the species. We used population viability analysis to estimate MVPs for 102 species. We define a minimum viable population size as one with a 99% probability of persistence for 40 generations. The models are comprehensive and include age-structure, catastrophes, demographic stochasticity, environmental stochasticity, and inbreeding depression. The mean and median estimates of MVP were 7316 and 5816 adults, respectively. This is slightly larger than, but in general agreement with, previous estimates of MVP. MVPs did not differ significantly among major taxa, or with latitude or trophic level, but were negatively correlated with population growth rate and positively correlated with the length of the study used to parameterize the model. A doubling of study duration increased the estimated MVP by approximately 67%. The increase in extinction risk is associated with greater temporal variation in population size for models built from longer data sets. Short-term studies consistently underestimate the true variances for demographic parameters in populations. Thus, the lack of long-term studies for endangered species leads to widespread underestimation of extinction risk. The results of our simulations suggest that conservation programs, for wild populations, need to be designed to conserve habitat capable of supporting approximately 7000 adult vertebrates in order to ensure long-term persistence. # 2003 Elsevier Science Ltd. All rights reserved.

Genetic adaptation to captivity in species conservation programs

As wild environments are often inhospitable, many species have to be captive‐bred to save them from extinction. In captivity, species adapt genetically to the captive environment and these genetic adaptations are overwhelmingly deleterious when populations are returned to wild environments. I review empirical evidence on (i) the genetic basis of adaptive changes in captivity, (ii) factors affecting the extent of genetic adaptation to captivity, and (iii) means for minimizing its deleterious impacts. Genetic adaptation to captivity is primarily due to rare alleles that in the wild were deleterious and partially recessive. The extent of adaptation to captivity depends upon selection intensity, genetic diversity, effective population size and number of generation in captivity, as predicted by quantitative genetic theory. Minimizing generations in captivity provides a highly effective means for minimizing genetic adaptation to captivity, but is not a practical option for most animal species. Population fragmentation and crossing replicate captive populations provide practical means for minimizing the deleterious effects of genetic adaptation to captivity upon populations reintroduced into the wild. Surprisingly, equalization of family sizes reduces the rate of genetic adaptation, but not the deleterious impacts upon reintroduced populations. Genetic adaptation to captivity is expected to have major effects on reintroduction success for species that have spent many generations in captivity. This issue deserves a much higher priority than it is currently receiving.

Contribution of inbreeding to extinction risk in threatened species

Wild populations face threats both from deterministic factors, e.g., habitat loss, overexploitation, pollution, and introduced species, and from stochastic events of a demographic, genetic, and environmental nature, including catastrophes. Inbreeding reduces reproductive fitness in naturally outbreeding species, but its role in extinctions of wild populations is controversial. To evaluate critically the role of inbreeding in extinction, we conducted realistic population viability analyses of 20 threatened species, with and without inbreeding depression, using initial population sizes of 50, 250, and 1000. Inbreeding markedly decreased median times to extinction by 28.5, 30.5, and 25% for initial populations of 50, 250, and 1000, respectively, and the impacts were similar across major taxa. The major variable explaining differences among species was initial population growth rate, whereas the impact of inbreeding was least in species with negative growth rates. These results demonstrate that the prospects for survival of threatened species will usually be seriously overestimated if genetic factors are disregarded, and that inappropriate recovery plans may be instituted if inbreeding depression is ignored.

Does Inbreeding and Loss of Genetic Diversity Decrease Disease Resistance

Inbreeding and loss of genetic diversity are predicted to decrease the resistance of species to disease. However, this issue is controversial and there is limited rigorous scientific evidence available. To test whether inbreeding and loss of genetic diversity affect a hosts resistance to disease, Drosophila melanogasterpopulations with different levels of inbreeding and genetic diversity were exposed separately to (a) thuringiensin, an insecticidal toxin produced by some strains of Bacillus thuringiensis, and (b) live Serratia marcescensbacteria. Inbreeding and loss of genetic diversity significantly reduced resistance of D. melanogasterto both the thuringiensin toxin and live Serratia marcescens. For both, the best fitting relationships between resistance and inbreeding were curvilinear. As expected, there was wide variation among replicate inbred populations in disease resistance. Lowered resistances to both the toxin and the pathogen in inbred populations were due to specific resistance alleles, rather than generalized inbreeding effects, as correlations between resistance and population fitness were low or negative. Wildlife managers should strive to minimise inbreeding and loss of genetic diversity within threatened populations and to minimise exposure of inbred populations to disease.