Supportive Breeding and Variance Effective Population Size

The practice of supporting weak, wild populations through release of individuals bred in captivity is becoming an increasingly important conservation measure. A frequently recommended form of such breeding-release activity refers to supportive breeding: a fraction of the target population is brought into captivity for reproduction, and the resulting progeny are released to mix with the wild segment of the population. We derived an expression for the variance effective size of a population managed through supportive breeding and discuss its relationship to previously published equations that are based on the assumption of random mating. We show that the effect of supportive breeding may be quite different on the inbreeding and the variance effective sizes. Whereas supportive breeding always results in a reduction of the inbreeding effective number, the variance effective number may either decrease, increase, or remain unchanged. We discuss these observations in relation to conservation management and suggest some general guidelines for supportive breeding situations. Our recommendations include making a distinction between inbreeding and variance effective numbers; taking particular care when dealing with organisms with high reproductive potential; assuring that the amount of drift be no larger than it would be without supportive breeding; and focusing primarily on the variance effective size of a population-that is, on the effective number directly related to the rate of loss of gene diversity.     

Genetically Effective Population Size of Large Mammals: An Assessment of Estimators

Calculamos tamuños de poblaciones genéticamente efectivas (N_e) para poblaciones simuladas del oso gris (Ursus arctos) trazando la péridida de heterozigasidad a tráves del tiempo, luego las comparamos con estimaciones de N_e Producidas aplicando fórmulas publicadas a los resultados demográficos de la simulación. Los valores de N_e calculados usando diferentes fórmula.s con datos idénticos, variaron mucho. Las ecuaciones publicadas por Hill (1972), y modificaciones de las usuadas por Ryman, et aL (1981) y Reed et al. (1986), proporcionaron los cálculos más precsios Fluctuaciones menores en las poblacionales tuvieron poco efecto sobre N_e pero la variación en el éxito repductivo por vida entre los machos V_km redujo tremendamente el N_e comparado con el valor esperado bajo exito reproductivo al azar. Todos los métodos para calcular N_e para poblaciones con demografias complejas requieren datos extensos, pero estimaciones para V_km en especies poligamas son especialmente dificiles de obtener. Sugerimos que modelos de simulación pueden proveer métodos alternutivos para calcular V_km y N_e.     

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.     

Effects of Habitat Fragmentation on Effective Population Size in the Endangered Rio Grande Silvery Minnow

Abstract:  We assessed spatial and temporal patterns of genetic diversity to evaluate effects of river fragmentation on remnant populations of the federally endangered Rio Grande silvery minnow ( Hybognathus amarus ). Analysis of microsatellite and mitochondrial DNA detected little spatial genetic structure over the current geographic range, consistent with high gene flow despite fragmentation by dams. Maximum-likelihood analysis of temporal genetic data indicated, however, that present-day effective population size ( N_eV ) of the largest extant population of this species was 78 and the ratio of effective size to adult numbers ( N_eV/N ) was ∼ 0.001 during the study period (1999 to 2001). Coalescent-based analytical methods provided an estimate of historical (river fragmentation was completed in 1975) effective size ( N_eI  ) that ranged between 10⁵ and 10⁶. We propose that disparity between contemporary and historical estimates of N_e and low contemporary N_e/N result from recent changes in demography related to river fragmentation. Rio Grande silvery minnows produce pelagic eggs and larvae subject to downstream transport through diversion dams. This life-history feature results in heavy losses of yearly reproductive effort to emigration and mortality, and extremely large variance in reproductive success among individuals and spawning localities. Interaction of pelagic early life history and river fragmentation has altered demographic and genetic dynamics of remnant populations and reduced N_e to critically low values over ecological time.     

Genetic Effects of Multiple Generations of Supportive Breeding

Abstract: The practice of supporting weak wild populations by capturing a fraction of the wild individuals, bringing them into captivity for reproduction, and releasing their offspring into the natural habitat to mix with wild ones is called supportive breeding and has been widely applied in the fields of conservation biology and fish and wildlife management. This procedure is intended to increase population size without introducing exogenous genes into the managed population. Previous work examining the genetic effects of a single generation of supportive breeding has shown that although a successful program increases the census population size, it may reduce the genetically effective population size and thereby induce excessive inbreeding and loss of genetic variation. We expand and generalize previous analyses of supportive breeding and consider the effects of multiple generations of supportive breeding on rates of inbreeding and genetic drift. We derived recurrence equations for the inbreeding coefficient and coancestry, and thereby equations for inbreeding and variance effective sizes, under three models for selecting captive breeders: at random, preferentially among those born in captivity, and preferentially among those born in the wild. Numerical examples indicate that supportive breeding, when carried out successfully over multiple generations, may increase not only the census but also the effective size of the supported population as a whole. If supportive breeding does not result in a substantial and continuous increase of the census size of the breeding population, however, it might be genetically harmful because of elevated rates of inbreeding and genetic drift.     

Effective Population Size and Lifetime Reproductive Success

The mean and variance of lifetime reproductive success, E_LRS and V_LRS, influence the ratio of effective to census population size, N_e/N_c. Because the complete data needed to calculate E_LRS and V_LRS are seldom available, we provide alternatives for estimating N_e/N_c from incomplete data. These estimates should be useful to conservation biologists trying to compute the effective size of a censused population. An analytical approach makes assumptions regarding the process influencing offspring survival. We provide a method for examining the validity of those assumptions and show that particular violations can result in either over- or underestimates. When the assumptions are violated or when more data are available, we suggest estimating N_e/N_c using computer simulations of models based on individuals. We examine how such simulations can be used to estimate N_e/N_c using an individual-based model for Lesser Snow Geese ( Anser caerulescens ). We demonstrate that such estimates can be biased unless the simulations are based on complete cohorts and samples of known age. We show that because the estimate of N_e/N_c depends on the stage of the reproductive cycle used as a point of reference in the model, the census population size N_c must be based on the same stage to provide unbiased estimates of N_e.     

Dimensionless Life Histories and Effective Population Size

The effective size ( N _e ) of a population can be estimated from demographic information. We evaluated a recent model, showing that N _e depends strongly on the relationship between age at reproductive maturity ( M ) and average adult lifespan ( A ). N _e converges on half the number of potentially reproducing adults ( N/2 ) as M decreases relative to A , but it increases linearly as M increases for a given value of A . Therefore, convergence of N _e on N/2 is more likely in organisms with a short sexual maturation period scaled to adult lifespan. To assess the generality of this convergence we asked whether most organisms are characterized by this requisite relationship between M and A . The dimensionless number M/A is approximately invariant within taxa, but it is markedly different across taxa. Previous work focused on birds and mammals, taxa with unusually small M/A (0.4 and 0.75). Other animal taxa take longer than most birds and mammals to reach maturity for a given reproductive lifespan, so they are characterized by larger M/A (e.g., fish, 2.0). In theory, these taxon-specific life histories strongly influence N _e . We conclude that N _e is expected to approach N/2 , provided that M/A is (unusually) small, and that N _e / N among poikilotherms may often exceed that of mammals and especially birds.     

Effective Population Size in Winter-Run Chinook Salmon

Winter-run chinook salmon from the Sacramento River, California, is federally listed as endangered. Since 1989 there has been aprogram to augment the natural population by capturing adults, artificially spawning them, raising tine young and releasing the smolt. Here we estimate the effective population size of these captive-raised fish, the natural run, and the combination of both groups over the three-year period from 1991 to 1993. We find that the most appropriate estimate of the effective population size of the captive-raised progeny is a variance estimate of effective population size standardized so that the number of released smolts returning to spawn was the same as the number of spawners used to produce the smolts originally. We have generated 10,000 random samples to simulate returns from these released progeny. The estimates of the effective population sizes in 1991, 1992, and 1993 were only 7.02, 19.07, and 7.74, respectively. We then determined limits on the effective population size of the natural run based on 0.1 and 0.333 of the run-size estimates. Using estimates of the captive proportion of the run, the minimum estimates of the effective population size of the overall run for the three years were 21.9, 127.3, and 39.0, and the maximum estimates were 61.6, 401.0, and 108.7. It does not appear that the hatchery program has reduced the overall effective population size. The run sizes in each year are extremely low, however, and it is possible that fish will be lost from this run in one of the years in the immediate future, making reestablishment of a healthy run even more difficult.     

Effective Population Size in Red-Cockaded Woodpeckers: Population and Model Differences

Loss of genetic variability in isolated populations is an important issue for conservation biology. Most studies involve only a single population of a given species and a single method of estimating rate of loss. Here we present analyses for three different Red-cockaded Woodpecker ( Picoides borealis ) populations from different geographic regions. We compare two different models for estimating the expected rate of loss of genetic variability, and test their sensitivity to model parameters. We found that the simpler model (Reed et al. 1988) consistently estimated a greater rate of loss of genetic variability from a population than did the Emigh and Pollak (1979) model. The ratio of effective population size (which describes the expected rate of loss of genetic variability) to breeder population size varied widely among Red-cockaded Woodpecker populations due to geographic variation in demography. For this species, estimates of effective size were extremely sensitive to survival parameters, but not to the probability of breeding or reproductive success. Sensitivity was sufficient that error in estimating survival rates in the field could easily mask true population differences in effective size. Our results indicate that accurate and precise demographic data are prerequisites to determining effective population size for this species using genetic models, and that a single estimate of rate of loss of genetic variability is not valid across populations.     

Effects of Population Size on Singing Behavior of a Rare Duetting Songbird

Although the genetic and ecological effects of population declines in endangered species have been well studied, little is known of the social consequences. Changes in signaling behavior may result in disrupted communication and affect both reproductive and conflict‐resolution activities. The North Island Kōkako (Callaeas wilsoni) is an endangered, duetting (i.e., alternating, coordinated singing by mated pairs) songbird endemic to New Zealand temperate rain forests. Scattered populations (approximately 1500 individuals in 13 surviving and 11 translocated populations) in isolated conservation areas of different sizes have been rescued from extirpation and are currently recovering. We examined key song attributes of the Kōkako to assess whether population size or growth rate are related to song complexity, the reduction of which may compromise effective communication. We analyzed song repertoire size and phrase‐type sharing (i.e., Jaccard index of similarity), vocal performance (singing rates, song switching rates, and diversity of phrase types), and song syntactical characteristics (i.e., unpredictability in sequences of phrase types) in surviving and translocated populations (populations of approximately 19–250 territorial individuals). Population size was positively correlated with a population's song repertoire, song diversity, and switching of song phrase types and negatively correlated with shared phrase types and variation in syntactical structure of songs. Population growth rate correlated positively with pair repertoire size, population repertoire size, and singing rates during song bouts. As for solo‐singing species in fragmented landscapes, songs in the fragmented populations of Kōkako appear to be undergoing microevolution as occurs in island colonization events. Our results suggest that vocal changes in small populations could affect population establishment and growth, particularly in multiple‐source translocations. We believe measurement of vocal behavior could be used as a supplement to periodic population censuses to allow more frequent monitoring of population size. Efectos de la Conducta de Canto sobre el Tamaño Poblacional de una Ave Canora Rara     

Understanding and Estimating Effective Population Size for Practical Application in Marine Species Management

Abstract: Effective population size (N_e) determines the strength of genetic drift in a population and has long been recognized as an important parameter for evaluating conservation status and threats to genetic health of populations. Specifically, an estimate of N_e is crucial to management because it integrates genetic effects with the life history of the species, allowing for predictions of a population's current and future viability. Nevertheless, compared with ecological and demographic parameters, N_e has had limited influence on species management, beyond its application in very small populations. Recent developments have substantially improved N_e estimation; however, some obstacles remain for the practical application of N_e estimates. For example, the need to define the spatial and temporal scale of measurement makes the concept complex and sometimes difficult to interpret. We reviewed approaches to estimation of N_e over both long‐term and contemporary time frames, clarifying their interpretations with respect to local populations and the global metapopulation. We describe multiple experimental factors affecting robustness of contemporary N_e estimates and suggest that different sampling designs can be combined to compare largely independent measures of N_e for improved confidence in the result. Large populations with moderate gene flow pose the greatest challenges to robust estimation of contemporary N_e and require careful consideration of sampling and analysis to minimize estimator bias. We emphasize the practical utility of estimating N_e by highlighting its relevance to the adaptive potential of a population and describing applications in management of marine populations, where the focus is not always on critically endangered populations. Two cases discussed include the mechanisms generating N_e estimates many orders of magnitude lower than census N in harvested marine fishes and the predicted reduction in N_e from hatchery‐based population supplementation.