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.     

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.     

Evaluating Genetic Diversity Associated with Propagation-Assisted Restoration of American Shad

Abstract: We investigated the conservation of genetic diversity during a restoration program for American shad ( Alosa sapidissima ) in Virginia ( U.S.A.). Restoration entailed capture of wild Pamunkey River shad broodstock followed by production and release of hatchery-reared fry to supplement the nearly extinct James River shad population. To assess the baseline genetic diversity of donor and recipient populations, we used five tri- and tetra-nucleotide microsatellite loci to test for genetic heterogeneity among yearly subsamples from both rivers and between early- and late-spawning shad from the donor population. Tests for allelic heterogeneity between James River and Pamunkey shad subsamples yielded no significant genetic differentiation (χ ² = 14.72, p = 0.132 and χ ² = 10.24, p = 0.440, respectively). We detected no significant genetic divergence between early- and late-spawning adults in Pamunkey River spawning aggregations in either year. The donor and recipient populations exhibited significant genetic differentiation (χ ² = 27.4, p = 0.003), however, indicating that the stocking program carries a risk of outbreeding depression. Because the two river populations are genetically divergent, replenishment of the James population with Pamunkey fry may be detectable in the future as heterozygote deficits and linkage disequilibria in the James River population. In an analysis of broodstock and their hatchery-reared progeny, microsatellites proved efficient for family analysis, unambiguously determining the parentage of 100% of the hatchery-reared fry studied. Genetic analysis indicated that breeding procedures may result in high levels of reproductive variance.     

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.     

Ammonia modeling for assessing potential toxicity to fish species in the Rio Grande, 1989-2002.

Increasing volumes of treated and untreated human sewage discharged into rivers around the world are likely to be leading to high aquatic concentrations of toxic, unionized ammonia (NH3), with negative impacts on species and ecosystems. Tools and approaches are needed for assessing the dynamics of NH3. This paper describes a modeling approach for first-order assessment of potential NH3 toxicity in urban rivers. In this study daily dissolved NH3 concentrations in the Rio Grande of central New Mexico, USA, at the city of Albuquerque's treated sewage outfall were modeled for 1989-2002. Data for ammonium (NH4+) concentrations in the sewage and data for discharge, temperature, and pH for both sewage effluent and the river were used. We used State of New Mexico acute and chronic NH3- N concentration values (0.30 and 0.05 mg/L NH3-N, respectively) and other reported standards as benchmarks for determining NH3 toxicity in the river and for assessing potential impact on population dynamics for fish species. A critical species of concern is the Rio Grande silvery minnow (Hybognathus amarus), an endangered species in the river near Albuquerque. Results show that NH3 concentrations matched or exceeded acute levels 13%, 3%, and 4% of the time in 1989, 1991, and 1992, respectively. Modeled NH3 concentrations matched or exceeded chronic values 97%, 74%, 78%, and 11% of the time in 1989, 1991, 1992, and 1997, respectively. Exceedences ranged from 0% to 1% in later years after enhancements to the wastewater treatment plant. Modeled NH3 concentrations may differ from actual concentrations because of NH3 and NH4+ loss terms and additive terms such as mixing processes, volatilization, nitrification, sorbtion, and NH4+ uptake. We conclude that NH3 toxicity must be considered seriously for its potential ecological impacts on the Rio Grande and as a mechanism contributing to the decline of the Rio Grande fish community in general and the Rio Grande silvery minnow specifically. Conclusions drawn for the Rio Grande suggest that NH3 concentrations may be high in rivers around the world where alkaline pH values are prevalent and sewage treatment capabilities are poorly developed or absent.     

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.     

Effects of Population Size, Mating System, and Evolutionary Origin on Genetic Diversity in Spiranthes sinensis and S. hongkongensis

Hong Kong once supported more than 109 species of wild orchids, of which approximately 30% were endemic. Most of the local wild orchids have now become rare or endangered. I conducted a comparative study of genetic diversity in two closely related terrestrial orchids, an allotetraploid, Spiranthes hongkongensis , and its diploid progenitor, S. sinensis , to assess the effects of the population bottleneck associated with the origin of the polyploid and to investigate the relationships between number of breeding individuals, mating system, and level of isozyme variation in their populations. Nearly complete genetic uniformity was observed both within and among populations of S. hongkongensis . In contrast, S. sinensis had high levels of genetic variation for all of the genetic parameters examined. Regression analysis of population size and several components of genetic diversity in S. sinensis revealed that, among various measures of within-population variation, the proportion of polymorphic loci ( P ) and average number of alleles per locus ( A ) or per polymorphic locus ( A _p ) were the most sensitive to population size ( R ² = 0.942, p = 0.001; R ² = 0.932, p = 0.002; and R ² = 0.923, p = 0.002 respectively). The highly negative correlation ( r = −0.999, p < 0.01) between population size and the mean frequency of private alleles in pairwise population comparisons, p (1), indicated that population size may also be used to predict the extent of population differentiation caused by random genetic drift. Conservation of genetic diversity in S. sinensis could be maximized by protecting several of both large and small populations, whereas fewer populations may be needed to achieve this goal for S. hongkongensis.     

Temporal Changes in Allele Frequencies and a Population's History of Severe Bottlenecks

Monitoring temporal changes in genetic variation has been suggested as a means of determining if a population has experienced a demographic bottleneck. Simulations have shown that the variance in allele frequencies over time ( F ) can provide reasonable estimates of effective population size ( N_e ). This relationship between F and N_e suggests that changes in allele frequencies may provide a way to determine the severity of recent demographic bottlenecks experienced by a population. We examined allozyme variation in experimental populations of the eastern mosquitofish ( Gambusia holbrooki ) to evaluate the relationship between the severity of demographic bottlenecks and temporal variation in allele frequencies. Estimates of F from both the fish populations and computer simulations were compared to expected rates of drift. We found that different methods for estimating F had little effect on the analysis. The variance in estimates of F was large among both experimental and simulated populations experiencing similar demographic bottlenecks. Temporal changes in allele frequencies suggested that the experimental populations had experienced bottlenecks, but there was no relationship between observed and expected values of F . Furthermore, genetic drift was likely to be underestimated in populations experiencing the most severe bottlenecks. The weak relationship between F and bottleneck severity is probably due to both sampling error associated with the number of polymorphic loci examined and the loss of alleles during the bottlenecks. For populations that may have experienced severe bottlenecks, caution should be used in making evolutionary interpretations or management recommendations based on temporal changes in allele frequencies.     

Flow regulation and fragmentation imperil pelagic-spawning riverine fishes.

Flow regulation and fragmentation of the world's rivers threaten the integrity of freshwater ecosystems and have resulted in the loss or decline of numerous fish species. Pelagic-spawning fishes (pelagophils) are thought to be particularly susceptible to river regulation because their early life stages (ichthyoplankton) drift until becoming free-swimming, although the extent of transport is largely unknown. Transport velocity and distance were determined for passively drifting particles, which mimicked physical properties of ichthyoplankton, in two large, regulated rivers (Rio Grande and Pecos River) of the arid Southwest United States. Particle drift data were incorporated into celerity-discharge equations (r2 > 0.90; P < 0.001), and reach-specific transport velocity was modeled as a function of discharge. Transport velocities of particles exceeded 0.7 m/s in all river reaches during typical spawning flows (i.e., reservoir releases or rainstorms) and were greatest in highly incised and narrow channel reaches. Mean transport distance of particles released in the Pecos River during sustained reservoir flows (141.1 km; 95% CI = 117.0-177.5 km) was significantly longer than during declining reservoir flows that mimicked a natural rainstorm (52.4 km; 95% CI = 48.8-56.5 km). Mean transport distance of particles in the Rio Grande during sustained reservoir flows was 138.7 km (95% CI = 131.0-147.2 km). There are 68 dams and 13 reservoirs that fragment habitats and regulate flow in the Rio Grande Basin (Rio Grande and Pecos River) in areas historically occupied by pelagophils. While the basin historically provided 4029 km of free-flowing riverine habitat, reservoir habitat now represents > 10% of the longitudinal distance. Only five unfragmented nonreservoir reaches > 100 km remain in the Rio Grande, and two remain in the Pecos River. Pelagophils were extirpated from all reservoirs and from nearly all short, fragmented reaches (< 100 km) of the Rio Grande Basin, but at least some fraction persisted in all longer reaches (> 100 km). The recovery and long-term persistence of pelagophils in regulated rivers, including those in this study, will likely depend on reestablishment and protection of long unfragmented reaches coupled with mimicry of the natural flow regime.     

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.     

Urban Plants as Genetic Reservoirs or Threats to the Integrity of Bushland Plant Populations

Abstract:  Remnant plants in urban fringes and native plants in gardens have the potential to contribute to the conservation of threatened plants by increasing genetic diversity, effective size of populations, and levels of genetic connectedness. But they also pose a threat through the disruption of locally adapted gene pools. At Hyams Beach, New South Wales, Australia, four bushland stands of the rare shrub, Grevillea macleayana McGillivray, surround an urban area containing remnant and cultivated specimens of this species. Numbers of inflorescences per plant, fruits per plant, and visits by pollinators were similar for plants in urban gardens and bushland. Urban plants represented a substantial but complex genetic resource, displaying more genetic diversity than bushland plants judged by H_e , numbers of alleles per locus, and number of private alleles. Of 27 private alleles in urban plants, 17 occurred in a set of 19 exotic plants. Excluding the exotic plants, all five stands displayed a moderate differentiation ( F_ST = 0.14 ± 0.02), although the urban remnants clustered with two of the bushland stands. These patterns may be explained by high levels of selfing and inbreeding in this species and by long-distance dispersal (several seeds in the urban stand were fathered by plants in other stands). Genetic leakage (gene flow) from exotic plants to 321 seeds on surrounding remnant or bushland plants has not occurred. Our results demonstrate the conservation value of this group of urban plants, which are viable, productive, genetically diverse, and interconnected with bushland plants. Gene flow has apparently not yet led to genetic contamination of bushland populations, but high levels of inbreeding would make this a rare event and difficult to detect. Remnant plants in urban gardens could successfully contribute to recovery plans for endangered and vulnerable species.     

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.     

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.     

Effects of Habitat Size and Patch Isolation on Reproductive Success of the Serpentine Morning Glory

Abstract: We examined the effects of habitat area and patch isolation on reproductive success in serpentine morning glory ( Calystegia collina [Convolvulaceae]), a primarily self-incompatible clonal plant endemic to serpentine outcrops in northern California's coast ranges. Within a 4000-km ² region, we compared the reproductive success of C. collina on 16 small (<5 ha) and 7 large ( >300 ha) outcrops. Flower and fruit production were significantly higher on large serpentine outcrops than on small outcrops. Fruit production also was positively correlated with the soil's ratio of calcium to magnesium. Successful pollination was positively affected by flower density and the number of other flowering patches within 100 m of a C. collina patch. The number of nearby flowering patches was considerably higher on large than on small outcrops. Flowers on large outcrops did not receive significantly more bee visitors than flowers on small outcrops, suggesting that pollination success is related to the quality rather than the quantity of pollen deposited. Fruit production by plants on both small and large outcrops was enhanced by the experimental addition of pollen from other patches, but not by the addition of pollen from the same patch. These findings demonstrate that the size of habitat may have strong effects on the reproductive success of locally endemic plants by enhancing opportunities for successful sexual reproduction. They also warn against the presumption that naturally patchy plant species are invulnerable to the effects of habitat fragmentation.     

Genetic Effects of Habitat Fragmentation on Common Species of Swiss Fen Meadows

Abstract:  The area of Caricion davallianae alliance in Switzerland has been considerably reduced and fragmented during the last 150 years. We assessed the genetic variability, inbreeding level, and among-population differentiation of two common habitat-specific plant species, Carex davalliana SM. and Succisa pratensis Moench, in 18 Caricion davallianae fen meadows subjected to fragmentation. We used a spatial field design of fen systems (six systems total), each consisting of one large habitat island and two small habitat islands. We used allozyme electrophoresis to derive standard genetic parameters ( A, P, H_O, H_E, F_IS, F_ST ). In Carex we identified a consistently lower A in isolated habitat islands; furthermore, H_E was lower in small habitat islands than in large habitat islands. In Succisa we identified a lower H_O in small habitat islands than in larger ones. Small habitat islands were marginally significantly differentiated (  F_ST ) from large islands for Succisa . For both species, no effects were evident for F_IS ; therefore, we argue that genetic drift rather than inbreeding is the main cause of the observed differences. The genetic structure of Carex and Succisa in small habitat islands differed from that in large habitat islands, but differences were small. It appears that the observed differences in genetic variability among fen meadows correspond to observed differences in fitness and demographic traits. We show that habitat fragmentation affects not only the rare species in an ecosystem but also reduces the survival probabilities of common species. One of the main goals of conservation should be to mitigate fragmentation of natural habitats in order to increase population sizes and connectivity.     

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.     

Insularization of Tanzanian Parks and the Local Extinction of Large Mammals

Island biogeography theory predicts that species will be lost on habitat "islands" created by the fragmentation of continental regions. Many Tanzanian parks are rapidly becoming habitat islands as a result of human settlement, agricultural development, and the active elimination of wildlife on adjacent lands. The rate of extinction of mammals in six Tanzanian parks over the last 35–83 years is significantly and inversely related to park area, suggesting that increasing insularization of the parks has been an important contributory factor in large mammal extinctions. I compared observed patterns of persistence of mammals in Tanzanian parks to predictions derived from earlier extinction models. The predictions of the S ¹ models of Soulé et al. (1979) and Burkey (1994) and the S ² and S ³ models of Soulé et al. (1979) match very closely the observed pattern of persistence of mammals in Tanzanian parks. The loss of mammal species will probably continue, particularly in the smaller parks. Establishment of wildlife corridors linking the parks in northern Tanzania could help to reduce the potential loss of species in the future.     

Clearance and Fragmentation of Tropical Deciduous Forest in the Tierras Bajas, Santa Cruz, Bolivia

Abstract: The Tierras Bajas is an area of 20,000 km ² of lowland deciduous forest in eastern Santa Cruz, Bolivia, that has undergone rapid change during the past two decades. As part of the largest remaining area of intact deciduous tropical forest in the world, it has been nominated a priority area for conservation by several environmental organizations. We quantified the spatial and temporal patterns of deforestation in the area by digital processing of high-resolution satellite imagery from 1975 through 1998. The estimated rate of deforestation was among the highest in the world for such a limited area, ranging from 160 km ²/year in the early1980s to almost 1200 km ²/year in the late 1990s. Although most deforestation up to 1984 was in Bolivian peasant and Mennonite colonies, most deforestation after 1984 was in non-Mennonite industrial soybean farms. The level of fragmentation of uncut forest, caused by the spatial patterns of deforestation, also differed among these broad land-use types. Deforestation in planned and spontaneous peasant colonies was complex in shape, forming relatively large areas of edge-affected forest, whereas that in Mennonite and other industrial farms was in large, rectangular increments, creating relatively less edge. But the distribution of these farms and the practice of initially clearing around the peripheries of properties resulted in the isolation of large areas of forest. In 1998 four-fifths of the remaining forest were either within 1 km of a clearance edge or in isolated fragments of <50 km². Compared with deforested areas, the areas of isolated and edge-affected forest were disproportionately large during the early stages of frontier colonization. These results imply that if the fragmentation effects of deforestation are to be minimized, conservation planning must occur at the earliest stages of frontier development.     

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.