Genetic Variation and Outcrossing Rate in Relation to Population Size in Gentiana pneumonanthe L

The amount of genetic variation in the rare perennial herb Gentiana pneumonanthe L. was determined to explore its relation to population size. Differences in isozyme variation between maternal plants and their offspring were used to investigate the relationship between population size and outcrossing rate. In 25 populations in The Netherlands, differing in size from 1 to more than 50,000 flowering individuals, 16 allozyme loci were analyzed on leaves of maternal plants and offspring grown in a greenhouse. Population size was significantly positively correlated with the proportion of polymorphic loci, but only marginally with heterozygosity and the mean effective number of alleles. Most of the studied populations were characterized by a complete absence of rare alleles, and F -statistics suggest relatively high levels of genetic differentiation among populations and thus a low level of gene flow. Leaf samples (maternal) were mostly in Hardy-Weinberg equilibrium, while several offspring samples showed an excess of homozygotes, which suggests selection favoring heterozygotes. Because most small populations consist only of adult survivors from formerly larger populations, this may partly explain the absence of a clear relationship between genetic variation of the maternal plants and population size. A significant positive correlation was found between the level of cross-fertilization and population size. From these results, we conclude that, to some degree, small populations have a reduced level of genetic variation, while their present isolation in nature reserves has resulted in a very limited interpopulational gene flow level. At present a higher level of inbreeding in small populations contributes to a further loss of genetic variation and may also result in reduced offspring fitness.     

Relationship of Genetic Variation to Population Size in Wildlife

Genetic diversity is one of three levels of biological diversity requiring conservation. Genetic theory predicts that levels of genetic variation should increase with effective population size. Soulé (1976) compiled the first convincing evidence that levels of genetic variation in wildlife were related to population size, but this issue remains controversial. The hypothesis that genetic variation is related to population size leads to the following predictions: (1) genetic variation within species should be related to population size; (2) genetic variation within species should be related to island size; (3) genetic variation should be related to population size within taxonomic groups; (4) widespread species should have more genetic variation than restricted species; (5) genetic variation in animals should be negatively correlated with body size; (6) genetic variation should be negatively correlated with rate of chromosome evolution; (7) genetic variation across species should be related to population size; (8) vertebrates should have less genetic variation than invertebrates or plants; (9) island populations should have less genetic variation than mainland populations; and (10) endangered species should have less genetic variation than nonendangered species. Empirical observations support all these hypotheses. There can be no doubt that genetic variation is related to population size, as Soulé proposed. Small population size reduces the evolutionary potential of wildlife species.     

Population Persistence and Offspring Fitness in the Rare Bellflower Campanula cervicaria in Relation to Population Size and Habitat Quality

Abstract: Data from several animal species and a few plant species indicate that small populations face an elevated risk of extinction. Plants are still underrepresented in these studies concerning the relation between population size and persistence. We studied the effect of population size on persistence among natural populations of the rare bellflower Campanula cervicaria in Finland. We monitored 52 bellflower populations for 8 years and found that the mean population size decreased from 24 to 14 during this period. Small populations with ≤5 individuals were more prone to losing all fertile plants than were larger ones. Reduction in population size was nevertheless unrelated to the degree of population isolation, measured as the distance to the nearest known population. To test the hypothesis that offspring fitness is lower in small populations, we germinated bellflower seeds from different-sized populations in a laboratory and found that seed germination ability was independent of population size. The seedlings raised from seeds of small populations grew faster than those taken from larger populations. Population size was negatively related to the amount of shade in the habitats. In conclusion, decreasing population sizes of C. cervicaria seemed not to be caused by lowered germination ability or growth rate in small populations; rather, population size reductions appeared to be due to closing of vegetation in the habitats.     

Inbreeding Depression, Environmental Stress, and Population Size Variation in Scarlet Gilia (Ipomopsis aggregata)

Despite a large body of theory, few studies have directly assessed the effects of variation in population size on fitness components in natural populations of plants. We conducted studies on 10 populations of scarlet gilia, Ipomopsis aggregata , to assess the effects of population size and year-to-year variation in size on the relative fitness of plants. We showed that seed size and germination success are significantly reduced in small populations (those 100 flowering plants) of scarlet gilia. Plants from small populations are also more susceptible to environmental stress. When plants from small and large populations were subjected to an imposed stress (combined effects of transplanting and experimental clipping, simulating ungulate herbivory) in a common garden experiment, plants from small populations suffered higher mortality and were ultimately of smaller size than plants from large populations. In addition, experimental evidence indicates that observed fitness reductions are genetic, due to the effects of genetic drift and/or inbreeding depression. When pollen was introduced from distant populations into two small populations, seed mass and percentage of germination were bolstered, while pollen transferred into a large population had no significant effect. Year-to-year variation in population size and its effects on plant fitness are also discussed. In one small population, for example, a substantial increase in size from within did not introduce sufficient new (archived) genetic material to fully overcome the effects of inbreeding depression.     

Genetic Diversity and Population Size in Four Rare Southern Appalachian Plant Species

Allozyme diversity was examined in four rare, high-montane plant species from the Appalachian Mountains of southeastern North America. These species may represent relictual members or descendants of an alpine community that was more widespread during the late Pleistocene. We sampled five populations of Geum radiatum (Rosaceae), Carex misera (Cyperaceae), Trichophorum cespitosum (Cyperaceae), and the four known populations of Calamagrostis cainii (Poaceae). Genetic diversity was low for all species but was typical of that found for plant species with limited ranges. Low genetic diversity may reflect historical events associated with changes in the species' biogeography. As the Pleistocene climate warmed, suitable habitat decreased in areal extent and became fragmented, probably resulting in smaller, more-isolated populations. In recent times these species, which co-occur in fragile rock outcrop habitats, have been adversely affected by human activities. Genetic analyses revealed reduced diversity in populations of decreasing size for three species. Estimates of gene flow were low ( Nm < 1.0) in all four species. Positive associations between genetic diversity and population size, evidence of recent population declines, and the low estimates of gene flow suggest that genetic drift may play a prominent role in shaping the present-day genetic composition of these species. Furthermore, these data suggest that the genetically depauperate populations are unlikely to regain genetic variation without human intervention.     

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, 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.     

Inbreeding and Loss of Genetic Variation in a Reintroduced Population of Mauritius Kestrel

Abstract:  Many populations have recovered from severe bottlenecks either naturally or through intensive conservation management. In the past, however, few conservation programs have monitored the genetic health of recovering populations. We conducted a conservation genetic assessment of a small, reintroduced population of Mauritius Kestrel ( Falco punctatus ) to determine whether genetic deterioration has occurred since its reintroduction. We used pedigree analysis that partially accounted for individuals of unknown origin to document that (1) inbreeding occurred frequently (2.6% increase per generation; N _eI= 18.9), (2) 25% of breeding pairs were composed of either closely or moderately related individuals, (3) genetic diversity has been lost from the population (1.6% loss per generation; N _eV= 32.1) less rapidly than the corresponding increase in inbreeding, and (4) ignoring the contribution of unknown individuals to a pedigree will bias the metrics derived from that pedigree, ultimately obscuring the prevailing genetic dynamics. The rates of inbreeding and loss of genetic variation in the subpopulation of Mauritius Kestrel we examined were extreme and among the highest yet documented in a wild vertebrate population. Thus, genetic deterioration may affect this population's long-term viability. Remedial conservation strategies are needed to reduce the impact of inbreeding and loss of genetic variation in this species. We suggest that schemes to monitor genetic variation after reintroduction should be an integral component of endangered species recovery programs.     

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.     

Effects of Population Size on Seed Production and Germinability in an Endangered, Fragmented Grassland Plant

Abstract: Fragmentation and isolation of plant populations are thought to affect demographic processes such as seed production and cause reductions in fitness. I followed seed set over a 3-year period in eight populations of the endangered Rutidosis leptorrhynchoides (Asteraceae) that differed in population size from 13 to over 5000 flowering plants. Germinability of the resultant seed was also examined to determine whether small populations had lower fitness than large populations. Seed set was significantly associated with population size in 2 of the 3 years. Small populations (<30 flowering plants) produced significantly fewer seeds per head in 1994 and 1995 than did large populations (500 to over 5000 flowering plants), which did not differ significantly from one another. There was, however, substantial variation within populations. In 1993 seed production did not follow any simple relationship with population size, possibly because environmental stress from low rainfall had an overriding impact. Differences in seed germinability between populations were largely not evident, suggesting that this aspect of fitness has not declined substantially in small populations relative to large populations. This study suggests that nongenetic, demographic factors are of immediate importance to the persistence of small populations of R. leptorrhynchoides because of their potential impacts on seedling recruitment.