Inbreeding levels and prey abundance interact to determine fecundity in natural populations of two species of wolf spider

Long-term effective population size is expected, and has been shown, to correlate positively with various measures of population fitness. Here we examine the interacting effects of population size (as a surrogate for genetic factors) and prey consumption rates (as a surrogate for environmental quality) on fecundity in two sympatric species of wolf spider, Rabidosa punctulata and Rabidosa rabida. Population size was estimated in each of seven genetically isolated populations in each of 3 years using mark-recapture methods. Fecundity was estimated as the mean number of live offspring produced by ∼15 females sampled from each population of each species each year for 3 years. Prey consumption rates were estimated by sampling ∼300 spiders per population per year and assaying the proportion of spiders with prey. Larger populations have higher fecundity and more genetic diversity than smaller populations. Variation among populations in fecundity for a given year could be attributed most strongly to differences in population size, with variation in prey consumption rates and the interaction between population size and prey consumption playing smaller but still important roles. During the most stressful environmental conditions, the smallest populations of both species experienced disproportionately low-fecundity rates, more than doubling the estimated number of lethal equivalents during those years. The evidence presented in this paper for inbreeding-environment interactions at the population level and further evidence for a log-linear relationship between population size and fitness have important implications for conservation.     

Loss of genetic variability in social spiders: genetic and phylogenetic consequences of population subdivision and inbreeding

The consequences of population subdivision and inbreeding have been studied in many organisms, particularly in plants. However, most studies focus on the short‐term consequences, such as inbreeding depression. To investigate the consequences of both population fragmentation and inbreeding for genetic variability in the longer term, we here make use of a natural inbreeding experiment in spiders, where sociality and accompanying population subdivision and inbreeding have evolved repeatedly. We use mitochondrial and nuclear data to infer phylogenetic relationships among 170 individuals of Anelosimus spiders representing 23 species. We then compare relative mitochondrial and nuclear genetic variability of the inbred social species and their outbred relatives. We focus on four independently derived social species and four subsocial species, including two outbred–inbred sister species pairs. We find that social species have 50% reduced mitochondrial sequence divergence. As inbreeding is not expected to reduce genetic variability in the maternally inherited mitochondrial genome, this suggests the loss of variation due to strong population subdivision, founder effects, small effective population sizes (colonies as individuals) and lineage turnover. Social species have < 10% of the nuclear genetic variability of the outbred species, also suggesting the loss of genetic variability through founder effects and/or inbreeding. Inbred sociality hence may result in reduction in variability through various processes. Sociality in most Anelosimus species probably arose relatively recently (0.1–2 mya), with even the oldest social lineages having failed to diversify. This is consistent with the hypothesis that inbred spider sociality represents an evolutionary dead end. Heterosis underlies a species potential to respond to environmental change and/or disease. Inbreeding and loss of genetic variability may thus limit diversification in social Anelosimus lineages and similarly pose a threat to many wild populations subject to habitat fragmentation or reduced population sizes.     

Allozymes and fitness: Evolution of a problem

It was expected that studies of electrophoretic variability in natural populations would resolve longstanding controversies concerning the form of natural selection and its effect on genetic variance in fitness. Recent studies of fitness components for allozymes in E. coli and Drosophila, where genetic backgrounds have been rigidly controlled, and experiments designed to detect small selection coefficients, suggest that selection is much weaker than earlier investigations would indicate. However, perturbing the metabolic background associated with specific loci often allows functional differences to be amplified to an experimentally measurable level. Frequencies of null activity variants in natural populations indicate that the fitness consequences of reduced activity in heterozygoles are probably very small. These results are supported by recent theoretical considerations suggesting that the activity variation associated with electrophoretic variation will have little effect on overall flux in many pathways.     

Chloroplast DNA Diversity among Trees, Populations and Species in the California Closed-Cone Pines (Pinus Radiata, Pinus Muricata and Pinus Attenuata)

The amount, distribution and mutational nature of chloroplast DNA polymorphisms were studied via analysis of restriction fragment length polymorphisms in three closely related species of conifers, the California closed-cone pines-knobcone pine: Pinus attenuata Lemm.; bishop pine: Pinus muricata D. Don; and Monterey pine: Pinus radiata D. Don. Genomic DNA from 384 trees representing 19 populations were digested with 9-20 restriction enzymes and probed with cloned cpDNA fragments from Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] that comprise 82% of the chloroplast genome. Up to 313 restriction sites were surveyed, and 25 of these were observed to be polymorphic among or within species. Differences among species accounted for the majority of genetic (haplotypic) diversity observed [G(st) = 84(+/-13)%]; nucleotide diversity among species was estimated to be 0.3(+/-0.1)%. Knobcone pine and Monterey pine displayed almost no genetic variation within or among populations. Bishop pine also showed little variability within populations, but did display strong population differences [G(st) = 87(+/-8)%] that were a result of three distinct geographic groups. Mean nucleotide diversity within populations was 0.003(+/-0.002)%; intrapopulation polymorphisms were found in only five populations. This pattern of genetic variation contrasts strongly with findings from study of nuclear genes (allozymes) in the group, where most genetic diversity resides within populations rather than among populations or species. Regions of the genome subject to frequent length mutations were identified; estimates of subdivision based on length variant frequencies in one region differed strikingly from those based on site mutations or allozymes. Two trees were identified with a major chloroplast DNA inversion that closely resembled one documented between Pinus and Pseudotsuga.     

Effects of life history traits and sampling strategies on genetic diversity estimates obtained with RAPD markers in plants

A compilation of studies using RAPD markers for evaluating population differentiation resulted in 78 estimates of AMOVA-derived Φ_ST and 31 estimates of Nei's G_ST, as well as in 41 estimates of Nei's within-population diversity. In outcrossing taxa, estimates of between-population diversity were closely correlated with maximum geographic distance between sampled populations. A corresponding association was not found in selfing taxa. These results suggest that RAPD can be a sensitive method for detection of genetic structuring according to the isolation-by-distance model. However, it also means that sampling strategies, as applied in individual studies, can seriously influence the resulting estimates of between-population diversity. Other sampling strategies, like number of plants per population and number of scored polymorphic markers, do not seem to impart any serious artefacts. As previously verified with allozyme data, RAPD markers showed that long-lived, outcrossing, late successional taxa retain most of their genetic variability within populations. By contrast, annual, selfing and/or early successional taxa allocate most of the genetic variability among populations. Estimates for between- and within-population diversity, respectively, proved to be negatively correlated, as previously reported for allozyme data. The only major discrepancy between allozymes and RAPD markers concerns geographic range; within-population diversity was strongly affected by distributional range of the investigated species in the allozyme data but not in the RAPD data. Moreover, RAPD-based values for between-population diversity increased with increasing distributional range whereas the opposite has been reported in a large allozyme data compilation. Contrary to allozymes, RAPD marker-derived within-population diversity is probably therefore not a very good predictor of total species genetic diversity.     

Genetic variability in a population of arbuscular mycorrhizal fungi causes variation in plant growth

Different species of arbuscular mycorrhizal fungi (AMF) alter plant growth and affect plant coexistence and diversity. Effects of within-AMF species or within-population variation on plant growth have received less attention. High genetic variation exists within AMF populations. However, it is unknown whether genetic variation contributes to differences in plant growth. In our study, a population of AMF was cultivated under identical conditions for several generations prior to the experiments thus avoiding environmental maternal effects. We show that genetically different Glomus intraradices isolates from one AMF population significantly alter plant growth in an axenic system and in greenhouse experiments. Isolates increased or reduced plant growth meaning that plants potentially receive benefits or are subject to costs by forming associations with different individuals in the AMF population. This shows that genetic variability in AMF populations could affect host-plant fitness and should be considered in future research to understand these important soil organisms.     

Post-colonization temporal genetic variation of an introduced fly, Rhagoletis completa

Evolutionary biologists have been puzzled by the success of introduced species: despite founder effects that reduce genetic variability, invasive species are still successful at colonizing new environments. It is possible that the evolutionary processes during the post-colonization period may increase the genetic diversity and gene flow among invasive populations over time, facilitating their long-term success. Therefore, genetic diversity and population structure would be expected to show greater temporal variation for successful introduced populations than for native populations. We studied the population genetics of the walnut husk fly, Rhagoletis completa, which was introduced into California from the Midwestern US in the early 1900s. We used microsatellites and allozymes to genotype current and historic fly populations, providing a rare perspective on temporal variability in population genetic parameters. We found that introduced populations showed greater temporal fluctuations in allele frequencies than native populations. Some introduced populations also showed an increase in genetic diversity over time, indicating multiple introductions had occurred. Population genetic structure decreased in both native and introduced populations over time. Our study demonstrates that introduced species are not at equilibrium and post-colonization processes may be important in ameliorating the loss of genetic diversity associated with biological invasions.     

Levels of genetic polymorphism: marker loci versus quantitative traits.

Species are the units used to measure ecological diversity and alleles are the units of genetic diversity. Genetic variation within and among species has been documented most extensively using allozyme electrophoresis. This reveals wide differences in genetic variability within, and genetic distances among, species, demonstrating that species are not equivalent units of diversity. The extent to which the pattern observed for allozymes can be used to infer patterns of genetic variation in quantitative traits depends on the forces generating and maintaining variability. Allozyme variation is probably not strictly neutral but, nevertheless, heterozygosity is expected to be influenced by population size and genetic distance will be affected by time since divergence. The same is true for quantitative traits influenced by many genes and under weak stabilizing selection. However, the limited data available suggest that allozyme variability is a poor predictor of genetic variation in quantitative traits within populations. It is a better predictor of general phenotypic divergence and of postzygotic isolation between populations or species, but is only weakly correlated with prezygotic isolation. Studies of grasshopper and planthopper mating signal variation and assortative mating illustrate how these characters evolve independently of general genetic and morphological variation. The role of such traits in prezygotic isolation, and hence speciation, means that they will contribute significantly to the diversity of levels of genetic variation within and among species.     

Reproductive fitness,population size and genetic variation in Muscari tenuiflorum (Hyacinthaceae): The role of temporal variation

In plant populations a positive correlation between population size, genetic variation and fitness components is often found, due to increased pollen limitation or reduced genetic variation and inbreeding depression in smaller populations. However, components of fitness also depend on environmental factors which can vary strongly between years. The dry grassland species Muscari tenuiflorum experiences long term habitat isolation and small population sizes. We analyzed seed production of M. tenuiflorum in four years and its dependence on population size and genetic variation. Genetic diversity within populations was high (AFLP: H_e = 0.245; allozymes: H_e = 0.348). An analysis of molecular variance revealed considerable population differentiation (AFLP: 26%; allozyme: 17%). An overall pattern of isolation by distance was found, which, however was not present at distances below 20 km, indicating stronger effects of genetic drift. Genetic diversity was positively correlated to population size. Self pollination reduced seed set by 24%, indicating inbreeding depression. Reproductive fitness was not correlated to genetic diversity and a positive correlation with population size was present in two of four study years. The absence of a general pattern stresses the importance for multi-year studies. Overall, the results show that despite long term habitat isolation M. tenuiflorum maintains seed production in many years independent of population size. The long term persistence of populations is thus expected to depend less on intrinsic genetic or demographic properties affecting seed production but on successful plant establishment and persistence, which latter are based on conservation and protection of suitable habitats.     

AN EXPERIMENTAL EVALUATION OF SELECTION ON COLOR MORPHS OF THE POLYMORPHIC SPIDER ENOPLOGNATHA OVATA (ARANEAE: THERIDIIDAE)

The annual theridiid spider Enoplognatha ovata exhibits a genetically based color polymorphism of red and nonred phenotypes. We evaluated fitness differences between red and nonred spiders by manipulating morph frequencies in a population in which red morphs were rare (≤5%). Broods from red females were introduced to open experimental plots from which natural aggregations of spiders had been repeatedly removed. Control plots in which spiders were removed but not replaced were used to estimate spider immigration from surrounding vegetation into experimental plots. Morph frequencies observed in experimental plots one year following the manipulation were adjusted by immigration estimates and tested against frequencies predicted with the hypothesis of no selection. We found no evidence of selection against red morphs: female morph frequencies in experimental plots did not differ significantly from expected frequencies assuming no selection; female frequencies did not change significantly between subadult and adult stages; and red and nonred spiders exhibited similar fecundities. We conclude that 1) selection on E. ovata color morphs is not likely to be detected easily within a single population because of the swamping effect of dispersal and 2) local patterns of morph-frequency variation may arise more from dispersal and drift than from selection on the color phenotypes.     

Interdemic variation of cannibalism in a wolf spider (Pardosa monticola) inhabiting different habitat types

Abstract.  1. Cannibalism was investigated in the wolf spider Pardosa monticola (Clerck) using spiders collected from four populations with varying densities, inhabiting two different coastal dune habitat types. Sampled individuals were paired randomly and tested immediately for their cannibalism propensity.
2. The occurrence of cannibalism was found to be influenced by the size (cephalothorax width) of both the smaller and the larger spider of a pair. Larger size differences enhanced cannibalism.
3. Cannibalism rates were not significantly different in spiders from high-density compared with low-density populations. Cannibalism rates showed, however, large variability between habitat types, with higher rates in spiders from dune grasslands than from dune slacks. This is suggested to result from differences in prey availability throughout the growing season between both habitat types.
4. Different size classes of spiders did not use different microhabitats, indicating that microhabitat segregation as a cannibalism-avoidance behaviour is absent in this species.     

Differentiation between natural and cultivated populations of Medicago sativa (Leguminosae) from Spain: analysis with random amplified polymorphic DNA (RAPD) markers and comparison to allozymes

The conservation of a crop's wild relatives as genetic resources requires an understanding of the way genetic diversity is maintained in their populations, notably the effect of crop-to-wild gene flow. In this study, the amount of differentiation between natural and cultivated populations of Medicago sativa was analysed using random amplified polymorphic DNA (RAPD) markers and an extension of the AMOVA procedure adapted to autotetraploid organisms. Simulations of structured populations were performed to test whether AMOVA provides estimates of population structure in autotetraploids that can be directly compared to those obtained for allozyme data. Simulations showed that straight phi-statistics allow a good estimation of population differentiation when unbiased allelic frequencies are used to correct the conditional expectations of squared genetic distances. But such unbiased estimates can not be practically guaranteed, and population structure is notably overestimated when some populations are fixed for the presence of amplified fragments. However, removing fixed loci from the data set improves the statistical power of the test for population structure. The genetic variation of 15 natural and six cultivated populations of M. sativa was analysed at 25 RAPD loci and compared to estimates computed with allozymes on the same set of populations. Although RAPD markers revealed less within-population genetic diversity than allozymes, the quantitative and qualitative patterns of population structure were in full agreement with allozymes. This confirmed the conclusions drawn from the allozymic survey: crop-to-wild gene flow occurred in many locations, but some other mechanisms opposed cultivated traits to be maintained into natural populations.     

FITNESS CONSEQUENCES OF OUTCROSSING IN A SOCIAL SPIDER WITH AN INBREEDING MATING SYSTEM

Inbreeding mating systems are uncommon because of inbreeding depression. Mating among close relatives can evolve, however, when outcrossing is constrained. Social spiders show obligatory mating among siblings. In combination with a female‐biased sex ratio, sib‐mating results in small effective populations. In such a system, high genetic homozygosity is expected, and drift may cause population divergence. We tested the effect of outcrossing in the social spider Stegodyphus dumicola. Females were mated to sib‐males, to a non‐nestmate within the population, or to a male from a distant population, and fitness traits of F1s were compared. We found reduced hatching success of broods from between‐population crosses, suggesting the presence of population divergence at a large geographical scale that may result in population incompatibility. However, a lack of a difference in offspring performance between inbred and outbred crosses indicates little genetic variation between populations, and could suggest recent colonization by a common ancestor. This is consistent with population dynamics of frequent colonizations by single sib‐mated females of common origin, and extinctions of populations after few generations. Although drift or single mutations can lead to population divergence at a relatively short time scale, it is possible that dynamic population processes homogenize these effects at longer time scales.     

Development of genetic differentiation and postzygotic isolation in experimental metapopulations of spider mites

We studied the development of genetic differentiation and postzygotic isolation in experimental metapopulations of the two-spotted spider mite, Tetranychus urticae Koch. A genetically diverse starter population was made by allowing six inbred sublines to interbreed. Then three migration patterns were tested: no migration, or one or three immigrants per subpopulation per generation. Variations in four traits were investigated: allozymes, acaricide resistance, diapause, and hatchability. In the allozymes, acaricide resistance, and diapause, genetic variation among subpopulations became high in metapopulations with no migration, but not in the others, which showed that one immigrant is enough to prevent genetic differentiation. Hatchability, which was decreased by interbreeding among the six sublines, gradually recovered in succeeding generations. In metapopulations with no migration, hatchability was reduced again after in-migration at the 15th generation. Different karyotypes or coadapted gene complexes can survive in different subpopulations by genetic drift, and both Wolbachia-infected and -noninfected subpopulations may be selected, which would lead to postzygotic isolation between isolated subpopulations. Our results indicate that sampling effects such as genetic drift or stochastic loss of Wolbachia produce postzygotic isolation in laboratory populations of spider mite.     

Bromeliads provide shelter against fire to mutualistic spiders in a fire‐prone landscape

1. A key challenge in the study of mutualistic interactions is understanding sources of variation that strengthen or weaken these interactions. In spider–plant mutualisms, spiders benefit plants by improving plant nutrition and protecting plants from herbivory. Although the benefits of plants to spider growth and survival are often claimed, they are rarely demonstrated. 2. In this study, empirical evidence is provided that bromeliads (Bromelia balansae, Bromeliaceae) are essential for the resilience of the mutualistic bromeliad‐living jumping spider populations (Psecas chapoda, Salticidae) after a fire event, sheltering spiders from the heat of the flames. 3. Spider populations were compared before and after a natural fire event and it was shown that spiders of different ages survived the fire. The survival of such individuals allowed the population of P. chapoda spiders to recover rapidly, returning to pre‐fire levels in 5 months. 4. Bromeliads reduced the susceptibility of P. chapoda spiders to burning, and this mutualistic relationship contributed to the resilience of the spider population after a fire event. It is suggested that frequent fires in fire‐prone landscapes may have strengthened this spider–plant relationship, contributing to the maintenance and evolution of this association.     

Mammalian herbivores reveal marked genetic divergence among populations of an endangered plant species

Quantitative genetically based traits in dominant and keystone tree species can have extended effects on other biota and also on ecosystem processes. This has direct implications for managed plant systems, where choice of genetic stock in conservation or commercial plantings will affect the ecological and evolutionary trajectory of the associated biotic communities. Hence an understanding of genetic variation in quantitative traits, especially those that relate directly to fitness, should be incorporated into the management of species. In plants, quantitative traits such as foliar defences that mediate the complexity of biotic interactions (e.g. herbivory), may be key fitness traits to consider in the management of gene pools of species that are of high conservation value. In this paper we examine the interactions of an endangered eucalypt species, Eucalyptus morrisbyi and a marsupial herbivore, the common brushtail possum Trichosurus vulpecula. We investigate the genetic variability of resistance of plants sourced from two populations and genetic variability in foliage defences as key quantitative traits that may be essential for survival of this eucalypt species. Trichosurus vulpecula detect clear genetic divergence in the two E. morrisbyi populations as evidenced by their browsing preferences in the field. In addition, trees from the more susceptible population (Calverts Hill) suffered fitness consequences with lower flowering than trees from the more resistant population (Risdon Hills). Field feeding preferences were confirmed in captive feeding trials arguing differences were due to foliar attributes consistent with the genetic‐based differences observed in key chemical and physical foliage traits. Biotic interactions such as herbivory may affect populations of rare plant species. Results of this study highlight the need to understand the degree of genetic differentiation of resistance to herbivores and in the quantitative traits mediating these interactions in species of high conservation value, as these traits affect the adaptive potential of populations.     

Impact of flower-dwelling crab spiders on plant-pollinator mutualisms

Indirect effects in interactions occur when a species influences a third species by modifying the behaviour of a second one. It has been suggested that indirect effects of crab spiders (Thomisidae) on pollinator behaviour can cascade down the food web and negatively affect plant fitness. However, it is poorly understood how different pollinator groups react to crab spiders and, thus, when a reduction in plant fitness is likely to occur. Using continuous video surveillance, we recorded the behaviour of pollinators on two flower species and the pollinators’ responses to three crab spider treatments: inflorescences (1) with a pinned dried spider, (2) with a spider model made of paper, and (3) without spiders (control). We found that pollinators avoided inflorescences with dried spiders only on one plant species (Anthemis tinctoria). Pollinators showed no significant avoidance of paper spiders. Honeybees and bumblebees did not react to dried spiders, but solitary bees and syrphid flies showed a strong avoidance. Finally, we found no evidence that inflorescences with dried spiders suffered from a decrease in fitness in terms of a reduced seed set. We hypothesise that top-down effects of predators on plants via pollinators depend on the degree of specialisation of pollinators and their tendency to avoid spiders.     

The clonal and population structure of a rare endemic plant, Wyethia reticulata (Asteraceae): allozyme and RAPD analysis

Genetic structure arises when limited gene flow between populations favours the development of distinct arrays of genetic characters within each population. Determining the spatial scale at which this differentiation occurs is critical to our understanding of population biology and microevolution of species. The genetic structure and spatial pattern of genetic variation in an endemic, clonal perennial, Wyethia reticulata E. Greene, was investigated using random amplified polymorphic DNA (RAPD) markers and allozyme alleles. Large stands (250–360 m²) were found to contain few genetic individuals. Despite the small population sizes and endemism of the species, W. reticulata was highly diverse genetically, with most of the variation (75–81%) distributed within populations. A population structure in full agreement with spatially defined populations was achieved only by combining RAPD and allozyme markers. Analysis using both types of markers appeared to provide estimates of genetic similarity between individuals that were most consistent with empirical data on plant distributions. We postulated that large, long-lived clones dominated genetic relationships within populations but also provided opportunities for gene flow between populations on a longer time scale. The two marker types yielded different estimates of between-individual similarity and revealed disparate patterns of population structure. This result will arise because allozymes and random DNA segments have dissimilar evolutionary dynamics with respect to mutation and selection.     

The importance of genetic variation for the viability of small avian populations—the Seychelles Warbler and the African Marsh Warbler

Eising, C.M., Blaakmeer, K.B. & Komdeur, J. 2000. The importance of genetic variation for the viability of small avian populations—the Seychelles Warbler and the African Marsh Warbler. Ostrich 71 (1 & 2): 304–306.

Nowadays, many natural populations have to face up to problems such as genetic drift and ‘forced’ inbreeding as a result of reduced numbers and population isolation. This is thought to have a major effect on their survival. Research on the isolated Seychelles warbler showed that fitness parameters are not negatively affected by a high inbreeding frequency and a low level of heterozygosity. However, one has to be careful to translate these results to other isolated species. It is hypothesised that the effect of ‘genetic erosion’ on fitness may be less disastrous for island species, which are used to go through narrow population bottlenecks, as compared to continental species. Conservationists should be aware of these differences between species and of the fact that the long-term prospects can be reduced by lack of adaptive genetic potential.