High Variance In Reproductive Success Research Articles

No evidence for temporally balanced selection on larval Pacific oysters Crassostrea gigas: a comment on Durland et al. (2021).

No evidence for temporally balanced selection on larval Pacific oysters Crassostrea gigas: a comment on Durland et al. (2021).

Effects of spawning group sex ratio and stocking density on the outcome of captive reproduction in golden cuttlefish Sepia esculenta

The supplemental release of hatchery-reared animals is important for maintaining or enhancing fishery resources. The golden cuttlefish, Sepia esculenta, is a commercially important cephalopod utilized in stock enhancement activities throughout East Asia. The long-term success of such activities requires proper breeding regimes to ensure the efficient use of resources and the preservation of genetic diversity in the offspring. For captive-bred cephalopods, information concerning the effects of different mating regimes on the paternity of offspring is scarce. However, it could be of particular importance for polyandric species such as golden cuttlefish. As previous studies have suggested that both the number and mating order of males are related to female fecundity, we investigated the effects of stocking density and sex ratio on female fecundity and determined the paternity of the offspring in golden cuttlefish. We used two experimental approaches. First, four different female (F): male (M) sex ratios (1F:1M, 1F:2.5 M, 1F:3M, 1F:6M) were set up to investigate the effects of operational sex ratio on mating behavior, female fecundity, and hatchling paternity. Next, we investigated female fecundity and hatchling paternity patterns under low and high stocking densities (5 or 10 individuals ∙ m−3), following a standard sex ratio setting that is widely employed in cuttlefish hatcheries (1F:2M). With an increasing proportion of males, the average fecundity of females and the participation rate of males in paternity decreased. With increasing stocking density, the females had lower average daily egg production, and the males had a high variance in reproductive success and decreased participation rate in spawning. The estimated effective number of breeders (Ne) was 19.29 (Ne/N ratio = 0.54) at high-density and 10.17 (Ne/N ratio = 0.57) at low density. To achieve high reproductive performance and assure genetic diversity of golden cuttlefish offspring, our study suggests that equal (1F:1M) or slightly male-biased (1F:2.5 M) sex ratios and a low stocking density may be useful when breeding golden cuttlefish in captivity.

Genetic characterization of wild, broodstock and seed samples of Polititapes rhomboides (Bivalvia: Veneridae): Implications for hatchery seed production

The banded carpet shell Polititapes rhomboides is a fishery resource and a suitable candidate for the development of native clam aquaculture in Europe. Here, we analyzed nine microsatellite loci in wild samples to provide estimates of genetic diversity and population differentiation for NW Spain, the main production area of this clam. We also analyzed wild-caught broodstock and hatchery-reared seed to investigate the genetic consequences of seed production for stock enhancement purposes by mass spawning. We examined the seed sample as a whole (total seed), but also subdivided it into three size classes to assess the genetic consequences of size grading. While wild samples, broodstock included, displayed minimal or no genetic differences, the total seed showed a reduction in allelic richness (12–24 %) and a significant level of differentiation (FST = 0.026−0.043) with respect to wild samples. After performing parentage assignment, we detected a drastic reduction in the effective number of breeders (Ne) compared to the census number (85 %), and an inbreeding rate of 0.036 per generation. The low Ne and high inbreeding rate were mainly due to high variance in reproductive success. The seed size classes also showed a decrease in allelic richness and significant genetic differentiation, but we did not find significant differences in parental contribution to each size class. Our results may help the genetic-assisted management of wild populations, give insight into the genetic composition of the seed produced by mass spawning and provide a basis for the development of more effective hatchery practices and sustainable stock enhancement programs in P. rhomboides.

Deciphering genetic mate choice: Not so simple in group-housed conservation breeding programs.

Incorporating mate choice into conservation breeding programs can improve reproduction and the retention of natural behaviors. However, different types of genetic‐based mate choice can have varied consequences for genetic diversity management. As a result, it is important to examine mechanisms of mate choice in captivity to assess its costs and benefits. Most research in this area has focused on experimental pairing trials; however, this resource‐intensive approach is not always feasible in captive settings and can interfere with other management constraints. We used generalized linear mixed models and permutation approaches to investigate overall breeding success in group‐housed Tasmanian devils at three nonmutually exclusive mate choice hypotheses: (a) advantage of heterozygous individuals, (b) advantage of dissimilar mates, and (c) optimum genetic distance, using both 1,948 genome‐wide SNPs and 12 MHC‐linked microsatellites. The managed devil insurance population is the largest such breeding program in Australia and is known to have high variance in reproductive success. We found that nongenetic factors such as age were the best predictors of breeding success in a competitive breeding scenario, with younger females and older males being more successful. We found no evidence of mate choice under the hypotheses tested. Mate choice varies among species and across environments, so we advocate for more studies in realistic captive management contexts as experimental or wild studies may not apply. Conservation managers must weigh up the need to wait for adequate sample sizes to detect mate choice with the risk that genetic changes may occur during this time in captivity. Our study shows that examining and integrating mate choice into the captive management of species housed in realistic, semi‐natural group‐based contexts may be more difficult than previously considered.

Genetic diversity and effective population size in successive mass selected generations of black shell strain Pacific oyster (Crassostrea gigas) based on microsatellites and mtDNA data

The Pacific oyster (Crassostrea gigas) is one of the highly fecund marine aquaculture species and has a large variance in reproductive success, these characteristics can result in loss of genetic diversity and inbreeding especially when conducting mass selection. In the current study, we investigated the level of genetic diversity and effective population size over three successive mass-selected generations in the black shell strain of C. gigas assessed by 11 microsatellites as well as mitochondrial COI region. The results showed that the genetic variation over three generations were maintained since no detectable depression of expected heterozygosity (He = 0.647–0.681), number of alleles (N = 5.6–6.0) and haplotypes (Nh = 2). The difference in alleles and haplotypes number between the base population (BP) and three mass selective generations (M5-M7) could be mainly due to the loss during preceding family selection process. Pair-wise FST values along with AMOVA analysis from both markers indicated unremarkable differentiation within each generation including BP. There was no deleterious effect on genetic diversity and population structure for mass selected generations imposed by our artificial breeding practice. The Neb estimation for M5, M6 and M7 was 29.8, 40.7 and 52.7, respectively. The different number of broodstocks used for each generation implied that using a balanced sex ratio and large size of broodstock as well as low selection pressure would help to increase the effective population size and avoid high level of inbreeding. The estimated linkage disequilibrium-based effective breeders size (Neb) was significantly lower than sex-ratio correction effective population size (Ne), which indicated high variance in family contribution. It is suggested during mass spawning to simply undertake several mini-spawn groups and then pool the embryos to produce next generation, which can be a hedge to the high variance in reproductive success, thus, the loss of effective populations size and diversity. Other practical strategies against the loss of genetic diversity were also discussed, however, it remains to be investigated how diverse selection and follow-up procedures will benefit retaining genetic variability. This study will provide an insight into the level of genetic diversity and effective population size within mass-selected black shell line and enable a better understanding of how efficient current breeding practices are at maintaining genetic variation. This information can be applied for future selective breeding program and in the design of suitable management guidelines for sustainable breeding of C. gigas.

Variance in Reproductive Success is Driven by Environmental Factors, not Mating System, in Bonytails.

Studying the reproductive ecology of aggregate broadcast spawning fishes is difficult because it generally is not feasible to sample all potential parents and unambiguously assign their offspring. We used molecular-based parentage analysis to gain insights into the reproductive ecology of the endangered Bonytail, and to evaluate whether protected off-channel habitats could be used as an alternative to hatchery production. By genotyping adults and offspring stocked (n = 4130) into two experimental backwaters across three years, we determined that most adults (82-97%) contributed to progeny production across years and backwaters, with one exception. Both sexes mated multiply and the number of mates and family size were positively correlated. There was also a positive correlation between adult size and metrics of reproductive success. There were strong interactions between sample years and backwaters suggesting that environmental factors are the primary driver of variance in reproductive success. Knowledge of mating systems and sources of variance in reproductive success is important for management of endangered fish because high variance in reproductive success leads to substantial losses of genetic variation when few individuals reproduce successfully. Although variance in reproductive success was observed, most adults contributed to genetically diverse progeny in experimental backwaters. These results support the use of predator-free, but otherwise natural, backwaters as an effective conservation tool for reintroducing Bonytail to its native habitat.

Current genetic status, temporal stability and structure of the remnant wild European flat oyster populations: conservation and restoring implications

The flat oyster (Ostrea edulis) is one of the most appreciated molluscs in Europe, but natural beds have been greatly reduced due to harvesting and the effects of the parasite Bonamia ostreae. Characterization of current wild populations is required to develop long-term bed restoration programmes by enhancing genetic diversity and tolerance to bonamiosis. Oysters from different locations corresponding to the main natural beds from Denmark, The Netherlands, England, Ireland, France and Spain, including two different cohorts per location were sampled in 2011 and 2013. Sixteen microsatellite loci were used to study temporal and geographical genetic structure. Temporal variation was low, although sometimes significant probably due to high variance in reproductive success among individuals. Conversely, samples from different countries showed much higher genetic divergence, and Ireland and France presented differences among locations within country. Clustering analyses grouped samples into three main geographical regions, associated with oceanic fronts: one group constituted by The Netherlands and Denmark; another by France, Ireland and England; and a third one exclusively by Spain. Effective population sizes (Ne) within regions were high (>1000), which reflects population stability and low levels of genetic drift. The presence of regional genetic structure shows the potential for local adaptation of O. edulis and suggests caution when transplanting individuals, especially between distant geographical regions.

Influences of population density on polyandry and patterns of sperm usage in the marine gastropod Rapana venosa.

Polyandry is a common mating strategy in animals, with potential for sexual selection to continue post-copulation through sperm competition and/or cryptic female choice. Few studies have investigated the influences of population density on polyandry and sperm usage, and paternity distribution in successive broods of marine invertebrates. The marine gastropod Rapana venosa is ideal for investigating how population density influences the frequency of polyandry and elucidating patterns of sperm usage. Two different population density (12 ind/m3 and 36 ind/m3) treatments with two replications were set to observe reproductive behaviors. Five microsatellite markers were used to identify the frequency of multiple paternity and determine paternal contributions to progeny arrays in 120 egg masses. All of the mean mating frequency, mean number of sires and mean egg-laying frequency were higher at high population density treatment relative to low population density treatment, indicating population density is an important factor affecting polyandry. The last sperm donors achieved high proportions of paternity in 74.77% of egg masses, which supported the “last male sperm precedence” hypothesis. In addition, high variance in reproductive success among R. venosa males were detected, which might have an important influence on effective population size.

Social monogamy vs. polyandry: ecological factors associated with sex roles in two closely related birds within the same habitat.

Why mainly males compete and females take a larger share in parental care remains an exciting question in evolutionary biology. Role-reversed species are of particular interest, because such 'exceptions' help to test the rule. Using mating systems theory as a framework, we compared the reproductive ecology of the two most contrasting coucals with regard to sexual dimorphism and parental care: the black coucal with male-only care and the biparental white-browed coucal. Both species occur in the same lush habitat and face similar ecological conditions, but drastically differ in mating system and sexual dimorphism. Black coucals were migratory and occurred at high breeding densities. With females being obligatory polyandrous and almost twice as heavy as males, black coucals belong to the most extreme vertebrates with reversed sexual dimorphism. Higher variance in reproductive success in fiercely competing females suggests that sexual selection is stronger in females than in males. In contrast, resident white-browed coucals bred at low densities and invariably in pairs. They were almost monomorphic and the variance in reproductive success was similar between the sexes. Black coucals were more likely to lose nests than white-browed coucals, probably facilitating female emancipation of parental care in black coucals. We propose that a combination of high food abundance, high population density, high degree of nest loss and male bias in the adult sex ratio represent ecological conditions that facilitate role reversal and polyandry in coucals and terrestrial vertebrates in general.

Condition Dependent Effects on Sex Allocation and Reproductive Effort in Sequential Hermaphrodites

Theory predicts the optimal timing of sex change will be the age or size at which half of an individual's expected fitness comes through reproduction as a male and half through reproduction as a female. In this way, sex allocation across the lifetime of a sequential hermaphrodite parallels the sex allocation of an outbreeding species exhibiting a 1∶1 ratio of sons to daughters. However, the expectation of a 1∶1 sex ratio is sensitive to variation in individual condition. If individuals within a population vary in condition, high-condition individuals are predicted to make increased allocations to the sex with the higher variance in reproductive success. An oft-cited example of this effect is seen in red deer, Cervus elaphus, in which mothers of high condition are more likely to produce sons, while those in low condition are more likely to produce daughters. Here, we show that individual condition is predicted to similarly affect the pattern of sex allocation, and thus the allocation of reproductive effort, in sequential hermaphrodites. High-condition sex-changers are expected to obtain more than half of their fitness in the high-payoff second sex and, as a result, are expected to reduce the allocation of reproductive effort in the initial sex. While the sex ratio in populations of sequential hermaphrodites is always skewed towards an excess of the initial sex, condition dependence is predicted to increase this effect.

Reproductive Variance of Brood Dams and Sires used in Restoration Enhancement of Spotted Seatrout in Texas Bays and Estuaries

AbstractSpawning patterns and reproductive variance of Spotted Seatrout Cynoscion nebulosus dams and sires at two restoration enhancement facilities in Texas were assessed across a spawning year by using parentage analysis based on 12 variable microsatellite loci. In total, 72.6% of all dams and sires contributed to at least one spawning event, although in contrasting patterns. Across all spawning events assayed, more sires contributed to each spawn, on average, than did dams; dams had considerably higher variance in reproductive success than sires. Dams alternatively had a higher average contribution to the number of progeny from a single spawn but also a much higher variance in the number of progeny produced per spawn. The variation in the number of progeny produced per dam and per sire and the number of actual mating combinations led to an average reduction of 64.3% in the genetic effective population size (Ne) per spawn relative to the maximum Ne that would be expected if (1) all possible dam × sire mating combinations occurred at random and (2) all families contained an equal number of progeny. Averaged over all spawns, the actual number of mating combinations accounted for approximately 83.6% of the total reduction in Ne, while variation in family size accounted for 16.4% of the total reduction in Ne. Results from this and other studies indicate that reductions in Ne of hatchery‐ or farm‐raised progeny stem primarily from noncontributing dams, suggesting that periodic identification and removal of low‐contributing dams from broodfish stocks constitute a critical step toward maximizing the Ne of hatchery offspring used in restoration enhancement.

Spatial and temporal patterns of larval dispersal in a coral‐reef fish metapopulation: evidence of variable reproductive success

Many marine organisms can be transported hundreds of kilometres during their pelagic larval stage, yet little is known about spatial and temporal patterns of larval dispersal. Although traditional population-genetic tools can be applied to infer movement of larvae on an evolutionary timescale, large effective population sizes and high rates of gene flow present serious challenges to documenting dispersal patterns over shorter, ecologically relevant, timescales. Here, we address these challenges by combining direct parentage analysis and indirect genetic analyses over a 4-year period to document spatial and temporal patterns of larval dispersal in a common coral-reef fish: the bicolour damselfish (Stegastes partitus). At four island locations surrounding Exuma Sound, Bahamas, including a long-established marine reserve, we collected 3278 individuals and genotyped them at 10 microsatellite loci. Using Bayesian parentage analysis, we identified eight parent-offspring pairs, thereby directly documenting dispersal distances ranging from 0km (i.e., self-recruitment) to 129km (i.e., larval connectivity). Despite documenting substantial dispersal and gene flow between islands, we observed more self-recruitment events than expected if the larvae were drawn from a common, well-mixed pool (i.e., a completely open population). Additionally, we detected both spatial and temporal variation in signatures of sweepstakes and Wahlund effects. The high variance in reproductive success (i.e., 'sweepstakes') we observed may be influenced by seasonal mesoscale gyres present in the Exuma Sound, which play a prominent role in shaping local oceanographic patterns. This study documents the complex nature of larval dispersal in a coral-reef fish, and highlights the importance of sampling multiple cohorts and coupling both direct and indirect genetic methods in order disentangle patterns of dispersal, gene flow and variable reproductive success.

Local genetic patchiness but no regional differences between Indo-West Pacific populations of the dogtooth tuna Gymnosarda unicolor

Physical barriers can have profound impacts on dispersal in marine species. Here, we investigate population structure and levels of relatedness among individuals of the coral reef associated fish, the dogtooth tuna Gymnosarda unicolor, collected from 15 sites across the Indo-West Pacific region. We screened 92 individuals for genetic variation at 13 nuclear microsatellite loci and the cytochrome c oxidase subunit 1 (COI) mitochondrial DNA (mtDNA) gene. We detected no genetic differentiation between ocean basins or between sites within ocean basins, suggesting G. unicolor possesses a highly mobile larval or juvenile stage. In addition, the lack of deep evolutionary mtDNA divergences suggests gene flow was also not limited historically. However, comparisons of relatedness between pairs of individuals revealed that individuals collected from the same site were more related to each other, on average, than individuals collected from different sites. Such patterns are consistent with chaotic genetic patchiness, a possible consequence of high variance in reproductive success and patchy, local recruitment.

High variance in reproductive success generates a false signature of a genetic bottleneck in populations of constant size: a simulation study

BackgroundDemographic bottlenecks can severely reduce the genetic variation of a population or a species. Establishing whether low genetic variation is caused by a bottleneck or a constantly low effective number of individuals is important to understand a species’ ecology and evolution, and it has implications for conservation management. Recent studies have evaluated the power of several statistical methods developed to identify bottlenecks. However, the false positive rate, i.e. the rate with which a bottleneck signal is misidentified in demographically stable populations, has received little attention. We analyse this type of error (type I) in forward computer simulations of stable populations having greater than Poisson variance in reproductive success (i.e., variance in family sizes). The assumption of Poisson variance underlies bottleneck tests, yet it is commonly violated in species with high fecundity.ResultsWith large variance in reproductive success (Vk ≥ 40, corresponding to a ratio between effective and census size smaller than 0.1), tests based on allele frequencies, allelic sizes, and DNA sequence polymorphisms (heterozygosity excess, M-ratio, and Tajima’s D test) tend to show erroneous signals of a bottleneck. Similarly, strong evidence of population decline is erroneously detected when ancestral and current population sizes are estimated with the model based method MSVAR.ConclusionsOur results suggest caution when interpreting the results of bottleneck tests in species showing high variance in reproductive success. Particularly in species with high fecundity, computer simulations are recommended to confirm the occurrence of a population bottleneck.

Fast evolutionary genetic differentiation during experimental colonizations

Founder effects during colonization of a novel environment are expected to change the genetic composition of populations, leading to differentiation between the colonizer population and its source population. Another expected outcome is differentiation among populations derived from repeated independent colonizations starting from the same source. We have previously detected significant founder effects affecting rate of laboratory adaptation among Drosophila subobscura laboratory populations derived from the wild. We also showed that during the first generations in the laboratory, considerable genetic differentiation occurs between foundations. The present study deepens that analysis, taking into account the natural sampling hierarchy of six foundations, derived from different locations, different years and from two samples in one of the years. We show that striking stochastic effects occur in the first two generations of laboratory culture, effects that produce immediate differentiation between foundations, independent of the source of origin and despite similarity among all founders. This divergence is probably due to powerful genetic sampling effects during the first few generations of culture in the novel laboratory environment, as a result of a significant drop in Ne. Changes in demography as well as high variance in reproductive success in the novel environment may contribute to the low values of Ne. This study shows that estimates of genetic differentiation between natural populations may be accurate when based on the initial samples collected in the wild, though considerable genetic differentiation may occur in the very first generations of evolution in a new, confined environment. Rapid and significant evolutionary changes can thus occur during the early generations of a founding event, both in the wild and under domestication, effects of interest for both scientific and conservation purposes.

Relationship between effective population size, inbreeding and adult fitness‐related traits in a steelhead (Oncorhynchus mykiss) population released in the wild

Inbreeding is of concern in supportive breeding programmes in Pacific salmonids, Oncorhynchus spp, where the number of breeding adults is limited by rearing space or poor survival to adulthood, and large numbers are released to supplement wild stocks and fisheries. We reconstructed the pedigree of 6602 migratory hatchery steelhead (Oncorhynchus mykiss) over four generations, to determine the incidence and fitness consequences of inbreeding in a northwest USA programme. The hatchery maintained an effective population size, Ñ(e) = 107.9 from F(0) to F(2), despite an increasing census size (N), which resulted in a decreasing N(e)/N ratio (0.35 in F(0) to 0.08 in F(2)). The reduced ratio was attributed to a small broodstock size, nonrandom transfers and high variance in reproductive success (particularly in males). We observed accumulation of inbreeding from the founder generation (in F(4), percentage individuals with inbreeding coefficients Δf > 0 = 15.7%). Generalized linear mixed models showed that body length and weight decreased significantly with increasing Δf, and inbred fish returned later to spawn in a model that included father identity. However, there was no significant correlation between Δf and age at return, female fecundity or gonad weight. Similarly, there was no relationship between Δf and reproductive success of F(2) and F(3) individuals, which might be explained by the fact that reproductive success is partially controlled by hatchery mating protocols. This study is one of the first to show that small changes in inbreeding coefficient can affect some fitness-related traits in a monitored population propagated and released to the wild.

Nest distribution affects behaviour and mating success in a marine fish

The distribution of breeding resources, such as nest sites, can have a pronounced impact on a population by affecting the proportion of individuals that succeed to breed and hence, the variation in reproductive success. Aggregation of important resources can lead to resource monopolisation by a limited number of individuals and thus affect the intensity of sexual selection. In this study, we tested, by contrasting two experimental treatments (dispersed vs. aggregated), how nest distribution affects: (1) mating behaviour, (2) male nest occupation and mating success, and (3) reproductive success and the opportunity for selection. We used the two-spotted goby (Gobiusculus flavescens), a small marine fish with a resource-based mating system, as our model species. When nests were aggregated, a larger proportion of the males behaved aggressively, fewer males succeeded in occupying a nest, fewer males became mated, and those males that mated received fewer eggs from spawning females. These effects resulted in a higher variance in reproductive success and hence, a higher opportunity for selection (Irs), in the aggregated treatment. We suggest that the results are a direct consequence of males defending a territory around their nest, preventing competitively inferior males from breeding. However, we found no significant selection differentials for body length or condition of males in either treatment. Our results support the hypothesis that aggregation of essential resources like nests promotes resource monopolisation. In species facing highly clumped nesting resources in the wild, monopolisation may negatively impact population productivity but could lead to strong selection on traits that promote male competitive ability.

Maintenance of genetic variation and panmixia in the commercially exploited western rock lobster (Panulirus cygnus)

Marine species with high fecundities and mortalities in the early life stages can have low effective population sizes, making them vulnerable to declines in genetic diversity when they are commercially harvested. Here, we compare levels of microsatellite and mitochondrial sequence variation in the western rock lobster (Panulirus cygnus) over a 14-year period to test whether genetic variation is being maintained. Panulirus cygnus is a strong candidate for loss of genetic variation because it is a highly fecund species that is likely to experience high variance in reproductive success due to an extended larval planktonic stage. It also supports one of the largest and most economically important fisheries in Australia, with landings of between 8,000 and 14,500 tons (~70 % of the total legal-sized biomass) being harvested in some years. We found remarkably high levels of genetic variation in all samples and no evidence of a decline in genetic diversity over the time interval we studied. Furthermore, there was no evidence of a recent genetic bottleneck, and effective population size estimates based on single sample and temporal methods were infinitely large. Analysis of molecular variance indicated no significant population structure along 960 km of coastline or genetic differentiation among temporal samples. Our results support the view that P. cygnus is a single, panmictic population, and suggest genetic drift is not strong enough to reduce neutral genetic diversity in this species if current management practices and breeding stock sizes are maintained.

Oceanographic drivers of offspring abundance may increase or decrease reproductive variance in a temperate marine fish

In species that reproduce into uncertain environments, the relationship between mean reproductive success (the abundance of new recruits) and the variance in reproductive success (whether adults contribute disproportionally more offspring) may not be straightforward because of stochastic environmental processes that create high variance in reproductive success among adults. In this study, we investigated the relationships between oceanography, reproductive success and reproductive variance in the black rockfish, Sebastes melanops, a long-lived temperate reef fish with pelagic larvae. We quantified black rockfish recruitment, genetic diversity and growth rates from otolith microstructure over 5 years (2005-2009) during which oceanographic conditions differed. We used cross-correlations to determine windows of time during which oceanographic variables were significantly correlated with the resulting abundance or genetic diversity of recruits. We found that warmer ocean temperatures were positively correlated with the abundance of recruits, as well as the effective number of breeders. In contrast, the strength of coastal upwelling during settlement was positively correlated with the annual abundance of new recruits, but was negatively correlated with the effective number of breeders. Larval growth rates were explained substantially more by temperature than by upwelling and suggested that temperature affected survival through growth, while upwelling affected survival through transport. Our results indicated that cold ocean temperatures and intense upwelling caused sweepstakes-like processes to operate on black rockfish populations, despite high abundances of recruits. We propose that a decoupling of the mean and variance in reproductive success may be characteristic of organisms that reproduce into uncertain environments.

Temporal Genetic Similarity Among Year-Classes of the Pacific Geoduck Clam (Panopea generosaGould 1850): A Species Exhibiting Spatial Genetic Patchiness

A previous study revealed genetic differences among collections of the commercially exploited geoduck clam (Panopea generosa) in Puget Sound, WA, but this heterogeneity did not follow an isolation-by-distance model. In this study, we investigated whether these differences were ephemeral or stable and tested predictions of the sweepstakes recruitment hypothesis, in which individuals show a high variance in reproductive success. We genotyped 11 allozyme and 7 microsatellite loci in 2,021 geoducks from 2 sites in Puget Sound and aged individuals by counting annuli in thin-section chondrophores under light microscopy. Genotypic data were then collated by year-class to test predictions of the sweepstakes recruitment hypothesis with allele count rarefaction, year-class relatedness, and 3 estimators of efective population size (Ne) using temporal shifts in allele frequencies. Although estimates of Ne were similar among year-classes, spatial shifts in allele frequencies and year-class strengths were detected among stations at 1 site, indicating that patchy settlement may be the result of an interaction between larval behavior during dispersal and hydrology.