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1.
In many marine invertebrates, long‐distance dispersal is achieved during an extended pelagic larval phase. Although such dispersal should result in high gene flow over broad spatial scales, fine‐scale genetic structure has often been reported, a pattern attributed to interfamilial variance in reproductive success and limited homogenization during dispersal. To examine this hypothesis, the genetic diversity of dispersing larvae must be compared with the postdispersal stages, that is benthic recruits and adults. Such data remain, however, scarce due to the difficulty to sample and analyse larvae of minute size. Here, we carried out such an investigation using the marine gastropod Crepidula fornicata. Field sampling of three to four larval pools was conducted over the reproductive season and repeated over 3 years. The genetic composition of larval pools, obtained with 16 microsatellite loci, was compared with that of recruits and adults sampled from the same site and years. In contrast to samples of juveniles and adults, large genetic temporal variations between larval pools produced at different times of the same reproductive season were observed. In addition, full‐ and half‐sibs were detected in early larvae and postdispersal juveniles, pointing to correlated dispersal paths between several pairs of individuals. Inbred larvae were also identified. Such collective larval dispersal was unexpected given the long larval duration of the study species. Our results suggest that each larval pool is produced by a small effective number of reproducers but that, over a reproductive season, the whole larval pool is produced by large numbers of reproducers across space and time.  相似文献   

2.

Background

Predicting the spatial and temporal patterns of marine larval dispersal and supply is a challenging task due to the small size of the larvae and the variability of oceanographic processes. Addressing this problem requires the use of novel approaches capable of capturing the inherent variability in the mechanisms involved.

Methodology/Principal Findings

In this study we test whether dispersal and connectivity patterns generated from a bio-physical model of larval dispersal of the crab Carcinus maenas, along the west coast of the Iberian Peninsula, can predict the highly variable daily pattern of wind-driven larval supply to an estuary observed during the peak reproductive season (March–June) in 2006 and 2007. Cross-correlations between observed and predicted supply were significant (p<0.05) and strong, ranging from 0.34 to 0.81 at time lags of −6 to +5 d. Importantly, the model correctly predicted observed cross-shelf distributions (Pearson r = 0.82, p<0.001, and r = 0.79, p<0.01, in 2006 and 2007) and indicated that all supply events were comprised of larvae that had been retained within the inner shelf; larvae transported to the outer shelf and beyond never recruited. Estimated average dispersal distances ranged from 57 to 198 km and were only marginally affected by mortality.

Conclusions/Significance

The high degree of predicted demographic connectivity over relatively large geographic scales is consistent with the lack of genetic structuring in C. maenas along the Iberian Peninsula. These findings indicate that the dynamic nature of larval dispersal can be captured by mechanistic biophysical models, which can be used to provide meaningful predictions of the patterns and causes of fine-scale variability in larval supply to marine populations.  相似文献   

3.
For marine invertebrates with a benthic adult form and a planktonic larva phase, the connectivity among populations is mainly based on larval dispersal. While an extended larval phase will promote gene flow, other factors such as an intensive fishery and geographical barriers could lead to changes in genetic variability. In this study, the population genetic structure of the commercial crab Metacarcinus edwardsii was analyzed along 700 km of the Chilean coast. The analysis, based on eight microsatellite loci genotyped from megalopae and adult crabs, considered temporal and spatial patterns of genetic variation. The results showed no evidence of spatial patterns in genetic structure, suggesting high connectivity among the sampling sites. The temporal analysis showed no evidence of changes in allele frequencies and no evidence of a recent bottleneck. The lack of spatial structure and allele variation over time could be explained by the interaction of factors such as i) low reproductive variance due to the capability of females to store sperm in the seminal receptacle, which can be used for successive broods, ii) high larval dispersal and iii) high individual reproductive output. Using our data as priors, a genetic modelling approach coincided, predicting this temporal and spatial stability. The same analysis showed that a reduction in population size leads to the loss of genetic variability in populations, as well as of the genetic cohesiveness between populations, pointing out the importance management for species under exploitation, such as M. edwardsii.  相似文献   

4.
Inbreeding may lead to morphological malformations in a wide variety of taxa. We used genetic markers to evaluate whether malformed urodeles were more inbred and/or had less genetic diversity than normal salamanders. We captured 687 adult and 1259 larval tiger salamanders (Ambystoma tigrinum tigrinum), assessed each individual for gross malformations, and surveyed genetic variation among malformed and normal individuals using both cytoplasmic and nuclear markers. The most common malformations in both adults and larvae were brachydactyly, ectrodactyly and polyphalangy. The overall frequency of adults with malformations was 0.078 compared to 0.081 in larval samples. Genetic diversity was high in both normal and malformed salamanders, and there were no significant difference in measures of inbreeding (f and F), allele frequencies, mean individual heterozygosity or mean internal relatedness. Environmental contaminants or other extrinsic factors may lead to genome alternations that ultimately cause malformations, but our data indicate that inbreeding is not a causal mechanism.  相似文献   

5.
The Native Fishes Work Group, formed in 1991, developed and implemented a protocol to enhance the dwindling razorback sucker population in Lake Mohave, Arizona-Nevada. This large, genetically diverse population is severely reduced in size as a result of recruitment failure associated with predation on larvae. To circumvent this problem, wild larvae are captured, reared in protective custody until they are large enough to escape predation, and then released back into the lake. We present results of a monitoring program designed to assess the effectiveness of the sampling design in transmitting the high genetic diversity found in wild adults. Variation in a fragment from the mitochondrial DNA gene cytochrome b was examined by analysis of single-stranded polymorphisms and direct sequencing. Samples were characterized from three life history stages. Characterization of wild adults verified previous results that identified considerable diversity and provided baseline data. Samples of larvae from several temporal collections from throughout the spawning season and four geographical areas were characterized for 7 years (1997-2003) to assess the transmission of genetic variation from wild adults to larvae. Several analyses identified significant differences among temporal collections, resulting from sampling errors associated with finite number of females spawning at a given time and place. Comparisons among areas and years failed to identify significant variation, indicating that pooled collections for each year possess the same levels and patterns of genetic variation. Examination of repatriates representing 11 years (1992-2002) also failed to identify significant differences among cohorts; however, some sample sizes were small and the amova may lack sufficient power to detect differences. Contrasts of wild adults, larvae, and repatriates identified statistically significant differences among collections within these three groups; however, levels of variation are small and not biologically meaningful. More importantly, this analysis failed to detect significant differences among adults, larvae, and repatriates indicating that the program has been achieving its goal of transmitting variation from adults through the larvae and into the repatriate population. The reproductive capability of repatriates has not been examined, so it is unknown if the program will maintain genetic variation found in the original adult population. This will be most easily achieved by periodic monitoring of genetic variation in larval samples. If levels of variation become reduced in repatriates, levels and patterns of diversity in larvae are also expected to become reduced, and deviations in estimates of genetic diversity may become larger and more frequent. If this is the case, intervention may be necessary to ensure that certain individuals are not over-represented in the repatriate population.  相似文献   

6.
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 0 km (i.e., self‐recruitment) to 129 km (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.  相似文献   

7.
Identifying the sex of larvae is important in social Hymenoptera. Until now for Polistes wasps it has been necessary to genotype larvae at microsatellite loci, and assign their sex based on homozygosity at these loci. In our study on the paper wasp Polistes dominulus we have found morphological differences between larval sexes that can be used for larvae from the 3rd instar on to easily and cheaply identify larval sex: the external gonopore and the shape and size of larval gonads. The robustness of these indicators was supported by genotype data at four microsatellite loci. Using gonopore and gonad features for sex assignment will assign diploid males as males, unlike techniques based on genetic loci or chromosomes. Received 12 July 2006; revised 4 January 2007; accepted 5 February 2007.  相似文献   

8.
Major histocompatibility complex (MHC)-dependent mating preferences have been observed across vertebrate taxa and these preferences are expected to promote offspring disease resistance and ultimately, viability. However, little empirical evidence linking MHC-dependent mate choice and fitness is available, particularly in wild populations. Here, we explore the adaptive potential of previously observed patterns of MHC-dependent mate choice in a wild population of Atlantic salmon (Salmo salar) in Québec, Canada, by examining the relationship between MHC genetic variation and adult reproductive success and offspring survival over 3 years of study. While Atlantic salmon choose their mates in order to increase MHC diversity in offspring, adult reproductive success was in fact maximized between pairs exhibiting an intermediate level of MHC dissimilarity. Moreover, patterns of offspring survival between years 0+ and 1+, and 1+ and 2+ and population genetic structure at the MHC locus relative to microsatellite loci indicate that strong temporal variation in selection is likely to be operating on the MHC. We interpret MHC-dependent mate choice for diversity as a likely bet-hedging strategy that maximizes parental fitness in the face of temporally variable and unpredictable natural selection pressures.  相似文献   

9.
Determining the conditions under which populations may persist requires accurate estimates of demographic parameters, including immigration, local reproductive success, and mortality rates. In marine populations, empirical estimates of these parameters are rare, due at least in part to the pelagic dispersal stage common to most marine organisms. Here, we evaluate population persistence and turnover for a population of orange clownfish, Amphiprion percula, at Kimbe Island in Papua New Guinea. All fish in the population were sampled and genotyped on five occasions at 2-year intervals spanning eight years. The genetic data enabled estimates of reproductive success retained in the same population (reproductive success to self-recruitment), reproductive success exported to other subpopulations (reproductive success to local connectivity), and immigration and mortality rates of sub-adults and adults. Approximately 50% of the recruits were assigned to parents from the Kimbe Island population and this was stable through the sampling period. Stability in the proportion of local and immigrant settlers is likely due to: low annual mortality rates and stable egg production rates, and the short larval stages and sensory capacities of reef fish larvae. Biannual mortality rates ranged from 0.09 to 0.55 and varied significantly spatially. We used these data to parametrize a model that estimated the probability of the Kimbe Island population persisting in the absence of immigration. The Kimbe Island population was found to persist without significant immigration. Model results suggest the island population persists because the largest of the subpopulations are maintained due to having low mortality and high self-recruitment rates. Our results enable managers to appropriately target and scale actions to maximize persistence likelihood as disturbance frequencies increase.  相似文献   

10.
As a consequence of free spawning in the unpredictable nearshore environment, marine species with large fecundities and high pre-reproductive mortality may be subject to extreme variance in reproductive success. If the unpredictability of the ocean results in only a small subset of the adult population contributing to each larval cohort, then reproduction may be viewed as a sweepstakes, with chance events determining which adults are successful each spawning season. Such a reproductive sweepstakes scenario may partially account for large reductions in effective population sizes relative to census population sizes in marine species. We evaluated two predictions of the sweepstakes reproductive success hypothesis by testing: (1) whether sea urchin recruits contain reduced genetic variation relative to the adult population; and (2) whether cohorts of sea urchin recruits are genetically differentiated. Mitochondrial DNA sequences were collected from 283 recently settled Strongylocentrotus purpuratus recruits from four annual cohorts spanning seven years in locations throughout California. Observed haplotype numbers and haplotype diversities showed little evidence of reduced genetic variation in the recruits relative to the diversity estimated from a previously reported sample of 145 S. purpuratus adults. Different cohorts of recruits were in some cases mildly differentiated from each other. A computer simulation of sweepstakes recruitment indicates that our sampling strategy had sufficient statistical power to detect large variances in reproductive success.  相似文献   

11.
Douek J  Amar KO  Rinkevich B 《Genetica》2011,139(11-12):1531-1542
Aspects of maternal-planula larval genetics in the monoecious scleractinian coral Stylophora pistillata (Red Sea, Eilat) were studied by amplified fragment length polymorphism (AFLP) methodology in two successive reproductive seasons. In total, 293 planulae and 10 adult colonies were analyzed. In June 2006, 147 planulae were collected from 10 shallow water colonies. In March, April and June 2007, 146 additional planulae were sampled from five of the ten 2006 sampled colonies. All AFLP products showed unalike band profiles indicating a fully sexual production pattern. We used 181 and 210 putative AFLP loci, of which the overall level of polymorphism in 2006 was 92 and 99 % in 2007 (respectively). Differences were also observed between 2006 and 2007 reproductive seasons in terms of total average gene diversity (0.191 vs. 0.247, respectively), suggesting fast turnover of sperm donor genotypes. In addition, increased numbers of potential sperm donor colonies in the vicinity of gravid females showed no impact on genetic differentiation levels in released larvae. UPGMA tree revealed clustering of maternal genotypes and their offspring, suggesting, as expected, high relatedness between planulae and their mothers. In addition, the average heterozygosity of each group of siblings was persistently lower than heterozygosity calculated for the respective maternal colony, suggesting the possibility of partial inbreeding. This trend of reduced genetic heterogeneity in Stylophora pistillata is an alarming sign for populations residing in the northern Red Sea coral reefs.  相似文献   

12.
Life history traits of highly vagile marine species, such as adult reproductive success and larval dispersal, are strongly determined by oceanographic and climatic forces. Nevertheless, marine organisms may show restricted dispersal in time and space. Patterns of isolation by distance (IBD) have been repeatedly observed in marine species. If spawning time is a function of geographical location, temporal and spatial isolation, can easily be confounded or misinterpreted. In this study, we aimed at discriminating between various forces shaping the genetic composition of recruiting juveniles of the European eel (Anguilla anguilla L.). By controlling for geographical variation, we assessed temporal variation and tested for possible isolation by time (IBT) between recruitment waves within and between years. Using 12 polymorphic allozyme and six variable microsatellite loci, we show that genetic differentiation was low (F(ST) = 0.01-0.002) and significant between temporal samples. Regression analysis between genetic and temporal distance, was consistent with a subtle interannual pattern of IBT. Our data suggest that the population dynamics of the European eel may be governed by a double pattern of temporal variance in genetic composition: (i) a broad-scale IBT of spawning cohorts, possibly as a consequence of the large migration loop in anguillids and strong variance in annual adult reproductive contribution; and (ii) a smaller-scale variance in reproductive success (genetic patchiness) within cohorts among seasonally separated spawning groups, most likely originating from fluctuating oceanic and climatic forces. The consistency of both mechanisms remains to be verified with fine-scale analyses of both spawning/migrating aged adults and their offspring to confirm the stochastic/deterministic nature of the IBT pattern in eel.  相似文献   

13.
长江口沿岸碎波带仔稚鱼的种类组成及其多样性特征   总被引:21,自引:0,他引:21  
为了进一步探明长江口沿岸碎波带仔稚鱼的种类组成和多样性特征,2006年7月-2007年6月的每月大潮期间,在长江口沿岸碎波带13个站位点的水深0.5-1.5m处,两人沿海岸平行方向步行拖曳小型拖网(1m×4m,网目1mm)采集仔稚鱼样本。周年共拖网397网次,采集到仔稚鱼49045尾。仔稚鱼隶属于31科,共84种,其中海洋性鱼类16种,河口性鱼类30种,洄游性鱼类4种,淡水性鱼类34种。洄游性鱼类刀鲚占总个体数的82.63%,为最优势种。仔稚鱼的物种数、个体数和丰度在冬季较低,以3月份最低,夏季最高。Margalef丰富度指数2006年11月—2007年4月相对较低,2006年10月最高。Shannon-Wiener多样性指数在10月最高,7月最低。Pielou均匀度指数全年较恒定,2006年7月和2007年6月相对较低。  相似文献   

14.
The life histories of oysters in the genus Crassostrea, like those of most marine bivalves, are typified by high fecundity and low survival in nature. Rearing conditions in hatcheries however ensure optimized density, diet, and temperature. Hatcheries are becoming increasingly important for the production of juveniles in aquaculture, and their culture practices often include culling of slow growing larvae to reduce and synchronize the time taken to reach settlement. Because previous studies have found substantial genetic variation for early life developmental traits in Crassostrea gigas, these culling practices are likely to cause highly different selective pressures in hatcheries from those in the natural environment. We studied the phenotypic and genetic impact of such culling practices in a factorial cross between 10 males and 3 females subjected to progressive culling of the smallest 50% of larvae, compared with a non-culled control. Measurements were made on larval growth, survival, time taken to attain pediveliger stage and settlement success. Culling had a larger effect on the variance of these larval traits than on their means. The larvae in culled cultures were approximately 10% larger than those in controls, whereas the coefficient of variation was reduced by 30-40%. Culling also reduced the mean time to settlement by 12% and its variance by 55%. Using a multiplexed set of microsatellite markers to trace parentage, we also estimated the variance in reproductive success in a controlled experiment to quantify the consequences of intensive hatchery rearing practices. We also focused on changes in effective population size and genetic structure over time (and developmental stages). Our results show a loss of genetic diversity following removal of the smallest larvae by culling, as well as temporally varying genetic structure of the larval population. This supports the existence of genetic variability in early life developmental traits in C. gigas. Culling in hatcheries, like size-related selective pressures in the wild, are likely to have a significant genetic impact, through their effects on the timing of settlement.  相似文献   

15.
Habitat fragmentation has been shown to disrupt ecosystem processes such as plant-pollinator mutualisms. Consequently, mating patterns in remnant tree populations are expected to shift towards increased inbreeding and reduced pollen diversity, with fitness consequences for future generations. However, mating patterns and phenotypic assessments of open-pollinated progeny have rarely been combined in a single study. Here, we collected seeds from 37 Eucalyptus incrassata trees from contrasting stand densities following recent clearance in a single South Australian population (intact woodland=12.6 trees ha−1; isolated pasture=1.7 trees ha−1; population area=10 km2). 649 progeny from these trees were genotyped at eight microsatellite loci. We estimated genetic diversity, spatial genetic structure, indirect contemporary pollen flow and mating patterns for adults older than the clearance events and open-pollinated progeny sired post-clearance. A proxy of early stage progeny viability was assessed in a common garden experiment. Density had no impact on mating patterns, adult and progeny genetic diversity or progeny growth, but was associated with increased mean pollen dispersal. Weak spatial genetic structure among adults suggests high historical gene flow. We observed preliminary evidence for inbreeding depression related to stress caused by fungal infection, but which was not associated with density. Higher observed heterozygosities in adults compared with progeny may relate to weak selection on progeny and lifetime-accumulated mortality of inbred adults. E. incrassata appears to be resistant to the negative mating pattern and fitness changes expected within fragmented landscapes. This pattern is likely explained by strong outcrossing and regular long-distance pollen flow.  相似文献   

16.
Biological invasions represent ideal systems for the study of evolutionary processes associated with colonization events. It has been hypothesized that the genetic diversity is generally decreasing from the centre of the range to the margins due to multiple founder events. Invasive populations offer the opportunity to test this hypothesis at a fine spatial and temporal scale. We analysed the genetic structure of a large expanding non-native population of the Common Wall Lizard (Podarcis muralis) in Passau (Germany) using thirteen microsatellite loci. We analyzed the genetic structure and levels of admixture across a transect reflecting the expansion process and tested for a loss of genetic diversity and an increase of genetic differentiation from the centre to the invasion front. Our results demonstrate that significant genetic population structure can emerge rapidly at a small spatial scale. We found a trend for an increase in genetic differentiation and a decrease in genetic diversity from the invasion centre to the expanding range margin, suggesting that genetic drift is the major factor causing this pattern. The correlation between genetic diversity and average genetic differentiation was significant among sites. We hypothesize that the territoriality of P. muralis generates sufficient rates of noncontiguous and stratified dispersal from longer established sites to maintain significant genetic diversity at the invasion front. Simultaneously, territoriality might restrict the colonization success of migrants at established sites, so that in combination with founder events a strong differentiation arises.  相似文献   

17.
Many marine organisms have pelagic larvae and these are often important agents of dispersal and establishment of exotic species. The larval phase and the hermaphroditic reproductive system of the invasive marine gastropod Crepidula fornicata are crucial in determining the success of this species along the European coasts. To analyse these factors, we developed seven microsatellite loci specific to C. fornicata. On the same set of individuals, the microsatellite loci exhibited a greater level of polymorphism than previously studied allozyme markers and, thus, will permit fine‐scale analysis and larval pool studies.  相似文献   

18.
Chaotic genetic patchiness denotes unexpected patterns of genetic differentiation that are observed at a fine scale and are not stable in time. These patterns have been described in marine species with free‐living larvae, but are unexpected because they occur at a scale below the dispersal range of pelagic larvae. At the scale where most larvae are immigrants, theory predicts spatially homogeneous, temporally stable genetic variation. Empirical studies have suggested that genetic drift interacts with complex dispersal patterns to create chaotic genetic patchiness. Here we use a coancestry model and individual‐based simulations to test this idea. We found that chaotic genetic patterns (qualified by global FST and spatio‐temporal variation in FST's between pairs of samples) arise from the combined effects of (1) genetic drift created by the small local effective population sizes of the sessile phase and variance in contribution among breeding groups and (2) collective dispersal of related individuals in the larval phase. Simulations show that patchiness levels qualitatively comparable to empirical results can be produced by a combination of strong variance in reproductive success and mild collective dispersal. These results call for empirical studies of the effective number of breeders producing larval cohorts, and population genetics at the larval stage.  相似文献   

19.
Population‐level consequences of dispersal ability remain poorly understood, especially for marine animals in which dispersal is typically considered a species‐level trait governed by oceanographic transport of microscopic larvae. Transitions from dispersive (planktotrophic) to nondispersive, aplanktonic larvae are predicted to reduce connectivity, genetic diversity within populations, and the spatial scale at which reproductive isolation evolves. However, larval dimorphism within a species is rare, precluding population‐level tests. We show the sea slug Costasiella ocellifera expresses both larval morphs in Florida and the Caribbean, regions with divergent mitochondrial lineages. Planktotrophy predominated at 11 sites, 10 of which formed a highly connected and genetically diverse Caribbean metapopulation. Four populations expressed mainly aplanktonic development and had markedly reduced connectivity, and lower genetic diversity at one mitochondrial and six nuclear loci. Aplanktonic dams showed partial postzygotic isolation in most interpopulation crosses, regardless of genetic or geographic distance to the sire's source, suggesting that outbreeding depression affects fragmented populations. Dams from genetically isolated and neighboring populations also exhibited premating isolation, consistent with reinforcement contingent on historical interaction. By increasing self‐recruitment and genetic drift, the loss of dispersal may thus initiate a feedback loop resulting in the evolution of reproductive isolation over small spatial scales in the sea.  相似文献   

20.
For organisms with great fecundity and high mortality in early life stages, such as shellfish or fishes, the need to match reproductive activity with environmental conditions conducive to spawning, fertilization, larval development and recruitment may result in extreme variance in reproductive success among individuals. The main objective of this study was to investigate evidence of large variance in the reproductive success of the striped bass Morone saxatilis in the Santee–Cooper system, South Carolina, USA. Seven microsatellite loci were analysed in 603 recruits representing three yearly cohorts from 1992 to 1994, and a group analysis was performed to identify full-sib families. Large variance in reproductive success was detected, with a few large, full-sib families contributing disproportionately to each of the cohorts. The severity of sweepstakes reproductive success varied among cohorts depending on environmentally imposed mortality. Estimations of the effective number of breeders in these long-lived fish ranged from 24 in 1992 to 44 in 1994. Furthermore, the estimated genetic effective population size ( N e = 93) is approximately four orders of magnitude lower than estimates of adult census size ( N  =   362 000). Furthermore, the presence of large full-sib families indicates that striped bass engage in pair mating in the wild. Heterogeneity in genetic composition was also observed among cohorts, suggesting that genetically different adults contribute to different cohorts and that chance rather than fitness variation determines reproductive success.  相似文献   

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