Variation In Reproductive Success Research Articles

Concurrent declines in nestling diet quality and reproductive success of a threatened seabird over 150 years

Successful conservation of threatened species is often hindered by a lack of long-term data required to identify the vital rates contributing to population decline and the extrinsic factors influencing those rates. Museum collections can provide a valuable resource for reconstructing the historic demography and diet of otherwise elusive species. Here, we used age ratios (the relative number of hatch-year to after-hatch-year individuals) to examine the hypothesis that population declines in a threatened seabird, the marbled murrelet Brachyramphus marmoratus, are due to declines in reproductive success over the past 150 yr. We also used stable-nitrogen isotopes to exam- ine the hypothesis that variation in reproductive success over this period is related to the quality of food received by young in the nest. Age ratios from at-sea surveys conducted from 1994 to 2001 were significantly lower than museum collection age ratios for the period 1860 to 1950. Stable-nitrogen isotope values indicated that the trophic feeding level of marbled murrelet nestlings declined signif- icantly (-2.6‰) from 1854 to 2008. Our results suggest that the reproductive success of marbeled murrelets breeding in the Salish Sea has declined over the past 150 yr and that declines in nestling diet quality may be partly responsible. Overall, our results support the idea that managers should consider the quality of both nesting and marine foraging habitat as they attempt to improve repro- ductive success and population growth rate in this threatened species.

Low and variable reproductive success of a neotropical tyrant-flycatcher, Chapada Flycatcher (Suiriri islerorum)

The decline of several bird populations has been related to reduced reproductive success. Here we used the logistic-exposure method to model the temporal (age, daily, annual) variation in reproductive success of a neotropical tyrant-flycatcher, the Chapada Flycatcher (Suiriri islerorum), which has been described only recently (2001). We monitored 56 nests, in a reserve in central Brazil, over four seasons from 2003 to 2006. Predation affected 66% of the monitored nests, but varied greatly across years. The probability of daily survival decreased 7% day−1 with the age of the nest and the reproductive success varied greatly between years (16.8, 75.4, 6.7 and 21.4%, 2003–06 respectively). Mean reproductive success (using three analytical methods) ranged from 21 to 25%, but from only 13 to 16% when results from 2004 were excluded. Such low and variable reproductive success is atypical for a neotropical bird. In combination with the small population size and known decline of this population, our findings raise conservation concerns for this isolated population.

Fishery‐induced demographic changes in the timing of spawning: consequences for reproductive success*

AbstractDemography can have a significant effect on reproductive timing and the magnitude of such an effect can be comparable to environmentally induced variability. This effect arises because the individuals of many fish species spawn progressively earlier within a season and may produce more egg batches over a longer period as they get older, thus extending their lifetime spawning duration. Inter‐annual variation in spawning time is a critical factor in reproductive success because it affects the early environmental conditions experienced by progeny and the period they have to complete phases of development. By reducing the average lifetime spawning duration within a fish stock, fishing pressure could be increasing the variability in reproductive success and reducing long‐term stock reproductive potential. Empirical estimates of selection on birth date, from experiments and using otolith microstructure, demonstrate that there is considerable variation in selection on birth date both within a spawning season and between years. The few multi‐year studies that have linked egg production with the survival of progeny to the juvenile stage further highlight the uncertainty that adults face in timing their spawning to optimize offspring survival. The production of many small batches of eggs over a long period of time within a season and over a lifetime is therefore likely to decrease variance and increase mean progeny survival. Quantifying this effect of demography on variability in survival requires a focus on lifetime reproductive success rather than year specific relationships between recruitment and stock reproductive potential. Modelling approaches are suggested that can better quantify the likely impact of changing spawning times on year‐class strength and lifetime reproductive potential. The evidence presented strengthens the need to avoid fishing severely age truncated fish stocks.

Variable reproductive success in fragmented populations

Marine habitats are naturally patchy and anthropogenic disturbance can further fragment them. Many marine animals are sessile as adults or obligate inhabitants of particular habitats, so populations living in isolated patches of habitat are linked largely by dispersal of planktonic larvae. Theoretically, larvae are more likely to find and settle into large patches of habitat than small patches, thus small habitat patches may experience a more discontinuous supply of recruits resulting in small populations with unusual size- or age-structures or odd sex ratios — conditions where Allee effects on reproductive success are likely. We tested this hypothesis for the Caribbean spotted spiny lobster ( Panulirus guttatus), an obligate inhabitant of coral patch reefs whose mating dynamics are size-dependent. We found that P. guttatus were less abundant on small reefs where their size structure and per capita reproductive success were significantly more variable, particularly among large females that are susceptible to sperm limitation that diminishes fertilization rates. These results are indicative of Allee effects and provide a mechanistic understanding of how size-dependent mating dynamics influence reproductive success in ways that alter population dynamics in patchy habitats.

Interannual variation in pup production and the timing of breeding in benthic foraging Australian fur seals

AbstractLarge interannual variations in reproductive success caused by fluctuations in oceanography and prey availability are common to many species of air breathing epipelagic predators. In contrast, little is known about variation in benthic foragers such as Australian fur seals (Arctocephalus pusillus doriferus). Between 1997 and 2007, pup production was assessed in 9 yr, while the timing of breeding and adult female condition was assessed in 5 yr at Kanowna Island in Bass Strait, southeastern Australia. Pup production was variable (= 1,726 ± 42, range = 1,386–2,301), but without temporal trend, as was median birth date (= 23 November ± 1, range = 21–25 November) and pupping synchrony (period of 90% births: = 28 ± 2 d, range = 23–31 d). Pup production was negatively correlated with median birth date and positively correlated with female condition, winter sea‐surface temperature (SST) and zonal wind strength within Bass Strait. Pup production was also negatively correlated with SST in the previous summer within Bass Strait and in the eastern Great Australian Bight upwelling region. The results suggest that the reproductive success of Australian fur seals is influenced by oceanography but less so than in otariids foraging epipelagically in major upwellings. Despite spanning several El Niño events, no correlation between pup production and the Southern Oscillation Index was observed.

Microsatellite pedigree analysis reveals high variance in reproductive success and reduced genetic diversity in hatchery-spawned northern abalone

The northern (or pinto) abalone, Haliotis kamtschatkana, is a broadcast-spawning marine gastropod that was recently listed as endangered in Canada. To aid in species recovery, a captive-breeding and supplementation program is underway in Barkley Sound, British Columbia. We genotyped first generation progeny for five microsatellite loci and used a pedigree reconstruction program (Pedigree 2.2) to identify their genealogical relationships in the absence of information on parental genotypes. We analyzed progeny from three separate group-spawning events and inferred considerable variation in the number of offspring produced by each parent; in the most severe case a single male sired all the progeny produced during one spawning event. After only one generation of captive-breeding we found a 55–60% reduction in allelic richness and a 17–18% reduction in heterozygosity relative to the diverse wild source population. This study illustrates the difficulty of managing genetic diversity in hatchery populations of a broadcast-spawning species, even when gametes are collected separately from each individual broodstock.

Maternal characteristics and the production and recruitment of sons in the eastern kingbird (Tyrannus tyrannus)

Sex allocation theory predicts that if variance in reproductive success differs between the sexes, females who are able to produce high-quality young should bias offspring sex ratio towards the sex with the higher potential reproductive success. We tested the hypothesis that high-quality (i.e., heavy) female eastern kingbirds (Tyrannus tyrannus) that bred early in the breeding season would produce male-biased clutches. A significant opportunity for sexual selection also exists in this socially monogamous but cryptically polygamous species, and we predicted that successful extra-pair (EP) sires would be associated with an excess of male offspring. Although population brood sex ratio did not differ from parity, it increased significantly with female body mass and declined with female breeding date, but was independent of the morphology and display (song) behavior (correlates of reproductive success) of social males and EP sires. Male offspring were significantly heavier than female offspring at fledging. Moreover, the probability that male offspring were resighted in subsequent years declined with breeding date, and was greater in replacement clutches, but lower when clutch size was large. Probability of resighting female offspring varied annually, but was independent of all other variables. Given that variance in reproductive success of male kingbirds is much greater than that of females, and that male offspring are more expensive to produce and have a higher probability of recruitment if fledged early in the season, our results support predictions of sex allocation theory: high-quality (heavy) females breeding when conditions were optimal for male recruitment produced an excess of sons.

Seasonal modulation of reproductive effort during early pregnancy in humans

Life history theory predicts that early pregnancy presents a relatively low cost, uncontested opportunity for a woman to terminate investment in a current reproductive opportunity if a conceptus is of poor quality and/or maternal status or environmental conditions are not propitious for a successful birth. We tested this hypothesis in rural Bolivian women experiencing substantial seasonal variation in workload and food resources. Significant risk factors for early pregnancy loss (EPL) included agropastoralism versus other economic strategies, conception during the most arduous seasons versus other seasons, and increasing maternal age. Anovulation rate (AR) was higher during the most arduous seasons and in older women. Breastfeeding and indicators of social status and living conditions did not significantly influence either risk of EPL or AR. Averaged over the year, anovulation occurred in about 1/4 of the cycles and EPL occurred in about 1/3 of the conceptions. This is the first evidence of seasonality of EPL in a non-industrialized population, and the first to demonstrate a relationship between economic activities and EPL. These findings suggest that both anovulation and EPL are potential mechanisms for modulating reproductive effort; such "failures" may also be nonadaptive consequences of conditions hostile to a successful pregnancy. In either case, variation in EPL risk associated with different subsistence activities can be expected to influence fertility levels and birth seasonality in both contemporary and past human populations. These consequences of variability in the risk of EPL can impact efforts to understand the sources of variation in reproductive success.

Modelling evolutionary processes in small populations: not as ideal as you think

Evolutionary processes are routinely modelled using 'ideal' Wright-Fisher populations of constant size N in which each individual has an equal expectation of reproductive success. In a hypothetical ideal population, variance in reproductive success (V(k)) is binomial and effective population size (N(e)) = N. However, in any actual implementation of the Wright-Fisher model (e.g., in a computer), V(k) is a random variable and its realized value in any given replicate generation (V(k)*) only rarely equals the binomial variance. Realized effective size (N(e)*) thus also varies randomly in modelled ideal populations, and the consequences of this have not been adequately explored in the literature. Analytical and numerical results show that random variation in V(k)* and N(e)* can seriously distort analyses that evaluate precision or otherwise depend on the assumption that N(e)* is constant. We derive analytical expressions for Var(V(k)) [4(2N - 1)(N - 1)/N(3)] and Var(N(e)) [N(N - 1)/(2N - 1) approximately N/2] in modelled ideal populations and show that, for a genetic metric G = f(N(e)), Var(G) has two components: Var(Gene) (due to variance across replicate samples of genes, given a specific N(e)*) and Var(Demo) (due to variance in N(e)*). Var(G) is higher than it would be with constant N(e) = N, as implicitly assumed by many standard models. We illustrate this with empirical examples based on F (standardized variance of allele frequency) and r(2) (a measure of linkage disequilibrium). Results demonstrate that in computer models that track multilocus genotypes, methods of replication and data analysis can strongly affect consequences of variation in N(e)*. These effects are more important when sampling error is small (large numbers of individuals, loci and alleles) and with relatively small populations (frequently modelled by those interested in conservation).

Reproductive Success and Sexual Selection in Wild Eastern Tiger Salamanders (Ambystoma t. tigrinum)

Variation in reproductive success is most pronounced in species with strongly biased operational sex ratios, prominent sexual dimorphisms, and where mate competition and choice are likely. We studied sexual selection in eastern tiger salamanders (Ambystoma t. tigrinum) and examined the role of body size on reproductive success. We genotyped 155 adults and 1,341 larvae from 90 egg masses at six microsatellite loci. Parentage analyses revealed both sexes engaged in multiple matings, but was more common among females (64%) than males (27%). However, the standardized variance in mating and reproductive success was higher in males. Bateman gradients were significant and nearly identical in both sexes, suggesting that sexual selection was roughly equal between sexes. Body size was not correlated with mating or reproductive success in either sex. The apparent lack of sexual selection on body size may be a result of sperm storage, sperm competition, alternative mating tactics, and/or random induction of spermatophores.

Temporal effective size estimates of a managed walleye Sander vitreus population and implications for genetic‐based management

The goal of this research was to use the long-term fishery data set and DNA from archived scales of walleye Sander vitreus in Escanaba Lake, WI, U.S.A., to improve the understanding of the underlying mechanism(s) influencing genetic diversity in naturally recruiting populations. The introduced population of S. vitreus in Escanaba Lake has a low mean effective population size (N(E)) between 124.6 and 185.5 despite a mean census size (N(C)) of 4659 (N(E)/N(C)c. 0.04), suggesting an accelerated rate of genetic drift between 1952 and 2002. These values are smaller than the median N(E) range of several studies suggesting typical N(E)/N(C) ratios of 0.11-0.16 in a wide range of taxa. N(E) increased steadily during the past two sampled decades (1992 and 2002) and was consistent with a lowering of the variance in S. vitreus reproductive success, possibly linked to a large, sustained exploitation (mean 28%) rate. Variance in reproductive success is one of the most important factors influencing N(E) in species, like S. vitreus, which have a potential for large fecundities and large juvenile mortalities (type III survivorship). The N(B) estimates across six sequential cohorts (age classes of S. vitreus, assayed from 1994 to 1999) was consistent with estimates of N(E) reported for 1992-2002. These results, coupled with in-depth census and exploitation data, show that the genetic characteristics of Escanaba Lake S. vitreus have changed substantially and that management activities, such as supplemental stocking and harvest practices, have profoundly influenced the genetic dynamics of S. vitreus in this lake.

The fitness advantage of a high-performance weapon

Weapons used in combat between males are usually attributed to sexual selection, which operates via a fitness advantage for males with weapons of better ‘quality’. Because the performance capacity of morphological traits is typically considered the direct target of selection, Darwin’s intrasexual selection hypothesis can be modified to predict that variation in reproductive success should be explained by variation in performance traits relevant to combat. Despite such a straightforward prediction, tests of this hypothesis are conspicuously lacking. We show that territorial male collared lizards with greater bite-force capacity sire more offspring than weaker biting rivals but exhibit no survival advantage. We did not detect stabilizing or disruptive selection on bite-force capacity. Taken together, these results support the hypothesis that superior weapon performance provides a fitness advantage through increased success in male contests. Sexual selection on weapon performance therefore appears to be a force driving the evolution and maintenance of sexual dimorphism in head shape. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96, 840‐845.

Diet specificity is not associated with increased reproductive performance of Golden EaglesAquila chrysaetosin Western Scotland

Amongst raptor species, individuals with specialized diets are commonly observed to have higher reproductive output than those with general diets. A suggested cause is that foraging efficiency benefits accrue to diet specialists. This diet specificity hypothesis thus predicts that diet breadth and reproductive success should be inversely related within species. We highlight, however, that a prey availability hypothesis also makes the same prediction in some circumstances. Hence, when high diet specificity results from high encounter rates with an abundant, preferred prey, then prey availability may affect reproductive success, with diet specialization as an incidental correlate. Using three insular study areas in western Scotland, we examine diet specificity and reproductive success in Golden EaglesAquila chrysaetos. Diet breadth and breeding productivity were not negatively related in any of our study areas, even though birds with specific diets did tend to have a higher incidence of preferred prey (grouse and lagomorphs) in the diet. Indeed, in two study areas there was evidence that diet generalists had higher breeding productivity. Our results therefore failed to support the diet specificity hypothesis but were consistent with the prey availability hypothesis. We highlight that although many other studies are superficially consistent with the diet specificity hypothesis, our study is not alone in failing to provide support and that the hypothesis does not provide a generic explanation for all relevant results. Diet specificity in predators can be at least partially a response to prey diversity, availability and distribution, and benefits associated with different prey types, so that being a generalist is not necessarily intrinsically disadvantageous. We suggest that the available evidence is more consistent with variation in prey abundance and availability as a more influential factor explaining spatial and temporal variation in breeding productivity of ‘generalist’ species such as the Golden Eagle. Under this argument, prey abundance and availability are the main drivers of variation in reproductive output. Diet specificity is a consequence of variation in prey availability, rather than a substantial cause of variation in reproductive success.

It takes two to tango: reproductive skew and social correlates of male mating success in a lek-breeding bird.

Variance in reproductive success among individuals is a defining characteristic of many social vertebrates. Yet, our understanding of which male attributes contribute to reproductive success is still fragmentary in most cases. Male-male reproductive coalitions, where males jointly display to attract females, are of particular interest to evolutionary biologists because one male appears to forego reproduction to assist the social partner. By examining the relationship between social behaviour and reproductive success, we can elucidate the proximate function of coalitions in the context of mate choice. Here, we use data from a 4-year study of wire-tailed manakins (Pipra filicauda) to provide molecular estimates of reproductive skew and to test the hypothesis that male-male social interactions, in the context of coordinated displays, positively influence a male's reproductive success. More specifically, we quantify male-male social interactions using network metrics and predict that greater connectivity will result in higher relative reproductive success. Our data show that four out of six leks studied had significant reproductive skew, with success apportioned to very few individuals in each lek. Metrics of male social affiliations derived from our network analysis, especially male connectivity, measured as the number of males with whom the focal male has extended interactions, were strong predictors of the number of offspring sired. Thus, network connectivity is associated with male fitness in wire-tailed manakins. This pattern may be the result of shared cues used by both sexes to assess male quality, or the result of strict female choice for coordinated display behaviour.

Sex-specific spatio-temporal variability in reproductive success promotes the evolution of sex-biased dispersal

Inbreeding depression, asymmetries in costs or benefits of dispersal, and the mating system have been identified as potential factors underlying the evolution of sex-biased dispersal. We use individual-based simulations to explore how the mating system and demographic stochasticity influence the evolution of sex-specific dispersal in a metapopulation with females competing over breeding sites, and males over mating opportunities. Comparison of simulation results for random mating with those for a harem system (locally, a single male sires all offspring) reveal that even extreme variance in local male reproductive success (extreme male competition) does not induce male-biased dispersal. The latter evolves if the between-patch variance in reproductive success is larger for males than females. This can emerge due to demographic stochasticity if the habitat patches are small. More generally, members of a group of individuals experiencing higher spatio-temporal variance in fitness expectations may evolve to disperse with greater probability than others.

The sex lives of parasites: Investigating the mating system and mechanisms of sexual selection of the human pathogen Schistosoma mansoni

The mating systems of internal parasites are inherently difficult to investigate although they have important implications for the evolutionary biology of the species, disease epidemiology, and are important considerations for control measures. Using parentage analyses, three topics concerning the mating biology of Schistosoma mansoni were investigated: the number of mates per adult male and female, variance in reproductive success among individuals, and the potential role for sexual selection on male body size and also mate choice for genetically dissimilar individuals. Results indicated that schistosomes were mostly monogamous, and evidence of only one mate change occurred over a period of 5–6weeks. One male was polygynous and contained two females in its gynecophoral canal although offspring were only detected for one of the females. Even though they were primarily monogamous and the sex ratio near even, reproductive success was highly variable, indicating a potential role for sexual selection. Male body size was positively related to reproductive success, consistent with sexual selection via male–male competition and female choice for large males. However, relatedness of pairs was not associated with their reproductive success. Finally, genetically identical individuals differed significantly in their reproductive output and identical males in their body size, indicating important partner and environmental effects on these traits.

Extra‐pair paternity in the socially monogamous mountain bluebird Sialia currucoides and its effect on the potential for sexual selection

Sexual selection theory posits that ornamental traits can evolve if they provide individuals with an advantage in securing multiple mates. That male ornamentation occurs in many bird species in which males pair with a single female is therefore puzzling. It has been proposed that extra‐pair mating can substantially increase the variance in reproductive success among males in monogamous species, thus increasing the potential for sexual selection. We documented the frequency of extra‐pair paternity and examined its effect on variation in male reproductive success in the mountain bluebird Sialia currucoides, a socially monogamous songbird in which males possess brilliant plumage ornamentation. Extra‐pair paternity was common in our Wyoming study population, with 72% of broods containing at least one extra‐pair offspring. The standardized variance in actual male reproductive success (i.e., the total number of within‐pair and extra‐pair offspring sired) was more than seven times higher than the variation in apparent success (i.e., success assuming that no extra‐pair mating occurred). Success at siring within‐pair and extra‐pair offspring both contributed to the variation in overall male reproductive success. Within‐pair success, however, did not predict a male's level of extra‐pair success, suggesting that males do not sacrifice within‐pair paternity to gain extra‐pair paternity. Calculation of the sexual selection (Bateman) gradient showed that males sire approximately two additional offspring for each extra‐pair mate that we identified. Thus, in this sexually dichromatic species, extra‐pair mating increases the variance in male reproductive success and provides the potential for sexual selection to act.

Founders, drift, and infidelity: the relationship between Y chromosome diversity and patrilineal surnames.

Most heritable surnames, like Y chromosomes, are passed from father to son. These unique cultural markers of coancestry might therefore have a genetic correlate in shared Y chromosome types among men sharing surnames, although the link could be affected by mutation, multiple foundation for names, nonpaternity, and genetic drift. Here, we demonstrate through an analysis of 1,678 Y-chromosomal haplotypes within 40 British surnames a remarkably high degree of coancestry that generally increases as surnames become rarer. On average, the proportion of haplotypes lying within descent clusters is 62% but ranges from 0% to 87%. The shallow time depth of many descent clusters within names, the lack of a detectable effect of surname derivation on diversity, and simulations of surname descent suggest that genetic drift through variation in reproductive success is important in structuring haplotype diversity. Modern patterns therefore provide little reliable information about the original founders of surnames some 700 years ago. A comparative analysis of published data on Y diversity within Irish surnames demonstrates a relative lack of surname frequency dependence of coancestry, a difference probably mediated through distinct Irish and British demographic histories including even more marked genetic drift in Ireland.

Male reproductive success in plateau pikas ( Ochotona curzoniae): A microsatellite analysis

In many social species, competition and cooperation between group members may lead to a large variance in reproductive success among individuals, especially for adult male. From April to August 2002, we studied the adult male reproductive success of plateau pikas in Haibei Alpine Meadow Ecosystem Research Station, Chinese Academy of Science, using microsatellite analysis of paternity, mark-recapture and behavioral observations. Our result indicated that the reproductive success of adult males had a large variance. Its average was 6 individuals and range was from 0 to 13 individuals. One-third of males sired 63.22% offspring. There was a hierarchy system in polygynandrous families. Although the reproductive success of dominant males was higher than that of subordinate males, subordinate males still play an important role in the reproductive success of the population.

An anomaly against a current paradigm – extremely low rates of individual fecundity variability of the Gordian worm (Nematomorpha: Gordiida)

Extreme variation in reproductive success (VRS) has been reported as a common feature of populations. Few individuals producing most of the offspring for the next generation has potential consequences for the population dynamics, genetics, and evolution of a group of organisms. High VRS has been described as a normal feature of helminth populations, although studies have focused largely on parasites of vertebrate hosts. Paragordius varius, a parasite of crickets, was used as a model system to study VRS. In this life cycle, worms absorb and store resources for reproduction from their hosts before being released into water. Egg output varied significantly with worm length, indicating that female length is an excellent predictor of fecundity. Analyses using the Lorenz curve and Gini coefficient suggest that there were no marked fecundity differences. This result was supported by data collected from a natural gordiid population, Gordius difficilis, suggesting that within gordiid populations the offspring of the next generation are contributed nearly equally by females. In addition, male body length appeared to be limited by intensity, whereas females showed no length limitation by crowding. These results contrast previous studies of parasites.