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Data from: Hidden genetic variance contributes to increase the short-term adaptive potential of selfing populations

Standing genetic variation is considered a major contributor to the adaptive potential of species. The low heritable genetic variation observed in self-fertilising populations has led to the hypothesis that species with this mating system would be less likely to adapt. However, a non-negligible amount of cryptic genetic variation for polygenic traits, accumulated through negative linkage disequilibrium, could prove to be an important source of standing variation in self-fertilising species. To test this hypothesis we simulated populations under stabilizing selection subjected to an environmental change. We demonstrate that, when the mutation rate is high (but realistic), selfing populations are better able to store genetic variance than outcrossing populations through genetic associations, notably due to the reduced effective recombination rate associated with predominant selfing. Following an environmental shift, this diversity can be partially remobilized, which increases the additive variance and adaptive potential of predominantly (but not completely) selfing populations. In such conditions, despite initially lower observed genetic variance, selfing populations adapt as readily as outcrossing ones within a few generations. For low mutation rates, purifying selection impedes the storage of diversity through genetic associations, in which case, as previously predicted, the lower genetic variance of selfing populations results in lower adaptability compared to their outcrossing counterparts. The population size and the mutation rate are the main parameters to consider, as they are the best predictors of the amount of stored diversity in selfing populations. Our results and their impact on our knowledge of adaptation under high selfing rates are discussed.

opencc-zeroAug 2020View details →

Variance in lifetime reproductive success of male polar bears

Despite the important role that population density plays in ecological and evolutionary processes, studies of solitary species that occur at low densities remain scarce. In the context of mating systems, density is expected to influence the ability of males to find and monopolize mates, in turn influencing variance in lifetime mating/reproductive success and the opportunity for selection. Herein we investigate variance in male lifetime mating success, lifetime reproductive success, and the mating system of a sexually dimorphic carnivore that occurs at low densities, the polar bear (Ursus maritimus). Across 17 cohorts, born from 1975 to 1991, male lifetime mating success ranged from 0 to10 mates and lifetime reproductive success from 0 to 14 cubs; 40% of known-age males were not known to have reproduced. The opportunity for sexual selection (Is = 1.66, range = 0.60-4.99) and selection (I = 1.76, range: 0.65-4.89) were low compared to species with similar levels of sexual size dimorphism. Skew in male lifetime reproductive success was also low but significant for most cohorts indicating non-random reproductive success. Age-specific reproductive success was biased toward males from 11-17 years of age, with variation in fecundity (54%) but not longevity (10%) playing an important role in male reproduction. Our results support a growing body of evidence that suggests that male-biased size dimorphism and polygynous mating systems need not be associated with high variance in male mating and/or reproductive success.

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Data from Koch et al. 2020 Evolution: Genetic variance in fitness and its cross-sex covariance predict adaptation during experimental evolution.

Abstract In presence of rapid environmental changes, it is of particular importance to assess the adaptive potential of populations, which is mostly determined by the additive genetic variation (VA) in fitness. In this study we used Tribolium castaneum (red flour beetles) to investigate its adaptive potential in three new environmental conditions (Dry, Hot, Hot-Dry). We tested for potential constraints that might limit adaptation, including negative genetic covariance between female and male fitness.  Based on VA estimates for fitness, we expected the highest relative fitness increase in the most stressful condition Hot-Dry and similar increases in single stress conditions Dry and Hot. High adaptive potential in females in Hot was reduced by a negative covariance with male fitness. We tested adaptation to the three conditions after 20 generations of experimental evolution and found that observed adaptation mainly matched our predictions. Given that body size is commonly used as a proxy for fitness, we also tested how this trait and its genetic variance (including non-additive genetic variance) were impacted by environmental stress. In both traits, variances were sex and condition dependent, but they differed in their variance composition, cross-sex and cross-environment genetic covariances, as well as in the environmental impact on VA.   Method Strain and environmental conditions We used the Tribolium castaneum Cro1 strain collected from a wild population in 2010 and adapted to lab standard conditions (33°C, 70% relative humidity) for more than 20 generations. Beetles were kept in 24h darkness on organic wheat flour mixed with 10% organic baker's yeast. We sterilized flour and yeast by heating them for 12h at 80°C before use. To test for adaptation to new environmental conditions we used replicate lines and exposed them to three treatments and Control conditions. The conditions in the treatments were: Dry: 33°C and 30% relative humidity; Hot: 37°C and 70% r. h.; Hot-Dry: 37°C and 30% r. h.   Crossing, fitness assay and measurement of body size In order to be able to estimate genetic variances, we applied a split-brood paternal half-sib breeding design.  We produced 147 half-sib families by mating virgin males to three virgin females. Half- as well as full-sib families were split across all conditions. Male and female offspring (four females and two males per full-sib family and condition) were separated at the pupal stage and transferred to 10 mL tubes with 1 g of medium and remained there until they were used for the fitness assay eight weeks later. To estimate fitness, we mated each virgin male with two unrelated virgin females from the same condition in 15mL tube with 1g medium. The male was removed after 24h and females transferred into two separate tubes. Females were removed from the tubes after one week of egg laying, and 9g medium was added to provide food for the developing offspring. After five weeks the number of adult offspring was counted. While we conducted the matings for the fitness assay, we followed a specific crossing design and always crossed two pairs of full-sib families. Individuals resulting from these crosses (the F2 genaration) were double first cousins. Body size was measured in the F2, i.e. in the offspring of beetles that were used for the fitness assay. To estimate body size, we used the centroid size of the abdominal segment IV as proxy for total size since it can be measured more accurately than dry weight in very small insects and shows a high correlation with body mass.   Experimental evolution We used ten replicate lines per condition originating from the same ancestral population (Cro1) and let them adapt for 20 generations. Each new generation was set up by randomly selecting 120 pupae and placing them into a new vial with 70g medium. One selection line in Dry became extinct. Adult beetles of generation 20 from all selection lines were transferred to control conditions, in which they stayed for one week to mate and lay eggs. After removal of the adults, we waited until their offspring had reached the pupal stage and separated males and females. These individuals (generation 21) developed completely in control conditions. When they had reached the adult stage, each virgin male was mated with a virgin female of the same selection line and their offspring was transferred to all four conditions in the egg stage, resulting in full-sib families split across all conditions . As soon as these offspring (generation 22) had reached the pupal stage, males and females were separated. To compare fitness of different selection lines and test for adaptation, a virgin male and a virgin female of the same selection line in the same condition, but from different families were mated and the number of adult offspring produced within four days of mating and egg laying was used as a fitness estimate.     Files This data publication contains the following files: Fitness_data.txt: Fitness data (offspring number) of the first generation in new environmental conditions. ANIMAL: ID of female individuals; male: ID of mating partner, father of the offspring; fitness: number of adult offspring per female produced within one week of egg laying; condition: condition under which the individuals grew up, mated, reproduced (CTL: control, D: dry, H: hot, HD: hot-dry, see Methods for details); MOTHER: mother of the female; Batch: samples where fitness assay was started on the same day   Size_data.txt: Centroid size of abdominal segment IV. ANIMAL: ID of measured individuals; condition: condition under which the individuals grew up (CT: control, D: dry, H: hot, HD: hot-dry, see Methods for details); sex: sex of individuals (f: female, m: male); AS: centroid size [Pixel] of abdominal segment IV; Mother: ID of their mother; batch: Batches represent individuals that grew up at the same time, and thus accounts for variations in the medium or lab temperature   Pedigree.txt: Pedigree of measured individuals. ANIMAL: individual ID ; MOTHER: mother ID; FATHER: father ID   Transplant_data.txt: Offspring number of different selection lines under different conditions after 20 generations of experimental evolution. Line: Selection-line ID; Selection: condition, in which the selection-line spent 20 generations (CT: Control; D: Dry; Hot; HD: Hot-Dry); Fam: each selection line consisted of several families; Treatment: condition, in which offspring number was measured; offspring: number of adult offspring that a female produced within four days of egg-laying

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opencc-zeroJan 2021View details →

Data from: Short-term effects of controlled mating and selection on the genetic variance of honeybee populations

Directional selection in a population yields reduced genetic variance due to the Bulmer effect. While this effect has been thoroughly investigated in mammals, it is poorly studied in social insects with biological peculiarities such as haplo-diploidy or the collective expression of traits. In addition to natural adaptation to climate change, parasites, and pesticides, honeybees increasingly experience artificial selection pressure through modern breeding programs. Besides selection, many honeybee breeding schemes introduce controlled mating. We investigated which individual effects selection and controlled mating have on genetic variance. We derived formulas to describe short-term changes of genetic variance in honeybee populations and conducted computer simulations to confirm them. Thereby, we found that the changes in genetic variance depend on whether variance is measured between queens (inheritance criterion), worker groups (selection criterion) or both (performance criterion). All three criteria showed reduced genetic variance under selection. In the selection and performance criteria, our formulas and simulations showed an increased genetic variance through controlled mating. This newly described effect counterbalanced and occasionally outweighed the Bulmer effect. It could not be observed in the inheritance criterion. A good understanding of the different notions of genetic variance in honeybees therefore appears crucial to interpret population parameters correctly.

Data from: Additive genetic variance and developmental plasticity in growth trajectories in a wild cooperative mammal

Individual variation in growth is high in cooperative breeders and may reflect plastic divergence in developmental trajectories leading to breeding vs. helping phenotypes. However, the relative importance of additive genetic variance and developmental plasticity in shaping growth trajectories is largely unknown in cooperative vertebrates. This study exploits weekly sequences of body mass from birth to adulthood to investigate sources of variance in, and covariance between, early and later growth in wild meerkats (Suricata suricatta), a cooperative mongoose. Our results indicate that (i) the correlation between early growth (prior to nutritional independence) and adult mass is positive but weak, and there are frequent changes (compensatory growth) in post-independence growth trajectories; (ii) among parameters describing growth trajectories, those describing growth rate (prior to and at nutritional independence) show undetectable heritability while associated size parameters (mass at nutritional independence and asymptotic mass) are moderately heritable (0.09 ≤ h2 < 0.3); and (iii) additive genetic effects, rather than early environmental effects, mediate the covariance between early growth and adult mass. These results reveal that meerkat growth trajectories remain plastic throughout development, rather than showing early and irreversible divergence, and that the weak effects of early growth on adult mass, an important determinant of breeding success, are partly genetic. In contrast to most cooperative invertebrates, the acquisition of breeding status is often determined after sexual maturity and strongly impacted by chance in many cooperative vertebrates, who may therefore retain the ability to adjust their morphology to environmental changes and social opportunities arising throughout their development, rather than specializing early.

Data from: The genetic structure of Asian corn borer, Ostrinia furnacalis, populations in China: haplotype variance in Northern populations and potential impact on management of resistance to transgenic maize

Asian corn borer, Ostrinia furnacalis (Guenée), is a severe pest that infests cultivated maize in the major production regions of China. Populations show genotype-by-environment variation in voltinism, such that populations with a single generation (univoltine) are fixed in Northern China where growing seasons are short. Low genetic differentiation was found among samples from 33 collection sites across China and one site from North Korea (n = 1,673) using variation at 6 nuclear microsatellite loci (ENA corrected global FST = 0.020; P-value < 0.05). Analysis of molecular variance (AMOVA) indicated that geographic region, number of generations or voltinism accounted for < 0.38% of the total genetic variation at nuclear loci and was corroborated by clustering of co-ancestries among genotypes using the program STRUCTURE. In contrast, a mitochondrial haplotype network identified four distinct clusters, where 70.5% of samples from univoltine populations were within a single group. Univoltine populations were also placed into a unique cluster using Population Graph and Principal Component analyses, which showed significant differentiation with multivoltine populations (ST = 0.400; P-value < 0.01). This study suggests that gene flow among O. furnacalis in China may be high among regions, with the exception of northeastern localities. Haplotype variation may be due to random genetic drift resulting from partial reproductive isolation between univoltine and multivoltine O. furnacalis populations. Such reproductive isolation might impact the potential spread of alleles that confer resistance to transgenic maize in China.

opencc-zeroDec 2015View details →

Data from: Genetic correlations and little genetic variance for reaction norms may limit potential for adaptation to pollution by ionic and nanoparticulate silver in a whitefish (Salmonidae)

For natural populations to adapt to anthropogenic threats, heritable variation must persist in tolerance traits. Silver nanoparticles, the most widely used engineered nanoparticles, are expected to increase in concentrations in freshwaters. Little is known about how these particles affect wild populations, and whether genetic variation persists in tolerance to permit rapid evolutionary responses. We sampled wild adult whitefish and crossed them in vitro full factorially. In total, 2896 singly raised embryos of 48 families were exposed to two concentrations (0.5 μg/L; 100 μg/L) of differently sized silver nanoparticles or ions (silver nitrate). These doses were not lethal; yet higher concentrations prompted embryos to hatch earlier and at a smaller size. The induced hatching did not vary with nanoparticle size and was stronger in the silver nitrate group. Additive genetic variation for hatching time was significant across all treatments, with no apparent environmental dependencies. No genetic variation was found for hatching plasticity. We found some treatment-dependent heritable variation for larval length and yolk volume, and one instance of additive genetic variation for the reaction norm on length at hatching. Our assessment suggests that the effects of silver exposure on additive genetic variation vary according to trait and silver source. While the long-term fitness consequences of low-level silver exposure on whitefish embryos must be further investigated to determine whether it is, in fact, detrimental, our results suggest that the evolutionary potential for adaptation to these types of pollutants may be low.

Data from: Genotypic diversity and trait variance interact to affect marsh plant performance

1. Intraspecific diversity can have important effects on population, community and ecosystem processes, yet we have little understanding of the relative importance of genetic- vs. trait-based measures of intraspecific diversity. 2. I conducted a manipulative field experiment of plant (Spartina alterniflora) genotypic diversity and trait diversity to examine their independent and interactive effects on plant performance and community structure. I focused on variation within and among genotypes in plant stem height, a trait that varies substantially across environmental gradients and can be an important predictor of plant competition intensity. 3. Trait and genotypic diversity interactively affected multiple metrics of plant performance. Both stem density and spatial spread increased with genotypic diversity in the low trait diversity combinations, yet there were negligible to weak negative effects in the high trait diversity treatments. S. alterniflora percent cover also varied with genotypic and trait diversity, but not in a clear linear pattern. 4. There were no effects of trait or genotypic diversity on associated macrofauna above-ground, yet they interactively affected below-ground measures. Infaunal abundance and sediment oxygen availability mirrored the idiosyncratic response of plant percent cover. 5. Despite the interactive effects of genotypic and trait diversity, high trait diversity consistently increased plant performance in genotypic monoculture. 6. Synthesis: The effects of intraspecific plant trait diversity on a range of plant and community responses in this study reinforce the premise that functional differences underlie ecological effects of genetic diversity and suggest that readily measured trait variance may serve as a valuable predictor of plant performance.

opencc-zeroDec 2011View details →

Data from: Oceanographic drivers of offspring abundance may increase or decrease variance in reproductive success 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.

opencc-zeroDec 2012View details →

Data from: Quantitative genetic divergence and standing genetic (co)variance in thermal reaction norms along latitude

Although the potential to adapt to warmer climate is constrained by genetic trade-offs, our understanding of how selection and mutation shape genetic (co)variances in thermal reaction norms is poor. Using 71 isofemale lines of the fly Sepsis punctum, originating from northern, central and southern European climates, we tested for divergence in juvenile development rate across latitude at five experimental temperatures. To investigate effects of evolutionary history in different climates on standing genetic variation in reaction norms, we further compared genetic (co)variances between regions. Flies were reared on either high or low food resources to explore the role of energy-acquisition in determining genetic trade-offs between different temperatures. Although the latter had only weak effects on the strength and sign of genetic correlations, genetic architecture differed significantly between climatic regions, implying that evolution of reaction norms proceeds via different trajectories at high versus low latitude in this system. Accordingly, regional genetic architecture was correlated to region-specific differentiation. Moreover, hot development temperatures were associated with low genetic variance and stronger genetic correlations compared to cooler temperatures. We discuss the evolutionary potential of thermal reaction norms in light of their underlying genetic architectures, evolutionary histories and the materialization of trade-offs in natural environments.

Data from: Genetic basis of between-individual and within-individual variance of docility

Between-individual variation in phenotypes within a population is the basis of evolution. However, evolutionary and behavioural ecologists have mainly focused on estimating between-individual variance in mean trait and neglected variation in within-individual variance, or predictability of a trait. In fact, an important assumption of mixed-effects models used to estimate between-individual variance in mean traits is that within-individual residual variance (predictability) is identical across individuals. Individual heterogeneity in the predictability of behaviours is a potentially important effect but rarely estimated and accounted for. We used 11,389 measures of docility behaviour from 1,576 yellow-bellied marmots (Marmota flaviventris) to estimate between-individual variation in both mean docility and its predictability. We then implemented a double hierarchical animal model to decompose the variances of both mean trait and predictability into their environmental and genetic components. We found that individuals differed both in their docility and in their predictability of docility with a negative phenotypic covariance. We also found significant genetic variance for both mean docility and its predictability but no genetic covariance between the two. This analysis is one of the first to estimate the genetic basis of both mean trait and within-individual variance in a wild population. Our results indicate that equal within-individual variance should not be assumed. We demonstrate the evolutionary importance of the variation in the predictability of docility, and illustrate potential bias in models ignoring variation in predictability. We conclude that the variability in the predictability of a trait should not be ignored, and present a coherent approach for its quantification.

Data from: Metapopulation patterns of additive and nonadditive genetic variance in the sea bass (Dicentrarchus labrax)

Describing and explaining the geographic within-species variation in phenotypes ("phenogeography") in the sea over a species distribution range is central to our understanding of a variety of eco-evolutionary topics. However, phenogeographic studies that have a large potential to investigate adaptive variation are overcome by phylogeographic studies, still mainly focusing on neutral markers. How genotypic and phenotypic data could covary over large geographic scales remains poorly understood in marine species. We crossed 75 noninbred sires (five origins) and 26 dams (two origins; each side of a hybrid zone) in a factorial diallel cross in order to investigate geographic variation for early survival and sex ratio in the metapopulation of the European sea bass (Dicentrarchus labrax), a highly prized marine fish species. Full-sib families (N = 1,950) were produced and reared in a common environment. Parentage assignment of 7,200 individuals was performed with seven microsatellite markers. Generalized linear models showed significant additive effects for both traits and pleiotropy between traits. A significant nonadditive genetic effect was detected. Different expression of traits and distinct relative performances were found for reciprocal crosses involving populations located on each side of the main hybrid zone located at the Almeria-Oran front, illustrating asymmetric reproductive isolation. The poor fitness performance observed for the Western Mediterranean population of sea bass is discussed as it represents the main source of seed hatchery production, but also because it potentially illustrates nonadaptive introgression and maladaptation.

opencc-zeroDec 2012View details →

Data from: From fine-scale foraging to home ranges: a semi-variance approach to identifying movement modes across spatiotemporal scales

Understanding animal movement is a key challenge in ecology and conservation biology. Relocation data often represent a complex mixture of different movement behaviors, and reliably decomposing this mix into its component parts is an unresolved problem in movement ecology. Traditional approaches, such as composite random walk models, require that the timescales characterizing the movement are all similar to the usually arbitrary data-sampling rate. Movement behaviors such as long-distance searching and fine-scale foraging, however, are often intermixed but operate on vastly different spatial and temporal scales. An approach that integrates the full sweep of movement behaviors across scales is currently lacking. Here we show how the semivariance function (SVF) of a stochastic movement process can both identify multiple movement modes and solve the sampling rate problem. We express a broad range of continuous-space, continuous-time stochastic movement models in terms of their SVFs, connect them to relocation data via variogram regression, and compare them using standard model selection techniques. We illustrate our approach using Mongolian gazelle relocation data and show that gazelle movement is characterized by ballistic foraging movements on a 6-h timescale, fast diffusive searching with a 10-week timescale, and asymptotic diffusion over longer timescales.

Data from: Evolution of genetic variance during adaptive radiation

Genetic correlations between traits can concentrate genetic variance into fewer phenotypic dimensions that can bias evolutionary trajectories along the axis of greatest genetic variance and away from optimal phenotypes, constraining the rate of evolution. If genetic correlations limit adaptation, rapid adaptive divergence between multiple contrasting environments may be difficult. However, if natural selection increases the frequency of rare alleles after colonisation of new environments, an increase in genetic variance in the direction of selection can accelerate adaptive divergence. Here, we explored adaptive divergence of an Australian native wildflower by examining the alignment between divergence in phenotype mean and divergence in genetic variance among four contrasting ecotypes. We found divergence in mean multivariate phenotype along two major axes represented by different combinations of plant architecture and leaf traits. Ecotypes also showed divergence in the level of genetic variance in individual traits, and the multivariate distribution of genetic variance among traits. Divergence in multivariate phenotypic mean aligned with divergence in genetic variance, with much of the divergence in phenotype among ecotypes associated with changes in trait combinations containing substantial levels of genetic variance. Overall, our results suggest that natural selection can alter the distribution of genetic variance underlying phenotypic traits, increasing the amount of genetic variance in the direction of natural selection and potentially facilitating rapid adaptive divergence during an adaptive radiation.

Data from: Complex sources of variance in female dominance rank in a nepotistic society

Many mammalian societies are structured by dominance hierarchies, and an individual's position within this hierarchy can influence reproduction, behaviour, physiology and health. In nepotistic hierarchies, which are common in cercopithecine primates and also seen in spotted hyaenas, Crocuta crocuta, adult daughters are expected to rank immediately below their mother, and in reverse age order (a phenomenon known as 'youngest ascendancy'). This pattern is well described, but few studies have systematically examined the frequency or causes of departures from the expected pattern. Using a longitudinal data set from a natural population of yellow baboons, Papio cynocephalus, we measured the influence of maternal kin, paternal kin and group size on female rank positions at two life history milestones, menarche and first live birth. At menarche, most females (73%) ranked adjacent to their family members (i.e. the female held an ordinal rank in consecutive order with other members of her maternal family); however, only 33% of females showed youngest ascendancy within their matriline at menarche. By the time they experienced their first live birth, many females had improved their dominance rank: 78% ranked adjacent to their family members and 49% showed youngest ascendancy within their matriline. The presence of mothers and maternal sisters exerted a powerful influence on rank outcomes. However, the presence of fathers, brothers and paternal siblings did not produce a clear effect on female dominance rank in our analyses, perhaps because females in our data set co-resided with variable numbers and types of paternal and male relatives. Our results also raise the possibility that female body size or competitive ability may influence dominance rank, even in this classically nepotistic species. In total, our analyses reveal that the predictors of dominance rank in nepotistic rank systems are much more complex than previously thought.

Data from: Female and male genetic effects on offspring paternity: additive genetic (co)variances in female extra-pair reproduction and male paternity success in song sparrows (Melospiza melodia)

Ongoing evolution of polyandry, and consequent extra-pair reproduction in socially monogamous systems, is hypothesised to be facilitated by indirect selection stemming from cross-sex genetic covariances with components of male fitness. Specifically, polyandry is hypothesised to create positive genetic covariance with male paternity success due to inevitable assortative reproduction, driving ongoing coevolution. However, it remains unclear whether such covariances could or do emerge within complex polyandrous systems. First, we illustrate that genetic covariances between female extra-pair reproduction and male within-pair paternity success might be constrained in socially monogamous systems where female and male additive genetic effects can have opposing impacts on the paternity of jointly reared offspring. Second, we demonstrate non-zero additive genetic variance in female liability for extra-pair reproduction and male liability for within-pair paternity success, modelled as direct and associative genetic effects on offspring paternity respectively, in free-living song sparrows (Melospiza melodia). The posterior mean additive genetic covariance between these liabilities was slightly positive, but the credible interval was wide and overlapped zero. Therefore, while substantial total additive genetic variance exists, the hypothesis that ongoing evolution of female extra-pair reproduction is facilitated by genetic covariance with male within-pair paternity success cannot yet be definitively supported or rejected either conceptually or empirically.

Data from: Collective dispersal leads to variance in fitness and maintains offspring size variation within marine populations

Variance in fitness is well known to influence the outcome of evolution but is rarely considered in the theory of marine reproductive strategies. In coastal environments, turbulent mesoscale eddies can collect larvae into 'packets' resulting in collective dispersal. Larvae in packets return to the coast or are lost offshore in groups, producing variance in fitness. Using a Markov process to calculate fixation probabilities for competing phenotypes, we examine the evolution of offspring size and spawning duration in species with benthic adults and pelagic offspring. The offspring size that provides mothers with the highest mean fitness also generates the greatest variance in fitness, but pairwise invasion plots show that bet-hedging strategies are not evolutionarily stable: maximizing expected fitness correctly predicts the unique evolutionarily stable strategy. Nonetheless, fixation can take a long time. We find that selection to increase spawning duration as a risk-avoidance strategy to reduce the negative impacts of stochastic recruitment success can allow multiple offspring sizes to coexist in a population for extended periods. This has two important consequences for offspring size: 1) coexistence occurs over a broader range of sizes and is longer when spawning duration is longer, because longer spawning durations reduce variation in fitness and increase the time to fixation, and 2) longer spawning durations can compensate for having a non-optimal size and even allow less optimal sizes to reach fixation. Collective dispersal and longer spawning durations could effectively maintain offspring size variation even in the absence of good and bad years or locations. Empirical comparisons of offspring size would, therefore, not always reflect environment-specific differences in the optimal size.