Local Adaptation and Genetics of Acid-Stress Tolerance in the Moor Frog, Rana arvalis

As potential to adapt to environmental stress can be essential for population persistence, knowledge on the genetic architecture of local adaptation is important for conservation genetics. We investigated the relative importance of additive genetic, dominance and maternal effects contributions to acid stress tolerance in two moor frog (Rana arvalis) populations originating from low and neutral pH habitats. Experiments with crosses obtained from artificial matings revealed that embryos from the acid origin population were more tolerant to low pH than embryos from the neutral origin population in embryonic survival rates, but not in terms of developmental stability, developmental and growth rates. Strong maternal effect and small additive genetic contributions to variation were detected in all traits in both populations. In general, dominance contributions to variance in different traits were of similar magnitude to the additive genetic effects, but dominance effects outweighed the additive genetic and maternal effects contributions to early growth in both populations. Furthermore, the expression of additive genetic variance was independent of pH treatment, suggesting little additive genetic variation in acid stress tolerance. The results suggest that although local genetic adaptation to acid stress has taken place, the current variation in acid stress tolerance in acidified populations may owe largely to non-genetic effects. However, low but significant heritabilities (h ² 0.07–0.22) in all traits – including viability itself – under a wide range of pH conditions suggests that environmental stress created by low pH is unlikely to lower moor frog populations' ability to respond to selection in the traits studied. Nevertheless, acid conditions could lower populations' ability to respond to selection in the long run through reduction in effective population size.     

Parental effects on embryonic viability and growth in Arctic charr Salvelinus alpinus at two incubation temperatures

The parental influences on three progeny traits (survival to eyed‐embryo stage, post‐hatching body length and yolk‐sac volume) of Arctic charr Salvelinus alpinus were studied under two thermal conditions (2 and 7° C) using a factorial mating design. The higher temperature resulted in elevated mortality rates and less advanced development at hatching. Survival was mostly attributable to maternal effects at both temperatures, but the variation among families was dependent on egg size only at the low temperature. No additive genetic variation (or pure sire effect) could be observed, whereas the non‐additive genetic effects (parental combination) contributed to offspring viability at 2° C. In contrast, any observable genetic variance in survival was lost at 7° C, most likely due to the increased environmental variance. Irrespective of temperature, dam and sire–dam interaction contributed significantly to the phenotypic variation in both larval length and yolk size. A significant proportion of the variation in larval length was also due to the sire effect at 2° C. Maternal effects were mediated partly through egg size, but as a whole, they decreased in importance at the high temperature, enabling a concomitant increase in non‐additive genetic effects. For larval length, however, the additive component, like maternal effects, decreased at 7° C. The present results suggest that an exposure to thermal stress during incubation can modify the genetic architecture of early developmental traits in S. alpinus and presumably constrain their short‐term adaptive potential and evolvability by increasing the amount of environmentally induced variation.     

VARIATION IN SEED CHARACTERS IN NEMOPHILA MENZIESII: EVIDENCE OF A GENETIC BASIS FOR MATERNAL EFFECT

A growing body of evidence indicates that phenotypic selection on juvenile traits of both plants and animals may be considerable. Because juvenile traits are typically subject to maternal effects and often have low heritabilities, adaptive responses to natural selection on these traits may seem unlikely. To determine the potential for evolutionary response to selection on juvenile traits of Nemophila menziesii (Hydrophyllaceae), we conducted two quantitative genetic studies. A reciprocal factorial cross, involving 16 parents and 1960 progeny, demonstrated a significant maternal component of variance in seed mass and additive genetic component of variance in germination time. This experiment also suggested that interaction between parents, though small, provides highly significant contributions to the variance of both traits. Such a parental interaction could arise by diverse mechanisms, including dependence of nuclear gene expression on cytoplasmic genotype, but the design of this experiment could not distinguish this from other possible causes, such as effects on progeny phenotype of interaction between the environmental conditions of both parents. The second experiment, spanning three generations with over 11,000 observations, was designed for investigation of the additive genetic variance in maternal effect, assessment of paternal effects, as well as further partitioning of the parental interaction identified in the reciprocal factorial experiment. It yielded no consistent evidence of paternal effects on seed mass, nor of parental interactions. Our inference of such interaction effects from the first experiment was evidently an artifact of failing to account for the substantial variance among fruits within crosses. The maternal effect was found to have a large additive genetic component, accounting for at least 20% of the variation in individual seed mass. This result suggests that there is appreciable potential for response to selection on seed mass through evolution of the maternal effect. We discuss aspects that may nevertheless limit response to individual selection on seed mass, including trade-offs between the size of individual seeds and germination time and between the number of seeds a maternal plant can mature and their mean size.     

Maternal genetic effects set the potential for evolution in a free-living vertebrate population

Heritable maternal effects have important consequences for the evolutionary dynamics of phenotypic traits under selection, but have only rarely been tested for or quantified in evolutionary studies. Here we estimate maternal effects on early-life traits in a feral population of Soay sheep (Ovis aries) from St Kilda, Scotland. We then partition the maternal effects into genetic and environmental components to obtain the first direct estimates of maternal genetic effects in a free-living population, and furthermore test for covariance between direct and maternal genetic effects. Using an animal model approach, direct heritabilities (h2) were low but maternal genetic effects (m2) represented a relatively large proportion of the total phenotypic variance for each trait (birth weight m2=0.119, birth date m2=0.197, natal litter size m2=0.211). A negative correlation between direct and maternal genetic effects was estimated for each trait, but was only statistically significant for natal litter size (ram= -0.714). Total heritabilities (incorporating variance from heritable maternal effects and the direct-maternal genetic covariance) were significant for birth weight and birth date but not for natal litter size. Inadequately specified models greatly overestimated additive genetic variance and hence direct h2 (by a factor of up to 6.45 in the case of birth date). We conclude that failure to model heritable maternal variance can result in over- or under-estimation of the potential for traits to respond to selection, and advocate an increased effort to explicitly measure maternal genetic effects in evolutionary studies.     

A genetic analysis of the mandible and maxilla in the rat

A quantitative genetic analysis of eight osteometric traits from the oral region together with femur length in the rat are described. The relevant questions being examined are (1) the heritability of each trait; (2) the relative contribution of genetic, maternal environmental, and residual environmental effects to the correlation between the oral traits, and (3) genetic and nongenetic components of the correlation between the oral traits and overall body size. Some quantitative genetic aspects of covarying traits is briefly reviewed with special emphasis on allometric variation. Phenotypic regression coefficients from log-transformed data (= allometry coefficients) of oral traits onto femur length are partitioned into components due to genetic, maternal environmental, residual environmental, and total environmental causes. All phenotypic regression coefficients and all but one based on an environmentally determined covariance component are significantly different from zero, suggesting a substantial body size effect in the oral region resulting from nonheritable causes. However, three genetic coefficients from regressions of oral traits to femur length are not different from zero, indicating a genetic correlation with body size in five traits but not in three others. A principal components analysis was carried out on the phenotypic, genetic, and environmental correlations of the eight oral traits to provide a multivariate depiction of the components of covariation in the maxilla and mandible of the rat. General multivariate effects due to body size together with group or special effects are demonstrated for each component of morphogenesis.     

The quantitative genetic basis of adaptive divergence in the moor frog (Rana arvalis) and its implications for gene flow

Knowledge on the relative contribution of direct genetic, maternal and environmental effects to adaptive divergence is important for understanding the drivers of biological diversification. The moor frog (Rana arvalis) shows adaptive divergence in embryonic and larval fitness traits along an acidification gradient in south-western Sweden. To understand the quantitative genetic basis of this divergence, we performed reciprocal crosses between three divergent population pairs and reared embryos and larvae at acid and neutral pH in the laboratory. Divergence in embryonic acid tolerance (survival) was mainly determined by maternal effects, whereas the relative contributions of maternal, additive and nonadditive genetic effects in larval life-history traits differed between traits, population pairs and rearing environments. These results emphasize the need to investigate the quantitative genetic basis of adaptive divergence in multiple populations and traits, as well as different environments. We discuss the implications of our findings for maintenance of local adaptation in the context of migrant and hybrid fitness.     

Natural selection on the genetical component of variance in body condition in a wild bird population

Although there is substantial evidence that skeletal measures of body size are heritable in wild animal populations, it is frequently assumed that the nonskeletal component of body weight (or ‘condition’) is determined primarily by environmental factors, in particular nutritional state. We tested this assumption by quantifying the genetic and environmental components of variance in fledgling body condition index (=relative body weight) in a natural population of collared flycatchers (Ficedula albicollis), and compared the strength of natural selection on individual breeding values with that on phenotypic values. A mixed model analysis of the components of variance, based on an ‘animal model’ and using 18 years of data on 17 717 nestlings, revealed a significant additive genetic component of variance in body condition, which corresponded to a narrow sense heritability (h²) of 0.30 (SE=0.03). Nongenetic contributions to variation in body condition were large, but there was no evidence of dominance variance nor of contributions from early maternal or common environment effects (pre‐manipulation environment) in condition at fledging. Comparison of pre‐ and post‐selection samples revealed virtually identical h² of body condition index, despite the fact that there was a significant decrease (35%) in the levels of additive genetic variance from fledging to breeding. The similar h² in the two samples occurred because the environmental component of variance was also reduced by selection, suggesting that natural selection was acting on both genotypic and environmental variation. The effects of selection on genetic variance were confirmed by calculation of the selection differentials for both phenotypic values and best linear unbiased predictor (BLUP) estimates of breeding values: there was positive directional selection on condition index both at the phenotypic and the genotypic level. The significant h² of body condition index is consistent with data from human and rodent populations showing significant additive genetic variance in relative body mass and adiposity, but contrasts with the common assumption in ecology that body condition reflects an individual’s nongenetic nutritional state. Furthermore, the substantial reduction in the additive genetic component of variance in body condition index suggests that selection on environmental deviations cannot alone explain the maintenance of additive genetic variation in heritable traits, but that other mechanisms are needed to explain the moderate to high heritabilities of traits under consistent and strong directional selection.     

Genetic variation in a female genital trait evolved by sexual coevolution

Understanding the patterns of genetic variation of traits subject to sexual selection is fundamental for explaining its evolutionary dynamics and potential for sexual coevolution. The signa of female Lepidoptera are sclerotized structures located on the inner surface of the genital receptacle that receives the spermatophore during copulation (the corpus bursae), whose main function is tearing the spermatophore envelope. Comparative data indicate that the evolution of signa has been influenced by sexually antagonistic coevolution with spermatophore envelopes. We looked for additive genetic variation in the size and shape of signa in females of the butterfly Callophrys xami (Lycaenidae) from two localities (BG and FC) in Mexico City. We also looked for genetic variation in female body size and in the size of corpus bursae. There were significant between-population differences in female body size, signa width and three signa shape traits. We found significant extranuclear maternal effects in one component of signa shape in the BG population, and in body weight, signa length and in one uniform component of signa shape in the FC population. Extranuclear maternal contributions could permit the evolution of female adaptations even if these reduce male fitness. We found additive genetic variation in signa length and width only in one population (BG); heritability estimates were high: 0.96 and 0.8, respectively. The existence of additive genetic variation in signa size could be, at least in part, a result of relaxed sexually antagonistic selection pressures due to the low level of polyandry exhibited by this species. Our results imply that there is currently potential for further sexual coevolution in this trait.     

Quantitative genetics of larval life-history traits in Rana temporaria in different environmental conditions

The degree to which genetic variation in a given trait varies among different populations of the same species and across different environments has seldom been quantified in wild vertebrate species. We investigated the expression of genetic variability and maternal effects in three larval life-history traits of the amphibian Rana temporaria. In a factorial laboratory experiment, five widely separated populations (max. 1600 km) were subjected to two different environmental treatments. Animal model analyses revealed that all traits were heritable (h(2) approximately 0.20) in all populations and under most treatment combinations. Although the cross-food treatment genetic correlations were close to unity, heritabilities under a restricted food regime tended to be lower than those under an ad libitum food regime. Likewise, maternal effects (m(2) approximately 0.05) were detected in most traits, and they tended to be most pronounced under restricted food conditions. We detected several cross-temperature genetic and maternal effects correlations that were lower than unity, suggesting that genotype-environment interactions and maternal effect-environment interactions are a significant source of phenotypic variation. The results reinforce the perspective that although the expression of genetic and maternal effects may be relatively homogeneous across different populations of the same species, local variation in environmental conditions can lead to significant variation in phenotypic expression of quantitative traits through genotype-environment and maternal effect-environment interactions.     

Female crickets trade offspring viability for fecundity

A growing number of studies are suggesting that females can improve the viability of their embryos by mating with multiple males. However, the reason why females should have low rates of embryo viability is puzzling. Here we conduct a quantitative genetic study of maternal effects on embryo viability in the field cricket Teleogryllus oceanicus. After controlling for female body size, we find significant additive genetic variance for ovary weight, a measure of fecundity, and egg hatching success, a measure of embryo viability. Moreover, we show a genetic trade-off between these traits that is predicted from life-history theory. High rates of embryo mortality in this highly fecund species might therefore be explained by selection favouring an optimum balance between fecundity and embryo viability that maximizes maternal fitness. Paternal effects on female fecundity and embryo viability are often seen as benefits driving the evolution of polyandrous behaviour. However, we raise the alternative possibility that paternal effects might shift females from their naturally selected optimum, and present some support for the notion that sexual conflict over a female's optimal fecundity and embryo viability might generate antagonistic coevolution between the sexes.     

Geographic variation in acid stress tolerance of the moor frog,Rana arvalis. II. Adaptive maternal effects

Abstract The knowledge about the relative contributions of additive genetic and maternal effects, as well as the proximate determinants of maternal effects variation, on population differentiation remains elusive. Likewise, although embryonic performance is often an important component of fitness, it has been relatively little explored in respect to population differentiation. By conducting reciprocal crosses between an acid and a neutral origin population of moor frogs ( Rana arvalis ), we investigated the relative importance of additive genetic versus maternal effects in local adaptation to acidity in embryonic traits. Furthermore, by performing removal experiments of gelatinous egg capsules (jelly), we evaluated the possibility that differences in the extraembryonic membranes might explain the interpopulation variation in embryonic acid tolerance found in this and earlier studies. Embryos were raised from fertilization to hatching at three different pH levels (pH 4.0, 4.25, and 7.5) in the laboratory, and acid stress tolerance was measured in terms of embryonic survival, growth and development (i.e., size and age at hatching). The results show that the higher acid tolerance of acid population embryos (in terms of survival) was maternally determined, indicating adaptive maternal effects. The jelly removal experiment revealed that adaptation to acidity in embryonic survival may arise through variation related to structure/composition of the egg capsules. There was no evidence for a genetic basis in acid tolerance in sublethal effects, but additive and nonadditive genetic effects were found in embryonic growth and development, independently of treatment. The results indicate a role for maternal effects in local adaptation to acidity in amphibians, and genetically based differences in early life-histories among the populations.     

Evolution of Adaptation and Mate Choice: Parental Relatedness Affects Expression of Phenotypic Variance in a Natural Population

Mating between relatives generally results in reduced offspring viability or quality, suggesting that selection should favor behaviors that minimize inbreeding. However, in natural populations where searching is costly or variation among potential mates is limited, inbreeding is often common and may have important consequences for both offspring fitness and phenotypic variation. In particular, offspring morphological variation often increases with greater parental relatedness, yet the source of this variation, and thus its evolutionary significance, are poorly understood. One proposed explanation is that inbreeding influences a developing organism’s sensitivity to its environment and therefore the increased phenotypic variation observed in inbred progeny is due to greater inputs from environmental and maternal sources. Alternatively, changes in phenotypic variation with inbreeding may be due to additive genetic effects alone when heterozygotes are phenotypically intermediate to homozygotes, or effects of inbreeding depression on condition, which can itself affect sensitivity to environmental variation. Here we examine the effect of parental relatedness (as inferred from neutral genetic markers) on heritable and nonheritable components of developmental variation in a wild bird population in which mate choice is often constrained, thereby leading to inbreeding. We found greater morphological variation and distinct contributions of variance components in offspring from highly related parents: inbred offspring tended to have greater environmental and lesser additive genetic variance compared to outbred progeny. The magnitude of this difference was greatest in late-maturing traits, implicating the accumulation of environmental variation as the underlying mechanism. Further, parental relatedness influenced the effect of an important maternal trait (egg size) on offspring development. These results support the hypothesis that inbreeding leads to greater sensitivity of development to environmental variation and maternal effects, suggesting that the evolutionary response to selection will depend strongly on mate choice patterns and population structure.     

Genetics of Mandible Form in the Mouse

The underlying determination of phenotypic variability and covariability is described for 14 traits that define the morphological size and shape of the mature mouse mandible. Variability is partitioned into components due to direct additive and dominance genetic effects, indirect maternal additive genetic effects, genetic covariance between direct additive and indirect maternal additive effects and common and residual environmental effects. Multivariate analyses of the dimensionality of genetic variability indicate several complex and independent genetic components underlie the morphological form of the mandible. The multidimensional nature of the genetic components suggests a complex picture with regard to the consequences of selection on mandibular form.     

The Effect of an Experimental Bottleneck upon Quantitative Genetic Variation in the Housefly

Effects of a population bottleneck (founder-flush cycle) upon quantitative genetic variation of morphometric traits were examined in replicated experimental lines of the housefly founded with one, four or 16 pairs of flies. Heritability and additive genetic variances for eight morphometric traits generally increased as a result of the bottleneck, but the pattern of increase among bottleneck sizes differed among traits. Principal axes of the additive genetic correlation matrix for the control line yielded two suites of traits, one associated with general body size and another set largely independent of body size. In the former set containing five of the traits, additive genetic variance was greatest in the bottleneck size of four pairs, whereas in the latter set of two traits the largest additive genetic variance occurred in the smallest bottleneck size of one pair. One trait exhibited changes in additive genetic variance intermediate between these two major responses. These results were inconsistent with models of additive effects of alleles within loci or of additive effects among loci. An observed decline in viability measures and body size in the bottleneck lines also indicated that there was nonadditivity of allelic effects for these traits. Several possible nonadditive models were explored that increased additive genetic variance as a result of a bottleneck. These included a model with complete dominance, a model with overdominance and a model incorporating multiplicative epistasis.     

Genetic parameter estimates for pre-weaning performance and reproduction traits in Markhoz goats

Genetic parameters for growth, mortality and reproductive performances of Markhoz goats were estimated from data collected during 1993–2010 at Markhoz goat Performance Testing Station in Sanandaj, Iran. For kid performance traits 3763 records were available for birth weight (BW), 2931 for weaning weight (WW), average daily gain (ADG) and Kleiber ratio (KR) (approximated as ADW/WW^0.75) and 3032 for pre-weaning mortality (PWM). For doe reproductive performance traits there were 2920 records available for litter size at birth (LSB), litter size at weaning (LSW), total litter weight at birth (TLWB) and litter mean weight per kid born (LMWKB), and 2182 for total litter weight at weaned (TLWW) and litter mean weight per kid weaned (LMWKW). Genetic parameters were estimated with univariate and bivariate models using restricted maximum likelihood (REML) procedures. Random effects were explored by fitting additive direct genetic effects, maternal additive genetic effects, maternal permanent environmental effects, the covariance between direct and maternal genetic effects, and common litter effects in different models for pre-weaning traits of kids. Also, in addition to an animal model, sire and threshold models, using a logit link function, were used for analyses of PWM. Models for LSB, LSW, TLWB, TLWW, LMWKB, and LMWKW included direct additive genetic effects, permanent environmental effects due to the animal as well as service sire effects. Estimated direct heritabilities were moderate for pre-weaning traits (0.22 for BW, 0.16 for WW, 0.21 for ADG, and 0.27 for KR and 0.29 for PWM), and low for reproduction traits (0.01 for LSB, 0.01 for LSW, 0.02 for TLWB, 0.03 for TLWW, 0.07 for LMWKB, and 0.06 for LMWKW). The estimates for the maternal additive genetic variance ratios were lower than direct heritability for BW (0.07) and KR (0.04). The estimate for the maternal permanent environmental variance ratios (c²) varied from 0.01 for KR to 0.07 for WW and ADG. The magnitude of common litter variance ratios (l²) was more substantial for BW (0.46) than the PWM (0.19) and KR (0.16). The estimate for the permanent environmental variance due to the animal (c²) ranged from 0.03 for LMWKB to 0.07 for TLWB and LMWKW, whereas service sire effects (s²) ranged from 0.02 to 0.04. The correlation between direct and maternal genetic effects were negative and high for BW (?0.51) and KR (?0.62). The genetic correlations between pre-weaning growth traits were positive and moderate to strong, as were genetic correlations between reproductive traits. Between BW and PWM the correlation was ?0.35. Phenotypic and environmental correlations for all traits were generally lower than genetic correlations.     

Maternal and genetic contributions to geographical variation in Rana temporaria larval life-history traits

The relative importance of genetic, environmental, and maternal effects as determinants of geographical variation in vertebrate life-histories has not often been explored. We examined the role of genetic and maternal effects as determinants of population divergence in survival and three important larval life-history traits (growth rate, age, and size at metamorphosis) using reciprocal crosses between two latitudinally separated populations of the common frog ( Rana temporaria Linnaeus). Genetic effects were important in all three traits as indicated by the significant effect of male origin, but there was also evidence for nonadditive genetic contributions on metamorphic size and growth rate. Likewise, maternal effect contributions to population divergence were large, partially environment dependent, and apparently acting primarily through egg size in two of three traits. These results suggest that both genetic and maternal effects are important determinants of geographical variation in amphibian life-histories, and that much of the differentiation resulting from maternal effects is mediated through variation in egg size. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 61–70.     

Ontogenetic patterns in heritable variation for body size: using random regression models in a wild ungulate population

Body size is an important determinant of fitness in many organisms. While size will typically change over the lifetime of an individual, heritable components of phenotypic variance may also show ontogenetic variation. We estimated genetic (additive and maternal) and environmental covariance structures for a size trait (June weight) measured over the first 5 years of life in a natural population of bighorn sheep Ovis canadensis. We also assessed the utility of random regression models for estimating these structures. Additive genetic variance was found for June weight, with heritability increasing over ontogeny because of declining environmental variance. This pattern, mirrored at the phenotypic level, likely reflects viability selection acting on early size traits. Maternal genetic effects were significant at ages 0 and 1, having important evolutionary implications for early weight, but declined with age being negligible by age 2. Strong positive genetic correlations between age-specific traits suggest that selection on June weight at any age will likely induce positively correlated responses across ontogeny. Random regression modeling yielded similar results to traditional methods. However, by facilitating more efficient data use where phenotypic sampling is incomplete, random regression should allow better estimation of genetic (co)variances for size and growth traits in natural populations.     

A QUANTITATIVE-GENETIC ANALYSIS OF LARVAL LIFE-HISTORY TRAITS IN HYLA CRUCIFER

We used a half-sib design to examine the genetic components of phenotypic variance in several life-history traits in Hyla crucifer. Egg viability, hatchling size, larval growth rate, length of larval period, and size at metamorphosis play critical roles in determining survivorship and are subject to persistent selection. Egg viability varied among families considerably, with most embryo mortality occurring between gastrulation and neurulation. Hatchling size was the only trait in which maternal effects were influential. Dominance genetic variance played the predominant role in determining phenotypic variance in hatchling size, growth rate, and length of larval period, accounting for, respectively, 70, 63, and 47% of the total variance. Size at metamorphosis displayed little dominance genetic variance and, unlike the other traits, displayed a high heritability. All additive genetic correlations between traits were positive. The directions of environmental correlations were the same as the directions of changes that have been induced in previous experimental work. The correlations due to dominance effects described a principal axis that independent ecological studies indicate to be directly correlated with fitness. These results agree with theoretical expectations for traits under consistent directional selection.     

Unravelling the effects of differential maternal allocation and male genetic quality on offspring viability in the dung beetle, Onthophagus sagittarius

Good genes models of mate choice assume heritability of fitness-related traits. However, maternal effects can inflate estimates of trait heritability, and genotype-environment interactions can have significant effects on good genes processes of evolution. Thus, partitioning genetic and maternal/environmental sources of variation in studies of good genes mate choice represents an empirical challenge. In this study, we used the dung beetle Onthophagus sagittarius to examine additive genetic and maternal effects on egg-to-adult offspring viability. We used a half-sib full-sib breeding design and manipulated the maternally provided environment by reducing or increasing the mass of the brood ball within which each offspring developed. We found evidence of differential allocation of investment by females in the brood balls they produced. However, experimental manipulations of maternal allocation to brood balls had only a weak and non-significant influence on the sire effects on offspring viability. Significant additive genetic effects on offspring viability were pervasive across our manipulations of the maternally provided larval environment. This finding indicates that although females do show differential allocation to offspring based on sire phenotype, ‘good genes’ benefits of mate choice are not dependent upon differential maternal allocation.     

Heritability and fitness-related consequences of squid personality traits

Dumpling squid, Euprymna tasmanica, show consistent individual differences in behaviour that can be classified according to indices reflecting shy-bold, activity and reactivity responses. Using crosses of wild-caught single males to multiple females with known behavioural phenotypes, this study estimated patterns of additive genetic and residual variance in these behavioural traits from offspring of squid in two contexts, a threat (antipredator) and feeding (foraging) test. Genetic contributions to behavioural expression were dependent on test context. Behaviours in antipredator contexts had significant heritabilities (h(2) = 0.2-0.8) while behaviours from foraging contexts had lesser additive genetic and greater residual components (h(2) = 0.05-0.08). Personality trait variation in females was not related to her fecundity. Female boldness in foraging situations, which co-varied with body size, explained small but significant variation ( approximately 21%) in brood hatching success, while successful fertilization was determined by positive assortion of mate pairs according to their shy-bold phenotype. These results are discussed in terms of the ecological and evolutionary significance of animal "personality" traits in wild populations of animals.