Current brood size and residual reproductive value predict offspring desertion in the burying beetle Nicrophorus vespilloides

Life-history theory suggests that offspring desertion can be an adaptive reproductive strategy, in which parents forgo the costly care of an unprofitable current brood to save resources for future reproduction. In the burying beetle, Nicrophorus vespilloides, parents commonly abandon their offspring to the care of others, resulting in female-only care, male-only care, brood parasitism, and the care of offspring sired by satellite males. Furthermore, when there is biparental care, males routinely desert the brood before larval development is complete, leaving females behind to tend their young. We attempted to understand these patterns of offspring desertion by using laboratory experiments to compare the fitness costs associated with parental care for each sex and the residual reproductive value of the 2 sexes. We also tested whether current brood size and residual reproductive value together predicted the incidence of brood desertion. We found that males and females each sustained fecundity costs as a consequence of caring for larvae and that these costs were of comparable magnitude. Nevertheless, males had greater residual reproductive value than females and were more likely than females to desert experimental broods. Our results can explain why males desert the brood earlier than females in nature and why female-only care is more common than male-only care. They also suggest that the tipping point from brood parasitism or satellite male behavior to communal breeding (and vice versa) depends on the value of the current brood relative to residual reproductive value.

Heritability of multiple mating by female field crickets, Gryllus integer (Orthoptera: Gryllidae)

The heritability of multiple mating in female Gryllus integer crickets was studied. Two preliminary experiments were conducted to determine when females first mate following the post-imaginal moult and to ascertain whether constant exposure to males affects female mating rate. Female Q. integer first mated at an average age of 3.6 days (S.D. = 2.3, Range = 0-8 days) . Exposing female crickets to courting males 24 hr daily did not significantly alter mating rates from those females in contact with males for only 5 hr per day. A heritability value of 0.690 ± 0.283 was calculated for multiple mating behavior in female Q. integer using a parent-offspring regression approach. Parental females mated between land 30 times (x 9.8, S . D. = 6. 6 ) and offspring matings ranged from 0 to 26 times (x 7 .3, S.D. = 3.4). Multiple mating is probably a sexually selected trait which functions as a mechanism of female choice and increases reproductive success through increased offspring production. Classical theory suggests that traits intimately related with fitness should exhibit negligible heritable variation. However, this study has shown that multiple mating, a trait closely linked with reproductive fitness, exhibits substantial heritability. These results are in concordance with a growing body of empirical evidence suggesting many fitness traits in natural populations demonstrate heritabilities far removed from zero. Various mechanisms which may maintain heritable variation for female multiple mating in wild, outbred Q. integer populations are discussed.

Sexual behavior and factors affecting female reproduction in house and field crickets

The objective of this investigation was to clarify the adaptive significance of female sexual behaviours in the house cricket, Acheta domesticus, and the Texas field cricket, Gryllus integer. Experiments were focussed primarily on: nutritional factors affecting female reproductive success; the ontogeny of female sexual behaviours; female mating frequency and progeny production; and the pattern of sperm competition. Reproduction of singly mated female A. domesticus assigned to 3 nutritional regimes was compared . Females fed a vitamin and protein-enriched mouse chow, cannibalistic females, and starved females produced on the average, 513 , 200 and 68 offspring, respectively. Cannibals probably could not obtain the same amounts of essential nutrients as females fed mouse chow. Reabsorption of oocytes was likely the major factor contributing to the decreased reproduction of starved females. In addition, female !. domesticus fed mouse chow, but allowed constant access to males produced 11 times as many offspring than did females fed corn meal. Females fed corn meal probably could not absorb or synthesize enough dietary lipids, thus resulting in poor ovariole growth. Female !. domesticus first mate at an average adult age of 7 days, closely corresponding to when they first exhibit positive phonotaxis. Females mate repeatedly and often consume the externally attached spermatophore. In ~. domesticus, females allowed constant access to males produced significantly more offspring than did single maters. Similarly, doubly mated G. integer females produced more offspring than did single maters. This difference resulted largely from the failure of many single maters to reproduce. Remating by female crickets partly functions in offsetting the possibility of a failed initial mating. Nymph production increased significantly with the time the spermatophore was attached in singly mated ~. domesticus. Spermatophore consumption by the female was not affected by male guarding behaviour, and the interval between mating and eating of the spermatophore may often be shorter than the time required for maximum insemination. Some degree of sperm depletion in singly mated !. domesticus and G. integer may have occurred. The patterns of daily offspring production of singly and multiplymated females suggests that a factor provided by a male during mating stimulates female oviposition and/or egg production. Female crickets also might acquire nutrition from spermatophore consumption, a benefit that is augmented by female multiple mating. The electrophoretic examination of various allozymes in ~. integer did not permit determination of a pattern of sperm competition. However, the possibility of last male sperm predominance is related to male guarding behaviour.

Costs of Reproduction in Breeding Female Mallards: Predation Risk during Incubation Drives Annual Mortality

The effort expended on reproduction may entail future costs, such as reduced survival or fecundity, and these costs can have an important influence on life-history optimization. For birds with precocial offspring, hypothesized costs of reproduction have typically emphasized nutritional and energetic investments in egg formation and incubation. We measured seasonal survival of 3856 radio-marked female Mallards (Anas platyrhynchos) from arrival on the breeding grounds through brood-rearing or cessation of breeding. There was a 2.5-fold direct increase in mortality risk associated with incubating nests in terrestrial habitats, whereas during brood-rearing when breeding females occupy aquatic habitats, mortality risk reached seasonal lows. Mortality risk also varied with calendar date and was highest during periods when large numbers of Mallards were nesting, suggesting that prey-switching behaviors by common predators may exacerbate risks to adults in all breeding stages. Although prior investments in egg laying and incubation affected mortality risk, most relationships were not consistent with the cost of reproduction hypothesis; birds with extensive prior investments in egg production or incubation typically survived better, suggesting that variation in individual quality drove both relationships. We conclude that for breeding female Mallards, the primary cost of reproduction is a fixed cost associated with placing oneself at risk to predators while incubating nests in terrestrial habitats.

Mammal reproductive strategies driven by offspring mortality-size relationships

Trade-offs have long been a major theme in life-history theory, but they have been hard to document. We introduce a new method that reveals patterns of divergent trade-offs after adjusting for the pervasive variation in rate of resource allocation to offspring as a function of body size and lifestyle. Results suggest that preweaning vulnerability to predation has been the major factor determining how female placental mammals allocate production between a few large and many small offspring within a litter and between a few large litters and many small ones within a reproductive season. Artiodactyls, perissodactyls, cetaceans, and pinnipeds, which give birth in the open on land or in the sea, produce a few large offspring, at infrequent intervals, because this increases their chances of escaping predation. Insectivores, fissiped carnivores, lagomorphs, and rodents, whose offspring are protected in burrows or nests, produce large litters of small newborns. Primates, bats, sloths, and anteaters, which carry their young from birth until weaning, produce litters of one or a few offspring because of the need to transport and care for them.

Effect of maternal protein restriction during pregnancy and postweaning high-fat feeding of diet-induced thermogenesis in adult mouse offspring

Purpose Prenatal undernutrition followed by postweaning feeding of a high-fat diet results in obesity in the adult offspring. In this study, we investigated whether diet-induced thermogenesis is altered as a result of such nutritional mismatch. Methods Female MF-1 mice were fed a normal protein (NP, 18 % casein) or a protein-restricted (PR, 9 % casein) diet throughout pregnancy and lactation. After weaning, male offspring of both groups were fed either a high-fat diet (HF; 45 % kcal fat) or standard chow (C, 7 % kcal fat) to generate the NP/C, NP/HF, PR/C and PR/HF adult offspring groups (n = 7–11 per group). Results PR/C and NP/C offspring have similar body weights at 30 weeks of age. Postweaning HF feeding resulted in significantly heavier NP/HF offspring (P < 0.01), but not in PR/HF offspring, compared with their chow-fed counterparts. However, the PR/HF offspring exhibited greater adiposity (P < 0.01) v the NP/HF group. The NP/HF offspring had increased energy expenditure and increased mRNA expression of uncoupling protein-1 and β-3 adrenergic receptor in the interscapular brown adipose tissue (iBAT) compared with the NP/C mice (both at P < 0.01). No such differences in energy expenditure and iBAT gene expression were observed between the PR/HF and PR/C offspring. Conclusions These data suggest that a mismatch between maternal diet during pregnancy and lactation, and the postweaning diet of the offspring, can attenuate diet-induced thermogenesis in the iBAT, resulting in the development of obesity in adulthood.

Female-specific intergenerational transmission patterns of the human corticolimbic circuitry

Parents have large genetic and environmental influences on offspring’s cognition, behavior, and brain. These intergenerational effects are observed in mood disorders, with particularly robust association in depression between mothers and daughters. No studies have thus far examined the neural bases of these intergenerational effects in humans. Corticolimbic circuitry is known to be highly relevant in a wide range of processes including mood regulation and depression. These findings suggest that corticolimbic circuitry may also show matrilineal transmission patterns. We therefore examined human parent-offspring association in this neurocircuitry, and investigated the degree of association in gray matter volume between parent and offspring. We used voxel-wise correlation analysis in a total of 35 healthy families, consisting of parents and their biological offspring. We found positive associations of regional grey matter volume in the corticolimbic circuit including the amygdala, hippocampus, anterior cingulate cortex, and ventromedial prefrontal cortex between biological mothers and daughters. This association was significantly greater than mother-son, father-daughter, and father-son associations. The current study suggests that the corticolimbic circuitry, which has been implicated in mood regulation, shows a matrilineal specific transmission patterns. Our preliminary findings are consistent with what has been found behaviorally in depression, and may have clinical implications for disorders known to have dysfunction in mood regulation such as depression. Studies such as ours will likely bridge animal work examining gene expression in the brains and clinical symptom-based observations, and provide promising ways to investigate intergenerational transmission patterns in the human brain.

Efficiency of uniparental male and female care against egg predators in two closely related syntopic harvestmen

Although the benefits of maternal care have been investigated in many species, the caring role of males in species with exclusive paternal care has received less attention. We experimentally quantified the protective role of paternal care in the harvestman Iporangaia pustulosa. Additionally, we compared the effectiveness of paternal care against predation in this species with a syntopic harvestman with maternal care, Acutisoma proximum. We demonstrated that nearly one-third of the unprotected Iporangaia clutches disappeared entirely in 12 days, while the other two-thirds suffered a mean reduction of 55% in egg number. Conversely, 50% of the control clutches did not suffer any reduction, and only one was entirely consumed by predators. We also demonstrated that the mucus coat that covers Iporangaia clutches has an important deterrent role against predation by conspecifics: 58.3% of the clutches without mucus were attacked and three of them were entirely consumed, whereas only three clutches with mucus were attacked, suffering a reduction of up to three eggs. Iporangaia males were as efficient as Acutisoma females in protecting eggs. However, unattended Acutisoma eggs were attacked 20% more frequently than unattended Iporangaia eggs. Unattended Iporangaia eggs are protected by a mucus coat that prevents or decreases predation rate, whereas Acutisoma eggs are more susceptible to predation, probably because they lack this mucus coat. Thus, besides the fact that Iporangaia males efficiently protect the offspring against egg predators, females also contribute to egg protection by providing a mucus coat that deters egg predators. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Song and female choice for extrapair copulations in the sedge warbler, Acrocephalus schoenobaenus

Although 90% of passerine birds live in socially monogamous pair bonds, molecular studies have revealed that genetic polygamy occurs in 86% of surveyed passerines, because individuals engage in copulations outside the pair bond (extrapair copulations; EPCs). Most explanations for the occurrence of EPCs involve female gaining indirect benefits from the extrapair male. The sedge warbler is a socially monogamous species in which some offspring result from EPCs (8% in this study). Complex song is a sexually selected male trait used by females which select mates based on a variety of male qualities. We used microsatellite DNA profiling to detect extrapair young and assign paternity. ‘Good genes’ theory predicts that females should engage in EPCs with males of higher quality than their social mate, with resulting fitness benefits. Extrapair males had smaller song repertoires and smaller territories than the social mate. This apparent preference for small-repertoire males as extrapair mates conflicts with the predictions from previous studies of this species. Sudden cessation of song after pairing may mean that song cues are unavailable for later extrapair matings and females may switch to other cues. Such behaviour may lead to different patterns of female choice during social and extrapair mating in the sedge warbler. We conclude that multiple reasons underlie patterns of female choice in this species.

Sperm competition and offspring viability at hybridization in Australian tree frogs, Litoria peronii and L. tyleri

Hybridization between closely related species often leads to reduced viability or fertility of offspring. Complete failure of hybrid offspring (post-zygotic hybrid incompatibilities) may have an important role in maintaining the integrity of reproductive barriers between closely related species. We show elsewhere that in Peron's tree frog, Litoria peronii, males more closely related to a female sire more offspring in sperm competition with a less related rival male. Observations of rare 'phenotypic intermediate' males between L. peronii and the closely related L. tyleri made us suggest that these relatedness effects on siring success may be because of selection arising from risks of costly hybridization between the two species. Here, we test this hypothesis in an extensive sperm competition experiment, which shows that there is no effect of species identity on probability of fertilization in sperm competition trials controlling for sperm concentration and sperm viability. Instead, there was a close agreement between a male's siring success in isolation with a female and his siring success with the same female in competition with a rival male regardless of species identity. Offspring viability and survival, however, were strongly influenced by species identity. Over a 14-day period, hybrid offspring suffered increasing mortality and developed more malformations and an obvious inability to swim and right themselves, leading to compromised probability of survival. Thus, hybridization in these sympatric tree frogs does not compromise fertilization but has a strong impact on offspring viability and opportunity for reinforcement selection on mate choice for conspecific partners.

Breeding ecology and bias in offspring sex ratio in little grassbirds (Megalurus gramineus)

Little grassbirds (Megalurus gramineus) are small, sexually monomorphic passerines that live in reed beds, lignum swamps and salt marshes in southern Australia. The breeding biology and patterns of sex allocation of the little grassbird were investigated over a single breeding season. Our observations of this species in the Edithvale Wetland Reserve revealed a highly male-biased population sex ratio, with some breeding territories containing several additional males. Nevertheless, there was little compelling evidence that little grassbirds breed cooperatively. The growth rates of male and female nestlings were similar and, as predicted by theory, there was no overall primary sex ratio bias. However, the primary sex ratio was female-biased early in the breeding season and became increasingly male-biased later in the breeding season.

Does female nuptial coloration reflect egg carotenoids and clutch quality in the Two-Spotted Goby (Gobiusculus flavescens, Gobiidae)?

Summary

1 - Carotenoid-based ornamentation has often been suggested to signal mate quality, and species with such ornaments have frequently been used in studies of sexual selection.

2 - Female Gobiusculus flavescens (Two-Spotted Goby) develop colourful orange bellies during the breeding season. Belly coloration varies among mature females, and previous work has shown that nest-holding males prefer females with more colourful bellies. Because males invest heavily in offspring during incubation, the evolution of this preference can be explained if colourful females provide males with eggs of higher quality.

3 - We tested this hypothesis by allowing males to spawn with ‘colourful’ and ‘drab’ females and comparing parameters including egg carotenoid concentration, clutch size, hatchability and larval viability between groups. We also investigated relationships between egg carotenoid concentration and clutch quality parameters.

4 - Eggs from colourful females had significantly higher concentrations of total carotenoids than drab females, and photographically quantified belly coloration was a good predictor of egg carotenoid concentration.

5 - Colourful females produced slightly larger clutches, but female belly coloration was not related to any measure of clutch quality. In addition, there were no significant relationships between egg carotenoids and clutch quality. Females with high levels of egg carotenoids spawned slightly earlier, however, possibly because they were more ready to spawn or because of male mate choice.

6 - Our results call into question the generality of a causal link between egg carotenoids and offspring quality.

Maternal stress to partner quality is linked to adaptive offspring sex ratio adjustment

Female birds have been shown to have a remarkable degree of control over the sex ratio of the offspring they produce. However, it remains poorly understood how these skews are achieved. Female condition, and consequent variation in circulating hormones, provides a plausible mechanistic link between offspring sex biases and the environmental and social stresses commonly invoked to explain adaptive sex allocation, such as diet, territory quality, and body condition. However, although experimental studies have shown that female perception of male phenotype alone can lead to sex ratio biases, it is unknown how partner quality influences female physiological state. Using a controlled within-female experimental design where female Gouldian finches (Erythrura gouldiae) bred with both high- and low-quality males, we found that partner quality directly affects female hormonal status and subsequent fitness. When constrained to breeding with low-quality males, females had highly elevated stress responses (corticosterone levels) and produced adaptive male-biased sex ratios, whereas when they bred with high-quality males, females had low corticosterone levels and produced an equal offspring sex ratio. There was no effect of other maternal hormones (e.g., testosterone) or body condition on offspring sex ratios. Female physiological condition during egg production, and variation in circulating hormones in particular, may provide a general mechanistic route for strategic sex allocation in birds.

Mothers adjust offspring sex to match the quality of the rearing environment

Theory predicts that mothers should adjust offspring sex ratios when the expected fitness gains or rearing costs differ between sons and daughters. Recent empirical work has linked biased offspring sex ratios to environmental quality via changes in relative maternal condition. It is unclear, however, whether females can manipulate offspring sex ratios in response to environmental quality alone (i.e. independent of maternal condition). We used a balanced within-female experimental design (i.e. females bred on both low- and high-quality diets) to show that female parrot finches (Erythrura trichroa) manipulate primary offspring sex ratios to the quality of the rearing environment, and not to their own body condition and health. Individual females produced an unbiased sex ratio on high-quality diets, but over-produced sons in poor dietary conditions, even though they maintained similar condition between diet treatments. Despite the lack of sexual size dimorphism, such sex ratio adjustment is in line with predictions from sex allocation theory because nutritionally stressed foster sons were healthier, grew faster and were more likely to survive than daughters. These findings suggest that mothers may adaptively adjust offspring sex ratios to optimally match their offspring to the expected quality of the rearing environment.

Detecting female precise natal philopatry in green turtles using assignment methods

It is well established that sea turtles return to natal rookeries to mate and lay their eggs, and that individual females are faithful to particular nesting sites within the rookery. Less certain is whether females are precisely returning to their natal beach. Attempts to demonstrate such precise natal philopatry with genetic data have had mixed success. Here we focused on the green turtles of three nesting sites in the Ascension Island rookery, separated by 5–15 km. Our approach differed from previous work in two key areas. First, we used male microsatellite data (five loci) reconstructed from samples collected from their offspring (N = 17) in addition to data for samples taken directly from females (N = 139). Second, we employed assignment methods in addition to the more traditional F-statistics. No significant genetic structure could be demonstrated with FST. However, when average assignment probabilities of females were examined, those for nesting populations in which they were sampled were indeed significantly higher than their probabilities for other populations (Mann–Whitney U-test: P < 0.001). Further evidence was provided by a significant result for the mAIC test (P < 0.001), supporting greater natal philopatry for females compared with males. The results suggest that female natal site fidelity was not sufficient for significant genetic differentiation among the nesting populations within the rookery, but detectable with assignment tests.