Local and neighboring patch conditions alter sex‐specific movement in banana weevils

Understanding the mechanisms underlying the movements and spread of a species over time and space is a major concern of ecology. Here, we assessed the effects of an individual's sex and the density and sex ratio of conspecifics in the local and neighboring environment on the movement probability of the banana weevil, Cosmopolites sordidus. In a “two patches” experiment, we used radiofrequency identification tags to study the C. sordidus movement response to patch conditions. We showed that local and neighboring densities of conspecifics affect the movement rates of individuals but that the density‐dependent effect can be either positive or negative depending on the relative densities of conspecifics in local and neighboring patches. We demonstrated that sex ratio also influences the movement of C. sordidus, that is, the weevil exhibits nonfixed sex‐biased movement strategies. Sex‐biased movement may be the consequence of intrasexual competition for resources (i.e., oviposition sites) in females and for mates in males. We also detected a high individual variability in the propensity to move. Finally, we discuss the role of demographic stochasticity, sex‐biased movement, and individual heterogeneity in movement on the colonization process.     

Sex differences in parental care: Gametic investment,sexual selection,and social environment

Male and female parents often provide different type and amount of care to their offspring. Three major drivers have been proposed to explain parental sex roles: (1) differential gametic investment by males and females that precipitates into sex difference in care, (2) different intensity of sexual selection acting on males and females, and (3) biased social environment that facilitates the more common sex to provide more care. Here, we provide the most comprehensive assessment of these hypotheses using detailed parental care data from 792 bird species covering 126 families. We found no evidence for the gametic investment hypothesis: neither gamete sizes nor gamete production by males relative to females was related to sex difference in parental care. However, sexual selection correlated with parental sex roles, because the male share in care relative to female decreased with both extra‐pair paternity and frequency of male polygamy. Parental sex roles were also related to social environment, because male parental care increased with male‐biased adult sex ratios (ASRs). Taken together, our results are consistent with recent theories suggesting that gametic investment is not tied to parental sex roles, and highlight the importance of both sexual selection and ASR in influencing parental sex roles.     

Condition and reproductive investment in the western mosquitofish (Gambusia affinis): little evidence for condition‐dependent sex‐biased investment

In sexually reproducing species, resources may theoretically be distributed with bias to the production of male or female offspring in response to the condition of the mother, commonly recognized as sex allocation. Using a recently characterized sex‐specific molecular marker, we tested for maternal sex allocation (i.e. maternal primary sex ratio bias and sex‐specific offspring investment) in captive laboratory‐bred western mosquitofish (Gambusia affinis) at early stages of offspring development. We found no statistical evidence to support sex allocation in G. affinis, based on maternal condition. In addition, we found little evidence for correlations between maternal condition and investment in the condition (mass) of individual offspring (of one sex or the other), although we did find that larger mothers tended to have higher fecundity.     

Age‐specific survival and movement among major African Penguin Spheniscus demersus colonies

Reliable estimates of survival and dispersal are crucial to understanding population dynamics, but for seabirds, in which some individuals spend years away from land, mortality and emigration are often confounded. Multistate mark–recapture methods reduce bias by incorporating movement into the process of estimating survival. We used a multistate model to provide unbiased age‐specific survival and movement probabilities for the Endangered African Penguin Spheniscus demersus based on 5281 nestlings and 31 049 adults flipper‐banded and resighted in the Western Cape, South Africa, between 1994 and 2012. Adult survival was initially high (≥ 0.74) but declined after 2003–2004 coincident with a reduction in the availability of Sardine Sardinops sagax and Anchovy Engraulis encrasicolus on the west coast of South Africa. Juvenile survival was poorly estimated, but was lower and more variable than adult survival. Fidelity to the locality of origin varied over time, but was high in adults at Robben and Dassen islands (≥ 0.88) and above 0.55 for juvenile and immature Penguins at all localities. Movement occurred predominantly during 1994–2003 and was indicative of immigration to Robben and Dassen islands. Our results confirm that a prolonged period of adult mortality contributed to the observed decline in the African Penguin population and suggest a need for approaches operating over large spatial scales to ensure food security for marine top predators.     

Sexual conflict in action: An antagonistic relationship between maternal and paternal sex allocation in the tammar wallaby,Notamacropus eugenii

Sex ratio biases are often inconsistent, both among and within species and populations. While some of these inconsistencies may be due to experimental design, much of the variation remains inexplicable. Recent research suggests that an exclusive focus on mothers may account for some of the inconsistency, with an increasing number of studies showing variation in sperm sex ratios and seminal fluids. Using fluorescent in‐situ hybridization, we show a significant population‐level Y‐chromosome bias in the spermatozoa of wild tammar wallabies, but with significant intraindividual variation between males. We also show a population‐level birth sex ratio trend in the same direction toward male offspring, but a weaning sex ratio that is significantly female‐biased, indicating that males are disproportionately lost during lactation. We hypothesize that sexual conflict between parents may cause mothers to adjust offspring sex ratios after birth, through abandonment of male pouch young and reactivation of diapaused embryos. Further research is required in a captive, controlled setting to understand what is driving and mechanistically controlling sperm sex ratio and offspring sex ratio biases and to understand the sexually antagonistic relationship between mothers and fathers over offspring sex. These results extend beyond sex allocation, as they question studies of population processes that assume equal input of sex chromosomes from fathers, and will also assist with future reproduction studies for management and conservation of marsupials.     

Determination of the optimal parasitoid‐to‐host ratio for efficient mass‐rearing of the parasitoid,Sclerodermus pupariae (Hymenoptera: Bethylidae)

Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae) is used as a potential biocontrol agent for several buprestid and cerambycid larvae. This study aimed to enhance the efficiency of mass‐rearing of this parasitoid by investigating the fitness gain of this bethylid wasp, including the proportion of successful parasitism and development, brood size, sex ratio, proportion of winged female offspring, body size and longevity of female offspring, under eight different maternal parasitoid density treatments using Thyestilla gebleri Faldermann as host in the laboratory. The results indicated that the foundress densities did not affect the parasitism or emergence rate of this parasitoid. Brood size of the parasitoids increased significantly when the number of maternal wasps ranged from one to four. However, further increases in foundress number did not affect the parasitoid brood size. The sex ratios of S. pupariae were always female‐biased. The proportions of male in the progeny colonies were <10% throughout all experimental treatments. The percentage of winged female progeny was not significantly influenced by the density of adult maternal parasitoids. Body sizes of parasitoids significantly declined with increasing maternal parasitoid densities. Although the parasitoid body size reduced when maternal wasp number was higher, it could be compromised by the relatively higher number of female offspring produced. Further, more than 70% of the parasitoids remained alive when they were stored at 12°C for four months throughout the experiments. These findings suggest that exposure of four female wasps to a single host larva would result in the highest fitness of S. pupariae. Our findings might provide a new approach to enhance the efficiency of mass‐rearing of this bethylid wasp.     

Late‐breeding Great Cormorants Phalacrocorax carbo sinensis produce fewer young of the more vulnerable sex

We examined the brood sex ratio and offspring body mass in relation to the timing of breeding and brood size in the Great Cormorant Phalacrocorax carbo sinensis. The brood sex ratio was not related to brood size but it was significantly related to the hatching date, with a decreasing proportion of males in the brood in the course of the season. Male chicks had significantly lower body mass if they hatched later in the season, whereas there was no such relationship for female offspring. Assuming that environmental conditions deteriorate with progress of the breeding season, and male offspring may be more vulnerable to poor environmental conditions, the observed decline in the proportion of male offspring late in the season may be adaptive.     

Variations in age‐ and sex‐specific survival rates help explain population trend in a discrete marine mammal population

Understanding the drivers underlying fluctuations in the size of animal populations is central to ecology, conservation biology, and wildlife management. Reliable estimates of survival probabilities are key to population viability assessments, and patterns of variation in survival can help inferring the causal factors behind detected changes in population size. We investigated whether variation in age‐ and sex‐specific survival probabilities could help explain the increasing trend in population size detected in a small, discrete population of bottlenose dolphins Tursiops truncatus off the east coast of Scotland. To estimate annual survival probabilities, we applied capture–recapture models to photoidentification data collected from 1989 to 2015. We used robust design models accounting for temporary emigration to estimate juvenile and adult survival, multistate models to estimate sex‐specific survival, and age models to estimate calf survival. We found strong support for an increase in juvenile/adult annual survival from 93.1% to 96.0% over the study period, most likely caused by a change in juvenile survival. Examination of sex‐specific variation showed weaker support for this trend being a result of increasing female survival, which was overall higher than for males and animals of unknown sex. Calf survival was lower in the first than second year; a bias in estimating third‐year survival will likely exist in similar studies. There was some support first‐born calf survival being lower than for calves born subsequently. Coastal marine mammal populations are subject to the impacts of environmental change, increasing anthropogenic disturbance and the effects of management measures. Survival estimates are essential to improve our understanding of population dynamics and help predict how future pressures may impact populations, but obtaining robust information on the life history of long‐lived species is challenging. Our study illustrates how knowledge of survival can be increased by applying a robust analytical framework to photoidentification data.     

Male‐specific mortality biases secondary sex ratio in Eurasian tree sparrows Passer montanus

Sex allocation theory predicts that parents bias the offspring sex ratio strategically. In avian species, the offspring sex ratio can be biased at multiple growth stages, although the mechanisms are not well known. It is crucial to reveal a cause and timing of biased offspring sex ratio. We investigated (i) offspring sex ratio at multiple growth stages, from laying to fledging; and (ii) the stage at which offspring sex ratio became biased; and (iii) the cause of biased offspring sex ratio in Eurasian tree sparrows Passer montanus. Sex determination of 218 offspring, including hatchlings and unhatched eggs from 41 clutches, suggested that the offspring sex ratio was not biased at the egg‐laying stage but was significantly female‐biased after the laying stage due to higher mortality of male embryos. Half of the unhatched eggs showed no sign of embryo development (37/74, 50.00%), and most undeveloped eggs were male (36/37, 97.30%). Additional experiments using an incubator suggested that the cause of embryo developmental failure was a lack of developmental ability within the egg, rather than a failure of incubation. This study highlights the importance of clarifying offspring sex ratio at multiple stages and suggests that offspring sex ratio is adjusted after fertilization.     

Reproductive senescence and parental effects in an indeterminate grower

Reproductive senescence is the decrease of reproductive performance with increasing age and can potentially include trans‐generational effects as the offspring produced by old parents might have a lower fitness than those produced by young parents. This negative effect may be caused either by the age of the father, mother or the interaction between the ages of both parents. Using the common woodlouse Armadillidium vulgare, an indeterminate grower, as a biological model, we tested for the existence of a deleterious effect of parental age on fitness components. Contrary to previous findings reported from vertebrate studies, old parents produced both a higher number and larger offspring than young parents. However, their offspring had lower fitness components (by surviving less, producing a smaller number of clutches or not reproducing at all) than offspring born to young parents. Our findings strongly support the existence of trans‐generational senescence in woodlice and contradict the belief that old individuals in indeterminate growers contribute the most to recruitment and correspond thereby to the key life stage for population dynamics. Our work also provides rare evidence that the trans‐generational effect of senescence can be stronger than direct reproductive senescence in indeterminate growers.     

Variation in offspring sex ratio of a long‐lived sexually dimorphic raptor,the Eastern Imperial Eagle Aquila heliaca

Sex ratio theory attempts to explain observed variation in offspring sex ratio at both the population and the brood levels. In the context of low‐fecundity organisms producing high‐investment offspring, the drivers of adaptive variation in sex ratio are incompletely understood. For raptors that display reverse sexual dimorphism (RSD), preferential allocation of resources to the putatively cheaper sex (male) may be a response to environmental, social or demographic stressors. To assess the extent of skew in offspring sex ratios and to evaluate possible dietary, environmental and demographic correlates of such skew to long‐lived RSD avian species, we evaluated the offspring sex ratio of 219 chicks from 119 broods in 30 territories of Eastern Imperial Eagles Aquila heliaca across 7 years and four regions at a nature reserve in Kazakhstan. Only in one region in 1 year of our study did the offspring sex ratio differ significantly from parity (10 males : 1 female in 11 territories). Whereas offspring sex ratios were independent of dietary diversity, precipitation, temperature and productivity, we found that year had a moderate effect on brood sex ratio within territories. Our results provide limited evidence of brood sex manipulation in these populations of Eastern Imperial Eagles, and no mechanistic insight into predictions associated with it. Stochastic variation is likely to explain much of the fluctuation we observed in sex ratios, but our observations are also consistent with the hypothesis that sex‐ratio manipulation may occur irregularly, in concurrence with atypical environmental or demographic conditions that fluctuate at a time scale longer than that of our 7‐year study.     

Lack of sex‐specific movement patterns in an alien species at its invasion front – consequences for invasion speed

Efficient targeting of actions to reduce the spread of invasive alien species relies on understanding the spatial, temporal, and individual variation of movement, in particular related to dispersal. Such patterns may differ between individuals at the invasion front compared to individuals in established and dense populations due to differences in environmental and ecological conditions such as abundance of conspecifics or sex‐specific dispersal affecting the encounter rate of potential mates. We assessed seasonal and diurnal variation in movement pattern (step length and turning angle) of adult male and female raccoon dog at their invasion front in northern Sweden using data from Global Positioning System (GPS)‐marked adult individuals and assessed whether male and female raccoon dog differed in their movement behavior. There were few consistent sex differences in movement. The rate of dispersal was rather similar over the months, suggesting that both male and female raccoon dog disperse during most of the year, but with higher speed during spring and summer. There were diurnal movement patterns in both sexes with more directional and faster movement during the dark hours. However, the short summer nights may limit such movement patterns, and long‐distance displacement was best explained by fine‐scale movement patterns from 18:00 to 05:00, rather than by movement patterns only from twilight and night. Simulation of dispersing raccoon dogs suggested a higher frequency of male–female encounters that were further away from the source population for the empirical data compared to a scenario with sex differences in movement pattern. The lack of sex differences in movement pattern at the invasion front results in an increased likelihood for reproductive events far from the source population. Animals outside the source population should be considered potential reproducing individuals, and a high effort to capture such individuals is needed throughout the year to prevent further spread.     

Pollen limitation and Allee effect related to population size and sex ratio in the endangered Ottelia acuminata (Hydrocharitaceae): implications for conservation and reintroduction

Small populations may suffer more severe pollen limitation and result in Allee effects. Sex ratio may also affect pollination and reproduction success in dioecious species, which is always overlooked when performing conservation and reintroduction tasks. In this study, we investigated whether and how population size and sex ratio affected pollen limitation and reproduction in the endangered Ottelia acuminata, a dioecious submerged species. We established experimental plots with increasing population size and male sex ratio. We observed insect visitation, estimated pollen limitation by hand‐pollinations and counted fruit set and seed production per fruit. Fruit set and seed production decreased significantly in small populations due to pollinator scarcity and thus suffered more severe pollen limitation. Although frequently visited, female‐biased larger populations also suffered severe pollen limitation due to few effective visits and insufficient pollen availability. Rising male ratio enhanced pollination service and hence reproduction. Unexpectedly, pollinator preferences did not cause reduced reproduction in male‐biased populations because of high pollen availability. However, reproductive outputs showed more variability in severe male‐biased populations. Our results revealed two component Allee effects in fruit set and seed production, mediated by pollen limitation in O. acuminata. Moreover, reproduction decreased significantly in larger female‐biased populations, increasing the risk of an Allee effect.     

Home range size of adult Indo‐Pacific bottlenose dolphins (Tursiops aduncus) in a coastal and estuarine system is habitat and sex‐specific

This study examined sex‐specific differences in home range size of adult Indo‐Pacific bottlenose dolphins off Bunbury, Western Australia. We applied a new kernel density estimation approach that accounted for physical barriers to movements. A Bayesian mixture model was developed to estimate a sex effect in home range size with latent group partitioning constrained by association data. A post hoc analysis investigated group partitioning relating to the proportion of time spent in open vs. sheltered waters. From 2007 to 2013, photographic‐identification data were collected along boat‐based systematic transect lines (n = 586). Analyses focused on adult dolphins of known sex (sighted ≥ 30 times; n = 22 males and 34 females). The 95% utilization distributions of males varied between 27 and 187 km² (; 94.8 ± 48.15) and for females between 20 and 133 km² (65.6 ± 30.9). The mixture model indicated a 99% probability that males had larger home ranges than females. Dolphins mostly sighted in open waters had larger home ranges than those in sheltered waters. Home ranges of dolphins sighted in sheltered waters overlapped with areas of highest human activity. We suggest that sex differences in home ranges are driven by male mating strategies, and home range size differences between habitats may be influenced by prey availability and predation risk.     

Interactions between cytoplasmic and nuclear genomes confer sex‐specific effects on lifespan in Drosophila melanogaster

Genetic variation outside of the cell nucleus can affect the phenotype. The cytoplasm is home to the mitochondria, and in arthropods often hosts intracellular bacteria such as Wolbachia. Although numerous studies have implicated epistatic interactions between cytoplasmic and nuclear genetic variation as mediators of phenotypic expression, two questions remain. Firstly, it remains unclear whether outcomes of cyto‐nuclear interactions will manifest differently across the sexes, as might be predicted given that cytoplasmic genomes are screened by natural selection only through females as a consequence of their maternal inheritance. Secondly, the relative contribution of mitochondrial genetic variation to other cytoplasmic sources of variation, such as Wolbachia infection, in shaping phenotypic outcomes of cyto‐nuclear interactions remains unknown. Here, we address these questions, creating a fully crossed set of replicated cyto‐nuclear populations derived from three geographically distinct populations of Drosophila melanogaster, measuring the lifespan of males and females from each population. We observed that cyto‐nuclear interactions shape lifespan and that the outcomes of these interactions differ across the sexes. Yet, we found no evidence that placing the cytoplasms from one population alongside the nuclear background of others (generating putative cyto‐nuclear mismatches) leads to decreased lifespan in either sex. Although it was difficult to partition mitochondrial from Wolbachia effects, our results suggest at least some of the cytoplasmic genotypic contribution to lifespan was directly mediated by an effect of sequence variation in the mtDNA. Future work should explore the degree to which cyto‐nuclear interactions result in sex differences in the expression of other components of organismal life history.     

Co‐foundress confinement elicits kinship effects in a naturally sub‐social parasitoid

Kinship among interacting individuals is often associated with sociality and also with sex ratio effects. Parasitoids in the bethylid genus Goniozus are sub‐social, with single foundress females exhibiting post‐ovipositional maternal care via short‐term aggressive host and brood defence against conspecific females. Due to local mate competition (LMC) and broods normally being produced by a single foundress, sex ratios are female‐biased. Contests between adult females are, however, not normally fatal, and aggression is reduced when competing females are kin, raising the possibility of multi‐foundress reproduction on some hosts. Here, we screen for further life‐history effects of kinship by varying the numbers and relatedness of foundresses confined together with a host resource and also by varying the size of host. We confined groups of 1–8 Goniozus nephantidis females together with a host for 5+ days. Multi‐foundress groups were either all siblings or all nonsiblings. Our chief expectations included that competition for resources would be more intense among larger foundress groups but diminished by both larger host size and closer foundress relatedness, affecting both foundress mortality and reproductive output. From classical LMC theory, we expected that offspring group sex ratios would be less female‐biased when there were more foundresses, and from extended LMC theory, we expected that sex ratios would be more female‐biased when foundresses were close kin. We found that confinement led to the death of some females (11% overall) but only when host resources were most limiting. Mortality of foundresses was less common when foundresses were siblings. Developmental mortality among offspring was considerably higher in multi‐foundress clutches but was unaffected by foundress relatedness. Groups of sibling foundresses collectively produced similar numbers of offspring to nonsibling groups. There was little advantage for individual females to reproduce in multi‐foundress groups: single foundresses suppressed even the largest hosts presented and had the highest per capita production of adult offspring. Despite single foundress reproduction being the norm, G. nephantidis females in multi‐foundress groups appear to attune sex allocation according to both foundress number and foundress relatedness: broods produced by sibling foundresses had sex ratios similar to broods produced by single foundresses (ca. 11% males), whereas the sex ratios of broods produced by nonsibling females were approximately 20% higher and broadly increased with foundress number. We conclude that relatedness and host size may combine to reduce selection against communal reproduction on hosts and that, unlike other studied parasitoids, G. nephantidis sex ratios conform to predictions of both classical and extended LMC theories.     

Sex ratio,sex‐specific pattern in vegetative growth and gemma production in an aquatic liverwort,Scapania undulata (Marchantiophyta: Scapaniaceae)

Scapania undulata is an aquatic dioicous liverwort growing in shallow streams in boreal to subtropical zones. We studied the expressed sex ratio, sex‐specific differences in shoot architecture and possible trade‐off between sexual and asexual reproduction in ten populations of S. undulata by surveying 100 plots in ten streams in southern Finland. The expressed sex ratio was male biased, in contrast with the sex ratio in most dioicous bryophytes. It was also highly variable between the streams, but individual plots frequently comprised shoots from only one sex. The overproduction of males might be a strategy to overcome sperm dilution and ensure fertilization over longer distances in water. No size differences between females and males were detected, but they differed in branching patterns. Evidence for a higher cost of sexual reproduction in females than males can be seen from the following: the male‐biased sex ratio; low number of sex‐expressing female shoots in female‐only plots; no co‐occurrence of gemmae and female sex organs on a single branch, and no more than one sexual branch per female shoot. In contrast, high gemma production of male and female sex‐expressing shoots indicates a minimal trade‐off between sexual and asexual reproduction. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 229–241.     

Telomere dynamics in a lizard with morph‐specific reproductive investment and self‐maintenance

Telomeres in human fibroblasts shorten progressively during in vitro culturing and trigger replicative senescence. Furthermore, shortened telomeres can be used as biomarkers of disease. These observations have led to the suggestion that telomere dynamics may also be associated with viability and selection for life history variation in non‐human taxa. Model systems to examine this suggestion would particularly benefit from the coexistence of multiple phenotypes within the same species with different life history trade‐offs, since those could be compared in terms of telomere characteristics. This scenario also provokes the classic question of why one morph does not have marginally higher fitness and replaces the others. One explanation is that different morphs have different reproductive tactics with equal relative fitness. In Australian painted dragons (Ctenophorus pictus), males differ in head color, the presence or absence of a gular bib, and reproductive expenditure. Red males out‐compete yellow males in dominance contests, while yellow males copulate quickly and have higher success in sperm competition than red males. Males with bibs better defend partners against rival matings, at the cost of loss of body condition. We show that yellow‐headed and bib‐less males have longer telomeres than red, blue and bibbed males, suggesting that telomere length is positively associated with higher investment into self‐maintenance and less reproductive expenditure.