Parental condition affects early life-history of a coral reef fish

Parents can exert a range of non-genetic effects on the growth and survival of their offspring. In particular, parents may modify the size or condition of their offspring depending on the amount of energy they have available for reproduction. In this study, the body condition of breeding pairs of the coral reef fish Acanthochromis polyacanthus was experimentally manipulated to test the effects of parental condition on reproductive output and offspring life history traits. Parents in good condition commenced breeding earlier, had higher reproductive output, and their eggs exhibited increased survival during embryogenesis, compared to parents in poorer condition. Increased reproductive output was attained through more reproductive bouts over the breeding season that contained both a greater number and an increased size of eggs. The offspring from parents in good condition were larger at hatching, with larger yolk reserves and increased survival on endogenous reserves. Larger size is expected to provide benefits to offspring through reduced susceptibility to size-selective mortality. The range of offspring characteristics modified by parental condition could result in a greater proportion of offspring from good condition parents recruiting to the population.     

Rain,prey and predators: climatically driven shifts in frog abundance modify reproductive allometry in a tropical snake

To predict the impacts of climate change on animal populations, we need long-term data sets on the effects of annual climatic variation on the demographic traits (growth, survival, reproductive output) that determine population viability. One frequent complication is that fecundity also depends upon maternal body size, a trait that often spans a wide range within a single population. During an eight-year field study, we measured annual variation in weather conditions, frog abundance and snake reproduction on a floodplain in the Australian wet-dry tropics. Frog numbers varied considerably from year to year, and were highest in years with hotter wetter conditions during the monsoonal season (“wet season”). Mean maternal body sizes, egg sizes and post-partum maternal body conditions of frog-eating snakes (keelback, Tropidonophis mairii, Colubridae) showed no significant annual variation over this period, but mean clutch sizes were higher in years with higher prey abundance. Larger females were more sensitive to frog abundance in this respect than were smaller conspecifics, so that the rate at which fecundity increased with body size varied among years, and was highest when prey availability was greatest. Thus, the link between female body size and reproductive output varied among years, with climatic factors modifying the relative reproductive rates of larger (older) versus smaller (younger) animals within the keelback population.     

Resource allocation in a simultaneously hermaphroditic slug with phally polymorphism

The theory of resource allocation assumes that a resource not allocated to one function may be reallocated to another. Thus, in hermaphroditic species, an individual that suppresses the use of one sex function may free resources for the other sex function. We determined the relative importance of male copulatory organs in terms of their fraction of the total dry body weight and tested whether in the pulmonate land slug Deroceras laeve (Müller), individuals that lack the male copulatory organs (aphallics) reallocate this resource towards the female structures and/or towards life-history traits. To this end, we raised 13 families under uniparental reproduction and compared growth, length of the juvenile period, number of eggs produced, percentage of hatched eggs and hatching time among a- and euphallics. We also measured the reproductive and sex allocation of all individuals. Six out of 13 families contained no euphallic individuals. In the other seven families, the proportion of euphallic individuals ranged from 0.13 to 0.43. There was an enormous variation in life-history traits and reproductive and sex allocation among individuals, even among individuals of the same family. Allocation to the male function was very low in euphallic slugs (i.e. 1.35% of the total body dry mass and 12.33% of the total reproductive dry mass). Our results did not reveal a reallocation from the lost male function towards the female function, nor towards one of the life-history traits. Finally, we propose a scenario that could explain the maintenance of phally polymorphism in D. laeve.     

Experience versus effort: what explains dynamic heterogeneity with respect to age?

Age‐related patterns of survival and reproduction have been explained by accumulated experience (‘experience hypothesis’), increased effort (‘effort hypothesis’), and intrinsic differences in phenotypes (‘selection hypothesis’). We examined the experience and effort hypotheses using a 40‐year data set in a population of Leach's storm‐petrels Oceanodroma leucorhoa, long‐lived seabirds for which the effect of phenotypic variation has been previously demonstrated. Age was quantified by time since recruitment (‘breeding age’). The best model of adult survival included a positive effect of breeding age (1, 2, 3+ years), sex (male > female), and year. Among‐individuals variation (fixed heterogeneity) accounted for 31.6% of the variance in annual reproductive success. We further examined within‐individual patterns in reproductive success (dynamic heterogeneity) in the subset of individuals with at least five breeding attempts. Three distinct phases characterized reproductive success – early increase, long asymptotic peak, late decline. No effect of early reproductive output on longevity was found, however, early success was positively correlated with lifetime reproductive success. Reproductive success was lower earlier than later in life. Among the few natally philopatric individuals in the population, age of first breeding had no effect on longevity, lifetime reproductive success, or early reproductive success. No support for the effort hypothesis was found in this population. Instead, age‐specific patterns of survival and reproduction in these birds are best explained by the experience hypothesis over and above the effect of intrinsic differences among individuals.     

Life-history evolution in Anodonta piscinalis (Mollusca,Pelecypoda)

Summary The correlations between certain life-history parameters (reproductive effort, reproductive life-span, age of first reproduction, general growth index, variation in juvenile survival, availability of resources) were studied in 13 populations of the mussel Anodonta piscinalis in south-western Finland in 1975 and 1976Reproductive life-span correlated positively (r _s=0,823, P<0.001) with variation in juvenile survival. The average availability of resources correlated negatively both with the reproductive life-span (r _s=-0.841, P<0.001 after the variation by juvenile survival had been deleted) and variation in juvenile survival (r _s=-0.676, P<0.05).The reproductive effort for female mussels at each site was computed by comparing body weight of reproductive females with body weights of non-reproductive individuals. Availability of resources was much higher in 1976 than in 1975. Consequently, the reproductive effort, an index of the strain of reproduction, was higher in 1975 than in 1976. In 1975 there was a significant correlation between reproductive effort and the length of the reproductive life-span (-0.727, -0.806) and also with the reproductive effort and the variation in juvenile survival (-0.718, -0.758) in females of the length of 60 mm and 70 mm respectively. In 1976, when availability of resources was better, such correlations were not found.Spatial and temporal change in the intake of resources complicates applicability of the principle of resource allocation in the theory of life-history evolution. Studying the mere allocation is not enough if the intake of resources varies in the groups studied. The ratio ovary weight/body weight is a dubious measure of reproductive effort in comparative studies when the input of resources can vary, and this possibility can be ruled out only exceptionally.Correlation between growth and reproductive effort was positive, obviously because both are important components in creating high reproductive capacity. In 1975 reproductive effort increased with size (age). Change in reproductive effort correlated with reproductive life-span (r _s=-0.633, P<0.05).The following parameters occurred together: short reproductive life-span, low age of first reproduction, high reproductive effort, rapid growth and high clutch size. They were realized at sites where the availability of resources was good and variation in juvenile survival was low-i.e. the environment was stable. The results conflict with the prediction of the theory of r and K-selection.     

Delayed timing of breeding as a cost of reproduction

Timing of breeding is a trait with considerable individual variation, often closely linked to fitness because of seasonal declines in reproduction. The drivers of this variation have received much attention, but how reproductive costs may influence the timing of subsequent breeding has been largely unexplored. We examined a population of northern wheatears Oenanthe oenanthe to compare three groups of individuals that differed in their timing of breeding termination and reproductive effort to investigate how these factors may carry over to influence reproductive timing and reproductive output in the following season. Compared to females that bred successfully, females that put in less effort and terminated breeding early due to nest failure tended to arrive and breed earlier in year 2 (mean advancement = 2.2 and 3.3 d respectively). Females that spent potentially more effort and terminated breeding later due to production of a replacement clutch after nest failure, arrived later than other females in year 2. Reproductive output (number of fledglings) in year 2 differed between the three groups as a result of group‐level differences in the timing of breeding in combination with the general seasonal decline in reproductive output. Our study shows that the main cost of reproduction was apparent in the timing of arrival and breeding in this migratory species. Hence, reproductive costs can arise through altered timing of breeding since future reproductive success (including adult survival) is often dependent on the timing of breeding in seasonal systems.     

Costs of reproduction in male lizards, Sceloporus virgatus

Models of life history evolution typically assume a balance between the benefits of current reproductive activity and the costs to future reproductive success or survivorship, but empirical studies often find positive correlations between such components of fitness in undisturbed animal populations. I examined possible survivorship costs of reproduction in free-ranging male lizards, Sceloporus virgatus , and found that males with low levels of mating success were less likely to survive to the following breeding season. I also investigated two possible indicators of reproductive effort, increase in ectoparasite load and decrease in body weight during the breeding season. Levels of parasitism with trombiculid mites at the end of the breeding season were not associated with any measure of fitness or body condition (mating success, survivorship to the following year, relative weight loss). Yearling males (which have low levels of mating success) usually gained weight during the breeding season, while older males generally lost weight during this period. This suggests that young males may have postponed reproduction in favor of body growth and that seasonal weight loss of older males might reflect reproductive effort. Within the group of older males, individuals with the highest levels of mating success did not have high levels of either weight loss or mortality. Mate guarding behavior, an alternative to the aggressive territorial behavior typical of many lizard species, may allow certain males to obtain mates without expending large amounts of energy or exposing themselves to great mortality risks.     

Breeding phenology,variation in reproductive effort and offspring size in a tropical population of the woodlouse Porcellionides pruinosus

Summary Most species of woodlice in temperate habitats have discrete breeding seasons. It is hypothesised that breeding synchronises with favourable environmental conditions to maximise offspring growth and survivorship (Willows 1984). We measured the breeding phenology of a species introduced to a tropical environment, primarily to consider the assumption that life histories in the tropics will differ fundamentally from those in temperate habitats. In addition to breeding phenology we considered variation in reproductive effort between individual females and the division of this effort between the size and number of young.A continuous breeding phenology was observed in a synanthropic population of Porcellionides pruinosus within the tropics. Reproductive effort varied between months, showed a weak relationship with female size and was independent of female fecundity. Female sizefecundity relationships varied between samples and when the proportion of reproductive females was high size-fecundity slopes were steeper than at other times. Mean offspring size varied between months and there was a wide range in offspring size within broods. Offspring size was not related to female body mass, reproductive effort or fecundity; consequently brood mass increased linearly with an increase in fecundity. Increased reproductive effort goes into more rather than larger offspring.We propose that the continuous breeding in this population was the result of the constant presence of an environmental cue to reproduction evolved in temperate habitats. Continuous breeding is not necessarily equivocal to high individual reproductive success even though overall population growth may be rapid. However, variation in reproductive effort suggests that individuals respond to current environmental conditions on short time scales.     

Growth or reproduction? Resource allocation by female frogs<Emphasis Type="Italic"> Rana temporaria</Emphasis>

The decision how to allocate marginal resources to reproduction and growth can have important effects on associated life-history parameters as well as on population dynamics. In addition to showing variation among individuals in a population, such allocation rules may be either condition-dependent or fixed in different individuals. While many studies on anuran amphibians have focused on egg numbers and egg sizes in females of different sizes, virtually no data exist on the relative allocation of marginal resources to growth versus reproduction. In the laboratory, we therefore offered female common frogs (Rana temporaria) low versus high food rations for a full reproductive cycle, and monitored their growth and later reproductive investment (egg number and egg size the following breeding season). Feeding rates had an effect both on female growth and on egg number and size. There was no trade-off found between the two forms of investment. A flexible allocation rule could not be supported as there was no significant effect of feeding rate on the relative allocation of resources to growth versus reproduction.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Determinants of age at first reproduction and lifetime breeding success revealed by full paternity assignment in a male ungulate

Age at first reproduction is an important determinant of individual variation in reproductive success in ungulates, but few studies have examined its relationship with later fitness‐related traits in males. We used a long‐term individual based study of a harvested moose population to quantify the individual reproductive performance and survival of males, as well as to examine the determinants of age at first reproduction and consequences of age at first reproduction on lifetime breeding success. The probability that a male successfully reproduced at the age of two was negatively related to the mean age of adult males in the population, but the relationship weakened with increasing population size. Large antlers and large body mass relative to other males in the population increased the number of calves sired at their first successful mating season. In addition, those that successfully reproduced as two year‐olds were more likely to sire calves the next year, making them more productive at a given age compared to those that first reproduced at the age of three or older. We emphasize the importance for males to start reproducing as soon as possible in a harvested population to gain lifetime fitness benefits, as surviving the hunt is a major determinant of reproductive success in this population. We found no costs of early reproduction in males, hence leading to high individual heterogeneity in male reproductive performance. The apparent lack of reproductive costs could partly be explained by the age distribution in the population, individual variation in early‐life body mass and antler size, and differences in probabilities of being hunted of successful and unsuccessful males.     

Fathers, fat, and maternal energetics in a biparental hamster: paternal presence determines the outcome of a current reproductive effort and adipose tissue limits subsequent reproductive effort

Djungarian hamster females, Phodopus campbelli, are severely constrained in their ability to reproduce successfully and lose 20% of their body weight by the time pups are weaned. In the wild and in the laboratory, biparental care improves maternal reproductive success. Two experiments quantified the effects of paternal presence and partial lipectomy [surgical depletion of parametrial white adipose tissue (PWAT) on day 8 of the 18-day gestation] on maternal energy balance, reproductive success, and investment in a subsequent reproductive attempt. Paired females reproduced successfully, maintained body weight, and invested in a second litter. Removal of the male decreased pup survival, growth, and readiness for dispersal by 18 days of age. Solitary females lost 10% of their body weight by the birth and a further 10% by day 18 after the birth. Thus, paternal presence balanced maternal energy budgets during reproduction and prevented a 20% loss in body weight. Equivalent weight loss occurs in response to other maternal stressors, therefore 20% may be the maximum tolerable weight loss in this species. Fresh weight of interscapular brown adipose tissue was predicted by the extent of maternal hyperthermia but not by maternal energy balance or lipectomy. Partial lipectomy did not adversely affect the female or the first litter but decreased the probability of investment in a second reproductive attempt and halved the size of the second litter. This effect may have been due to the 0.1% of body weight amount of lipid removed or may reflect a specialized role for PWAT in adjusting maternal investment.     

Individual quality, survival variation and patterns of phenotypic selection on body condition and timing of nesting in birds

Questions about individual variation in quality and fitness are of great interest to evolutionary and population ecologists. Such variation can be investigated using either a random effects approach or an approach that relies on identifying observable traits that are themselves correlated with fitness components. We used the latter approach with data from 1,925 individual females of three species of ducks (tufted duck, Aythya fuligula; common pochard, Aythya ferina; northern shoveler, Anas clypeata) sampled on their breeding grounds at Engure Marsh, Latvia, for over 15 years. Based on associations with reproductive output, we selected two traits, one morphological (relative body condition) and one behavioral (relative time of nesting), that can be used to characterize individual females over their lifetimes. We then asked whether these traits were related to annual survival probabilities of nesting females. We hypothesized quadratic, rather than monotonic, relationships based loosely on ideas about the likely action of stabilizing selection on these two traits. Parameters of these relationships were estimated directly using ultrastructural models embedded within capture-recapture-band-recovery models. Results provided evidence that both traits were related to survival in the hypothesized manner. For all three species, females that tended to nest earlier than the norm exhibited the highest survival rates, but very early nesters experienced reduced survival and late nesters showed even lower survival. For shovelers, females in average body condition showed the highest survival, with lower survival rates exhibited by both heavy and light birds. For common pochard and tufted duck, the highest survival rates were associated with birds of slightly above-average condition, with somewhat lower survival for very heavy birds and much lower survival for birds in relatively poor condition. Based on results from this study and previous work on reproduction, we conclude that nest initiation date and body condition covary with both reproductive and survival components of fitness. These associations lead to a positive covariance of these two fitness components within individuals and to the conclusion that these two traits are indeed correlates of individual quality.     

Does loss of mass during breeding correlate with reproductive success? A study on Blue Tits Parus caeruleus

Substantial loss of mass of female Blue Tits Parus cueruleus during breeding is commonly explained by three not mutually exclusive explanations: shrinking of gonadal tissues, cost of reproduction and adaptation to save energy during flight. This study showed that loss of body mass was inversely correlated with reproductive success of individual females. Female tarsus length and timing of breeding correlated with loss of body mass, whereas ectoparasite loading had no significant effect on body mass. Loss of body mass during the rearing of young could be a useful measure of cost of reproduction by accounting for individual variation in female quality.     

Group Structure,Nest Size and Reproductive Success in the Cooperatively Breeding Cichlid Julidochromis ornatus: A Correlation Study

The effect of group size on reproductive success has long been studied in cooperatively breeding species, as it might provide an adaptive explanation for group‐living in social species. Numerous studies have shown positive effects of subordinates on reproductive success (‘helper effect’), but these studies have also revealed the importance of controlling statistically, or experimentally, for the effect of other factors that might affect reproductive success. Here, we first examine the relationships between group size, body size of group members and nest size in the cooperatively breeding cichlid Julidochromis ornatus, in which unrelated helpers frequently participate in reproduction and their breeding nests inside rock crevices may be crucial for reproduction and survival of all group members. Then, we subsequently investigate the relationship between group size and reproductive success, while controlling for these factors. The results showed that group size was significantly related to body size of group members rather than nest size; and larger breeders had larger helpers. It was found that group size significantly increased group reproductive output. More importantly, reproductive success of male breeders did not depend on the presence of mature helpers, whereas female reproductive success increased when two males assisted her and tended to decrease when two females bred cooperatively. We conclude that breeding groups of J. ornatus have size hierarchical societies that relate to group size, and group composition of genetically unrelated and co‐breeding members affects their reproductive success.     

Age and terminal reproductive attempt influence laying date in the thorn‐tailed rayadito

Age‐specific variation in reproductive effort can affect population dynamics, and is a key component of the evolution of reproductive tactics. Late‐life declines are a typical feature of variation in reproduction. However, the cause of these declines, and thus their implications for the evolution of life‐history tactics, may differ. Some prior studies have shown late‐life reproductive declines to be tied to chronological age, whereas other studies have found declines associated with terminal reproduction irrespective of chronological age. We investigated the extent to which declines in late life reproduction are related to chronological age, terminal reproductive attempt or a combination of both in the thorn‐tailed rayadito Aphrastura spinicauda, a small passerine bird that inhabits the temperate forest of South America. To this end we used long‐term data (10 years) obtained on reproductive success (laying date, clutch size and nestling weight) of females in a Chilean population. Neither chronological age nor terminal reproductive attempt explained variation in clutch size or nestling weight, however we observed that during the terminal reproductive attempt older females tended to lay later in the breeding season and younger females laid early in the breeding season, but this was not the case when the reproductive attempt was not the last. These results suggests that both age‐dependent and age‐independent effects influence reproductive output and therefore that the combined effects of age and physiological condition may be more relevant than previously thought.     

Adult survival and reproductive rate are linked to habitat preference in territorial,year‐round resident Song Sparrows Melospiza melodia

Individual animal fitness can be strongly influenced by the ability to recognize habitat features which may be beneficial. Many studies focus on the effects of habitat on annual reproductive rate, even though adult survival is typically a greater influence on fitness and population growth in vertebrate species with intermediate to long lifespans. Understanding the effects of preferred habitat on individuals over the annual cycle is therefore necessary to predict its influences on individual fitness. This is particularly true in species that are resident and territorial year‐round in the temperate zone, which may face potential trade‐offs between habitat that maximizes reproduction and that which maximizes non‐breeding season (‘over‐winter’) survival. We used a 37‐year study of Song Sparrows Melospiza melodia residing territorially year‐round on a small island to examine what habitat features influenced adult over‐winter survival, how site‐specific variation in adult survival vs. annual reproductive rate influenced long‐term habitat preference, and if preferred sites on average conferred higher individual fitness. Habitat features such as area of shrub cover and exposure to intertidal coastline predicted adult over‐winter survival independent of individual age or sex, population size, or winter weather. Long‐term habitat preference (measured as occupation rate) was better predicted by site‐specific annual reproductive rate than by expected over‐winter survival, but preferred sites maximized fitness on average over the entire annual cycle,. Although adult over‐winter survival had a greater influence on population growth (λ) than did reproductive rate, the influence of reproductive rate on λ increased in preferred sites because site‐specific variation in reproductive rate was higher than variation in expected over‐winter survival. Because preferred habitats tended to have higher mean site‐specific reproductive and adult survival rates, territorial birds in this population do not appear to experience seasonal trade‐offs in preferred habitat but are predicted to incur substantial fitness costs of settling in less‐preferred sites.     

The cost of reproduction in the glaucous-winged gull

Summary Experimental enlargement of brood size in the glaucous-winged gull (Larus glaucescens) resulted in increased adult foraging time, decreased adult body weight at the end of the breeding season, and decreased over-winter adult survival. The decreased survival of breeding adults was associated with reduced body condition at the end of breeding (resulting from physiological costs of reproduction). Decreased survival was not due to an increased risk of injury or predation during the breeding season. Brood size did not directly affect the fecundity of surviving birds in the subsequent year. However, brood size may have an indirect effect on subsequent fecundity because the probability of mate loss increased among birds with large broods and the reproductive performance of birds with new mates was reduced. Based on estimates of life-time fitness calculated from fecundity and survivorship, birds with two- or three-chick broods (the normal brood size) have higher fitness than birds with one- or four-chick broods. However, the decreased fitness of birds with four-chick broods was slight, and probably not a sufficient explanation for the absence of natural four-chick broods in the glaucouswinged gull.     

Phenotypic variation in the breeding phenology of the woodlouse Armadillidium vulgare

Summary The breeding phenology of temperate wood-lice is strongly seasonal, the result of physiological constraints and precise environmental cues for reproduction. The adaptive value of such mechanisms is that the release of offspring coincides with favourable conditions for growth and survival (Willows 1984). We recorded the breeding phenology of Armadillidium vulgare (Latreille) on two grassland sites in Great Britain and found between-site and between-year variation in the onset of reproduction, the duration of reproductive activity, the release of offspring, the size of reproductive females and the number of broods per female. Between 82.7 and 97.7% of gravid females sampled were semelparous at 23 months, with the remainder iteroparous, producing a second brood after 35 months. On one site (Weeting Health) improved growth conditions during 1984 allowed some females (19.3% of gravid females sampled in that year) to produce a brood after 11 months. There was also an increase in the number of 3-year-old females found to be gravid. An experimental manipulation of the same habitat confirmed that such changes in life history tactics could be phenotypic responses. The observed phenotypic variation was sufficient to produce a range of life history tactics within a population. Mixtures of life history tactics within a population may be typical of invasive species and populations at the edge of the species range. Our results support the idea that phenotypic plasticity can be an appropriate tactic to maximise fitness in a fluctuating environment (Caswell 1983, 1989).     

荒漠草原土壤相对湿度对猪毛蒿表型可塑性的影响

猪毛蒿(Artemisia scoparia)为菊科蒿属草本植物,是一种适应性较强的广幅种。研究荒漠草原不同土壤相对湿度条件下猪毛蒿的表型可塑性,对认识异质生境下猪毛蒿的生存适应策略具有重要的生态学意义。结果表明:株高、茎粗、根长、根重和单株生物量均表现出随土壤相对湿度的增大而增加的趋势,对异质生境具有较强的可塑性,而根冠比则表现出相对的稳定性。植株不同部位生物量大小排序为:上部中部下部,且植株下部显著大于上部生物量(P0.05)。土壤相对湿度40%生境下的头状花序数量和重量显著高于土壤相对湿度30%和30%—40%生境。繁殖器官绝对投入量(lg R)随着个体大小(lg V)的增大呈极显著的增加(P0.001),繁殖阈值介于1.868—2.006 g。随着土壤相对湿度的增加,繁殖分配比例极显著增大(P0.001)。营养器官和繁殖器官生物量、头状花序重量和数量、地下生物量和地上生物量均呈极显著线性正相关关系(P0.001),存在正向权衡。单个头状花序重量并不随个体大小和头状花序数量的增加而发生显著变化(P0.05),且在不同土壤相对湿度和不同部位间均无显著差异(P0.05)。由此可见,猪毛蒿在异质生境下产生的可塑性是其生存繁殖的重要反应机制之一。