Experimental evidence that age-specific reproductive success is independent of environmental effects

An age-specific improvement in reproductive performance has been reported in many iteroparous breeders. However, whether this is a consequence of intrinsic differences in competence amongst age classes or extrinsic differences in the environment they experience is unclear since the timing of breeding within a season generally also differs with age. To disentangle these effects, we experimentally manipulated the timing of breeding in shags, Phalacrocorax aristotelis. Old and young individuals thus reared their chicks at the same time both early and late in the breeding season. When breeding in the same environmental conditions, old pairs performed consistently better than young pairs. These data clearly demonstrate that the age-related differences in reproductive performance are not a result of environmental effects, but rather a consequence of intrinsic differences in brood rearing capacity.     

Infanticide in the evolution of reproductive synchrony: effects on reproductive success

Synchronous breeding in animals and plants has stimulated both a theoretical and empirical examination of the possible benefits of active synchronization. The selective pressures of predation and infanticide are the strongest candidates proposed to explain the evolution of reproductive synchrony. Alternatively, breeding asynchronously with conspecifics may ensure a greater availability of resources per breeder. However, the possible fitness benefits resulting from active asynchronization have not yet received attention in evolutionary ecology. Here we present a hypothesis, based on a graphical model, illustrating the costs and benefits of the two modes of reproduction. We tested the hypothesis empirically using a 2 x 2 full factorial study design, where reproductive synchrony and infanticide tactics were manipulated in enclosed populations of the bank vole. The results reveal a relationship between infanticide tactics and breeding synchrony as illustrated by our hypothesis. In general, female reproductive success (number and size of offspring surviving to weaning) was significantly lower in infanticidal populations. Moreover, an asynchronous breeding pattern proved to be advantageous in the noninfanticidal population but this advantage of asynchrony was lost as infanticide became common in the population. Our findings support the idea that synchronous reproduction could have evolved as a counterstrategy against infanticide.     

Age-specific reproductive success: evidence for the selection hypothesis

Age-specific reproductive success has been demonstrated in many species. Three hypotheses have been raised to explain this general phenomenon: the experience hypothesis based on age-specific reproductive experience, the effort hypothesis based on age-specific reproductive effort, and the selection hypothesis based on progressive disappearance of phenotypes due to variation in individual productivity and survival. We used data from a long-term study of Leach's storm-petrels (Oceanodroma leucorhoa) to present a single test of mutually exclusive predictions about the relationship between early breeding success and longevity. There should be no correlation between early breeding success and longevity under the experience hypothesis, a negative correlation under the effort hypothesis, and a positive correlation under the selection hypothesis. We found a significant (P < 0.0001) positive relationship between success in the first two breeding attempts and longevity in this population of long-lived seabirds, strongly suggesting that low-productivity parents were also less likely to survive early breeding. These data provide some of the strongest support to date for the selection hypothesis.     

Experimental manipulation of breeding density and litter size: effects on reproductive success in the bank vole

1. Reproductive success of individual females may be determined by density-dependent effects, especially in species where territory provides the resources for a reproducing female and territory size is inversely density-dependent.
2. We manipulated simultaneously the reproductive effort (litter size manipulation: ± 0 and + 2 pups) and breeding density (low and high) of nursing female bank voles Clethrionomys glareolus in outdoor enclosures. We studied whether the reproductive success (number and quality of offspring) of individual females is density-dependent, and whether females can compensate for increased reproductive effort when not limited by saturated breeding density.
3. The females nursing their young in the low density weaned significantly more offspring than females in the high density, independent of litter manipulation.
4. Litter enlargements did not increase the number of weanlings per female, but offspring from enlarged litters had lower weight than control litters.
5. In the reduced density females increased the size of their home range, but litter manipulation had no significant effect on spacing behaviour of females. Increased home range size did not result in heavier weanlings.
6. Mother's failure to successfully wean any offspring was more common in the high density treatment, whereas litter manipulation or mother's weight did not affect weaning success.
7. We conclude that reproductive success of bank vole females is negatively density-dependent in terms of number, but not in the quality of weanlings.
8. The nursing effort of females (i.e. the ability to provide enough food for pups) seems not to be limited by density-dependent factors.     

Positive effects of helpers on reproductive success in the brown treecreeper and the general importance of future benefits

1. Numerous studies of cooperatively breeding species have tested for effects of helpers on reproductive success to evaluate hypotheses for the evolution of cooperation, but relatively few have used experimental or statistical approaches that control for the confounding effects of breeder and territory quality. 2. In the brown treecreeper Climacteris picumnus, most helpers are male offspring of the breeding pair that have delayed dispersal. We analysed 5 years of data (97 territory-years) using hierarchical linear modelling to test for effects of helpers on reproductive success while controlling for confounding factors. 3. The number of helpers was related positively to reproductive success even after controlling for differences between territories and breeders. A threshold effect was observed, with success increasing most with the presence of a second helper (i.e. at group size of four). 4. Feeding at the nest was one mechanism responsible for this effect, as larger groups had higher total feeding rates at all nesting stages. Higher total feeding rates, as well as higher feeding rates by helpers, were correlated in turn with greater reproductive success. 5. An analysis of the effects of helper feeding rate on reproductive success in groups with just one helper produced only weak support for a positive effect of helpers. Controlled comparisons of this kind utilize only a small fraction of the total data available and thus have limited statistical power compared to hierarchical or mixed-modelling. 6. A number of hypotheses to explain the evolution and maintenance of helping behaviour are consistent with our results for brown treecreepers including kin selection and hypotheses based on future direct benefits. 7. A previous synthesis of studies of helper effects that controlled for confounding factors suggested a pattern in which male helpers rarely have positive effects on reproductive success. However, revising that synthesis to include recent hierarchical or mixed-modelling studies suggests that helpers of both sexes usually have positive effects, and that the relative importance of future direct benefits may have been underestimated.     

Seasonal reproductive success of the Moorhen Gallinula chloropus: the importance of male weight

During a three-year study of 82 pairs of Moorhens Gallinula chloropus, seasonal reproductive success (SRS) varied greatly between pairs. Pairs on big territories had a higher SRS than pairs on small territories, probably because clutch survival was higher on big territories. Pairs that began breeding early had a higher SRS than pairs that began late because early breeders laid larger clutches and could attempt more broods. Pairs with heavy males had bigger territories and began breeding earlier than pairs with light males. Heavy males therefore had a higher SRS than light males. Female weight did not affect territory size, timing of breeding, or female SRS. Assuming SRS to be a good predictor of lifetime reproductive success, selection should favour heavy males but not heavy females. Male Moorhens are significantly heavier than female Moorhens. Finally, pairs with breeding experience had a higher SRS than pairs without, because they had bigger territories and began breeding earlier. However, males in inexperienced pairs were not lighter than males in experienced pairs, so it is not clear why they defended smaller territories or laid later.     

Age-specific variation in survival, reproductive success and offspring quality in red squirrels: evidence of senescence

Individual performance is expected to decrease with age because of senescence. We analyzed long-term data collected on a North American red squirrel population to assess the influence of age on body mass, survival and reproductive performance, and to study the effects of sex and of environmental conditions during early life on senescence patterns. Mass of males and females did not decrease at the end of life, possibly because body mass mostly reflects overall size in income breeders such as red squirrels. On the other hand, we found evidence of senescence in survival of both sexes and, to a lesser extent, in female reproductive traits. When compared to females, males had both higher survival and delayed decrease in survival, suggesting a weaker senescence in males. The offspring survival from weaning to one year of age also decreased with increasing mother age. This suggests that older females produce juveniles of lower quality, providing evidence of an intergenerational effect of mother's age on juveniles' fitness. Finally, our results indicate that variations in food conditions during early life influenced the reproductive tactics of females in the first years of their life, but not senescence patterns.     

Variable nursery pollinator importance and its effect on plant reproductive success

Nursery pollination, in which insects use as hosts the very plants they pollinate, ranges from obligate mutualism to parasitism. In the non-obligate interaction between Greya moths and the host Lithophragma sp., the relative density of nursery pollinators and copollinators, which do not use plant tissues for larval development, is a key determinant of the interaction’s outcome. Silene (Caryophyllaceae) nursery pollination by Hadena moths (Lepidoptera: Noctuidae), studied primarily in Europe, is considered antagonistic because copollinators comprise a substantial proportion of the pollinator community. However, there are few studies that ascertain the direction of the Silene–Hadena interaction by taking into account both pollinator service and seed predation. Here, we report a novel comprehensive evaluation of the direction of the interaction between North American Hadena ectypa on Silene stellata, by comparing the relative contributions of nursery and copollinators to S. stellata pollination and relate this to variation in fruit predation and reproductive success of S. stellata across multiple sites and years. Hadena ectypa pollinator importance (pollen deposited/visit/h) varied between years, resulting from variable visitation rate. Copollinator importance was higher than H. ectypa in 1 year and equivalent in another. In two of three sites, lowered H. ectypa activity was not correlated with a significant decrease in plant reproductive success, indicating a negative interaction. Although pollinator service by H. ectypa is substantial in this system, copollinators’ service is at least as great, and when the cost of fruit predation is factored in, the net effect of the interaction is parasitism of host plants.     

Some effects of mastectomy on reproductive success in the guinea-pig

Virgin guinea-pigs were mastectomized in two stages between 11 and 18 weeks of age and then mated, starting 19 weeks after final surgery. In the subsequent first pregnancy, the incidence of still-births and neonatal deaths was significantly higher in the mastectomized animals (6 out of 12 mothers (50%) and 14 out of 49 young (29%) compared with intact guinea-pigs (1 out of 15 mothers (7%) and 1 out of 58 young (2%)). There was no significant effect of mastectomy on litter size and weight or on gestation period. The still-born were not significantly different in weight from those born alive. A significant relation was found between maternal weight changes in the period 20 to 5 days before parturition and the occurrence of still-births and neonatal deaths; still-births were associated with a period of reduced weight gain. No effect of mastectomy on the length of the oestrous cycle was apparent but a significant increase in the incidence of non-pregnancy was found. The results provide further evidence that mastectomy influences reproductive success in the guinea-pig and suggest that parturition is a key process affected.     

Inbreeding effects on fertility in humans: evidence for reproductive compensation.

The effects of inbreeding on prereproductive mortality have been demonstrated in many natural populations, including humans. However, little is known about the effects in inbred individuals who survive to adulthood. We have investigated the effects of inbreeding on fertility among inbred adult Hutterites and demonstrate significantly reduced fecundity among the most inbred Hutterite women, as evidenced by longer interbirth intervals (P=.024) and longer intervals to a recognized pregnancy (P=.010) but not by increased rates of fetal loss (P>.50). These data suggest the presence of recessive alleles that adversely affect fecundity among the population. In contrast, completed family sizes do not differ among the more and the less-inbred Hutterite women who were born after 1920, suggesting that reproductive compensation is occurring among the more-inbred and less-fecund women. This recent reproductive strategy would facilitate the maintenance of recessive alleles and contribute to an overall decline in fertility in the population.     

Human longevity at the cost of reproductive success: evidence from global data

A trade‐off between reproduction and somatic maintenance and hence survival is fundamental to life‐history theory. We investigated the relationship between female fecundity and longevity in Homo sapiens using data from 153 countries located all over the world. The raw correlation between life span and fecundity was highly significant with a negative trend. After longevity and fecundity estimates were controlled for by confounding factors such as historical (i.e. human ethnic groups), religious, geographical, socio‐economical and parasitological components, we still observed a negative relationship between the mean female fecundity and the mean longevity in a country. These findings support the hypothesis for the existence of a trade‐off between these two key life‐history traits in humans, as also reported by a recent single longitudinal study in England.     

The effects of introduced plants on songbird reproductive success

The effects of non-native plants on habitat use are well studied; however, whether introduced plants negatively influence reproductive output of animal populations has received much less attention. We conducted a systematic literature review to evaluate the influence of non-native plants on reproductive performance in songbirds. Our global search resulted in 32 relevant articles, from which we compiled a dataset of 133 songbird responses to nesting in or around non-native vegetation. Reproductive metrics examined included measures of nest survival/mortality, productivity, fledgling survival, adult survival, nestling condition, and brood parasitism. Thirty-five percent of songbird reproductive responses were negative (n = 47), with 31% positive (n = 41) and 34% neutral (n = 45) responses found. Only 15% of responses were statistically significant effects (n = 20), and of these, negative effects were reported three times as often as positive effects. Non-significant trends were more prevalent (51% of responses), and the frequency of negative and positive trends was similar. The probability of finding a negative response (significant effect or non-significant trend) was higher for birds using introduced shrubs and wetland habitats. Mechanisms underlying responses were diverse, though similar drivers, such as differences in vegetation characteristics, predation pressure, and resource availability, were offered to explain both positive and negative effects. We found evidence for non-native plants as ecological traps in 39% of articles that assessed these phenomena (n = 16). This review highlights the sparsity of research targeting reproductive responses to plant invasion and synthesizes existing information to enhance our understanding of how birds respond to non-native plants. Our findings could be used to inform future research priorities in a world increasingly dominated by novel ecosystems.     

Mechanisms for delayed density-dependent reproductive traits in field voles, Microtus agrestis: the importance of inherited environmental effects

Reproductive traits of voles vary with the phases of the population density fluctuations. We sought to determine whether the source of this variation resides in the individuals or in their environment. Overwintering field voles ( Microtus agrestis ) from two cyclic out-of-phase populations (increase and peak phases) were sampled in early spring and bred in the laboratory for two generations under standardised conditions with ambient light and temperature. Monitoring of the source populations by capture-mark-recapture showed large differences in reproductive performance. In the increase area, reproduction started six weeks earlier, the probability of maturation of young-of-the-year was more than ten times higher during mid-summer, and reproduction continued nearly two months later in the autumn than in the peak area. These differences were not found to be associated with a difference in age structure of overwintered animals between the two areas (assessed by the distribution of eye lens masses from autopsy samples). Although the population differences in reproductive traits were to some degree also present among the overwintered animals in the laboratory, we found no difference in reproductive traits in the laboratory-born generations. There was a strongly declining seasonal trend in probability of sexual maturation both in the field and in the laboratory under ambient light conditions. However, in the field there were large population differences in the steepness of the seasonal decline that were not seen under the standardised laboratory conditions. We conclude that seasonal decline in maturation rates is governed by change in photoperiod, but that the population level variation in the shape of the decline is caused by a direct response to the environment and not due to variation in any intrinsic state of the individuals reflecting the environment experienced by the previous generation(s).     

Assessing the contributions of multiple interacting traits to plant reproductive success: environmental dependence

The reproductive success of sibling cocklebur plants (Compositae: Xanthium strumarium) was monitored after growth at different levels of availability of water and nutrient resources. Variation in reproductive success among individual plants was related to physiological, structural, and phenological characteristics. Reproductive success increased with increased availability of resources, but the relative contribution of particular traits to reproductive success varied with resource availability. Allocation of biomass to different vegetative tissues, time to seedling emergence, degree of branching, transpiration rates, water use efficiency, the rate of decline in height growth after seedling emergence and final plant size all varied significantly with resource availability. However, the changes in each of these phenotypic traits across three garden environments did not always correlate with reproductive success. The shifts across environments in the apparent importance of somatic traits for reproductive success were attributed to plastic changes in the traits but also to changes in the phenotypic correlations of the traits with reproductive success.     

Maternal and environmental influences on reproductive success of a viviparous grassland lizard

Understanding the factors that drive population persistence and growth is fundamental to both conservation management and evolutionary biology. Internal (maternal) and external (environmental) factors can affect female reproductive output, and in oviparous reptiles both may strongly influence offspring phenotype and quality. However, the link between these effects, their importance for reproductive output and offspring characteristics of live-bearing lizards, and whether population declines are linked to these factors in modified versus native habitats are unknown. We used a common New Zealand skink species, Oligosoma maccanni (McCann?s skink), found in grazed native and exotic grassland to test whether differences in environmental or maternal characteristics influenced birth date, pregnancy success and offspring phenotype. In both grassland types the date of birth was c. 8 days earlier at lower altitudes (altitudinal range = 564?719 m a.s.l), and small females were less likely to have successful pregnancies. However, larger females had more weak or deformed offspring, suggesting that reproductive senescence may exist in this species. While other research shows that exotic grasslands do not support as many skinks as native tussock grassland, reproductive success of pregnant skinks (viable litter size) was not affected by habitat modification. However, neonates had greater body condition (mass for length) when from females with higher post-partum body condition, and these females were from the native tussock grasslands. In conjunction with previously published data on McCann?s skink, our data suggest that reduced offspring quality may contribute to the lower population numbers in the exotic habitats.     

Seasonal patterns in reproductive success of temperate‐breeding birds: Experimental tests of the date and quality hypotheses

For organisms in seasonal environments, individuals that breed earlier in the season regularly attain higher fitness than their late‐breeding counterparts. Two primary hypotheses have been proposed to explain these patterns: The quality hypothesis contends that early breeders are of better phenotypic quality or breed on higher quality territories, whereas the date hypothesis predicts that seasonally declining reproductive success is a response to a seasonal deterioration in environmental quality. In birds, food availability is thought to drive deteriorating environmental conditions, but few experimental studies have demonstrated its importance while also controlling for parental quality. We tested predictions of the date hypothesis in tree swallows (Tachycineta bicolor) over two breeding seasons and in two locations within their breeding range in Canada. Nests were paired by clutch initiation date to control for parental quality, and we delayed the hatching date of one nest within each pair. Subsequently, brood sizes were manipulated to mimic changes in per capita food abundance, and we examined the effects of manipulations, as well as indices of environmental and parental quality, on nestling quality, fledging success, and return rates. Reduced reproductive success of late‐breeding individuals was causally related to a seasonal decline in environmental quality. Declining insect biomass and enlarged brood sizes resulted in nestlings that were lighter, in poorer body condition, structurally smaller, had shorter and slower growing flight feathers and were less likely to survive to fledge. Our results provide evidence for the importance of food resources in mediating seasonal declines in offspring quality and survival.