Density-dependent age at first reproduction in the eastern kingbird

Theory predicts that maximal fitness is obtained by individuals who begin to breed immediately upon reaching sexual maturity. However, delayed breeding occurs regularly in some taxa, and in birds and mammals is most often associated with long lifespan and/or limited access to suitable habitats. Delayed breeding is not expected among relatively short-lived species such as migratory passerine birds, but this assumption remains untested in many species. Here we quantify age at first reproduction in an eastern kingbird Tyrannus tyrannus population breeding in an ecological island, and through both observational and experimental approaches, investigate the potential causes for the high frequency of delayed breeding that occurs in this population. Nearly half of the fledged nestlings that returned to the breeding grounds did not breed in their first potential breeding season. Some non-breeders occupied territories, for at least some period, but most remained as non-territorial 'floaters'. Parentage analysis failed to show any reproductive success for female floaters, and only limited success for male floaters, indicating that floating was not a successful reproductive tactic. On the other hand, a strong negative relationship existed between population size and the proportion of young birds that bred in their first year, and non-territorial birds of both sexes quickly filled territory vacancies created by experimental adult removals. Limited breeding habitat and territorial behavior of older birds thus appear to be the main causes of delayed breeding in kingbirds. The frequency of delayed breeding in most species is unknown but of potential significance because failure to incorporate accurate estimates of age at first reproduction in population models may lead to flawed population projections.     

Hatching date influences age at first reproduction in the black-headed gull

In long-lived colonial birds, age at recruitment is an important life-history character. Variation in this parameter may reflect differences in several factors, including competitive ability and breeding strategies. Further, these differences may be due to timing of hatching (for instance through differences in competitive ability). We investigated the age of first-time breeders in relation to hatching date in a black-headed gull Larus ridibundus colony situated in central France, from 1979 to 1993. Age at first breeding was estimated for four groups of individuals (total n=550) according to their hatching date, using a recent capture-recapture methodology which allowed us to estimate recruitment rate without the limiting assumptions of methods relying on simple return rates. The age at first breeding was negatively correlated with the hatching date of individuals: individuals hatched earlier in the season started breeding at a younger age than individuals born later. Proportionally more 2-year-old late-hatched individuals were seen breeding on small peripheral colonies than young early-hatched individuals. This difference disappeared after age 3 years. These results strongly suggest that individuals hatched late in the season start to breed on peripheral colonies before recruiting to their natal colony. A difference of few weeks in hatching date has consequences which can last for several years.     

Evolutionarily stable age at first reproduction in a density-dependent model.

We develop a new model of life history evolution to investigate the evolution of age at first reproduction. Density dependence is taken into account. For a given "species", age of maturity, offspring survival, immature survival, adult survival, fecundity, immature age-classes entering in competition with adults and immature competitive ability are traits adjustable by natural selection, and constitute a particular strategy. On the contrary, the type of intraspecific competition (scramble or contest), strength of competition and inherent net reproductive rate Ro(inh) are fixed (specific) characteristics. As a consequence of fixing Ro(inh), the evolution of any trait will affect trade-offs between others. Evolutionarily stable strategies are determined numerically by using the mathematical concept of Lyapunov exponents. Altogether, we consider 960 different hypothetical "species" (i.e. different combinations of fixed traits). Corresponding ESSs are analyzed with respect to their age at first reproduction, adult survival and immature competitive ability components. They appear to be gathered in three groups. One is intuitive and characterized by a reduction of immature competitive ability and a correlation of age of maturity with adult survival; populations reach mainly equilibria. The two other groups respectively include "species" with low age of maturity but high adult survival, and "species" close to semelparity with delayed maturity; immature competitive ability may not be minimized, and populations possibly exhibit complex dynamics. In conclusion, the hypothesis that the evolution of a demographic parameter modifies trade-offs between others turns out to have important consequences. We argue that life history theory cannot ignore the source and mode-of-operation of density dependence and must regard potential short-term instability as essential.     

Life history invariants,age at maturity and the ferox trout

Summary Data on brown trout in Norway and on Arctic char in Norway, Iceland, Greenland or Canada are used to make three points about life-history invariants. First, invariants constructed from adaptive programmes of development that are conditioned on growth and mortality rates are more complicated than those constructed solely on the basis of dimensional analysis, but are more consistent with the data. Second, if one allows the possibility of escaping a size ceiling (e.g. by switching from planktivory to piscivory) ferox trout — brown trout that are exceptionally large and long lived — are predicted with the theory only if mortality is size dependent and the growth rate on food sources with different asymptotic sizes differs. Third, a successful empirical approach for finding life-history invariants can be more fully understood as the result of adaptive programmes of development.     

Variation in age at first reproduction of male Japanese fluvial sculpin induced by the timing of parental reproduction

The relationships between age and size at reproduction and lifetime reproductive output of male Japanese fluvial sculpin Cottus pollux were estimated by a mark-recapture study. Although all males were physiologically capable of breeding at age 2 years, age at first successful reproduction varied amongst individuals. Males with delayed reproduction had lower net reproductive rate than males that bred at age 2 years on average suggesting that age at first reproduction was a conditional strategy. Males that delayed reproduction were significantly smaller at age 1 and 2 years than males that bred at age 2 years. Despite no significant difference in body size of hatched yolk-sac larvae between the early and late phase of the breeding season, by May of the first year of life, progeny from nests in the early phase had hatched earlier and were larger than those from the nests in the late phase. The results suggested an important effect of timing of reproduction of parents on the growth and subsequent age at first reproduction of their progeny.     

Sociality, age at first reproduction and senescence: comparative analyses of birds

Evolutionary theories of senescence suggest that aging evolves as a consequence of early reproduction imposing later viability costs, or as a consequence of weak selection against mutations that act late in life. In addition, highly social species that live in sites that are protected from extrinsic mortality due to predation should senesce at a slower rate than solitary species. Therefore, species that start reproducing late in life should senesce at a slower rate than species that start reproducing early. In addition, social species should senesce more slowly than solitary species. Here I investigate the rate of senescence using an extensive data set on longevity records under natural field conditions to test predictions about the evolution of senescence among 271 species of birds. Longevity records increased with sampling effort and body mass, but once these confounding variables were controlled statistically, there was a strongly positive relationship between relative longevity and relative adult survival rate. Relative longevity after controlling statistically for sampling effort, body mass and adult survival rate, increased with age at first reproduction, but not with degree of breeding sociality. These findings suggest that the evolution of senescence is related to timing of first reproduction, but that the evolution of breeding sociality has played a negligible role in the evolution of senescence.     

Selection on age at first and at last reproduction in the long‐lived Alpine Swift Apus melba

The way an organism spreads its reproduction over time is defined as a life‐history trait, and selection is expected to favour life‐history traits associated with the highest fitness return. We use a long‐term dataset of 277 life histories to investigate the shape and strength of selection acting on the age at first reproduction and at last reproduction in the long‐lived Alpine Swift. Both traits were under strong directional selection, but in opposite directions, with selection favouring birds starting their reproductive career early and being able to reproduce for longer. There was also evidence for stabilising selection acting on both traits, suggesting that individuals should nonetheless refrain from reproducing in their first 2 years of life (i.e. when inexperienced), and that reproducing after 7 years of age had little effect on lifetime fitness, probably due to senescence.     

Individual heterogeneity in life-history trade-offs with age at first reproduction in capital breeding elephant seals

Recruitment age plays a key role in life-history evolution. Because individuals allocate limited resources among competing life-history functions, theory predicts trade-offs between current reproduction and future growth, survival and/or reproduction. Reproductive costs tend to vary with recruitment age, but may also be overridden by fixed individual differences leading to persistent demographic heterogeneity and positive covariation among demographic traits at the population level. We tested for evidence of intra- and inter-generational trade-offs and individual heterogeneity relating to age at first reproduction using three decades of detailed individual life-history data of 6,439 capital breeding female southern elephant seals. Contrary to the predictions from trade-off hypotheses, we found that recruitment at an early age was associated with higher population level survival and subsequent breeding probabilities. Nonetheless, a survival cost of first reproduction was evident at the population level, as first-time breeders always had lower survival probabilities than prebreeders and experienced breeders of the same age. However, models accounting for hidden persistent demographic heterogeneity revealed that the trade-off between first reproduction and survival was only expressed in “low quality” individuals, comprising 35% of the population. The short-term somatic costs associated with breeding at an early age had no effect on the ability of females to allocate resources to offspring in the next breeding season. Our results provide strong evidence for individual heterogeneity in the life-history trajectories of female elephant seals. By explicitly modeling hidden persistent demographic heterogeneity we show that individual heterogeneity governs the expression of trade-offs with first reproduction in elephant seals.     

Postmetamorphic growth,survival, and age at first reproduction of the salamander,hynobius nebulosus tokyoensis Tago in relation to a consideration on the optimal timing of first reproduction

The postmetamorphic growth and survival of the salamander Hynobius nebulosus tokyoentisTago were surveyed in the study site located in Habu village of Hinodemachi, a suburb of Tokyo City, during 1975–1981. A laboratory experiment on the growth rate of juveniles was conducted in parallel with the field survey. The result indicated that this salamander grew at the rate of 8,mm in s.v.l. per year during the juvenile stage, but its growth rate decreased markedly as low as 1.8 mm for males and 1.1 mm for females, once it had attained sexual maturity. According to the “capture-recapture” procedure the annual survival rate after metamorphosis was found to be quite high; that is, approximately 0.7. By using the growth rate of juveniles and the difference between the sizes at metamorphosis and sexual maturity, the age at first reproduction was estimated to be 4 year for males and 5 year for females. From the data obtained in this study, the intrinsic rates of increase (r) were calculated for various values of age at first reproduction under different survival schedules, and the relationship between the age at first reproduction and fitness as measured by r was examined. The result indicated that an optimal age maximizing fitness always existed under respective survival schedules, and the observed age at first reproduction of this salamandei was found to coincide well with the predicted optimal age.     

Life history of the moose Alces alces: relationship between growth and reproduction

We studied the relationship between increase in body weight and reproductive performance in different populations of Norwegian moose to evaluate costs associated with early onset of reproduction, viz. whether early onset of reproduction was correlated with low adult body weight or reduced adult fecundity. The mean carcass weight of non-ovulating yearlings was significantly lower than for ovulating yearlings. However, those 2.5 yr old females that conceived as yearlings were lighter than non-reproducing females of the same age. Thus, to begin to reproduce as a yearling was assumed to be expensive because it reduces the possibilities for further growth. The cost associated with reproduction was further illustrated by the fact that the difference in mean carcass weight from age 2.5 to 3.5 yr of females that produced calves in both years, was less for the females from regions with lowest mean yearling weights, i.e. regions with probably the lowest resource availability. In populations with high mean yearling carcass weights, the proportion of cows with calf and the number of calves per pregnant female in the early reproductive phase (2.5 or 3.5 yr old) were higher than in populations in which the mean yearling weights were low. There was a negative correlation between growth rate in the population after 1.5 yr of age and the mean yearling carcass weight. Thus, low yearling weight was associated with a prolonged period of growth and thereby a reduced reproductive output during the first year of the female's life. For old females (≥ 9.5 yr old) the number of calves produced per pregnant female was highest in populations where yearling carcass weights were highest. Furthermore, mean yearling weight and the mean adult female weight were positively correlated in those regions. This relationship suggests that within this species early onset of reproduction is not related to retarded reproduction or lower weight later in life. We suggest that the moose has been selected for an early onset of reproduction.     

Factors affecting reproduction in rehabilitant female orangutans: young age at first birth and short inter-birth interval

This study investigated the reproductive parameters of free-ranging rehabilitant female orangutans. We aimed to assess the factors that influence these parameters and provide information that could assist with the management of orangutan reintroduction programs. We analyzed the birth records of free-ranging female rehabilitants at Bukit Lawang, Bukit Tigapuluh, Sepilok, Camp Leakey, Kaja Island, Sungai Wain, and Meratus and compared them with reproductive parameters reported in wild and zoo populations. Females’ ages at first birth were 10.6–14.7 years, significantly earlier than those of wild and zoo orangutans. Computed inter-birth intervals (IBIs) calculated by the Kaplan–Meier method were 65.1–90.1 months; the values for Camp Leakey and Bukit Lawang rehabilitants were significantly shorter than those reported for wild Sumatran orangutans. Infant mortality rates were 18–61%; the values for Bukit Lawang and Sepilok were significantly higher than those reported for wild Sumatran and zoo orangutans. In rehabilitants, young ages at first birth and shorter IBIs may result from the high energy intake enabled by provisioning, although the possibility exists that they reflect underestimations of age on arrival at rehabilitation centers. The observed high infant mortality rate may reflect poor mothering skills due to human rearing and/or increased disease transmission. This study demonstrates that accelerated reproductive rates (younger age at first birth and shorter IBI) are common in provisioned rehabilitant females on both Sumatra and Borneo.     

Maternal risk of breeding failure remained low throughout the demographic transitions in fertility and age at first reproduction in Finland

Radical declines in fertility and postponement of first reproduction during the recent human demographic transitions have posed a challenge to interpreting human behaviour in evolutionary terms. This challenge has stemmed from insufficient evolutionary insight into individual reproductive decision-making and the rarity of datasets recording individual long-term reproductive success throughout the transitions. We use such data from about 2,000 Finnish mothers (first births: 1880s to 1970s) to show that changes in the maternal risk of breeding failure (no offspring raised to adulthood) underlay shifts in both fertility and first reproduction. With steady improvements in offspring survival, the expected fertility required to satisfy a low risk of breeding failure became lower and observed maternal fertility subsequently declined through an earlier age at last reproduction. Postponement of the age at first reproduction began when this risk approximated zero-even for mothers starting reproduction late. Interestingly, despite vastly differing fertility rates at different stages of the transitions, the number of offspring successfully raised to breeding per mother remained relatively constant over the period. Our results stress the importance of assessing the long-term success of reproductive strategies by including measures of offspring quality and suggest that avoidance of breeding failure may explain several key features of recent life-history shifts in industrialized societies.     

Trade-off between age of first reproduction and survival in a female primate

Trade-offs are central to life-history theory but difficult to document. Patterns of phenotypic and genetic correlations in rhesus macaques, Macaca mulatta—a long-lived, slow-reproducing primate—are used to test for a trade-off between female age of first reproduction and adult survival. A strong positive genetic correlation indicates that female macaques suffer reduced adult survival when they mature relatively early and implies primate senescence can be explained, in part, by antagonistic pleiotropy. Contrasts with a similar human study implicate the extension of parental effects to later ages as a potential mechanism for circumventing female life-history trade-offs in human evolution.     

Life history and age at maturity of an anadromous smelt, the eulachon Thaleichthys pacificus (Richardson)

Trace element and fork length ( L _F) frequency analyses of eulachon Thaleichthys pacificus otoliths were used to determine age at maturity and repeat spawning potential, two aspects of eulachon life history that are not known but are important for successful management of this species. The L _F-frequency analysis for ocean caught and spawning eulachon was used to estimate age at maturation. Two size classes of eulachon were caught in the ocean and spawning eulachon were consistently the largest fish indicating that spawners from mid-coast of British Columbia were 3 years old. Laser-ablation inductively-coupled plasma mass spectrometry (LA-ICP-MS) was also used to reconstruct the Ba:Ca and Sr:Ca molar ratios deposited spatially into the otolith to estimate spawner age for five populations of eulachon. Age at maturation differed among populations examined. Based on the seasonal fluctuations in Ba:Ca molar ratios caused by seasonal upwelling of deep waters, it was determined that more southerly populations spawned at a younger age than the northern populations examined. Southern populations of eulachon, Columbia River, Washington, U.S.A., spawn after 2 years. Eulachon from the Fraser, Kemano and Skeena Rivers in British Columbia, Canada, generally mature after 3 years. Some Skeena River eulachon and most of the eulachon from the Copper River, Alaska, U.S.A., matured after 4 years. In contrast to the Ba:Ca molar ratios in the otolith, Sr:Ca molar ratios maintained a relatively flat profile over the life of the eulachon. The lack of a change in Sr:Ca ratios within the otolith, the single size class of spawners across all systems and the single age class within most populations strongly suggest that eulachon in the present study are semelparous.     

Determination of age, longevity and age at reproduction of the frogMicrohyla ornata by skeletochronology

Skeletochronological estimation of age, longevity, age at sexual maturity and breeding of Microhyla ornata was done. Frogs (n=62) were collected locally in August (rainy season) 1997 and brought to the laboratory. Body mass and snout-vent-length (SVL) of each frog was recorded; the 4th toe of both the hind limbs was clipped under anaesthesia, fixed in 10% formalin, demineralized in 5% nitric acid and processed for histology. Limb bones (femur, humerus, tibiofibula and radioulna) of 6 large sized frogs were also processed for skeletochronology in order to study the rate of resorption. Gonads of 25 frogs (belonging to different body size ranges) were processed for histology in order to ascertain the gametogenic status of individual frogs. One to four growth rings consisting of growth zones and lines of arrested growth (LAGs) were noticed in frogs of different body sizes; the number of LAGs remained identical in all the limb bones and phalanges in 5 out of 6 frogs. Back calculation indicated that the resorption rate is very low in this frog. Male frogs possessed sperm bundles in seminiferous tubules in the 1st year, while females showed yolky follicles in the ovary in the 2nd year. Frogs found in amplexus were 3 5 years old. The results suggest that this frog may live for a maximum of 5 years in the natural population.     

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.     

Life history implications of allocation to growth versus reproduction in dynamic energy budgets

We compare the implications of determinate vs. indeterminate growth of a parthenogenetic iteroparous ectotherm at constant food density in the context of the dynamic energy budget theory, which specifies the tight links between life history traits, such as feeding, aging, growth and reproduction. We do a comparative analysis using, as measure of fitness, the life span reproduction, the population growth rate, and the conversion efficiency of food to biomass. When extrinsic mortality is constant, indeterminate growth cannot maximize fitness if measured by the population growth rate or the conversion efficiency, except when mortality is low, in which case both types of animals are similar. If the fitness measure is life span reproduction, indeterminate growth maximizes fitness even with constant mortality, provided it is not very high. When mortality decreases with size, indeterminate growth maximizes fitness for almost all measures of fitness. Finally, we suggest an evolutionary link between allocation strategies and expected life span. In populations of long living species, each type of animal can establish in the population of the other. In populations of short living species, determinate growers can invade, and displace, a population of indeterminate ones. However, when the mortality risk of organisms with small size is much higher than those of large size, indeterminate growers can be superior.