Reproduction is adapted to survival characteristics across geographically isolated medfly populations

We propose the hypothesis that individual longitudinal trajectories of fertility are closely coupled to varying survival schedules across geographically isolated populations of the same species, in such a way that peak reproduction takes place before substantial increases in mortality are observed. This reproductive adaptation hypothesis is investigated for medflies through a statistical analysis of biodemographic data that were obtained for female medflies from six geographically far apart regions. The following results support the hypothesis: (i) both survival and reproductive schedules differ substantially between these populations, where early peaks and subsequently fast declining reproduction are observed for short-lived and protracted reproductive schedules for long-lived flies; (ii) when statistically adjusting reproduction for the observed differences in survival, the differences in reproductive schedules largely vanish, and thus the observed differences in fertility across the populations can be explained by differences in population-specific longevity; and (iii) specific survival patterns of the medflies belonging to a specific population predict the individual reproductive schedule for the flies in this population. The analysis is based on innovative statistical tools from functional data analysis. Our findings are consistent with an adaptive mechanism whereby trajectories of fertility evolve in response to specific constraints inherent in the population survival schedules.     

A DEMOGRAPHIC TRANSITION ALTERED THE STRENGTH OF SELECTION FOR FITNESS AND AGE‐SPECIFIC SURVIVAL AND FERTILITY IN A 19TH CENTURY AMERICAN POPULATION

Modernization has increased longevity and decreased fertility in many human populations, but it is not well understood how or to what extent these demographic transitions have altered patterns of natural selection. I integrate individual‐based multivariate phenotypic selection approaches with evolutionary demographic methods to demonstrate how a demographic transition in 19th century female populations of Utah altered relationships between fitness and age‐specific survival and fertility. Coincident with this demographic transition, natural selection for fitness, as measured by the opportunity for selection, increased by 13% to 20% over 65 years. Proportional contributions of age‐specific survival to total selection (the complement to age‐specific fertility) diminished from approximately one third to one seventh following a marked increase in infant survival. Despite dramatic reductions in age‐specific fertility variance at all ages, the absolute magnitude of selection for fitness explained by age‐specific fertility increased by approximately 45%. I show that increases in the adaptive potential of fertility traits followed directly from decreased population growth rates. These results suggest that this demographic transition has increased the adaptive potential of the Utah population, intensified selection for reproductive traits, and de‐emphasized selection for survival‐related traits.     

Determinants of family size in rural Glacia (Spain)

The determinants of complete family size and offspring survival were assessed by means of family reconstitution data from a 19th & 20th century rural population of Northwest Spain. The reproductive performances of families were submitted to a stepwise regression analysis in which the number of children born alive and those surviving to reproductive age per family were taken as dependent variables, while 13 other variables were defined as independent. Infant mortality appears to explain most fertility variation; an earlier marital age and at first maternity leads to an increased family size. Survival seems to be the result of the apposite interaction of both fertility and mortality in early childhood.     

The interaction between reproductive lifespan and protandry in seasonal breeders

The timing and duration of reproductive activities are highly variable both at the individual and population level. Understanding how this variation evolved by natural selection is fundamental to understanding many important aspects of an organism's life history, ecology and behaviour. Here, we combine game theoretic principles governing reproductive timing and the evolutionary theory of senescence to study the interaction between protandry (the earlier arrival or emergence of males to breeding areas than females) and senescence in seasonal breeders. Our general model applies to males who are seeking to mate as frequently as possible over a relatively short period, and so is relevant to many organisms including annual insects and semelparous vertebrates. The model predicts that protandry and maximum reproductive lifespans should increase in environments characterized by high survival and by a low competitive cost of maintaining the somatic machinery necessary for survival. In relatively short seasons under these same conditions, seasonal declines in the reproductive lifespans of males of equivalent quality will be evolutionarily stable. However, over a broad range of potential values for daily survival and maintenance cost, reproductive lifespan is expected to be relatively short and constant throughout a large fraction of the season. We applied the model to sockeye (or kokanee) salmon Oncorhynchus nerka and show that pronounced seasonal declines in reproductive lifespan, a distinctive feature of semelparous Oncorhynchus spp., is likely part of a male mating strategy to maximize mating opportunities.     

Local adaptation of reproductive performance during thermal stress

Considerable evidence exists for local adaptation of critical thermal limits in ectotherms following adult temperature stress, but fewer studies have tested for local adaptation of sublethal heat stress effects across life‐history stages. In organisms with complex life cycles, such as holometabolous insects, heat stress during juvenile stages may severely impact gametogenesis, having downstream consequences on reproductive performance that may be mediated by local adaptation, although this is rarely studied. Here, we tested how exposure to either benign or heat stress temperature during juvenile and adult stages, either independently or combined, influences egg‐to‐adult viability, adult sperm motility and fertility in high‐ and low‐latitude populations of Drosophila subobscura. We found both population‐ and temperature‐specific effects on survival and sperm motility; juvenile heat stress decreased survival and subsequent sperm motility and each trait was lower in the northern population. We found an interaction between population and temperature on fertility following application of juvenile heat stress; although fertility was negatively impacted in both populations, the southern population was less affected. When the adult stage was also subject to heat stress, the southern population exhibited positive carry‐over effects whereas the northern population's fertility remained low. Thus, the northern population is more susceptible to sublethal reproductive consequences following exposure to juvenile heat stress. This may be common in other organisms with complex life cycles and current models predicting population responses to climate change, which do not take into account the impact of juvenile heat stress on reproductive performance, may be too conservative.     

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.     

Reproductive pattern in agrarian and immigrant receptor populations: a survey of El Ejido (SE Spain)

This paper deals with the fertility pattern of the El Ejido population, an agricultural Spanish community characterised by the rapid development of its modern agrarian economy. Consequently, the arrival of immigrants has sharply increased throughout the second half of the twentieth century, with important demographic consequences as well as reproductive changes. The analyses of the age-specific fertility rate (fx) and the total fertility rate (TFR) were used in order to describe the reproductive pattern of this population in 2000. The main characteristics were the following: a) Regarding the temporal change, an important decrease of fx has been observed in all age groups for the last twenty years, as a consequence of progressive birth control. However, the reproductive pattern has kept almost invariable and has been characterised by a maximum fertility at age group of 25-29 years old. b) Regarding the general Spanish fertility, the comparison of fx in both populations suggests a younger maternity in the agricultural population than in the national, the maximum fertility delayed to the 30-34 age group. c) Moreover, the El Ejido population showed a clear higher offspring per woman (TFR = 1.42) than the national (TFR = 1.24), according to the agrarian character of the El Ejido population. d) Finally, this greater reproductive level of El Ejido is also due to the arrival of women at fertile age, who come mainly from Africa, and above all from Morocco.     

Understanding contributions of cohort effects to growth rates of fluctuating populations

1. Understanding contributions of cohort effects to variation in population growth of fluctuating populations is of great interest in evolutionary biology and may be critical in contributing towards wildlife and conservation management. Cohort-specific contributions to population growth can be evaluated using age-specific matrix models and associated elasticity analyses. 2. We developed age-specific matrix models for naturally fluctuating populations of stoats Mustela erminea in New Zealand beech forests. Dynamics and productivity of stoat populations in this environment are related to the 3-5 year masting cycle of beech trees and consequent effects on the abundance of rodents. 3. The finite rate of increase (lambda) of stoat populations in New Zealand beech forests varied substantially, from 1.98 during seedfall years to 0.58 during post-seedfall years. Predicted mean growth rates for stoat populations in continuous 3-, 4- or 5-year cycles are 0.85, 1.00 and 1.13. The variation in population growth was a consequence of high reproductive success of females during seedfall years combined with low survival and fertility of females of the post-seedfall cohort. 4. Variation in population growth was consistently more sensitive to changes in survival rates both when each matrix was evaluated in isolation and when matrices were linked into cycles. Relative contributions to variation in population growth from survival and fertility, especially in 0-1-year-old stoats, also depend on the year of the cycle and the number of transitional years before a new cycle is initiated. 5. Consequently, management strategies aimed at reducing stoat populations that may be best during one phase of the beech seedfall cycle may not be the most efficient during other phases of the cycle. We suggest that management strategies based on elasticities of vital rates need to consider how population growth rates vary so as to meet appropriate economic and conservation targets.     

On the reproductive value and the spectrum of a population projection matrix with implications for dynamic population models

The eigenvalues of a population projection matrix-except for the Lotka coefficient-are uniquely determined by the reproductive values and the survival. This relation (proposed earlier, but not really well known in western literature) follows from another useful relation between fertility, reproductive values, survival, and Lotka’s coefficient. These results are applied to provide demographic interpretations to the intrinsically dynamic and metastable population models by Schoen and co-workers.     

Mass- and density-dependent reproductive success and reproductive costs in a capital breeder

For capital breeders, mass may affect reproductive potential. Reproductive expenditure may reduce future reproductive potential, particularly when resources are scarce. To test the hypothesis that reproductive success and the costs of reproduction vary according to mass and population density, we analyzed 25 yr of data on bighorn ewes (Ovis canadensis). The number of adult females was first limited by yearly removals, then allowed to triple. We found no survival costs of reproduction for ewes aged 4-7 yr. For ewes aged 8-14 yr, survival was density dependent for barren ewes but not for ewes that weaned lambs. Failure to lamb was rare and negatively correlated with fertility the following year. At low population density, lactation had a negative effect on mass gain but had a limited reproductive cost. At high density, heavy ewes had higher reproductive success than light ewes, and the reproductive cost and somatic costs of reproduction increased. The cost of reproduction was greater for light than for heavy ewes. Survival of weaned lambs to 1 yr was affected by population density but not by maternal mass or previous reproductive success. In large mammals, manipulations of reproductive effort are problematic, but long-term monitoring of individual mass and reproductive success under varying conditions of resource availability can provide insights into the evolution of life histories.     

Evolution of delayed reproduction in uncertain environments: a life-history perspective

Environmental uncertainty alone can select for delayed reproduction; however, its relative role in the evolution of delayed reproduction across life histories is not known. Along a life-history spectrum from low-survival/high-fertility species to high-survival/low-fertility species, we show that the latter are more likely to evolve delayed reproduction if fertility varies over time. By contrast, if survival varies over time, low-survival life histories are more likely to evolve delays. If there is variation in both survival and fertility, and if this variation is positively associated, the evolutionarily stable reproductive delay is decreased (relative to independent variation in survival and fertility). Conversely, if variation in survival and fertility is negatively associated, the evolutionarily stable reproductive delay is increased. We further show that environmental uncertainty can drive the evolution of delayed reproduction in an iteroparous organism but only in the special case where juvenile survival is greater than adult survival. For common iteroparous life histories (adult survival > juvenile survival), environmental uncertainty does not select for delayed reproduction. Thus, any benefits that delayed reproduction might have on reproduction or survival could be especially important in explaining the common observation of delayed reproduction in many vertebrates and perennial plants.     

Assessing seasonal demographic covariation to understand environmental‐change impacts on a hibernating mammal

Natural populations are exposed to seasonal variation in environmental factors that simultaneously affect several demographic rates (survival, development and reproduction). The resulting covariation in these rates determines population dynamics, but accounting for its numerous biotic and abiotic drivers is a significant challenge. Here, we use a factor‐analytic approach to capture partially unobserved drivers of seasonal population dynamics. We use 40 years of individual‐based demography from yellow‐bellied marmots (Marmota flaviventer) to fit and project population models that account for seasonal demographic covariation using a latent variable. We show that this latent variable, by producing positive covariation among winter demographic rates, depicts a measure of environmental quality. Simultaneously, negative responses of winter survival and reproductive‐status change to declining environmental quality result in a higher risk of population quasi‐extinction, regardless of summer demography where recruitment takes place. We demonstrate how complex environmental processes can be summarized to understand population persistence in seasonal environments.     

Did food increase fertility? Evaluation of !Kung and Herero history.

Low fertility among nomadic !Kung foragers of the northern Kalahari Desert of Botswana has been hypothesized to be an adaptation to scarcity of food. However, a comparison of !Kung fertility before and after a transition to a more sedentary lifestyle indicates that more food did not increase fertility. An examination of the fertility of neighboring sedentary Bantu-speaking Herero pastoralists during the same period also indicates that low female reproductive rates in this region are not unique to the !Kung. Herero fertility has increased dramatically in recent decades, probably in response to the control of sexually transmitted diseases in northwestern Botswana. More food appears to have substantially increased !Kung reproductive success by reducing infant and child mortality rates to levels observed among the Herero. These findings suggest that low !Kung fertility and mortality reflect contact with the Herero, who began expanding in large numbers into !Kung territory in the 1950s. This study emphasizes the need for a more rigorous comparative perspective in anthropology to understand better the significance of findings from restricted populations, and I suggest that evolutionary ecology would benefit from shifting some of its current focus on fertility to mortality.     

Birth seasonality and calf mortality in a large population of Asian elephants

In seasonal environments, many species concentrate their reproduction in the time of year most likely to maximize offspring survival. Asian elephants (Elephas maximus) inhabit regions with seasonal climate, but females can still experience 16‐week reproductive cycles throughout the year. Whether female elephants nevertheless concentrate births on periods with maximum offspring survival prospects remains unknown. We investigated the seasonal timing of births, and effects of birth month on short‐ and long‐term mortality of Asian elephants, using a unique demographic data set of 2350 semicaptive, longitudinally monitored logging elephants from Myanmar experiencing seasonal variation in both workload and environmental conditions. Our results show variation in birth rate across the year, with 41% of births occurring between December and March. This corresponds to the cool, dry period and the beginning of the hot season, and to conceptions occurring during the resting, nonlogging period between February and June. Giving birth during the peak December to March period improves offspring survival, as the odds for survival between age 1 and 5 years are 44% higher for individuals born during the high birth rate period than those conceived during working months. Our results suggest that seasonal conditions, most likely maternal workload and/or climate, limit conception rate and calf survival in this population through effects on maternal stress, estrus cycles, or access to mates. This has implications for improving the birth rate and infant survival in captive populations by limiting workload of females of reproductive age. As working populations are currently unsustainable and supplemented through the capture of wild elephants, it is imperative to the conservation of Asian elephants to understand and alleviate the effects of seasonal conditions on vital rates in the working population in order to reduce the pressure for further capture from the wild.     

Life history of a neotropical marsupial: Evaluating potential contributions of survival and reproduction to population growth rate

The relative effect of survival and reproductive rates to population growth rate is expected to be similar across species with similar life-histories. We employed a matrix population model and sensitivity and elasticity analysis to assess the absolute and relative importance of age-specific survival and fertility to population growth rate of Didelphis aurita (Didelphimorphia, Didelphidae) in a rural area of Rio de Janeiro, southeastern Brazil. The results were compared to expectations for mammals that mature early and have short generation time, such as D. aurita. Prospective analysis showed that changes in pouch young and juveniles survival would have large effects on population growth rate, relative to other vital rates, being the most critical time periods in the life cycle of D. aurita, whereas the effect of fertilities were always low. These findings do not fit to the observed pattern in mammals that mature early, where reproductive parameters have the largest relative influence on population growth rate. Although reproductive rates were characterized by a relatively small influence on population growth rate, they are still relevant because of their high variability and response to potential environmental disturbances. The first application of matrix population models to a neotropical rainforest marsupial provides information on marsupial demography and life-history strategy, increasing comprehension of this unknown group.     

Life history,resting egg formation,and hatching may explain the temporal-geographical distribution ofArtemia strains in the Mediterranean basin

Life history traits are presented for the sexual species (Artemia tunisiana) and for parthenogenetic diploid and tetraploid strainsA. parthenogenetica reared at 15 °, 24 ° and 29.5 °C.In laboratory cultures, we present evidence that the seasonal appearance of the sexualArtemia tunisiana (the dominant winter-spring population), and of two parthenogenetic populations ofArtemia (the dominant spring-summer populations) in certain Spanish saltworks is controlled by temperature through its effect on reproductive and survival traits. Minimum and maximum reproductive output and survival for the sexual and parthenogenetic populations, respectively, occurred at a typical temperature (24 °C) of the late-spring season when the sexual population is replaced by parthenogenetic forms. Furthermore, the high production and hatchability of cysts from the sexual population at low temperatures (15 °C), and of the parthenogenetic populations at middle temperature (24 °C), indicate the role of dormancy as an adaptation regulating seasonal occurrence.     

Seasonal changes in reproductive development in male spiny mice (Acomys spinosissimus) from South Africa

Seasonal reproduction is a common characteristic of many small mammals which inhabit seasonal environments in temperate regions, the sub-tropics as well as the tropics. It is important for an animal to reproduce during the most favourable time of the year to ensure the survival of the young and maximize reproductive success. In southern Africa, female spiny mice (Acomys spinosissimus) breed during the warm and wet spring and summer months, whereas the reproductive pattern of males is unknown although an opportunistic breeding pattern has been implicated. We investigated testes mass and volume, seminiferous tubule diameter, spermatogenesis and plasma testosterone concentrations in a South African population of male spiny mice on a 2-monthly basis over one year. Testes mass and volume started to increase in July/August and was high from September until December. Seminiferous tubule diameter and spermatogenesis increased during the same months. Plasma testosterone concentration was elevated from July/August to November/December. Development of the reproductive characteristics of male spiny mice was correlated with high rainfall and high ambient temperatures, but reproductive development had already started during the dry season and the coldest months. This shows that reproductive development in males may not be dependent on climatic conditions, and other factors, such as photoperiod, may trigger the onset of reproduction. The data, however, suggest that A. spinosissimus is a true seasonal breeder with reproduction confined to the spring and summer months in southern Africa.     

Temporal variation in reproductive costs and payoffs shapes the flowering strategy of a neotropical milkweed, <Emphasis Type="Italic">Asclepias curassavica</Emphasis>

A central goal of evolutionary ecology is to understand the factors that select for particular life history strategies, such as delaying reproduction. For example, environmental variation and reproductive costs to survival and growth often select for reproductive delays in semelparous and iteroparous species. In this study, we examine how variation in reproductive cost, which we define as a reduction to growth, survival, or future reproduction after a reproductive event, may select for reproductive delay in an iteroparous Neotropical milkweed with no obvious reproductive season. We analyzed demographic data collected every 3 months for 3 years from four populations of Asclepias curassavica in Monteverde, Costa Rica. We detected costs of flowering to survival and growth that varied in magnitude between our 12 transition periods without a seasonal pattern. The populations also exhibited temporal variation in reproductive payoffs measured as seedling establishment. We incorporated these reproductive costs into demographic projection models, which predicted a delayed flowering strategy only when we included temporal variation in costs and payoffs. Temporal variation in reproductive costs and payoffs is an important selective force in the evolution of delayed flowering in iteroparous species. Further, a lack of predictable seasonal pattern to reproductive costs and payoffs may contribute to the lack of seasonal reproductive patterns observed in our study species and other Neotropical species.     

Women’s work,child care,and helpers-at-the-nest in a hunter-gatherer society

Considerable research on helpers-at-the-nest demonstrates the positive effects of firstborn daughters on a mother’s reproductive success and the survival of her children compared with women who have firstborn sons. This research is largely restricted to agricultural settings. In the present study we ask: “Does ‘daughter first’ improve mothers’ reproductive success in a hunting and gathering context?” Through an analysis of 84 postreproductive women in this population we find that the sex of the first- or second-born child has no effect on a mother’s fertility or the survival of her offspring. We conclude that specific environmental and economic factors underlay the helpers-at-the-nest phenomenon.     

Senescence in a short-lived migratory bird: age-dependent morphology, migration, reproduction and parasitism

1. Senescence reflects age-dependent changes in residual reproductive value. Annual survival rates of the barn swallow Hirundo rustica L. increased from 1- to 2-year-old individuals, but decreased among 5 years old or older individuals. Estimates of age-dependent reproductive value showed a similar pattern.
2. Longitudinal data from two long-term population studies were used to test whether a number of different measures of performance (condition-dependent morphological traits, migratory performance, reproductive success, intensity of parasitism) changed among individuals when reaching old age.
3. The length of the outermost tail feathers (a secondary sexual character) decreased among old individuals, while two measures of individual developmental instability increased with age. Migratory performance decreased in old barn swallows as reflected by a delay in spring arrival at the breeding grounds. Reproductive performance measured as seasonal reproductive success decreased with age. The intensity of infestations with an haematophagous mite and a mallophagous ectoparasite increased among old barn swallows.
4. These results suggest that the condition-dependent secondary sexual character, developmental stability, and measures of migratory and reproductive performance deteriorated, and the frequency of parasitism increased among old individuals. Ageing was thus associated with a general deterioration of performance.