Free housing for declining populations: Optimizing the provision of artificial breeding structures

The improvement of habitat quality and reproductive success through the implementation of artificial breeding structures is one of the most widespread in situ conservation strategies applied to the recovery of declining wildlife populations. Several past studies have monitored the use of artificial breeding structures in the wild, but virtually none of them have investigated which demographic and environmental factors actually determine their effectiveness in facilitating reproductive success. Therefore, the aim of the present study was to identify those factors influencing breeding success in artificial structures. With this purpose we surveyed a declining population of a keystone species of Mediterranean ecosystems, namely the European wild rabbit. In Doñana National Park, we sampled during the course of two years the breeding success of wild rabbits from 47 artificial warrens, experimentally provided under different demographic and environmental conditions. In order to determine the relative importance of such factors, we applied an AIC model selection procedure to alternative biological hypothesis groups, postulating that breeding success in artificial structures would primarily depend on population density, food availability and/or predator limitation. We found that the efficiency of artificial warrens in facilitating rabbit reproductive success can vary, with an increase associated with greater pasture availability. Predator limitation by fencing also positively affected rabbit reproduction, but only in combination with high pasture availability. On their own, fencing treatment and population reinforcements had low and null effectiveness facilitating reproduction, respectively, as well as entailing significant economic and biological costs. The best way to improve breeding success in artificial structures, therefore, was through their strategic placement in potential high-quality habitats, i.e. with suitable pasture availability. We suggest that vegetation management can effectively increase carrying capacity and consequently the target species’ reproductive success, especially in fast breeding species like the European wild rabbit. The provision of artificial breeding structures for conservation purposes may significantly benefit from the elaboration of concurrent experimental studies similar to the present research, which can help establish useful guidelines for optimizing future conservation efforts.     

Dying like rabbits: general determinants of spatio-temporal variability in survival

1. Identifying general patterns of how and why survival rates vary across space and time is necessary to truly understand population dynamics of a species. However, this is not an easy task given the complexity and interactions of processes involved, and the interpopulation differences in main survival determinants. 2. Here, using European rabbits (Oryctolagus cuniculus) as a model and information from local studies, we investigated whether we could make inferences about trends and drivers of survival of a species that are generalizable to large spatio-temporal scales. To do this, we first focused on overall survival and then examined cause-specific mortalities, mainly predation and diseases, which may lead to those patterns. 3. Our results show that within the large-scale variability in rabbit survival, there exist general patterns that are explained by the integration of factors previously known to be important at the local level (i.e. age, climate, diseases, predation or density dependence). We found that both inter- and intrastudy survival rates increased in magnitude and decreased in variability as rabbits grow old, although this tendency was less pronounced in populations with epidemic diseases. Some causes leading to these higher mortalities in young rabbits could be the stronger effect of rainfall at those ages, as well as, other death sources like malnutrition or infanticide. 4. Predation is also greater for newborns and juveniles, especially in population without diseases. Apart from the effect of diseases, predation patterns also depended on factors, such as, density, season, and type and density of predators. Finally, we observed that infectious diseases also showed general relationships with climate, breeding (i.e. new susceptible rabbits) and age, although the association type varied between myxomatosis and rabbit haemorrhagic disease. 5. In conclusion, large-scale patterns of spatio-temporal variability in rabbit survival emerge from the combination of different factors that interrelate both directly and through density dependence. This highlights the importance of performing more comprehensive studies to reveal combined effects and complex relationships that help us to better understand the mechanisms underlying population dynamics.     

Home range of the European rabbit (Oryctolagus cuniculus) in three contrasting French populations

Despite their pest status in numerous areas throughout the World, the populations of European rabbits (Oryctolagus cuniculus) have strongly decreased in South Western Europe since the mid-20th century. Such a decrease constitutes a major threat on top predators and calls for a better understanding of its mechanisms to provide suitable management responses. Infectious diseases have been invoked as the main responsible factors, but they cannot by themselves explain the magnitude of this decrease. Habitat fragmentation may indeed act as a synergetic factor, and habitat use studies are needed to better understand the impact of fragmentation on rabbit population dynamics. We investigated the variability of home range size with respects to age, sex and season in three wild populations of rabbits using telemetry. Home ranges were smaller in the highest density populations (7333 and 6878 vs. 20,492 m²) suggesting differences in habitat quality between the populations. In addition, home range sizes were larger during the reproductive season for both sexes, and adults tended to have smaller home ranges than juveniles. Clearly, the home range sizes reported here were smaller than those previously reported in rabbits.     

The Edible Dormouse (<Emphasis Type="Italic">Glis glis</Emphasis>, Gliridae,Rodentia) in the Periphery of Its Distribution Range: Body Size and Life History Parameters

The body size, longevity, growth rate, and the intensity of breeding in a peripheral population of the edible dormouse from the Zhiguli Mountains are analyzed from the standpoint of life cycle theory. A breeding peculiarity of the species lies in periodically repeated years of reproductive failure, i.e., the total absence of young, seen throughout the species range. In reproductively successful years, anticipatory reproduction is observed, when the birth of posterity precedes a period of maximum food abundance. In the optimum of the distribution range, in Central and Western Europe, the mechanism of reproductive control in unfavorable years is based on the suppressing of the male reproductive activity at the beginning of the active season. In the Zhiguli population, natality regulation is rooted in mass resorption of embryos in the vast majority of females. In that population, the body size and weight are the minimum, while the difference from individuals from other populations studied may reach three times. The peculiarities of the edible dormouse biology in the peripheral population are related to the life cycle parameters. In contrast to the Central European populations, reproduction in the Zhiguli occurs every year, but in pregnant females, the death of embryos is observed in lean years. In males of that population, which are reproductively active every year, no significant costs of reproduction related to weight loss have been identified. The lower growth rate in comparison with more western European populations is shown in four age groups conventionally distinguishable based on morphometric parameters. In yearling males whose growth is not yet finished, the onset of the mating period depends on the body mass. Typically, the lifespan in the study population does not exceed four years, this being much shorter than in other populations where it lasts seven to nine years.     

Climate variation and regional gradients in population dynamics of two hole-nesting passerines

Latitudinal gradients in population dynamics can arise through regional variation in the deterministic components of the population dynamics and the stochastic factors. Here, we demonstrate an increase with latitude in the contribution of a large-scale climate pattern, the North Atlantic Oscillation (NAO), to the fluctuations in size of populations of two European hole-nesting passerine species. However, this influence of climate induced different latitudinal gradients in the population dynamics of the two species. In the great tit the proportion of the variability in the population fluctuations explained by the NAO increased with latitude, showing a larger impact of climate on the population fluctuations of this species at higher latitudes. In contrast, no latitudinal gradient was found in the relative contribution of climate to the variability of the pied flycatcher populations because the total environmental stochasticity increased with latitude. This shows that the population ecological consequences of an expected climate change will depend on how climate affects the environmental stochasticity in the population process. In both species, the effects will be larger in those parts of Europe where large changes in climate are expected.     

Sex in space: population dynamic consequences

Sex, so important in the reproduction of bigametic species, is nonetheless often ignored in explorations of the dynamics of populations. Using a growth model of dispersal-coupled populations we can keep track of fluctuations in numbers of females and males. The sexes may differ from each other in their ability to disperse and their sensitivity to population density. As a further complication, the breeding system is either monogamous or polygamous. We use the harmonic mean birth function to account for sex-ratio-dependent population growth in a Moran–Ricker population renewal process. Incorporating the spatial dimension stabilizes the dynamics of populations with monogamy as the breeding system, but does not stabilize the population dynamics of polygamous species. Most notably, in populations coupled with dispersal, where the sexes differ in their dispersal ability there are rarely stable and equal sex ratios. Rather, a two-point cycle, four-point cycle and eventually complex behaviour of sex-ratio dynamics will emerge with increasing birth rates. Monogamy often leads to less noisy sex-ratio dynamics than polygamy. In our model, the sex-ratio dynamics of coupled populations differ from those of an isolated population system, where a stable 50:50 sex ratio is achievable with equal density-dependence costs for females and males. When sexes match in their dispersal ability, population dynamics and sex-ratio dynamics of coupled populations collapse to those of isolated populations.     

Effects of climate on reproduction in the European wild rabbit (Oryctolugus cuniculus)

Reproduction in a population of European wild rabbits occupying parkland habitat in East Anglia showed an annual seasonality in terms of numbers of emergent young, male testes size and an index of female oestrous status. Weather conditions showed marked variation over the four-year study period with significantly higher minimum ground temperatures and hours of daily sunshine, and less snowfall recorded during the second two years. Parailel shifts in reproductive seasonality were found with male testes size peaking in January and February as opposed to May and April, first female oestrus recorded one month earlier (January compared to February), and young emerging over an average of 5.5 instead of 4.5 months each year and their first appearance above ground advanced by two months from April to February. Mean annual productivity increased from 7.2 and 6.1 emergent young per female, to 10.1 and 9.1 over the four years, while the number of adults present in the population at the start of the breeding season increased from 22 to 45, with females always outnumbering males. The usefulness is demonstrated of a noninvasive index of vulva appearance, and morphometric measures of testes to determine female oestrous status and estimates of male testicular weight in wild populations.
The reported changes in reproduction are probably mediated through the influence of climatic variables on the quality or quantity of food available to the rabbits. These data demonstrate the possible effects of predicted global warming on productivity in the rabbit and highlight the major influence of climate on reproductive seasonality in this important pest species.     

Contrasting Effects of Climate Change on Rabbit Populations through Reproduction

Background

Climate change is affecting many physical and biological processes worldwide. Anticipating its effects at the level of populations and species is imperative, especially for organisms of conservation or management concern. Previous studies have focused on estimating future species distributions and extinction probabilities directly from current climatic conditions within their geographical ranges. However, relationships between climate and population parameters may be so complex that to make these high-level predictions we need first to understand the underlying biological processes driving population size, as well as their individual response to climatic alterations. Therefore, the objective of this study is to investigate the influence that climate change may have on species population dynamics through altering breeding season.

Methodology/Principal Findings

We used a mechanistic model based on drivers of rabbit reproductive physiology together with demographic simulations to show how future climate-driven changes in breeding season result in contrasting rabbit population trends across Europe. In the Iberian Peninsula, where rabbits are a native species of high ecological and economic value, breeding seasons will shorten and become more variable leading to population declines, higher extinction risk, and lower resilience to perturbations. Whereas towards north-eastern countries, rabbit numbers are expected to increase through longer and more stable reproductive periods, which augment the probability of new rabbit invasions in those areas.

Conclusions/Significance

Our study reveals the type of mechanisms through which climate will cause alterations at the species level and emphasizes the need to focus on them in order to better foresee large-scale complex population trends. This is especially important in species like the European rabbit whose future responses may aggravate even further its dual keystone/pest problematic. Moreover, this approach allows us to predict not only distribution shifts but also future population status and growth, and to identify the demographic parameters on which to focus to mitigate global change effects.     

Temporal Variation in Population Size of European Bird Species: Effects of Latitude and Marginality of Distribution

In the Northern Hemisphere, global warming has been shown to affect animal populations in different ways, with southern populations in general suffering more from increased temperatures than northern populations of the same species. However, southern populations are also often marginal populations relative to the entire breeding range, and marginality may also have negative effects on populations. To disentangle the effects of latitude (possibly due to global warming) and marginality on temporal variation in population size, we investigated European breeding bird species across a latitudinal gradient. Population size estimates were regressed on years, and from these regressions we obtained the slope (a proxy for population trend) and the standard error of the estimate (SEE) (a proxy for population fluctuations). The possible relationships between marginality or latitude on one hand and slopes or SEE on the other were tested among populations within species. Potentially confounding factors such as census method, sampling effort, density-dependence, habitat fragmentation and number of sampling years were controlled statistically. Population latitude was positively related to regression slopes independent of marginality, with more positive slopes (i.e., trends) in northern than in southern populations. The degree of marginality was positively related to SEE independent of latitude, with marginal populations showing larger SEE (i.e., fluctuations) than central ones. Regression slopes were also significantly related to our estimate of density-dependence and SEE was significantly affected by the census method. These results are consistent with a scenario in which southern and northern populations of European bird species are negatively affected by marginality, with southern populations benefitting less from global warming than northern populations, thus potentially making southern populations more vulnerable to extinction.     

Demography of cape petrels in response to environmental changes

Predicting the responses of populations in changing environments is an important task for ecologists. Understanding the population dynamics of high-latitude breeding species is critical given the particularly rapid environmental changes that occur in these regions. Using long-term mark–resighting data acquired over 53-years in Pointe Géologie, Terre Adélie, Antarctica, we estimated age-specific demographic parameters and evaluated the effect of the environment on survival of a poorly known species, the cape petrel Daption capense. We then modeled the dynamics of this population using a life-history model and performed prospective and retrospective analyses to estimate the sensitivity of the population growth rate to demographic parameters, and to quantify their relative contribution. Survival of cape petrel increased with age, being 0.610 (±0.193) for juveniles, 0.739 (±0.158) for individuals from 2 to 4, and 0.920 (±0.031) for older individuals. Minimum age at first reproduction was 3 years old, the age at which all birds were recruited was 14 years, and mean age at first reproduction was 9.05 (±2.06) years. Adult survival increased over time and was positively correlated with the southern annular mode (SAM). The stochastic population growth rate was estimated at 1.019, and adult survival over age 5 made the largest contribution to variance of the population growth rate. Sensitivity analyses revealed that population regulation was mainly driven by the SAM. Our results suggest that despite the decrease in breeding success, the population of cape petrels at Pointe Géologie increased due to the increase in immature and adult survival.     

Environmental variability and population dynamics: do European and North American ducks play by the same rules?

Density dependence, population regulation, and variability in population size are fundamental population processes, the manifestation and interrelationships of which are affected by environmental variability. However, there are surprisingly few empirical studies that distinguish the effect of environmental variability from the effects of population processes. We took advantage of a unique system, in which populations of the same duck species or close ecological counterparts live in highly variable (north American prairies) and in stable (north European lakes) environments, to distinguish the relative contributions of environmental variability (measured as between‐year fluctuations in wetland numbers) and intraspecific interactions (density dependence) in driving population dynamics. We tested whether populations living in stable environments (in northern Europe) were more strongly governed by density dependence than populations living in variable environments (in North America). We also addressed whether relative population dynamical responses to environmental variability versus density corresponded to differences in life history strategies between dabbling (relatively “fast species” and governed by environmental variability) and diving (relatively “slow species” and governed by density) ducks. As expected, the variance component of population fluctuations caused by changes in breeding environments was greater in North America than in Europe. Contrary to expectations, however, populations in more stable environments were not less variable nor clearly more strongly density dependent than populations in highly variable environments. Also, contrary to expectations, populations of diving ducks were neither more stable nor stronger density dependent than populations of dabbling ducks, and the effect of environmental variability on population dynamics was greater in diving than in dabbling ducks. In general, irrespective of continent and species life history, environmental variability contributed more to variation in species abundances than did density. Our findings underscore the need for more studies on populations of the same species in different environments to verify the generality of current explanations about population dynamics and its association with species life history.     

Optimizing the genetic management of reintroduction projects: genetic population structure of the captive Northern Bald Ibis population

Many threatened species are bred in captivity for conservation purposes and some of these programmes aim at future reintroduction. The Northern Bald Ibis, Geronticus eremita, is a Critically Endangered bird species, with recently only one population remaining in the wild (Morocco, Souss Massa region). During the last two decades, two breeding programs for reintroduction have been started (in Austria and Spain). As the genetic constitution of the founding population can have strong effects on reintroduction success, we studied the genetic diversity of the two source populations for reintroduction (‘Waldrappteam’ and ‘Proyecto eremita’) as well as the European zoo population (all individuals held ex situ) by genotyping 642 individuals at 15 microsatellite loci. To test the hypothesis that the wild population in Morocco and the extinct wild population in the Middle East belong to different evolutionary significant units, we sequenced two mitochondrial DNA fragments. Our results show that the European zoo population is genetically highly structured, reflecting separate breeding lines. Genetic diversity was highest in the historic samples from the wild eastern population. DNA sequencing revealed only a single substitution distinguishing the wild eastern and wild western population. Contrary to that, the microsatellite analysis showed a clear differentiation between them. This suggests that genetic differentiation between the two populations is recent and does not confirm the existence of two evolutionary significant units. The European zoo population appears to be vital and suitable for reintroduction, but the management of the European zoo population and the two source populations for reintroductions can be optimized to reach a higher level of admixture.     

A basic equation for population dynamics with destruction of breeding habitats and its application to outbreak of cyprinid herpesvirus 3

In reproduction, many animal species migrate to local habitats that are appropriate for reproduction and for growth of newly born offspring. The examples are ubiquitous among crabs, freshwater fishes, amphibians, migratory birds, and sea animals. We propose a basic equation for population dynamics of such animals, assuming that the number of offspring is proportional to the area of the local breeding habitats as a first approximation. This equation is very simple to be solved analytically, and useful for representing environmental issues of habitat destruction and degradation. According to the equation, the adult density in breeding habitats increases temporarily during habitat destruction and returns to the original density afterwards. The temporal peak value is higher for a larger proportion of area with destruction, a higher temporal rate of destruction, and a higher survival probability of the adults. In contrast, habitat degradation results simply in a decrease of the adult density in breeding habitats. Using this equation, we will discuss the vulnerability of populations to epidemic diseases due to temporal local high densities with decreasing breeding habitats by human activities, exemplifying an outbreak of cyprinid herpesvirus 3 for wild carps in Lake Biwa.     

Sex-specific costs of reproduction on survival in a long-lived seabird

Costs of reproduction on survival have captured the attention of researchers since life history theory was formulated. Adults of long-lived species may increase survival by reducing their breeding effort or even skipping reproduction. In this study, we aimed to evaluate the costs of current reproduction on survival and whether skipping reproduction increases adult survival in a long-lived seabird. We used capture–mark–recapture data (1450 encounters) from two populations of Bulwer''s petrel (Bulweria bulwerii), breeding in the Azores and Canary Islands, North Atlantic Ocean. Using a multi-event model with two different breeding statuses (breeders versus non-breeders), we calculated probabilities of survival and of transitions between breeding statuses, evaluating potential differences between sexes. Females had lower survival probabilities than males, independent of their breeding status. When considering breeding status, breeding females had lower survival probabilities than non-breeding females, suggesting costs of reproduction on survival. Breeding males had higher survival probabilities than non-breeding males, suggesting that males do not incur costs of reproduction on survival and that only the highest quality males have access to breeding. The highest and the lowest probabilities of skipping reproduction were found in breeding males from the Azores and in breeding males from the Canary Islands, respectively. Intermediate values were observed in the females from both populations. This result is probably due to differences in the external factors affecting both populations, essentially predation pressure and competition. The existence of sex-specific costs of reproduction on survival in several populations of this long-lived species may have important implications for species population dynamics.     

Reintroduction of the European Capercaillie from the Capercaillie Breeding Centre in Wis?a Forest District: Genetic Assessments of Captive and Reintroduced Populations

The Western capercaillie (Tetrao urogallus) is a specific bird species, which, despite its very broad distribution and large global population size, is highly endangered in many Western and Central European countries. According to the species situation, in many countries (including Poland), breeding and reintroduction programmes have been started. One of the most complex and large-scale reintroduction programmes was started in Bory Dolnośląskie Forest, and the Capercaillie Breeding Centre in Wisła Forest District was used as one of the sources of individuals for reintroduction. As genetic tools provide essential knowledge about species biodiversity, which is crucially important during the breeding process and reintroduction, both captive and reintroduced grouse populations were genetically analysed. We were particularly interested in genetic diversity of the individuals in both populations and the genetic relationship between them, as well as between them and other capercaillie representatives from their current range. To fulfil these goals we determined nine microsatellite loci along with a fragment of the mitochondrial control region. Genetic diversity parameters were moderate to high compared to populations from other Central and Western European countries. Both populations were clustered into three distinct genetic clades based on microsatellites. Phylogenetic analysis placed all mitochondrial haplotypes we revealed in the Eurasian clade. The present results will play an important role as they will help to preserve and maximize genetic diversity in captive populations, and will provide a basis for future monitoring of the reintroduction process.     

Beyond average: an experimental test of temperature variability on the population dynamics of <Emphasis Type="Italic">Tribolium confusum</Emphasis>

The relationship between ectotherm ecology and climatic conditions has been mainly evaluated in terms of average conditions. Average temperature is the more common climatic variable used in physiological and population studies, and its effect on individual and population-level processes is well understood. However, the intrinsic variability of thermal conditions calls attention to the potential effects that this variability could have in ecological systems. Regarding this point, two hypotheses are proposed. From the allocation principle, it may be inferred that if temperature variability is high enough to induce stress in the organisms, then this extra-cost should reduce the energetic budget for reproduction, which will be reflected in population parameters. Moreover, a mathematical property of non-linear functions, Jensen’s inequality, indicates that, in concave functions, like the temperature–reproduction performance function, variability reduces the expected value of the output variable, and again modifies population parameters. To test these hypotheses, experimental cultures of Tribolium confusum under two different thermal variability regimens were carried out. With these data, we fitted a simple population dynamics model to evaluate the predictions of our hypothesis. The results show that thermal variability reduces the maximum reproductive rate of the population but no other parameters such as carrying capacity or the nonlinear factor in a nonlinear version of the Ricker model, which confirms our hypotheses. This result has important consequences, such as the paradoxical increase in population variability under a decrease in thermal variability and the necessary incorporation of climatic variability to evaluate the net effect of climate change on the dynamics of natural populations.     

Impact of population age structure on Wolbachia transgene driver efficacy: ecologically complex factors and release of genetically modified mosquitoes

Wolbachia symbionts hold theoretical promise as a way to drive transgenes into insect vector populations for disease prevention. For simplicity, current models of Wolbachia dynamics and spread ignore ecologically complex factors such as the age structure of vector populations and overlapping vector generations. We developed a model including these factors to assess their impact on the process of Wolbachia spread into populations of three mosquito species (Anopheles gambiae, Aedes aegypti and Culex pipiens). Depending on the mosquito species, Wolbachia parameters, released mosquito life stage and initial age structure of the target population, the number of Wolbachia-infected mosquitoes that we predict would need to be released ranged from less than the threshold calculated by the simple model to a 10-30-fold increase. Transgenic releases into age-structured populations, which is an expectation for wild mosquitoes, will be difficult and depending on the circumstances may not be economically or logistically feasible due to the large number of infected mosquitoes that must be released. Our results support the perspective that understanding ecological factors is critical for designing transgenic vector-borne disease control strategies.     

Mortality rates of wild boar Sus scrofa L. in central Europe

In many parts of Europe, wild boar Sus scrofa population increase, and thus, high densities and dispersal into new areas are accompanied by economic problems. Due to many factors like insufficient hunting strategies as well as underestimation of population densities and reproduction rates, harvest rates seem to be insufficient. Thus, we calculated mortality rates of several wild boar populations from 1998 to 2009, to show the efficiency of hunting within several studies distributed over eight European states. For calculating mortality rates, the daily probability of survival of radio telemetrically observed wild boar was analysed according to Mayfield (Wilson Bull 73:255-261, 1961) and with survival analysis in R for three age classes (0, 1, ≥2 years) and both sexes. The mortality rates of wild boar per annum, especially piglets, were comparably low (about 0.5 for piglets and similar for total population). About three third of all observed animals survived at least until the next period of reproduction. Mortality rates differed between some study areas, the sexes and age classes. The sex ratio of the shot piglets equals the sex ratio of captured piglets; there seems to be no sex-biased hunting in this age class, but in an older age. Shooting was the main cause of death; only very few animals died by natural causes, e.g. diseases. The comparative analysis of all studies reflects a low mortality of wild boar in highly productive populations. Our results certified the findings of several studies that predation, natural mortality, and road mortality have only small impact on wild boar populations, whereas especially, nutrition or hunting are mainly decisive. Assuming net reproduction rates of more than 200 % according to literature data, our results indicate that harvest rates are not sufficient at our study sites. In all our studies, mortality rates and, thus, harvest rates are less than the assumed total net reproduction. Especially, the harvest rate of piglets seems to be insufficient. Thus, the population will increase further. High reproduction has to be counteracted by regulating mainly the reproductive animals. For regulating a population, combined and effective hunting methods have to be conducted to harvest at least the net reproduction. Thus, we recommend higher hunting rates of piglets (80 % of the offspring should be harvested) and of adult females. Intensified hunting of piglets by drive hunts and at an early age as well as intensified single hunt on adult females might help regulating wild boar populations.     

Reproductive senescence in a cooperatively breeding mammal

1. Senescence (or 'ageing') is a widespread and important process in wild animal populations, but variation in ageing patterns within and between species is poorly understood.
2. In cooperatively breeding species, the costs of reproduction are shared between breeders and one or more helpers. The effects of ageing in breeders may therefore be moderated by the presence of helpers, but there have been very few studies of senescence patterns in natural populations of cooperative breeders.
3. Here, we use 13 years of data from a long-term study population of wild meerkats ( Suricata suricatta ) to investigate age-related changes in several traits known to be key components of reproductive success in females of this species.
4. Four of the six traits studied exhibited significant declines with age, indicating senescence. Litter size, the number of litters produced per year and the number of pups that survived to emergence from the natal burrow per year all increased with female age up to a peak at c. 4 years, and declined steeply thereafter; the mean pup weight at emergence in a given litter declined steadily from age zero.
5. These results provide the first evidence of reproductive senescence in a wild population of a cooperatively breeding vertebrate. Breeding success declined with age despite the sharing of reproductive costs in this species, but further study is needed to investigate whether helping affects other aspects of senescence, including survival.     

Variation in probability of first reproduction of Weddell seals

1. For many species, when to begin reproduction is an important life-history decision that varies by individual and can have substantial implications for lifetime reproductive success and fitness. 2. We estimated age-specific probabilities of first-time breeding and modelled variation in these rates to determine age at first reproduction and understand why it varies in a population of Weddell seals in Erebus Bay, Antarctica. We used multistate mark-recapture modelling methods and encounter histories of 4965 known-age female seals to test predictions about age-related variation in probability of first reproduction and the effects of annual variation, cohort and population density. 3. Mean age at first reproduction in this southerly located study population (7.62 years of age, SD=1.71) was greater than age at first reproduction for a Weddell seal population at a more northerly and typical latitude for breeding Weddell seals (mean=4-5 years of age). This difference suggests that age at first reproduction may be influenced by whether a population inhabits the core or periphery of its range. 4. Age at first reproduction varied from 4 to 14 years, but there was no age by which all seals recruited to the breeding population, suggesting that individual heterogeneity exists among females in this population. 5. In the best model, the probability of breeding for the first time varied by age and year, and the amount of annual variation varied with age (average variance ratio for age-specific rates=4.3%). 6. Our results affirmed the predictions of life-history theory that age at first reproduction in long-lived mammals will be sensitive to environmental variation. In terms of life-history evolution, this variability suggests that Weddell seals display flexibility in age at first reproduction in order to maximize reproductive output under varying environmental conditions. Future analyses will attempt to test predictions regarding relationships between environmental covariates and annual variation in age at first reproduction and evaluate the relationship between age at first reproduction and lifetime reproductive success.