Fitness costs of reproduction play a key role in understanding the evolution of reproductive tactics. Nevertheless, the detection and the intensity of costs of reproduction vary according to which life-history traits and species are studied. We propose an evolutionary model demonstrating that the chance of detecting a cost of reproduction should be lower when the fitness component studied has a low rather than high variance. Consequently, the fitness component that is affected the most by costs of reproduction should vary with life speed. Since long-lived species have developed a strategy that avoids jeopardizing their survival and short-lived species favour current reproduction, variance in survival is smaller and variance in reproduction higher in long-lived vs. short-lived species. We review empirical studies of costs of reproduction in free-ranging mammals, comparing evidence of costs reported among species and focal traits. In support of our model, more studies reported evidence of reproductive costs of reproduction in ungulates than in rodents, whereas survival costs of reproduction were more frequent in rodents than in ungulates. The life-history model we propose is expected to apply to any species, and hence provides a better understanding of life-history variation, which should be relevant to all evolutionary ecologists. KeywordsEnvironmental canalization, investment, mammals, reproductive tactics, resource acquisition, resource allocation, rodents, ungulates, van Noordwijk and de JongÕs model, variable environment.Ecology Letters (2010) 13: 915-935 Positive or negative co-variations among fitness components are central to life-history theory (Stearns 1992). The notion of negative co-variations originates from the principle of allocation (Williams 1966), which posits that resources available for an individual are limited, so that individuals should allocate resources optimally among growth, survival, and reproduction in order to maximize their fitness (Stearns 1992). As reproduction requires high energy allocation (Robbins 1993), fitness costs of reproduction (see Box 1 for definitions) should occur and lead to negative co-variations between current reproduction and future survival and ⁄ or reproduction (Stearns 1992;Roff 2002). Williams (1966) was the first to introduce this cost-benefit concept of reproduction, which was further developed by numerous authors (e.g. Gadgil & Bossert 1970;Bell 1980;Reznick et al. 2000). Costs of reproduction have received a particular attention in the literature because they play a key role in the evolution of reproductive tactics (Roff 2002). They can also have fundamental demographic consequences on population dynamics by decreasing reproduction and survival, and thereby changing the population age structure and growth rate (Silvertown & Dodd 1999;Proaktor et al. 2008).As energy allocation is particularly important during gestation and lactation (Gittleman & Thompson 1988), mammals are relevant model species to assess costs of reproduction. In his review of ...

Summary 1.Although life-history theory predicts substantial costs of reproduction, individuals often show positive correlations among life-history traits, rather than trade-offs. The apparent absence of reproductive costs may result from heterogeneity in individual quality. 2. Using detailed longitudinal data from three contrasted ungulate populations (mountain goats, Oreamnos americanus ; bighorn sheep, Ovis canadensis ; and roe deer, Capreolus capreolus ), we assessed how individual quality affects the probability of detecting a cost of current reproduction on future reproduction for females. We used a composite measure of individual quality based on variations in longevity (all species), success in the last breeding opportunity before death (goats and sheep), adult mass (all species), and social rank (goats only). 3. In all species, high-quality females consistently had a higher probability of reproduction, irrespective of previous reproductive status. In mountain goats, we detected a cost of reproduction only after accounting for differences in individual quality. Only low-quality female goats were less likely to reproduce following years of breeding than of nonbreeding. Offspring survival was lower in bighorn ewes following years of successful breeding than after years when no lamb was produced, but only for low-quality females, suggesting that a cost of reproduction only occurred for low-quality females. 4. Because costs of reproduction differ among females, studies of life-history evolution must account for heterogeneity in individual quality.

Variations among individuals in phenotypic quality and fitness often confound analyses of life-history strategies assessed at the population level. We used detailed long-term data from three populations of large herbivores with generation times ranging from four to nine years to quantify heterogeneity in individual quality among females, and to assess its influence on mean annual reproductive success over the lifetime (MRS). We also determined how environmental conditions in early life shaped individual quality and tested A. Lomnicki's hypothesis that variance in individual quality should increase when environmental conditions deteriorate. Using multivariate analyses (PCA), we identified one (in sheep and deer) or two (in goats) covariations among life-history traits (longevity, success in the last breeding opportunity, adult mass, and social rank) as indexes of individual quality that positively influenced MRS of females. Individual quality was reduced by unfavorable weather, low resource availability, and high population density in the year of birth. Early-life conditions accounted for 35-55% of variation in individual quality. In roe deer, we found greater variance in individual quality for cohorts born under unfavorable conditions as opposed to favorable ones, but the opposite was found in bighorn sheep and mountain goats. Our results demonstrate that heterogeneity in female quality can originate from environmental conditions in early life and can markedly influence the fitness of females in species located at different positions along the slow-fast continuum of life-history strategies.

Summary 1.In recent years, the Normalized Difference Vegetation Index (NDVI) has been used to assess the relationships between habitat quality and animal life-history traits. Since numerous ecological studies now use NDVI rather than perform direct vegetation assessments, field validations are essential to provide confidence in the biological significance of NDVI estimates. While some studies have compared NDVI with plant biomass, very few have examined the relationship between NDVI and changes in vegetation quality. 2. Using data from two long-term studies of alpine ungulates, we assessed the relationship between two NDVI indices and the date of peak in faecal crude protein (FCP), which represents temporal variability in the availability of high-quality vegetation. We also evaluated if NDVI data could predict annual variation in the timing of spring green-up. 3. In both populations, integrated NDVI in June was negatively correlated with the date of the peak in FCP, indicating that high integrated NDVI values corresponded to early springs in alpine habitats. Maximum NDVI increase during spring green-up was positively correlated with the timing of peak FCP, illustrating that rapid increases in NDVI represented delayed springs. 4. Predicted values of date of peak FCP estimated each year from NDVI data satisfactorily fitted observed values, and prediction intervals included all observed values. These results suggest that NDVI can reliably predict variation over years in the timing of spring. 5. Synthesis and applications. Our long-term studies demonstrate that a multi-year time series of Normalized Difference Vegetation Index (NDVI) can reliably measure yearly changes in the timing of the availability of high-quality vegetation for temperate herbivores. This finding therefore supports the use of NDVI as a proxy for vegetation attributes in population ecology and wildlife management studies.

Reproduction should reduce resources available for somatic investment and result in fundamental trade-offs among life-history traits. Using 18 years of longitudinal data from marked mountain goats (Oreamnos americanus), we assessed whether reproductive status affected female survival and future reproduction when accounting for parity, age, individual quality, population density, and environmental conditions. Reproduction reduced the probability of parturition and offspring survival in the following year. Female survival, however, was independent of previous reproduction, suggesting that females favored their own survival over that of their offspring. The lower probability of parturition in females that had a kid the previous year was only detected at high population density and among young and prime-aged females, suggesting that fitness costs of reproduction can be masked by variations in resource availability and individual characteristics. Primiparous females were less likely than multiparous females to reproduce in the subsequent year. Offspring survival was reduced at high density and after severe winters. Environmental conditions mainly influenced offspring survival, whereas female survival and fecundity were principally modulated by female characteristics. Our study highlights how different intrinsic and environmental factors can affect the probability of future reproduction and also underlines the value of long-term monitoring of known individuals.

Summary1. The development of camera-traps has provided an opportunity to study ecological relationships and population dynamics of species that are rare, difficult to observe or capture. Their use has seen a major increase recently, particularly with the recent progress in methods adapted to species for which individuals cannot be identified. 2. We took advantage of extensive camera-trap data sets from large spatiotemporal-scale studies of a diverse assemblage of avian and mammalian scavengers in subarctic/arctic tundra to determine sampling designs that minimize detection errors (false-negative) and to evaluate the influence of sampling design on estimation of site occupancy. 3. Results showed that raw error rates in daily presence varied between 5 and 30% among species when using time-triggered cameras with a 5-min interval. Using movement-triggered cameras resulted in larger raw error rates, between 30 and 70%, as well as a lower number of daily presences detected. Increasing the time interval from 5 to 20 min greatly increased the raw error rate in daily presence, but it had negligible impacts on estimates and precision of occupancy and detection probability. 4. Occupancy estimates were mostly influenced by variation in the number of days included during the sampling period. For most species, a threshold of between 20 and 30 problem-free days (i.e. without camera-related technical problems) was required to stabilize occupancy and detection probability, as well as to maximize their precision. 5. Based on the results, we discuss guidelines for establishing sampling designs according to the different ecological questions researchers might want to answer. To our knowledge, our study is the first to directly test the influence of sampling design in camera-trap studies, providing guidelines that are likely to be directly applicable to a large range of species and ecosystems.

Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection.

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